Key Considerations for Managing the

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The impact of Whitebark Pine mortality on Clark's Nutcracker demography: Key Considerations for Managing the Will restoration work? Clark's Nutcracker-Whitebark Pine Mutualism Taza Schaming tazaschaming@gmail.com

Whitebark Pine & Clark s Nutcrackers L. Rentz

How is widespread habitat loss impacting Clark s Nutcrackers? Population status? Behavior? Life history?

Research Objectives 1) How stable & resilient is this keystone mutualism? -> How is nutcracker demography & behavior associated with habitat type, quality & abundance? Breeding ecology & reproductive success Home range size Seasonal habitat selection Seasonal habitat use Foraging ecology Emigration/dispersal behavior 2) Suggest management strategies to promote stability & resilience L. Rentz T. Schaming

Methods (2009-2016) STUDY SITE 23,500km 2 N = 7 satellite-tracked N = 187 trapped N = 76 radio-tracked N = 3,135 10 min Ψ surveys, @ 238 random points N = 74.5 hours fledgling surveys Jada Schaming Taza Schaming Pocholo Martinez Rob Curtis

Highlighted Results Breeding ecology & reproductive success Home range size Seasonal habitat selection Seasonal habitat use Foraging ecology Emigration/dispersal behavior M. Ouchida

(Schaming 2015) Is reproductive success related to whitebark pine cone crop? No breeding occurred population-wide in 2 years! -> If increasing numbers of low whitebark pine cone crops leads to fewer breeding years, this could lead to serious population-level consequences

Is fledging success associated with habitat? 33 active nests 35% fledged Habitat nest site local area landscape scale 1 st predictions 2 nd classification tree: 20 fixed effects 3 rd 16 mixed models: 4 fixed effects; AICc N = 29 (2012) -> Lots of whitebark pine & only small amount of Douglas-fir optimal -> BUT high whitebark pine mortality surrounding nest counteracts benefits of nesting in landscape w/whitebark habitat (Schaming 2016 Dissertation)

Highlighted Results Breeding ecology & reproductive success Home range size Knowing home range size helps ensure management is carried out at the appropriate scale. Seasonal habitat selection Seasonal habitat use Foraging ecology Emigration/dispersal behavior M. Ouchida

Home Range Size JAN FEB MAR APR MAY JUN JUL AUG SEPT OCT NOV DEC -> Nutcrackers use an extensive area! -> Is this normal in different years? In different quality habitats? ~100 ha (n = 19) ~200 ha (n = 25; 56) ~340 ha (n = 35) ~1,300 ha (n = 20) ~17,440 ha (n = 7) (Schaming 2016 Dissertation)

Highlighted Results Breeding ecology & reproductive success Home range size Seasonal habitat selection Habitat selection Seasonal = habitat behavioral use process by which individuals choose certain habitats to use from what is available. Foraging ecology Emigration/dispersal behavior M. Ouchida

Habitat Selection: Home range within available habitat -> Optimal habitat mosaic includes whitebark pine & Douglas-fir & limber pine (Schaming 2016 PLOS ONE, Dissertation)

(Schaming 2016 PLOS ONE, Dissertation) Habitat Selection: Locations within home range Satellite-tagged birds: 86% (n = 6) selected Douglas-fir in a disproportionately high amount -> Optimal habitat mosaic includes whitebark pine & Douglas-fir

Highlighted Results Breeding ecology & reproductive success Home range size Seasonal habitat selection Seasonal habitat use Habitat use choice, & observed pattern may be driven by external constraints Foraging ecology (e.g. competition). Emigration/dispersal behavior M. Ouchida

Seasonal Habitat Use: Occupancy Models Which resources were important drivers of occurrence? Nutcracker occurrence vs. Whitebark pine - cone crop: presence/absence and density - at local site: presence/absence and density - at landscape scale Douglas-fir - at local site: presence/absence and density - at landscape scale Detection vs. Tree density Local whitebark pine OR (Schaming in review)

Results: Detectability (Schaming in review)

Habitat Use (Fall harvest season) -> As long as seeds were present, high probability of nutcracker occurrence, even at low cone crop densities & in high mortality stands -> Manage for presence of cone bearing trees, NOT higher density -> Manage whitebark pine at a landscape scale, not a stand scale (Schaming in review)

Habitat Use (when whitebark pine cones not available) BREEDING SEASON POST-HARVEST SEASON Whitebark pine (ha) Minimum Mean Maximum -> Optimal = habitat mosaic w/whitebark pine & low abundance of Douglas-fir -> Preference & prevalence yield different conclusions (Schaming in review)

Highlighted Results Breeding ecology & reproductive success Home range size Seasonal habitat selection Seasonal habitat use Foraging ecology Understanding how foraging behavior varies under different environmental conditions & in different Emigration/dispersal habitats enables better behavior predictions of how animals will respond to environmental change. M. Ouchida

Foraging Events ( 5%) WINTER LATE SUMMER BREEDING SEASON FALL HARVEST -> Whitebark pine & Douglas-fir are important foods -> Cached seeds are small portion of diet, but slightly higher after high cone crop -> More invertebrates in diet not correlated w/increased breeding EARLY SUMMER POST-HARVEST (Schaming 2016 PLOS ONE, Dissertation)

