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

Not Bob Marley Heat map photo of & by Tom Whitham

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

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

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.

Illustration by Paul Heinrich

Geographical Distribution of Ecotypes

Central California Ecotype Utah High Plateau Ecotype Sonoran Desert Ecotype

Geographical adapted ecotypes have evolved in response to environmental differences across the range of P. fremontii.

Ikeda et al. 2017 Global Change Biology Structure Analysis based upon molecular marker

Central California Ecotype Utah High Plateau Ecotype Sonoran Desert Ecotype

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).

Bio.15 – Precipitation seasonality Bio.6 - Max temp of warmest month Bio.11 – Mean temp coldest quarter Most Predictive Environmental Variables

Genetics-based models are up to 12x better at predicting ecoregion test points

Genetics No Genetics

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California Central Valley Sonoran Desert Utah High Plateau

Binomial Probability (ln) Ecoregion Central California Valley Sonoran Desert Utah High Plateau

Ecoregion Predictive accuracy

Ikeda et al. 2017 Global Change Biology

Genetics No Genetics

Reciprocal common gardens show finer scale local adaptation within the Sonoran desert ecotype 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.)

Reciprocal Common Gardens

TNC Dugout Ranch, Canyonlands Cooler Garden – MAT 10.7 °C AZG&F Horseshoe Ranch Intermediate – MAT 17.2 °C BLM Mittry Lake, Yuma Hot Garden – MAT 22.8 °C

4600 tree common garden on Arizona Game & Fish Dept. lands at Horseshoe Ranch surrounded by Agua Fria National Monument.

March 8, 2017 – Photos by Tom Whitham Populus fremontii field trial Indian Creek Agua Fria Creek

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).

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)

Fremont cottonwood

• P. fremontii

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.

Emergence trap photo by Zacchaeus Compson

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

Axis 1

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0.0 0.5 1.0 1.5

Axis 2

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0.0 0.2 0.4 0.6 0.8 1.0 996 1008 1012 1017 1020 RM2 T15

Different genotypes of P. angustifolia support different stream macro-invertebrates. Compson 2016 Ecosphere

Community 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

AFLP genetic markers

Ecosystem Phenotypes

Aphid survival

Frequency

Traditional Phenotype

• f Resistance

Genetic Linkage Map of Populus

Woolbright et al. 2008 Heredity Matt Zinkgraf et al. 2016 J Insect Physiology Woolbright et al. 2018 Ecology & Evolution

lg8 lg8 lg6

Leaf Chemistry Phenology

lg6

Architecture

lg1 lg12

QTL Mapping

Condensed Tannin Salicortin Spring Leaf Flush

Second Year Branch Number

Insect Resistance

lg1 lg8

Resistance to

• P. betae

lg18

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.

Illustration by Victor Leshyk

San Francisco Peaks & Wupatki National Monument view from Navajo Nation Photo by Tom Whitham

Communities are far more linked than previously envisioned.

Lichen 2010 EM Fungi 2006 Leaf Pathogens 2010

Lichen 2010 Ectomycorrhizal Fungi 2006 Fungal Leaf Pathogens 2010

Lichen 2010 EM Fungi 2006 Soil Bacteria 2004 Soil Fungi 2004 Endophytes 2006 Leaf Modifiers 2010 Leaf Pathogens 2009 Leaf Pathogens 2010 P ! 0.05 0.05<P<0.1 P ! 0.1 Phyllosphere Trunk Soil

Lichen 2010 Ectomycorrhizal Fungi 2006 Soil Bacteria 2004 Soil Fungi 2004 Twig Endophyes 2006 Leaf Modifying Arthropods 2010 Fungal Leaf Pathogens 2009 Fungal Leaf Pathogens 2010

B A

The network of correlated communities is defined by individual tree genotypes – the importance of maintaining network structure in restoration. Community-genetic correlations - changes in the composition of one community among plant genotypes that are mirrored by changes in the composition of another community.

Populus angustifolia

Endophytes

Lamit et al. 2015 Journal of Ecology

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.

In just 25 years plant hardiness zones have shifted northward and upward in elevation by one zone.

Climate change results in mis-matches with the local environment.

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?

Photos by Hillary Cooper (top) & Tom Whitham (bottom) Cottonwood mortality on Bill Williams River National Wildlife Refuge – March 28, 2017

Photo by Tom Whitham Sept 15, 2004 - North of San Francisco Peaks, AZ

Record drought of 2002 – An evolutionary event that changed the genetic structure of the tree population.

Sthultz et al. 2009 Global Change Biology

Diverse arthropod community on pinyons negatively affected by drought

Stress Index

0.0 0.5 1.0 1.5 2.0 2.5

Total richness per tree

10 20 30 40

Stress Index

0.0 0.5 1.0 1.5 2.0 2.5

Total abundance per tree

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5 10 15 20 25

Species Richness r2=0.53, p<0.0001 Abundance r2=0.53, p<0.0001

Within a site, increased stress negatively affects arthropod diversity.

Stress Index = (standardized branch dieback + number of needle cohorts + radial trunk growth) Stone et al. 2010 Oecologia 155 arthropod species

Part IV - Solutions to the mis-match

EVOLVE, ACQUIRE BETTER MUTUALISTS, MOVE or DIE

Painted Desert from SP Crater Photo by Tom Whitham

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 (VP) = VG + VE + VGxE. From Cooper et al. 2018 Global Change Biology (in press).

