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Whitebark Pine Status and the Potential
Role of Biotechnology in Restoration
Diana F. TombackDept. Integrative Biology
University of Colorado Denver
Webinar, Committee on Forest Health and
Biotechnology, NASEM, April 2, 2018.
• Four case histories illustrating the threat posed by
Cronartium ribicola
• Restoration approaches
• How biotechnology can expedite restoration efforts
• The National Whitebark Pine Restoration Plan
Willmore Wilderness Park, Alberta, Canada
• Distribution
• ESA status review
• Ecology
• Foundation and
keystone roles
• Threats and trends.
Outline of presentation
Taxonomy: Pinus albicaulis Engelm., whitebark pine
Family Pinaceae, Genus Pinus, Subgenus Strobus, Section Quinquefoliae.*
• Subsect. Strobus -“five-needle pines” (revised)*.
• Most recent phylogenies for subgenus Strobus constructed from nuclear,
mitochondrial, and chloroplast gene sequences show diverse affinities between
P. albicaulis and species native to North America, Asia, or Europe (Hao et al. 2015).
• Hao et al. (2015)—“…ancient and relatively recent introgressive hybridization
events…particularly in northeastern Asia and northwestern North America.”
Genome of whitebark pine characterized as extremely large and highly repetitive.
*New Subsect. Strobus from combined subsects. Strobus and Cembrae, Gernandt et al. 2005;
Syring et al. 2007.
Whitebark pine range
• Upper subalpine and treeline forest zones.
• Western U.S. and Canada.
• 96% of the U.S. distribution is on federally
owned/managed lands.
• 37o to 55o N lat.
• 107o to 128o W long.
• Elevation: 900-3,660 m
• Estimated areal coverage:
Keane et al. 2012, Table 4.1
-ca. 5,770,000 ha
-ca. 14,252,000 acres
Whitebark pine range by Government Jurisdiction
Total estimated area ~ 5,770,000 ha
0 500,000 1,000,000 1,500,000 2,000,000 2,500,000 3,000,000
Forest Service
Wilderness (all agencies)
National Park Service
Private and State Lands
Native American Tribal Lands
Bureau of Land Management
Other (misc.)
Range area (ha)
47%
10%1%
1%
3%
38%
0%
Forest Service
National Park Service
Bureau of Land Management
Native American Tribal Lands
Private and State Lands
Wilderness (all agencies)
Other (misc.)
Data from Keane et al. 2012, Table 4.1
Events leading to ESA status review
Whitebark pine paradox: How can a species that inhabits remote locations and so widely-distributed be declining?
• December 8, 2008: Natural Resources Defense Council (NRDC) petition to U.S. Fish & Wildlife Service to list whitebark pine under the Endangered Species Act.”
• July 19, 2011: Fish & Wildlife Service 12-month finding: “…we find that the listing of P. albicaulis as threatened or endangered is warranted. However, currently listing P. albicaulis is precluded by higher priority actions to amend the Lists of Endangered and Threatened Wildlife and Plants.” “.…we will add P. albicaulis to our candidate species list.” Federal Register, Vol. 76, NO. 138, July 19, 2011.
Threats cited: Fire suppression and advancing succession, climate change and its interactions with mountain pine beetle and fire, white pine blister rust, and mountain pine beetles.
White Calf Mountain, Glacier National Park, MT
View: Blackfeet Reservation
Ecology
Foundation and keystone roles
Whitebark pine community types
• Successional communities on favorable sites, upper subalpine zone.
• Climax (self-replacing) communities on exposed upper subalpine sites.
• Treeline communities on cold sites in the alpine treeline ecotone.
Beartooth Plateau, WYBanff National Park, AB
Grand Teton NP, WY
Adaptations of whitebark pine for seed dispersal by nutcrackers
• Large, wingless seeds.
• Cones remain closed after seeds ripen: obligate
mutualism.
• Horizontally-oriented cones on upswept branches.
• Seeds adapted for caching: viable for
several years under soil. Don Pigott
Seed dispersal by nutcrackers
Nutcrackers
• Place seeds in caches of 1 to 15 seeds.
