Earthwatch 2018 Field Report RESTORING FIRE, WOLVES, AND BISON TO THE

CANADIAN ROCKIES

PI: Dr. Cristina Eisenberg

REPORT COMPLETED BY: PI Dr. Cristina Eisenberg, and Co-PI Dr. David E. Hibbs PERIOD COVERED BY THIS REPORT: January 2018 – December 2018

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Dear Earthwatchers,

Thank you for joining us afield in 2018 on our research project, Restoring Fire, Wolves, and Elk to the Candian Rockies. While this research has been ongoing since 2006, it became an Earthwatch project in 2015. Several of you joined us previously, and having you with us felt like being with family and good friends. Those who joined us for the first time became valued new friends, and everyone contributed much. As you know, our work takes place in Waterton Lakes National Park, Alberta (WLNP), a UNESCO International Biosphere Reserve, a World Heritage Site, and an International Peace Park. This park lies in the Crown of the Continent Ecosystem, one of the most intact landscapes in North America. It contains all the ecological elements present in the early 1800s, except for free-ranging bison. You can think of this ecosystem as a three-legged stool. The “legs” are keystone ecological forces: fire, predation by wolves, and bison herbivory. European settlers eliminated all of these in 1880, but today this system is gradually being rewilded. Wolves returned on their own in the early 1990s. In 2003 park managers began setting controlled prescribed burns, including since 2008, large fires in two sites: the Y-Camp and Eskerine Complex. But according for the International Union for the Conservation of Nature (IUCN), fire and wolves are not enough to stabilize our ecological three-legged stool. Accordingly, a bison reintroduction has occurred in Banff to the north, and is being planned by the Iinnii Initiative for the Badger-Two Medicine area and Chief Mountain, just south of our study site, with an additional bison herd to be established on Kainai First Nation (Blackfoot) land, just east of the park. All of this creates an exciting opportunity for us to study the ecological impacts of restoring fire, wolves, and bison. However, this involves collecting a huge amount of data—and that’s why Earthwatch volunteers are essential! This past field season, you worked alongside my co-PI David Hibbs , graduate student Chris Anderson, our field technicians, and me as we continued our work on Kainai First nation historic tribal timber land, and also collected data in the park. The large, high-quality dataset you helped us collect enabled the analyses in this field report and directly supported ecological restoration at both sites. You took action to enable us to collect crucial fire-severity data for the Kenow wildfire, a large, lightning- caused fire that unexpectedly burned our entire study site in 2017 with mostly extreme severity, changing our research landscape and creating new “burning” questions for 2018—and beyond. Because wildfires are increasing in severity globally, understanding the ecological impacts of a fire such as the Kenow wildfire is crucially important to science. Thanks to you and other Earthwatch citizen scientists, our project will be contributing significantly to humanity’s understanding big, severe fires, and of how to restore landscapes and create more resilient ecosystems in their aftermath. Looking ahead to 2019, we’ll be asking how the effects of wildfire differ from the effects of very carefully controlled prescribed burn set by humans—and how and animal community responses to wildfire and prescribed burns might differ. Our 2018 data show many interesting things, most notably the vigorous positive response and resilience of this ecosystem to extreme-severity wildfire. Together we learned that prescribed burning creates more open aspen stands, and that elk browsing on burned aspen was negligible, likely due to a variety of factors, including plant defenses and risk of predation by wolves. To underscore this, within aspen stands we found multiple carcasses of fresh, wolf-killed elk. You helped us discover that the

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Eskerine prairie is so healthy that it continues to be nearly completely composed of native grass . You also helped us find Pleistocene bison and wolf bones in our study site, near a buffalo jump, and Blackfoot artifacts, such as stone tools and arrowheads, which demonstrate that the relationships we’re studying are ancient. In 2018 we entered the third year of the Kainai First Nation Community Fellows Program, and hosted 34 fellows. We were delighted to have teachers and community members from the tribe join us as Earthwatch team members. They shared important Traditional Ecological Knowledge insights and provided much inspiration for all, supported by our Kainai field technicians, Elliot Fox, Justin Bruised Head, Alex Shade, and Dustin Fox. We’e thrilled that this program will be growing significantly in 2019. So until next field season, our heartfelt thanks to each of you for all you contributed to our project. Wherever life leads you, may you find opportunities to advance conservation, to take action to help restore landscapes and their species, and in so doing have amazing adventures. My colleagues and I feel fortunate to have crossed paths with you and hope to see some of you again afield to work together to do science that helps save nature.

Warm Regards,

Chief Scientist Earthwatch Institute

Earthwatch participants heading into the field at the start of the field season • Photo by Cristina Eisenberg

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Wildflower superbloom after the Kenow wildfire • Photo by Earthwatch participant Angela Chan

Earthwatch participants in the field with us on the Blood Timber Limit • Photo by Elliot Fox.

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Earthwatch participants get a mapping lesson • Photo by Cristina Eisenberg

Earthwatch Community Fellows tracking wildlife • Photo by Cristina Eisenberg

Kainai Community Fellow Austin Fox Earthwatching • Photo by Claire Doe

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Earthwatch participants, end of the field season, Waterton, Alberta • Photo by Elliot Fox

Pleistocene bison skull found by Earthwatch participants, Team 1 • Photo by Cristina Eisenberg

SUMMARY In 2018 we had several accomplishments and discoveries:

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1. We found that the Kenow wildfire burned our study site with 75.4% extreme and 12.6% high severity, which is unprecedented for any large fire in North America, and even perhaps globally. We wrote and submitted a journal article about this surprising finding. 2. Per our microhistological elk and bison diet analysis, we found elk and bison eating almost no aspen. Elk browsing on aspen dropped after fire, but we didn’t expect it to decline to nearly nothing as it did. The extremely low amount of browsing on aspen, an important elk food, is perhaps due to predation risk or to the defense compounds aspen produce to make them less edible after a fire. 3. After the Kenow wildfire, the proportion of native grass on the prairie increased, and the proportion of invasive non-native grasses decreased, demonstrating the benefits of even extreme-severity wildfire. 4. Wolf activity was moderate in the park and high on tribal land. Wolves shifted their activity to portions of WLNP closed to the public because of fire damage. 5. On the Timber Limit we found deer the dominant herbivore by number, but not by biomass. This differs from WLNP where elk are the dominant herbivore by number and biomass. 6. We implemented the third year of the Kainai First Nation Community Fellows Program and continued our research on tribal land adjacent to WLNP to gather data to prepare for the return of bison to this landscape.

Special thanks Elliot Fox, for providing exemplary leadership and guidance as our lead Kainai Technician from 2016-2018.