Location of Aboveground Insect Foraging Events -> Douglas-fir, aspens & snags are important foraging locations -> Negative association between dead trees & fledging, BUT snags are great for foraging (Schaming 2016 Dissertation)

Highlighted Results Breeding ecology & reproductive success Home range size Seasonal habitat selection Seasonal habitat use Understanding Foraging helps ecology evaluate population viability in a region as a function of habitat, & determine the scale at which to focus management efforts. Emigration/dispersal behavior M. Ouchida

Results: Dispersal When do nutcrackers disperse? Where do they go? 71% 7% 71% -> A high proportion disappeared during (Haroldson et al. 2015) moderate to high cone crops -> Moved over considerable distance 300 km (Schaming 2016 Dissertation)

FLEW SOUTH (btwn 8/24/15 & 9/26/15) & BACK TO WY mid-june 2016 FLEW SOUTH (btwn 8/8/15 & 1/10/16) -> Dispersed during (n = 2-4) or after (n 2) the whitebark pine seed harvest *Not retrieving seeds overwinter -> n = 4/5 which left, returned to GYE (4 overwintered in Utah) FLEW SOUTH (btwn 9/5/15 & 11/18/15) -> Availability & health of alternative habitats is important to GYE nutcrackers! *Focus on metapopulation stability & resilience! FLEW SOUTH (btwn 8/10/15 & 9/9/15) & BACK TO WY btwn 6/11/16 and 7/3/16 & BACK TO WY btwn 6/30/16 and 7/3/16

Broad Recommendations for effective conservation of the Clark s nutcracker-whitebark pine mutualism 1 Long-term, range-wide studies (preference & prevalence) Understand metapopulation stability as conditions change 2 Adaptive management approach Behavior & population vary w/density of species & habitats Monitor as conditions change & management implemented 3 Managing for persistent, stable local populations Conservation tool: present = available to disperse seeds ( dispersing seeds & persistence) Resident birds disperse seeds further 4 Effective conservation may depend on protection of a network of key habitats (Schaming 2016 Dissertation)

Thank you! Acknowledgements Advisor: Janis Dickinson, Committee: André Dhondt, Evan Cooch, John Fitzpatrick Wesley Hochachka, Irby Lovette, Gus Axelson, Bert Raynes, Susan Patla, Chuck Schneebeck, Dick Qua, Andy Royle, Lisa Robertson, Erica Mudrak Teresa Lorenz, Bob Keane, Anya Tyson, Kori Blakely, Tyler Stuart, Ned Corkran, Lee Tafelmeyer, Sean Beckett, Asya Rahlin, Jerry Cole, Tim Perez, & Jaime Ervin Dept. of Natural Resources & Lab of Ornithology students & faculty, Jackson Hole Bird Club, Nature Mapping Jackson Hole, Lighthawk Bridger-Teton National Forest, Wyoming Game and Fish, Grand Teton National Park, Northern Rockies Conservation Cooperative Mia Davis, Laura Martin, Stephanie Leonard, Dawn Webster, Nina Chaopricha, Jen Shirk, Nini Clayton, Lois and David McDonald, Ayano Hamilton Pocholo Martinez, Peter & Teal & Jada Schaming Funding (40 grants + NASA & CLO fellowships) Athena Fund of the Cornell Lab of Ornithology Meg and Bert Raynes Wildlife Fund Cornell Lab of Ornithology NASA Harriett G. Jenkins Pre-doctoral Fellowship Mazamas Graduate Student Research Grant American Ornithologists Union Cornell Lab of Ornithology Halberstadt Graduate Fellowship Cornell University Research Travel Grant Fischer Equipment Grant Cornell University Sigma Xi Grants-in-Aid of Research Wilson Ornithological Society Paul A. Stewart Research Award Havahart Equipment Grant Cornell Lab of Ornithology Samuel and Linda Kramer Award Andrew W. Mellon Student Research Grant Explorers Club Exploration Fund Conservation and Research Foundation Western Bird Banding Association American Philosophical Society Lewis and Clark Fund Cornell Lab of Ornithology Kramer Graduate Fellowship Wyoming Wildlife The Foundation, Wyoming Community Foundation The Garden Club of America Frances M. Peacock Scholarship Charles Redd Center for Western Studies Summer Award for Upper Division and Graduate Students

Questions? M. Meyers L. Rentz T. Schaming

Seed Energy A nutcracker needs ~12,000 Cal for winter survival Species Cal/seed # seeds/winter Whitebark pine 1.23 ~10,000 Limber pine 0.58 ~20,000 Douglas-fir 0.06 ~200,000 Engelmann spruce 0.24 NA Lodgepole pine 0.02 NA Subalpine fir 0.92 NA

Conclusions * When designing a conservation plan for plant-animal seed disperser mutualisms, consider seed dispersers : 1: Population status & behavioral plasticity at appropriate scale 2: In relation to all of the habitats on which they depend 3: Under variable & changing environmental conditions 4: Preference & prevalence