• Bury seed caches 1 to 3 cm under substrate.
• Carry seeds from a few meters to >32 km.
• Store >35,000 whitebark pine seeds per year per bird.
• Retrieve caches using highly accurate spatial memory.
Unretrieved caches germinate, leading to regeneration.
Seed dispersal by nutcrackers determines:
• The distribution of whitebark pine on the landscape—elevation and topography.
• Where whitebark pine grows locally—nutcracker cache site selection and environmental suitability.
• Rise of treeline with climate change—because nutcrackers cache seeds above tree limits.
• Fine-scale population genetic structure.
• Watershed and regional population structure. Tomback and Linhart 1990, Tomback 2001
Rogers et al. 1999, Tomback 2005
Stanley Glacier, Kootenay NP, BC, CA
• Wide spectrum of community types.
• 7 recognized SAF cover types.
• High elevation wildlife habitat, shelter,
and nest sites.
• Seeds provide wildlife food (birds, small
mammals, bears,. & foxes)
Whitebark pineKeystone species
Promotes biodiversity
Willmore Wilderness Park, AB, CA
Whitebark pine
Foundation species (Dayton 1972, Ellison et al. 2005)
“…a single species that defines much of the structure of a community
by creating locally stable conditions for other species and by
modulating and stabilizing fundamental ecosystem processes.”
Defines ecosystem structure and function• Early establishment after disturbance.
• Fosters community development through
mitigation of harsh conditions and facilitation.
• Nurse tree on harsh sites (facilitation).
• Tree island initiator (facilitation).
Grand Teton National Park
Whitebark Pine Ecosystem Services
• High elevation forests and treeline communities
redistribute and retain snow.
• Shade from these forests slows summer snowmelt,
regulating downstream flow.
• Roots stabilize soil, reducing soil erosion.
• Trees stabilize snow, reducing avalanche hazard.
Wind River Mtns., WYBlackfeet Indian Reservation, MT
Role of treeline communities in snow redistribution and retention (Fig. 6 from Tomback et al. 2016)
The four major threats to whitebark pine
• Cronartium ribicola—exotic fungal pathogen that causes white pine blister rust.
• Mountain pine beetle (Dendroctonusponderosae) outbreaks.
• Altered fire regimes—successional replacement from fire exclusion actions.
• Climate warming—driving bark beetle outbreaks, drought stress and mortality, larger, more frequent, and severe fires.
https://en.wikipedia.org/wiki/Mountain_pine_beetle
White Pine Blister rust (WPBR): an exotic disease naturalizedto North America
• Accidental introduction(s) to the Northwest around 1910.
• First detected in PNW in1921.
• Conditions (pine hosts, alternate host Ribes spp., and climate) highly favorable to its spread.
• Infects and kills all age classes.
• Continues to spread geographically and intensify locally.
• Now in regions once believed to be too cold, warm, or dry.
• Spread facilitated by wave years.
Life Cycle of white pine blister rust
R. Hunt 1983.
Canadian Forestry
Service
These spores may blow
500 km or farther
Only infects five-needle white
pines (subgenus Strobus taxa).
Most common alternate hosts:
Ribes spp.
• Five spore stages in life cycle.
• Aeciospores—transmission
from pines to alternate hosts.
• Basidiospores—transmission
from alternate hosts to pines.
U. S. distribution of WPBR
U S Forest Service, Forest Health ProtectionU.S. & Canadian
pines impacted:
• Whitebark pine
• Limber pine
• Southwestern
white pine
• Sugar pine
• Western white
pine
• Foxtail pine
• Rocky Mountain
bristlecone pine
Not yet infected:
• Great Basin
bristlecone pine
Mountain pine beetle MPB mortality in whitebark pine
• Major losses of mature, cone-bearing trees over two decades.
• Loss of trees resistant to WPBR.
• Some research shows preference by MPB for trees weakened by WPBR.
• Outbreak still active: diminishing in Rockies, active in the western distribution.