Lead Kainai technician Elliot Fox • Photo by Cristina Eisenberg

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Kenow Wildfire, Waterton • Photo by Ryan Peruniak

GOALS, OBJECTIVES, AND RESULTS

Background Since 2008, we have been studying the complex dynamics of aspen (Populus tremuloides), elk (Cervus elaphus), fire regimes, and wolves (Canis lupus) in aspen and grassland communities in the elk winter range of Waterton Lakes National Park (WLNP) in the Crown of the Continent Ecosystem. Our work takes place in two WLNP sites: the Eskerine prescribed burns, which burned 852 ha in 2014 and 772 ha in May 2017; and the Y-Camp prescribed burns, which burned 835 ha in April 2008, and 762 ha in April 2015 (Collingwood 2016, 2017). As an unburned control site, we have been using the Crooked Creek area, which lies just north of the Chief Mountain highway from the Y-Camp. In 2017 we added Beebe Flats on the Blood Timber Limit as a second unburned control site. These control sites are very similar ecologically to each other and to our Y-Camp and Eskerine Complex Study sites. All contain high elk density, an active, but variable, wolf population, and similar plant communities and weather. They all lie in the foothills-parkland ecoregion (Achuff et al. 2005). The Kainai First Nation owns the Blood Timber Limit, a 1,974 ha southwest Alberta forest tract established in 1888. It lies within the Blood Reserve, which is located in the Oldman River watershed, with the Belly River demarcating its eastern boundary (Fox 1998). WLNP borders the Blood Timber Limit to the west, south, and east. The purpose of this traditional land-use site is to provide the Kainai First Nation with hunting and access to timber for home construction, firewood, and lodgepoles. In the 1960s, several gas wells were established on the Blood Reserve, but development has been minimal. The land has also been used for cattle (Bos taurus) grazing, which ended in 1988.

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The ecology of the Blood Timber Limit is similar to that of WLNP. Ecoregions within the Blood Timber Limit include foothills-parkland, montane, subalpine, and alpine. Forests are mainly coniferous, dominated by lodgepole pine (Pinus contorta), Douglas fir (Pseudotsuga menziesii), and white spruce (Picea glauca). Aspen comprises 25% of the forest cover (Fox 1998; Achuff et al. 2005). Belly River outflow created an alluvial fan that contains wetlands and a 3-ha grassland bordered by aspen, called Beebe Flats, an important elk range. On the Blood Timber Limit, our sampling is taking place in Beebe Flats. This site contains diverse vegetation, including rare species such as mountain brome (B. carinatus), ecologically important native grasses—Idaho fescue (Festuca idahoensis) and Parry’s oatgrass (Danthonia parryi)—and noxious weeds such as smooth brome (B. inermis) and Timothy grass (Phleum pratense). In the early 1990s, Elliot Fox, working with park ecologists, found the first wolf den in southwest AB since this species was extirpated in the 1920s. The den was located adjacent to Beebe Flats, in WLNP (Fox and Van Tighem 1994). Wolves have had a strong presence in this area since Fox and park ecologists found this den. Until 2008, none of the sites we are studying had burned since 1910 (Barrett 1996). In 2002, WLNP began a prescribed burn program. This program has involved setting the Eskerine and Y-Camp prescribed burns (among others) and has resulted in several mixed-severity fires (Schwanke 2008; Murphy 2016). Additionally, in September 2017, the Kenow wildfire, caused by a lightning strike in British Columbia, burned WLNP, including our Eskerine Complex and Y-Camp study sites, as well as a portion of our Crooked Creek control site. It did not burn the Blood Timber Limit. Per park differenced Normalized Burn Ratio (dNBR) analysis, the Kenow wildfire burned with mostly extreme severity (Fig. 1, Collingwood 2017), creating an opportunity to compare the impacts of prescribed burns to those of wildfire. Fire has been a part of western North American ecosystems during the Holocene and Anthropocene geological epochs—the past 12,000 years (Whitlock and Bartlein 1997). In southern Alberta (AB) and British Columbia (BC), using fossil pollen found within sediment cores, palynologists have identified Holocene plant community composition. They have linked climatic shifts to shifts in fire regimes and shifts in plant communities. For example, working in Kootenay National Park, BC, Hallett and Walker (2000) found fire frequency increased and reached its maximum during the early to mid- Holocene, from 8200 to 4000 BP, and that this corresponds to the dry, hot Hypsithermal period. In Crowsnest Pass, AB, since 7500 BP, periods of increased grass pollen (vs. arboreal pollen) have been correlated to warm periods (Driver 1978). In Kananaskis, AB, Mott, and Jackson (1982) found pollen cores containing grass and aspen occurring in a similar temporal pattern as other studies have observed. Taking a closer spatiotemporal look, in WLNP Barrett (1996) linked multiple drought periods with large- fire activity in the late 1600s to late 1700s in grassland and dry Douglas-fir communities. In North America, ecologists have found evidence of both wildfire and anthropogenic fire since the late Glacial Maximum and the beginning of the Holocene geological epoch. Wildfires originated mostly due to lightning strikes, with a fire-return interval of 150-300 years in the northern Rocky Mountains (Amoroso et al. 2011; Hallett and Walker 2000; Johnson and Fryer 1986). However, Barrett (1996) found a shorter fire-return interval in WLNP, linked to severe drought concurrent with Anthropocene Euro-American expansion. In his dendrochronology tree fire-scar study, Barrett found that fire frequency declined after 1940, due to efficient fire suppression. He also found fire-return

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intervals shorter in foothills-parkland plant communities than in montane and sub-alpine plant communities. Indigenous people frequently set fires to improve wildlife habitat. These fires created early-seral plant communities, increased berry-producing shrub yield for deer (Odocoileus spp) and elk, and kept grasslands open for bison (Barrett and Arno 1982; Turner, Ignace, and Ignace 2000; Boyd 2002), thereby improving subsistence resources for humans. In southwest Alberta, Indigenous people set fires in the late-spring dormant growing season, but in other regions (e.g., the Midwestern United States) set them later in the year (Boyd 2002). Dormant-season fires occur when there is less above-ground annual growth of perennial grasses and shrubs to burn, resulting in lower fire severity and improved forage (Boyd 2002). Late growing-season fires occur when grasses and shrubs are tall and dry. Due to greater biomass of above-ground fuels at that time, these fires can be higher in severity than dormant-season fires and do more damage to plant communities, (Brockaway, Gatewood, and Paris 2002). The WLNP prescribed burn program is an attempt to return the foothills-parkland ecoregion within the park to historic conditions (e.g., more open grassland, smaller, more structurally complex aspen stands), based in part on Indigenous fire regimes in this landscape and Traditional Ecological Knowledge (TEK), defined as “a cumulative body of knowledge and beliefs, handed down through generations by cultural transmission, about the relationship of living beings (including humans) with one another and with their environment (Berkes, Colding, and Folke 2000).” The lightning-caused Kenow wildfire falls within the historic fire-return interval in southwest Alberta (Jonnson and Fryer 1987; Barret 1996). The Kenow wildfire was 35,000 ha in size. Of the 19,302 ha of this fire within WLNP, 75.4% burned with extreme severity, 12.6% burned with high severity, 6.4% burned with moderate severity, and 5.6% burned with low severity (Fig. 1). It burned the Eskerine Complex portion of our study site with 97% extreme severity. Nearly a megafire, defined as one ≥40,000 ha in size, it may have differed from historic fires in this system due to its mostly extreme severity (Fig. 1). Wildfires in North America, including western Canada, have a history of being very heterogeneous in severity, creating a fire mosaic in a landscape (Turner et al. 1994; Amoroso et al. 2011; Halofsky et al. 2011; Morgan et al. 2014; Dunn and Bailey 2016). Globally, due to climate change, wildfires have been burning with greater severity and possibly increasing in size (Parisien et al. 2011; Krawchuck, and Moritz 2011; Dennison et al. 2014). Fire ecologists characterize fires as either fuel-limited or climate-limited. The Kenow wildfire was a climate-limited fire, in which a moist spring in 2017 created abundant fuel to carry fire, and unusually hot, dry summer conditions caused this lightning fire to grow rapidly (Krawchuk and Moritz 2011; Johnston et al. 2017). In North America, aspen have been widely observed to be declining over the past 70 years (Kashian, Romme, and Regan 2007; Hogg 2008; Worrall et al. 2010), but in WLNP, the opposite has occurred, with aspen cover increasing significantly over the past 70 years, to the point that it has encroached on ecologically important, at-risk habitat, such as fescue grassland (Levesque 2005; Eisenberg, Hibbs, and Donato 2014). Aspen declines have been observed in sites where fire is used as a treatment to restore this fire-adapted species in the absence of wolves (Bartos, and Mueggler 1981; Canon, Urness, and DeByle 1987; Hessl, and Graumlich 2002). In systems that contain wolves, elk spend less time in habitat that makes it difficult to escape wolves, thereby reducing browsing pressure on aspen (Hebblewhite and Merrill 2007; Kuijper 2013; Eisenberg, Hibbs, and Ripple 2015). However, in the