Genetics-based responses to climate change

Plasticity Genetic Variation in Plasticity Genetic

10 20 30 40 50 60 70 80 % mortality

Drought Susceptible Pinyons Drought Resistant Pinyons

Rapid Evolution with Drought

Drought susceptible trees were >3X more likely to die during the 2002 record drought than drought resistant trees.

Gehring et al. 2014 Molecular Ecology

A 2500 tree pinyon pine common garden shows that 1) drought tolerance is a heritable trait and 2) mycorrhizae on drought tolerant trees are better mutualists.

Seedling from Drought Intolerant Mother Seedling from Drought Tolerant Mother

Figure 4

Drought Tolerant Drought Intolerant

Adults Seedlings Seedlings Adults

Drought Tolerant Drought Intolerant

• 1. Mycorrhizal communities

are heritable plant traits.

• 2. Drought tolerant

mycorrhizal communities are better mutualists.

Gehring et al. 2017 PNAS

Mycorrhizal Community Pinyon Seedling Performance

Photo by Lloyd A. Whitham, circa 1950, nursery established 1863

Assisted gene flow/migration has been used inappropriately for a long time; we must get smarter in its use.

Common gardens allow us to predict the best source populations for a given level of climate change

Cooper et al. 2018 Global Change Biology (in press).

Hot Garden Intermediate Cool Garden

Long Growing Season Short Growing Season

Assisted Migration Assisted Migration No Assisted Migration Options

Phenotypic plasticity defines limits to assisted migration

Home - Keams Canyon, AZ Yuma Garden - ∆MAMT = 12.7 °C hotter transfer distance Chevelon Garden - ∆MAMT = 3.2 °C hotter transfer distance

Photos and data by Jackie Parker

A fundamental issue in assisted migration is if you move plants to mitigate the impacts of climate change, will plants acquire the communities of their home sites? In other words,

if you build it will they come?

Odgen Nature Center restoration site Photo by Tom Whitham

Up to a point, if you built it they will come. With transfers of 18 and 48 km, garden and wild trees support similar communities, but at 90 km they are quite different (Keith et al. unpublished data).

Marble Canyon dust storm “haboob” Photo by Tom Whitham

Part I. Plants can be very locally adapted. negative effects with climate change Part II. Plant genotype can define communities. negative effects with climate change Part III. Climate change creates a mis-match of once locally adapted plants and the new environment. negative effects with climate change Part IV. Solutions to the mis-match. passive - genetic variation & plasticity; human

intervention - assisted gene migration & ecosystem engineering; inoculate with better mutualists

Summary

Collaborators in Community Genetics and Genetics-Based Restoration

Rachel Adams – plant ecology Gery Allan – molecular ecology Petter Axelsson – transgenic trees Joe Bailey – community ecology Randy Bangert – biogeography Rebecca Best – ecology & evolution Davis Blasini – ecophysiology Helen Bothwell – phylogeography Posy Busby – ecological plant pathology Abraham Cadmus – ecophysiology Aimée Classen – soil ecology Zacchaeus Compson – aquatic ecology Hillary Cooper – phylogenetics Sam Cushman – landscape genetics Steve DiFazio – molecular ecology Rodolfo Dirzo – community ecology Luke Evans – population ecology Sharon Ferrier – conservation ecology Dylan Fischer – ecophysiology Paul Flikkema – systems engineering Kevin Floate – insect ecology Catherine Gehring – microbial ecology Heather Gillette – molecular ecology Kevin Grady – restoration Steve Hart – ecosystem/soil ecology Erika Hersch – ecological genetics Joakim Hjältén – ecology Lisa Holeski – genetics & chemistry Kevin Hultine – invasive species Dana Ikeda – climate modeling Julia Hull – endophytes Nathalie Isabel – molecular ecology Karl Jarvis – phylogeny Art Keith – insect community ecology George Koch – ecophysiology Tom Kolb – plant physiology Lela Andrews - molecular ecology Jamie Lamit – microbial ecology Matthew Lau – network modeling Carri LeRoy – aquatic ecology Rick Lindroth – chemical ecology Jane Marks – aquatic ecology Lisa Markovchick – microbial ecology Tamara Max – molecular ecology Richard Michalet - facilitation & ecology Nashelly Meneses – ecological genetics George Newcombe – plant pathology Emily Palmquist – hydrology Jackie Parker – plant ecology Brad Potts – quantitative genetics Jen Schweitzer – ecosystems David Smith – landscape ecology Steve Shuster – theoretical genetics Chris Sthultz – plant ecology Amy Whipple – ecological genetics Tom Whitham – community ecology Gina Wimp – community ecology Todd Wojtowicz – litter arthropods Troy Wood – ecology Scott Woolbright - molecular genetics Adam Wymore – aquatic ecology Matt Zinkgraf – molecular genetics GO & NGO collaborators: Mary McKinley – Ogden Nature Center, Gregg Garnett – Bureau of Reclamation, Kris Haskins – The Arboretum at Flagstaff, Paul Burnett – Utah Dept. of Natural Resources, Billy Cordasco – Babbitt Ranches Outreach – Lara Schmit, Victor Leshyk - NAU

Macrosystems MRI