45,891 85,257 91,037
218,121
679,372
939,450
1,088,748
-
200,000
400,000
600,000
800,000
1,000,000
1,200,000
NV CA OR WA ID MT WY
1997-2016 Cumulative Whitebark Pine MPB Footprint: Total 3,147,876 Acres (~25% range)
Fire exclusion leads to advancing succession
• Aggressive fire exclusion since early
20th century.
• Altered fire regimes have led to
successional replacement of whitebark
pine in several regions.
(Warwell et al. 2007)
Climate change CC
and whitebark pine
• Predictions based on
Species Distribution Models
(Bioclimatic Envelope
Models): WP upwards and
northwards.
We need to add cc mitigation
to restoration:
• Rely on resilience in
established whitebark pine.
• Find local refugia.
• Use genetic diversity.
Four case histories
• Northern U.S. Rocky Mountains
• The Greater Yellowstone Area
• Treeline and northern edge communities
• The southern Sierra Nevada
All case histories demonstrate that Cronartium ribicola is still spreading geographically and intensifying within communities.
Currently, WPBR is an existential threat to whitebark pine.
Northern Rocky Mountains
The northern Rocky Mountains, US, and southern Rocky Mountains, Canada, are the epicenter of whitebark pine decline. Climate eminently suited to the survival and spread of Cronartium ribicola (BR). Brief history (McDonald and Hoff 2001):
• BR introduced to Pacific NW around 1910.
• Climatic “Wave years”: 1913, 1917, 1921, 1923, 1927, 1936.
• First detected on WP in 1926 in coastal range, BC.
• Idaho 1923 in western white pine.
• Northern Idaho 1938.
• Continental Divide, Glacier National Park 1939.
The Northern Rocky Mountains has many areas with little to no living whitebark pine or trees so damaged, the communities are non-functional.
Northern Rocky
Mountains, USForest Health Protection
Risk Map
Surveys Living trees, Mean percent BR
Smith et al. 2008:
Glacier NP 67% (1-100%)
Waterton Lakes 71.5 (22-97%)
Elk, Flathead Valleys 67.4% (41-95%)
Zeglen 2002:
Cranbrook region 44.9%
Keane et al. 1994:
Bob Marshall WA 48% (10-99%)
Keane & Arno 1993:
Western Montana 61% (20-90%)
FHP National Risk Maps
Greater Yellowstone Area
Yellowstone was originally considered to be too cold and dry to support BR (Kendall and Asebrook 1998)
• BR found on Ribes in the Gallatin NF in 1937; found at Mammoth Hot Springs in YNP in 1944.
• BR began to spread through the YNP.
• Survey in 1961 found 7% infection.
• BR management through Ribes control 1945-1977.
• 8 million Ribes shrubs removed over 175,000 acres.
• $2,420,238 (>$11,112,000 in 1994 dollars).
Monitoring in the GYE since 2007
Greater Yellowstone Whitebark Pine
Monitoring Working Group
2017 report (2016 data), based on
Panel 1 (of 4 panels), 43 transects,
809 trees tagged in 2012:
• 20-30% of whitebark pine infected
by BR
• 6% increase in BR since Panel 1
was surveyed in 2012.
• Distribution of infected (yellow) and
dead trees (black) across the GYE,
monitored since 2013.
Treeline and the northern edge
Willmore Wilderness Park, AB
Divide Mountain, Blackfeet Tribal Lands
The conditions at treeline and at the northern
edge of whitebark pine’s distribution (54.5° N)
should be even less conducive than the GYE to
the establishment and spread of WPBR.
Since 2006, we (L. M. Resler, Virginia Tech,
students) have been studying whitebark pine’s
ecological role at treeline throughout the Rocky
Mountains.
Blister rust at treeline and at the northern edge
Study area
Infection
percent
Willmore Wilderness, AB 1.1
Gibbon Pass, Banff NP 0
Stanley Glacier, Kootenay NP 16.2
Divide Mountain, Glacier NP 23.6
Line Creek, Beartooth Plateau 19.2
Tibbs Butte, Beartooth Plateau 0.6
Paintbrush Divide, Grand Teton NP 18.1
Hurricane Pass, Grand Teton NP 14.1
Christina Mountain, Wind River Range 2.0
The southern Sierra Nevada
BR has been detected in the southern
Sierra Nevada since the 1960’s.