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absence of a wolf population, elk browse aspen sprouts to death, resulting in little to no recruitment of aspen into the forest canopy (Romme et al. 1995; Hobbs 1996). Stand dynamics and prescribed burns also influence the density of snags, which provide important avian habitat (Lee 1998; McComb et al. 2010).

Figure 1. Kenow wildfire, September 2017, dNBR. Map credit: Parks Canada, Adam Collingwood.

Native grasslands have been declining in North America in general and in the WLNP foothills- parkland ecoregion since the 1880s (Samson, and Knopf 1994; Achuff et al. 2005; Hop et al. 2007; Johnson, and Fryer 1986). Ecologists attribute this to elimination of bison (Bison bison) and fire (Larson 1940; Campbell et al. 1994; Knapp et al. 1999), introduction of non-native agronomic grass species, and suppression of fires set by Indigenous people (Johnson, and Fryer 1986; Turner, Ignace, and Ignace 1999). WLNP contains a large elk herd that between 1990 and 2015 fluctuated from 528 to 1,170 individuals, and is currently estimated at around 1,000 individuals (Johnston 2016). Fire increases resources for elk, improving forage as well as browse species by creating early-seral habitat types (Debyle, Urness, and Blank 1989; Singer, and Harter 1996; Swanson et al. 2010). Fire could be used to increase carrying capacity for elk and bison in WLNP, since both species are drawn to post-fire forage, and in the case of elk, post-fire browse (Hobbs et al. 1981; Baker 2009, pp. 170-187; McComb 2015, pp. 181-187).

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Restoring free-ranging, wild, plains bison to the Crown of the Continent Ecosystem has been under consideration for nearly two decades, but more recently has become a possibility, supported by the Buffalo Treaty (Little Bear 2014). After completing an Environmental Assessment, in early 2017 Parks Canada reintroduced a herd of 16 plains bison to the eastern slopes of Banff National Park. In early 2018, the herd was released into the 1,189 km2 reintroduction zone in the park’s eastern valleys.

Bison Treaty celebration • Photo by Cristina Eisenberg

The Iinnii Initiative is a partnership between the Wildlife Conservation Society, the Blackfoot Confederacy, and several Canadian and US government agencies that seeks to restore bison to Blackfoot Confederacy lands, which span the US-Canadian border (Iinnii means bison in the Blackfoot language). The Iinnii Initiative has been making strong progress toward establishing the policy and partnerships that would enable bison reintroduction on the Blackfeet Reservation in Montana (where a currently captive herd of what were originally 88 animals sourced from Elk Island National Park, AB, is being held) and in the Chief Mountain area, which would include WLNP and the Blood Timber Limit, and the Kainai Band Ranch—a protected rangeland owned by the Kainai First Nation. This initiative is encouraging a shift in the management status of this species at provincial and state levels in Canada and the US from “livestock” to “wildlife.” Currently the IUCN gives this North American species “Near Threatened” Red List status (IUCN 2016).

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Captive bison in Waterton • Photo by Ryan Peruniak WLNP has had a captive bison herd since 1952. They are kept in the summer bison paddock and the winter bison paddock. In WLNP, Barb Johnston conducted a comprehensive feasibility analysis of establishing a semi-free-ranging bison population in the park. She found that “Establishing free-ranging bison in the park would not be feasible because the benefits to the ecological integrity of the park and to the experience of visitors would not be significant enough relative to the cost at this small geographic scale.” Although many logistical challenges could be overcome, the ecological repercussions of confining a wide-ranging species to an area where it has been replaced by a large herd of elk as the dominant grazer, and the resulting potential impacts of increased year-round grazing pressure on rare and sensitive fescue grasslands would outweigh the benefits of bison reintroduction (Johnston 2008). WLNP and the Blood Timber Limit lie within an ecological corridor from Banff National Park to the Badger-Two Medicine area identified by the IUCN American Bison Specialist Group as ideal for a landscape-scale, free-ranging, wild bison reintroduction (IUCN ABSG 2016), in keeping with recommendations about scale and scope of such efforts by Sanderson et al. (2008), and by the Kainai First Nation about historical use of their lands and TEK (Fox 1998). Resilient fescue grassland and fire are important historical features of bison habitat (England and Devos 1969; Feden et al. 1974; Shaw, and Carter 1990; Shaw 1995; Binnema 1996; Eisenberg 2018). As such, and per WLNP management requests, we are contextualizing our research with the potential to inform bison conservation. This report presents our progress and findings from the 2018 field season. For further information on our methods and other results, please refer to our 2014 final report of the first five years of sampling of the 2008 Y-Camp prescribed fire and our 2015, 2016, and 2017 annual reports (Eisenberg, Hibbs, and Donato 2014, 2015; Eisenberg, Hibbs, and Edson 2016; Eisenberg and Hibbs 2017). Our research supports the WLNP Management Plan. Specifically, our research helps WLNP administrators “restore and monitor natural heritage, while showcasing innovative activities.” It further provides an opportunity for the park to engage a variety of interested stakeholders to collaborate in implementing solutions to restore the prairie and increase the ecological resiliency and biodiversity of this landscape. Given the cultural significance to the Kainai First Nation of the prairie, fire, wolves, and bison, our work helps increase the profile of this historic United Nations Educational, Scientific, and Cultural Organization (UNESCO) International Biosphere Reserve site and improve the condition of its cultural resources (WLNP 2010; Parks Canada 2011). Research efforts such as ours have been identified as a critical component of adaptive management in order to better understand fire behavior, impacts, and risk (Thompson, Dunn, and Calkin 2015).