• Primarily in sugar pine and western
white pine.
• A puzzle why Cronartium ribicola
has been slow to infect whitebark
pine, but this is changing.
• Kliejunas and Adams 2003: North
to south spread of BR in the Sierra
Nevada.
• MPB-killed and BR-infected
whitebark pine at Minaret Summit,
near Mammoth Mountain, Inyo
National Forest, southern Sierra,
2016.
Jon Nesmith, NPS
D. F. Tomback
0
5000
10000
15000
20000
25000
30000
35000
40000
45000
1997 1998 1999 2000 2001 2002 2003 2004 2005 2006 2007 2008 2009 2010 2011 2012 2013 2014 2015 2016
Whitebark Pine MPB ADS Acres by Year 1997-2016
California Nevada Oregon Washington
Surveys of whitebark pine
Duriscoe & Duriscoe 2002
Sequoia and Kings Canyon National Park
• Infection on whitebark pine = 0%
Dunlap 2012, n = 49 plots
Southern Sierra Nevada; scattered plots
in northern California
• Overall mean percent BR = 12% (0-76)
• Northern Sierra plots had higher rust
incidence than southern = 24%
• Southern Sierra Nevada = 4%
FHP National Risk Maps
Whitebark pine restoration• Speed up natural selection by developing and
planting blister-rust resistant seedlings.
• Protect against MPB; reset succession; mitigate
climate change.
Steps in developing blister rust resistance
Protect ripening cones.
Harvest cones.
Grow
seedlings.
Screen
seedlings for resistance.
Protect resistant seed
sources against
mountain pine beetles.
Plant
seedlings.
R. Sniezko
Restoration actions that can benefit from biotechnology*
Developing rust resistance: high priority
Current protocol
• Select putative resistant trees
• Harvest cones
• Grow seedlings until 2-3 years of age
• Screen for genetic resistance through WPBR inoculations
• Score resistance responses for 1 to 5 years
• Outplant field trials
• Goal: Establish seed orchards
(*Tomback and Sniezko. 2017 Western Five-needle White Pine Knowledge Gaps/Priority Needs Informational Brief for WWETAC)
Genomic approaches would shortcut this process
• Blister rust resistance in whitebark pine is highly polygenic.
• Resistance varies geographically (Sniezko et al. 2011; Mahalovich and Hipkins 2011).
• Identification of the genes and their interaction; understanding variation across whitebark pine’s geographic distribution.
Need a high quality genome reference sequence for whitebark pine, likePineRefSeq Project funded by USDA/NRI in 2011.
• Consortium effort to provide high quality genome reference sequences for conifers: Loblolly pine, sugar pine, Douglas-fir.
• Cost to construct a reference sequence for whitebark pine, building on the sugar pine sequence: estimated at about $1 million by D. Neale.
(Mangold, R. 2014. Expansion of whitebark pine restoration methods through tree genomics. Forest Service Briefing Paper. Pacific Northwest Research Station, October 28, 2014.)
Other genomic/biotech applications to facilitate restoration and climate change mitigation
Products expected • Shortcuts for identifying parent trees with blister rust resistance--for example,
rapid tests for resistance, bar-coding.
• Possible ways to identify parent trees with resistance to mountain pine beetle.
• Refined seed transfer zones.
• Determining which genes vary with environment and which environmental factors.
• Determining adaptive potential within populations.
• Determining appropriate genotypes for assisted genotype migration or transfer to mitigate climate change in appropriate steps.
• Identifying genes for resistance in Eurasian stone pines to increase options.
• Strategic breeding programs to develop durable resistance without losing adaptation.
Genomics/biotechnology should be applied to identifying and sorting natural genotypes to attain better adapted phenotypes
Caveats in relation to genomic manipulation:
• Hybridization could result in loss of ESA protection.
• May not have public support.
• May have unintended consequences because of pleiotropic effects.
R. Sniezko, Dorena Genetic Resource Center
Canadian-U.S. collaboration
Interaction and collaboration with Canadian colleagues concerning whitebark pine management and restoration has taken multiple forms.