Sampling strategy We measured wildlife use of the Timber Limit (ungulates and large carnivores). We also measured the impacts of the Kenow wildfire within Y-Camp and Eskerine prescribed burn units. The 2017 Eskerine prescribed burn occurred in May 2017 (Fig. 2). The 2014 Eskerine prescribed burn was a subset of the Red Rock Complex prescribed burn (Fig. 3). From 2003 – 2014, the park set several prescribed burns in the Eskerine Complex (Fig. 4). All fires lie within elk winter range in WLNP (Fig. 5). The Kenow wildfire, a 35,000 ha wildfire, burned our Eskerine and Y-Camp study sites, as well as our

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Crooked Creek control site, mostly with high severity per the dNBR analysis done by WLNP geospatial scientist Adam Collingwood (Fig. 1). We have been measuring each prescribed burn one year post-fire, applying an alternate-year sampling strategy (Table 1). Additionally, we have been collecting pellet transect data (number of ungulate pellet piles, carnivore scats, carcasses) two years after each prescribed burn (Table 1), to produce an index of elk and deer density (Neff 1968; Collins, and Urness 1981; Fuller 1991; St. Laurent, and Ferron 2008; Sanchez et al. 2009; Eisenberg, Hibbs, and Ripple 2015). This temporal sampling strategy is necessary, because an aspen/grassland community’s response to fire develops over a multi- year period (Baker 1997; Baker 2009, pp. 170-187). We currently have a five-year time series of complete surveys for the Y-Camp prescribed burn (2008-2012), with a survey of the 2015 Y-Camp prescribed burn done in 2016, and aspen parkland edge surveys done in 2013, 2014, and 2017. For the Eskerine prescribed burn site, we have a complete pre-fire dataset gathered in 2008 and data from the full post-fire survey completed in 2015 (Table 1). We have adapted our sampling intervals to capture the ecological impacts of a shorter fire-return interval (e.g., every 3 years vs. 8 years, as with the 2008 and 2015 Y-Camp prescribed burns).

Figure 2. Eskerine prescribed burn unit, May 2017. Map credit: Parks Canada, Adam Collingwood.

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Figure 3. Red Rock Complex/Eskerine prescribed burn unit, April 2014. Map credit: Parks Canada, Adam Collingwood.

Figure 4. Eskerine Fire History 1833 – 2017.Map credit: Adam Collingwood.

In 2018, we conducted the sampling depicted in Table 1.

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Table 1. WLNP and Blood Timber Limit spatiotemporal sampling strategy. Abbreviations: DS = dormant season; LS = late growing season; A = aspen; P = prairie; P=planned.

2008 2009 2010 2011 2012 2013 2014 2015 2016 2017 2018 2019 2020

Y-Camp Fire Prescribed Prescribed Wildfire History burn (DS) burn (DS) (LS) Aspen X X X X X X X Pellets X X P Stand X X X X X X X X X X X Edge Survey Grass (A) X X Grass (P) X X X Fire X Severity Year 2008 2009 2010 2011 2012 2013 2014 2015 2016 2017 2018 2019 2020 Eskerine Prescribed Prescribed Complex burn (DS) burn (DS) Wildfire (LS) Aspen X P Pellets X Stand X X X Edge Grass (A) X X Grass (P) X X X Grass (BP) X X Mapping X X X (ground) Mapping X (UAS) Fire X Severity

2008 2009 2010 2011 2012 2013 2014 2015 2016 2017 2018 2019 2020 Blood Timber Limit Aspen X Grass (A) X Grass (P) X Pellets X P

2008 2009 2010 2011 2012 2013 2014 2015 2016 2017 2018 2019 2020 Control Wildfire Crooked (LS) Creek Aspen X X X X P Grass (A) X X Grass (P) X X Fire X Severity

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Goals and Objectives Our principal goal is to investigate the effect of fire treatments on elk resource selection (browsing) and how this influences aspen encroachment onto fescue grassland in a system that has very active wolf predation on elk (the dominant ungulate herbivore by number and biomass). As this research develops, and with graduate students to our project, we are expanding our questions to take a landscape-scale as well as patch-scale look at how fire affects aspen and grass communities and their diversity of flora and fauna. Managers strategize that by using fire to kill aspen trees and stimulate sprouting, the combination of fire and subsequent elk browse on aspen sprouts will diminish the extent of aspen cover and increase native grass cover, as has been observed widely in other sites, such as in Yellowstone pre-wolves (Romme et al. 1995; White, Olmsted, and Kay 1998; Hessl, and Graumlich 2002; Binkley 2007). Dormant-season burning increases grass biomass and is the preferred method of restoring grasslands where fire has been present historically but has been excluded for a prolonged period (Brockway et al. 2002). However, given that wolves are apex predators that affect elk density and behavior, wolf presence may influence elk impacts on aspen and the effectiveness of fire in restoring fescue grassland (White, Feller, and Bayley 2003; Fortin et al. 2005; Hebblewhite et al. 2005; Halofsky and Ripple 2008). Presence of additional large carnivore species influence these trophic relationships further, as may presence of non-native, ruderal grasses, which respond very positively to fire, and which include invasive species such as smooth brome that are less nutritious to elk than native grass species (Constan 1972; Hobbs, and Huenneke 1992) and degrade the prairie by excluding native species, reducing biodiversity and carrying capacity for wild ungulates (Alberta Fish and Game 2016). However, these predator-driven (top-down) and resources-driven (bottom-up) relationships are complex (Strong 1992; Power et al. 1992; Kay 1998; Eisenberg 2017).

Hypotheses, Research Design, and Analysis Given the Kenow wildfire, in 2019 we will be revising our research questions and hypotheses to add questions about the interaction between wildfire and prescribed burn treatments. In 2018, we worked from the following research questions and hypotheses, which evolved from the questions and hypotheses we began to ask in 2008 when we initiated our study of the Y-Camp prescribed burn:

Q1 How has the extent of aspen cover changed?

Q2 How have fire and herbivory by elk changed aspen demography (e.g., the ability of aspen to grow into the forest canopy)?

Q3 How does the presence of an apex predator (the wolf) influence elk consumption of aspen?

Q4 How do repeat fires affect extent of canopy cover and aspen demography in an elk-wolf system?

Global Hypothesis: Post-fire aspen sprout persistence and recruitment vary based on fire severity and elk herbivory. Elk herbivory varies spatially due to wolf presence via fear of predation, a behaviorally- mediated trophic cascade (BMTC).

Research Hypotheses:

H1 Aspen sprouting is positively related to fire severity.

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H0 Aspen sprouting is not positively related to fire severity.

H2 Aspen recruitment is affected by elk browsing.

H0 Aspen recruitment is not affected by elk browsing.

H3 Elk spend more time in sites of highest fire severity, which have high aspen growth and food resources. H0 Elk do not spend more time in sites of highest fire severity, which have high aspen growth and food resources.

H4 Repeat fires reduce aspen sprouting and reduce aspen recruitment. H0 Repeat fires do not reduce aspen sprouting and do not reduce aspen recruitment.

For analysis purposes, we created research questions based on the above research hypotheses. Below are these questions and the hypotheses we used to test them for the 2008-2014 Y-Camp time series of data, expressed as statistical multivariate models.

QA What factors influence elk browse on aspen? Browse = f(distance to stand edge, fire severity, site index*, shrub %, sprouts per ha) Elk Density = f(distance to stand edge, fire severity, site index, shrub %, sprouts per ha)

QB What factors influence aspen recruitment? Aspen recruitment = f(browse, fire severity, site index, sprouts per ha, distance to stand edge) Aspen recruitment = f(elk density, fire severity, site index, sprouts per ha, distance to stand edge) *A measure of the productivity of a site

For the Y-Camp re-burn, we will replicate the above analyses. For the Eskerine Complex site, we will refine our hypotheses to incorporate additional factors of stand size and heat load (a measure of local solar exposure and aridity driven by slope and aspect) (McCune and Keon 2002). These factors did not exist in the Y-Camp prescribed burn site, because it is an aspen parkland (vs. discrete aspen stands). Furthermore, the slope we detected in the Y-Camp was insufficient to support creating a heat-load index. However, in the Eskerine Complex, we may find that limited range of aspect may influence our ability to add heat load as an index. We are achieving our overarching goal, to learn more about trophic relationships and the degree to which fire and wolves can help us create more resilient grassland and aspen forest communities that are closer to their historic condition, via the following objectives: 1. Test the effects of prescribed burn on aspen regeneration and recruitment; 2. Measure elk presence and feeding behavior in WLNP; 3. Investigate the effects of elk herbivory on post-fire aspen regeneration and recruitment; and 4. Examine the potential for wolves to influence elk feeding choices.

Geospatial Questions and Methods In 2016, we began geospatial sampling, to help WLNP managers determine whether prescribed burning meets the following management objectives: • Decrease the size of aspen stands; • Increase grassland area (reduce woody species including aspen and shrubs); and

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• Decrease aspen recruitment. Accordingly, we added the following research questions:

Q5 Do distance to stand edge, aspen regeneration density, and the amount of standing (dead or alive) aspen or downed woody debris influence elk browse?

Q6 Does prescribed burn impact above-ground biomass significantly?

In 2016, we collected our first geospatial dataset with mapping-grade Trimble Geo7X global navigation satellite system (GNSS) receivers to help answer these above questions. This sub-foot precision instrument is capable of receiving US and foreign GNSS satellite signals with integrated technology for improved accuracy under forest canopy. With it we mapped area (polygon) perimeters of mature aspen stands, aspen regeneration, and associated shrubs on the perimeter of stands and extending outward (called “shrub avulsion”). In 2017 we collected unmanned Aerial System (e.g., UAS or drone) data for Chris Anderson’s MSc research on our project.

Data Collected: During the 2018 field season, we made strong progress on our stated objectives. Prior to the Kenow wildfire, we conducted aspen and grass plot surveys in our two control sites: Crooked Creek and Beebe Flats, on the Blood Timber Limit, using the same methods we have been employing in surveying aspen and grasses in our Y-Camp and Eskerine prescribed burn study sites. These methods are described in detail, with definitions of each variable, in Eisenberg, Hibbs, and Donato 2014 and in Eisenberg 2012. We collected pellet transect data as an index of elk presence within Y-Camp aspen parkands and on the Y-Camp prairie. We measured the aspen parkland edge in the Y-Camp prescribed burn, and stand edge in the Eskerine Complex, and mapped the Eskerine aspen stands with Trimble GEO 7X units. This complemented the 2017 UAS data. To examine predator impacts on prey, each week we conducted noninvasive surveys of wolf activity, utilizing traditional wildlife tracking methods. Noninvasive methods are increasingly used tools to survey carnivore communities (Gomper et al. 2006; Heinemeyer 2008; McComb et al. 2010). We based our surveys on sites identified by WLNP ecologists as hotspots of wolf activity using camera-trap data and park staff observations. Camera traps are widely used to identify and quantify activity patterns (Stolle et al. 2015) and have been employed in WLNP for over a decade. Upon completion of field data collection, we built several databases, entered the data, conducted preliminary statistical analyses, and graphed our findings.

Overview: Our 2018 data primarily focused on 1) wildlife use of the Timber Limit; and 2) impacts of the Kenow wildfire. On the Timber Limit, we found a mean elk density of 2.55 ±0.17 (95% CI) piles of pellets per 100 m2 plot, and a mean deer density of 4.16 ±0.22 (95% CI) piles of pellets per 100 m2 plot. This indicates that the primary herbivore on the Timber limit, by number, is deer. However, given that elk are ~4.5 times greater in biomass than deer, elk are the dominant herbivore on the Timber Limit by biomass. This differs from WLNP, where the primary herbivore is elk, by both biomass and number. In WLNP, per differenced Normalized Burn Ratio (dNBR) analysis, the Kenow wildfire burned with 75.4% extreme, 12.6% high, 6.4% moderate, and 5.6% low fire severities (Fig. 1). In our study area we found a Fire Severity Index (on a scale of 0 – 4) of 3.91 ±0.03, or extreme. 97% of Eskerine plots burned with extreme severity. This matched the WLNP dNBR for the Eskerine Complex. Our rebar

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transect markers indicate that the Kenow wildfire and post-fire wind removed up to 0.7 m of topsoil (although soil removal patterns were very patchy), revealing bison bones and artifacts such as arrowheads. In Eskerine Complex prairie grass plots, proportion of grass cover dropped from 75.56% ± 0.88 (95% CI) in 2016 to 23.89% ±1.14 in 2018 (t=-35.42, 338 df, p=<0.00001)—a shift in dominant cover type (Fig. 5). Post-2014 and 2017 prescribed burns, we detected no mineral soil in prairie grass plots. In 2018, proportion of mineral soil in prairie grass plots was 50.84% ±1.42, and became the dominant cover type (Fig. 5). Between 2016 and 2018, proportion of shrub cover was unchanged, due to re-sprouting.

Figure 5. Change in cover proportion by type and site on the Eskerine Complex prairie in WLNP. Abbreviations: E = Eskerine prairie plots; SBP = Summer Bison Paddock prairie plots; WBP = Winter Bison Paddock prairie plots; 18 = the year 2018; 16 = the year 2016. These sites are all located within the Eskerine Complex prairie.