• Participation in joint professional meetings and exchange of information annually.
• Invitations for U.S. scientists to share expertise, collaborate on projects, and participate in graduate education in Canada, and reciprocal invitations from the U.S.
-- Beginning: 2003 Parks Canada Whitebark Pine and Limber Pine Workshop, Calgary.
--Transboundary: Crown of the Continent High Five Working Group, chartered in 2016.
• Shared participation through the Whitebark Pine Ecosystem Foundation and the establishment of the Whitebark Pine Ecosystem Foundation of Canada.
--Annual Whitebark Pine Ecosystem Foundation Science and Management Workshop, held in Canada every third year (Jasper National Park, Alberta, 2017)
• Coauthorship on U.S. reports and research.
• U.S. scientist and manager review of provincial and Environmental Canada documents related to whitebark pine (and limber pine, which is approved for endangered status under COSEWIC), and requested contribution to U.S. F&WS status review.
Canada has taken leadership in providing protected status for whitebark pine at provincial (Alberta) and national levels (SARA).
• Canada has also been at the conceptual forefront on both application of genomics and forest management under climate change (e.g., Aitken et al. 2008, McLane and Aitken 2012).
The U.S. Forest Service has nearly 70 years’ experience in developing planting stock with blister rust resistance, dating to the early 1950s.
• The U.S. Forest Service has the expertise, infrastructure, capacity, and protocols.
• Primarily two western U.S. facilities which also engage in operational production of seedlings for planting: Dorena Genetic Resource Center and Coeur d’Alene Nursery.
• Canadian agencies, including Parks Canada and BC Forestry are now utilizing our expertise and screening facilities.
• They are at the early stages in developing their own capacity.
• Previously, they relied on field trials, which is more definitive but restricts their capacity. Canada’s natural resource agencies face funding limitations similar to the U.S.
Canadian-U.S. differences in strengths
National Whitebark Pine
Restoration Plan Strategic Plan Approach and Vision
Inyo National Forest, CA
The U.S. Forest Service, Washington Office, American Forests, the Whitebark Pine Ecosystem Foundation are partnering to develop a core area restoration plan for the U.S. distribution of whitebark pine.
Collaborators include all federal and state agencies and tribes with whitebark pine under their jusrisdiction.
This strategic plan will identify selected areas within the U.S. range of whitebark pine for priority restoration, focusing resources.
Components to plan: tentative 10-20 year timeline
Core area restoration
plan
Nominated core areas from each
jurisdiction
Priority designation within each nominated
area
Criteria used to identify core areas
Proposed restoration actions within core areas
Implementation costs for restoration
action across nominated area
Monitoring and adaptive
management strategies
Cost of restoration
One deliverable from the National Whitebark Pine Restoration Plan is estimation of restoration costs for implementation. Funding restoration is anticipated as the outcome of a partnership between government and NGO organizations.
• American Forests develops corporate sponsorships and can work towards restoration goals.
• Restoration on the rangewide scale for whitebark pine will entail costs for:
--Accelerated identification of resistance to WPBR (currently at $1200/family.)
--Seed orchard establishment and maintenance ($15,000/acre).
--Better characterization of adaptive genetic variation across the range.
--Development of restoration strategies considering climate change mitigation.
--Infrastructure development to support resistance development.
--Infrastructure support for operational seedling production.
• Efforts will absorb as much funding as we can raise (several $ million per year).
• Estimated costs per management/restoration action are available from me.
Rob Mutch
The overarching goal of whitebark pine
conservation and restoration is to develop
and sustain healthy and resilient whitebark
pine communities in the face of current and
future challenges.
The National Whitebark Pine Restoration Plan
Thanks to
• Gregg DeNitto, Annalisa Ingegno, Frank Sapio – USFS, R1 Forest Health
Protection
• Jeanine Paschke – USFS, Forest Health Assessment & Applied Sciences
Team, Ft. Collins
• Bob Keane, Lisa Holsinger, Molly Retzlaff – USFS, RMRS, Missoula Fire
Sciences Lab
• Lynn Resler-Virginia Tech, George Malanson-University of Iowa.
• Many research colleagues and students over the years.