Dominant native and non-native grasses in 2016 and 2018 were the same, except for sheep fescue (F. ovina). Frequency of native-species detection (presence in plot) by year was similar, except for Columbia needlegrass (increased from 30 to 149 detections), and Parry’s oatgrass (decrease from 164 to 111 detections). Three of four non-native species decreased: Timothy grass from 25 to 18 detections; smooth brome from 15 to 4 detections; and sheep fescue from 9 to 0 detections. Reedgrass (Calamagrostis spp) and lamb’s quarters (Chenopodium album), previously seldom detected here, became common in aspen stands, seed-sprouting from mineral soil.

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Proportion of native grass cover was equivalent between 2016 and 2018 (Fig. 4). In 2016, Eskerine prairie plots had a higher proportion of native grass cover, 95.56% ±1.12, than Summer Bison Paddock (SBP) prairie plots, 83.76% ±3.59 (t=3.63, 118 df, p=0.0004), and Winter Bison Paddock (WBP) prairie plots, 85.92% ±3.70 (t=2.87, 119 df, p=0.005), attributable to 1960s-1970s inadvertent introduction of non-native agronomic species. In 2018, Eskerine prairie plots had a higher proportion of native grass, 90.08% ±2.58, than SBP prairie plots, 79.44% ±4.51 (t=2.19, 119 df, p=0.03). Between 2016 and 2018, mean number of native-grass species decreased significantly as follows: Eskerine prairie 5.28 ±0.16 to 4.51 ±0.13 (t=-3.77, 140 df, p=0.002); SBP prairie 6.02 ±0.25 to 5.20 ±0.25 (t=-2.38, 98 df, p=0.02); WBP prairie 6.16 ±0.18 to 4.54 ±0.28 (t=-4.92, 98 df, p=<0.00001). In 2016, the mean number of native grass species was lower in the Eskerine prairie than the SBP prairie (t=-2.64, 119 df, p=0.009) and WBP prairie (t=-3.57, 119 df, p=0.0005). In 2018, the Eskerine prairie had fewer native-grass species than the SBP prairie (t=-2.71, 119 df, p=0.01), but no difference existed between Eskerine and WBP prairie plots. When we evaluated the influence of the number of prescribed burns (0-3 prescribed burns) on grass-plot ecological response to prescribed burning, we found the proportion of native grass higher in plots that had 2 prescribed burns, versus 0 prescribed burns (t=-2,19, 70 df, p=0.03). This suggests that prescribed burning benefits native grasses. From January through April 2017, we sampled elk and bison fecal matter monthly in elk winter range. Samples were collected in the field, and were all freshly deposited specimens. We froze the specimens and sent them to the microhistology lab at the University of Washington for analysis. The specimens were pooled by month of sampling for analysis. The lab analysis showed that elk total diet during the months of January through April consisted of between 3.9 to 9.3% woody material (e.g., shrubs and trees). Elk only consumed aspen in February 2017 (1.4% of their diet) and April 2017 (0.5% of their diet). Bison consumed 0% aspen, and 0% woody species of any kind, during all months sampled. Elk primarily ate grass (81.8% - 87.7% of their diet), as did bison (85.3% - 93.5% of their diet). Rough fescue (F. campestris) was the primary grass species consumed by elk (21.8% - 28.6% of their diet) and bison (21.5% - 34.2% of their diet). The above findings indicate high similarity of elk and bison diets in this ecosystem, and also the conservation importance of rough fescue, given that it is such a high component of elk and bison diets. In mapped aspen stands, between 2016 and 2018 total canopy area decreased 98.09% from 57.60 ha to 1.10 ha; regeneration area increased 123.11% from 38.22 ha to 88.27 ha; shrub avulsion area increased 89.28% from 8.74 ha to 16.55 ha; and total aspen area decreased 9.8%, from 95.82 ha to 86.37 ha, but these changes were insignificant. The only significant change between 2016 and 2018 was canopy area per stand, which decreased from 1.92 ± 0.88 ha to 0.03 ± 0.02 ha (t= 2.13, 58 df, p=0.037) (Fig. 6).

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Figure 6. Change in Aspen Stand Structure. Map credit: Christopher Anderson and Curtis B. Edson.

Eskerine Complex aspen sprout density was 621.26 ±586.94 ha in 2008; 16,120.07 ±1057.55 ha in 2016, and 32,324.48 ±2755.87 ha in 2018. The highest density measured in 2018 was 481,443.88 sprouts ha (Fig. 7). Difference in density was significant between 2008 and 2015 (t=5.08, 735 df, p=<0.00001); 2015 and 2018 (t=-3.92, 783 df, p=<0.00001); and 2008 and 2018 (t=6.26, 734 df, p=<0.00001). For all years, aspen sprout density was patchy, with no evidence of sprouting from seed. Figure 7 below depicts the plot where we found a sprout density of 481,443.88 sprouts ha. This plot burned with extreme severity. This vigorous sprouting response demonstrates the resilience of aspen communities to prescribed burns and extreme-severity wildfire. This sprout density is one of the highest sprout densities reported for aspen in North America.

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Figure 7. Plot with highest sprout density, WLNP • Photo by Cristina Eisenberg

Aldo Leopold famously wrote, “The last word in ignorance is the man who says of an animal or plant, ‘What good is it?’ If the land mechanism as a whole is good, then every part is good, whether we understand it or not. If the biota, in the course of eons, has built something we like but do not understand, then who but a fool would discard seemingly useless parts? To keep every cog and wheel is the first precaution of intelligent tinkering” (Leopold, 1986, 190). This statement is critical to conserve ecosystems, and manage them for restoration and resilience. Bison were long part of WLNP, and humans hunted them. An entry from Peter Fiddler’s journal (Dec. 28, 1792) describes a hunt in Shepard Creek (near what is today WLNP), while a grass fire burned nearby (Haig 1934). By 1880, bison had been mostly extirpated from North America (Flores, 1991). Today the Buffalo Treaty, Iinni Initiative, and reintroduction efforts (Banff National Park 2017, pending Badger-Two Medicine, Montana, Blood Band Ranch, AB) are advancing bison recovery (Little Bear, 2014).

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However, Aldo Leopold’s statement did not acknowledge relationships inherent in social- ecological systems such as the Eskerine Complex. WLNP contains many archeological sites, including buffalo-drivelines and pounds—features spatially related to prairie fire, used for communal hunting, including a buffalo jump on the SW Eskerine Complex edge (Langemann, and Perry 1985; Reeves 2007). Re-establishing historical social dynamics here by relying on TEK primarily transferred via oral histories rather than practice, due to colonial removal of Indigenous people, may be more challenging than restoring this system’s physical components. Yet, relationships among the Kainai, landscapes such as the Eskerine Complex, and the broader Crown of the Continent Ecosystem arguably retain more TEK than other systems with similar post-colonization social-ecological change. How can TEK inform our understanding of wildfire as this site responds to potentially unprecedented disturbances? Based on the many bones and artifacts the Kenow wildfire revealed, bison and human hunters were long part of this system (Hills et al. 1985; Roos et al. 2018). These relationships had trophic impacts on human wellbeing and survival then, as now. The important role of people tending ecosystems with frequent fire has been recognized (Reyes-Garcia et al. 2018). These TEK land- use patterns can be replicated today to restore ecological processes with which flora and fauna co- evolved. This seems especially important in the Eskerine, where without anthropogenic burning and bison grazing, aspen might take over (Campbell et al. 1994). The same stewardship principles might apply to other frequent-fire ecosystems globally. The long-term perspectives afforded by this research project, in addition to the unexpected impacts of the Kenow wildfire, illustrate an important concept in disturbance ecology. Specifically, fires in the WLNP foothills-parkland ecoregion are not individual events easily bounded in space and time; rather, multiple fires overlap, and their effects represent the synergy among fire frequency and magnitude, ecological productivity, and adaptive traits that confer resilience to disturbance. Conceptualizing disturbance as a synergistic process builds on the classic definition of disturbance as a relatively discrete event (Pickett, and White 1985). Because fire promotes persistence of fire-adapted species at the expense of others, recurring fire derived from human ignitions aims to push the system toward a desired condition that benefits both social and ecological components. Temporally dispersed events, such as 1910 and 2017 wildfires, are too infrequent to induce a process-based response indicative of historical dynamics, highlighting the importance of continuing prescribed burn programs. Prescribed burn objectives (diminishing area of aspen cover on the prairie) were not fully realized after 3 prescribed burns over 11 years, likely because this system had significantly deviated from historical conditions and developed a contemporary ecological inertia to overcome. Understanding fire as “process” rather than “event” demonstrates the importance of managing near-term expectations as we bring ecosystems slowly back into alignment with historical relationships that persisted longer than the 100+ years of fire exclusion here. A tangible benefit of WNP prescribed burns was that they did not increase the proportion of non-native grass, which decreased post-Kenow wildfire. Given its extreme behavior, can the Kenow wildfire be described as “catastrophic?” Not from a plant-community perspective (Falk 2017). Our data illustrate this plant community’s strong resiliency to extreme wildfire (Fig. 7). We do not consider these responses a state shift (transition from one ecological state to another; Stringham et al. 2003). This landscape’s fundamental characteristics remained unchanged, indicating ecological stability (Suding et al. 2004). This suggests that because

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patchy, moderate fires are part of the foothills-parkland historical fire regime (Roos et al. 2018), prescribed burns can help maintain native grass, particularly in landscapes prone to extreme fire. The 2017 Prescribed burn and the Kenow wildfire differed in severity and scale. The Kenow wildfire significantly shifted ecosystem structure and grass diversity, while prescribed burns removed little aspen canopy and exposed no mineral soil. Prescribed burns occurred in late spring, with little fuel available, while the Kenow wildfire occurred in late summer, with abundant fuel—amplifying this difference (Knapp et al. 2009). As in other climate-limited fire regimes, prescribed burn treatments did not mitigate the severity of the Kenow wildfire (Stephens et al. 2009).

Management and Ecological Restoration Recommendations We suggest several widely applicable management principles that emerged from the ashes of the Kenow wildfire, to help improve ecological and cultural resiliency to extreme fire events:

1. Do not assume that extreme disturbance is an ecological catastrophe.

2. Plan for future change by restoring historical ecological processes when possible.

3. Ecological plant legacies can help restore degraded systems.

4. Collaborate across disciplines and cultures, including TEK, to restore degraded systems.

Applying these principles to adaptive management and ecological restoration and looking more deeply at the earth’s capacity to heal after large, extreme fires will help societies realize a more ecologically resilient future.

Impacts 1. INCREASING SCIENTIFIC KNOWLEDGE

MOS 1.1 Total Volunteer Research Hours: Each volunteer contributed 50 hours per week to data collection. Volunteers with us for 10-day teams contributed 80 hours of time to data collection.

MoS 1.2 Peer Reviewed Publications: In 2018/2019, the Kainai Community Conservation Fellows Program generated and inspired publications in a spectrum of genres, from peer-reviewed scientific journal articles to literary journal articles and book chapters. Peer-reviewed publications (listed below) serve to do outreach about this program and advance public policy that can improve bison and wolf conservation and acceptance of fire.

Peer-reviewed journal articles:

Published: McDonald, T., G. Gann, J. Jonson, J. Aronson, J. G. Hallet, C. R. Nelson, B. Walder, C. Echeverria, F. Hua, J. Liu M. Guariguata, C. Eisenberg, and K. W. Dixon. 2019. International Principles and Standards for the Practice of Ecological Restoration. Restoration Ecology.

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Eisenberg, C. 2019. “Conclusion: Cultural Keystone Species and Traditional Ecological Knowledge on the Northern Plains and Beyond,” In Grace Morgan, Beaver, Bison, Horse: An Indigenous Ecology of the Prairies (Regina, Sask: University of Regina Press). [peer-reviewed book chapter]

McDonald, T., J. Aronson, G. Gann, and C. Eisenberg. 2019. The SER Standards, cultural ecosystems, and the nature-culture nexus—a reply to Evans & Davis. Restoration Ecology.

Eisenberg, C. 2018. Revelations of the Kenow wildfire. About Place 7.1

Eisenberg, C. 2018. The eco-cultural roots of environmental and ecological literacy. Ecology 99(11).

In Review: Eisenberg, C., C. Anderson, A. Collingwood, C. Dunn, G. Meigs, R. Sissons, D. E. Hibbs, L. Little Bear, C. Edson, and B. Johnston. 2019. Effects of Extreme Wildfire, Prescribed Burns, and Indigenous Burning on Ecosystem Structure and Diversity. 2019. Frontiers in Ecology and Evolution [in review]

In Preparation: Aronson, J., C. Eisenberg, and D. Falk. 2019. Finding common ground in a rapidly changing world. [in preparation] Jubilee article. Restoration Ecology [invited paper].

MoS 1.3 Grey Literature and Other Dissemination of Your Results: Gray literature: The PI and co-PIs for this project produced four Parks Canada internal reports and a report for the Kainai First Nation, as follows: final report of the first time-series of sampling of the Y- Camp prescribed burn (Elk, Fire and Wolf Ecology in Aspen and Grassland Communities in Waterton Lakes National Park of Canada, 2010-2013 Y-Camp Prescribed Burn Final Investigators’ Report, Eisenberg, Hibbs, and Donato 2014), and reports of 2015, 2016, and 2017 work (Elk, Fire and Wolf Ecology in Aspen and Grassland Communities in Waterton Lakes National Park of Canada: Red Rock Complex Prescribed Burn and Y-Camp Prescribed Burn Investigators’ Annual Progress Report 2015; Eisenberg, Hibbs, and Donato 2015; Elk, Fire and Wolf Ecology in Aspen and Grassland Communities in Waterton Lakes National Park of Canada: Red Rock Complex Prescribed Burn and Y-Camp Prescribed Burn Investigators’ Annual Progress Report 2016; Eisenberg, Hibbs, and Edson; Elk, Fire and Wolf Ecology in Aspen and Grassland Communities in Waterton Lakes National Park of Canada: Red Rock Complex Prescribed Burn and Y-Camp Prescribed Burn Investigators’ Annual Progress Report 2017; Eisenberg, Hibbs, and Edson; and Blackfoot Science: Using Traditional Ecological Knowledge and Community Ecology to Assess Blood Timber Limit Health and Sustainability, Investigators’ Annual Progress Report 2017, Eisenberg, Hibbs, and Edson).

For a general audience, the PI documents work on this project on her website: www.cristinaeisenberg.com. She has written about this project in her books, The Wolf’s Tooth: Keystone Species, Trophic Cascades, and Biodiversity (Washington, DC: Island Press, 2010) and The Carnivore Way: Coexisting with and Conserving America’s Predators (Washington, DC: Island Press, 2014). She is currently at work on a book about climate change, Taking the Heat: Climate Change, Extinction, and Wildlife in which some of our findings in WLNP will be discussed. In 2017 the PI secured a contract from

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Oregon State University to write a book about bison restoration, entitled Bison Homecoming: Repatriating an Icon, Rewilding the American West, with a foreword by Dr. Leroy Little Bear. This book will include substantial material about our project. The PI will be donating all of her royalties from this book to the Kainai First Nation.

Lectures and Presentations: In the past year, we gave the following 12 presentations (Table 2) about our project:

Table 2. Presentations given about our project in 2018. Event Presenter Date Location Presentation type

Environmental Sciences Cristina Eisenberg 5/15/18 Corvallis, Oregon plenary address Symposium, Oregon State University

Buffalo Treaty Celebration Cristina Eisenberg 5/22/18 Polson, MT invited lecture

Association for Fire Ecology Cristina Eisenberg 5/24/18 Missoula, MT oral presentation Conference

KEPA Summit David Hibbs 6/6/18 WLNP invited presentation

KEPA Summit Elliot Fox 6/6/18 WLNP invited lecture

KEPA Summit Chris Anderson and Elliot Fox 6/6/18 WLNP and Timber Limit field trip

Place-Based Citizen Science Elliot Fox 9/9/18 Calgary, Alberta invited lecturer

Society for Ecological Cristina Eisenberg 9/10/18 Reykjavik, Iceland invited panelist Restoration Europe

Society for Ecological Cristina Eisenberg 10/16/18 Spokane, WA oral presentation Restoration North America

Fort Peck Reservation Cristina Eisenberg 11/8/18 Wolf Point, MT Invited lecture

Cochrane Ecology Institute Cristina Eisenberg 11/10/18 Cochrane, AB keynote speech

Alaska Wildlife Alliance, Cristina Eisenberg 12/5/18 Jueau, AK Invited lecture University of Alaska

2. DEVELOPING ENVIRONMENTAL LEADERS

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MoS 2.1 Education: In 2016 we launched a pilot program to teach Kainai First Nation community members field ecology and ecological restoration methods within a TEK context. Our objective was to help band members develop natural resources job skills. To that end, all Kainai First Nation Community Conservation Fellows assisted in field data collection, working directly with us and our field crew. In 2016 we fielded 4 Kainai Community Fellows; in 2017 we successfully fielded 21 Kainai Community Fellows; in 2018 we successfully fielded 34 Kainai Community Fellows, and in 2019 we have funding to field 52 Kainai Community Fellows. This partnership, which is funded by the Kainai Board of Education, the AGL Foundation, the Davin Family Trust, and the Eisenberg Family Trust, is significantly contributing to development of environmental leaders. For example, Diandra Bruised Head, a Kainai Community Fellow in 2016, completed her BSc subsequent to joining us in the field, after having been out of school for some years. After completion of her degree in May 2017, we hired her as a full-time field technician (Fig. 8). She worked for us through September 2017, and then secured a highly competitive position as the climate change intern for Blood Tribal Land Management, a 2-year, federally funded appointment. In 2018 we employed Elliot Fox, Justin Bruised Head, Alex Shade, and Dustin Fox as field technicians. Dustin Fox has since applied to law school and Justin Bruised Head is in the process of completing his BSc in spring 2019.

Figure 8. Former Kainai Community Fellow Diandra Bruised Head teaches 2017 Kainai Community Fellow Monroe Fox grass identification on the Timber Limit • Photo by Cristina Eisenberg

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3. PARTNERSHIPS

MoS 3.1 Organizations Actively Engaged: In addition to Earthwatch Institute, the partners listed below (Table 3) have generously supported our project over the years. Additionally, we consider the ~80 or so technicians, interns, and volunteers who joined our project before it was an Earthwatch project, between 2006-2014, as project partners. This work would not have been possible without their contributions and those of 188 Earthwatch participants (2015-2018), which include community fellows.

Table 3. Project partners, funding sources, and years of association.

Partner Support Type Years of Association Waterton Lakes National Park Funding, housing, data, logistics, permits 2006-present Earthwatch Institute Funding 2014-present Kainai First Nation Funding, data, permits 2015-present Davin Family Trust Funding 2017-present Oregon State University Funding, logistical and technical support 2006-present Whitefish Community Foundation Funding 2015-present Society for Ecological Restoration, IUCN Funding, collaboration 2017-present Michigan Technological University Funding, logistical and technical support; 2016-2019 Funding for graduate students on this project Shell Canada Funding 2006-2012 Boone & Crocket Club Funding 2008-2012 Alberta Fish and Wildlife Data, logistical support, permits 2006-present Glacier National Park Data, logistical support, permits 2006-2012 US Fish Wildlife & Parks Data, logistical support, permits 2007-2012 Montana Fish, Wildlife & Parks Data, logistical support, permits 2007-2013 US Forest Service Data, logistical support, permits 2006-present Smithsonian Institution Collaboration, academic support 2012-present Wildlife Conservation Society Collaboration, data exchange 2016-present Yellowstone to Yukon Collaboration, data exchange 2016-present

4. CONTRIBUTIONS TO POLICIES OR MANAGEMENT PLANS

MoS 4.1 Informing Policies or Management Plans: We are directly contributing to public policy and land management with our research as follows:

Community-Level Impacts: Our project’s findings are influencing local landowners and the Kainai First Nation (who ranch and harvest timber) in their natural resources practices. By having community members and tribal elders join us in the field, we are increasing awareness of and encouraging coexistence with both fire and wolves in mixed-use landscapes. This is a reciprocal relationship, as landowner insights are contributing to our understanding of the system we are studying.

Local, Regional, and National Level Contributions and Impacts: Our work directly supports the WLNP Management Plan. Specifically, our research helps WLNP administrators “restore and monitor natural heritage, while showcasing innovative activities.” It further provides an opportunity for WLNP to engage a variety of stakeholders to collaborate in implementing solutions to restore the prairie and increase the ecological resiliency and biodiversity of this landscape. Given the cultural significance to the Kainai of

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the prairie, fire, wolves, and bison, our work helps increase the profile of this historic International Biosphere Reserve site and improves the condition of its cultural resources (WLNP 2010). We annually report our findings to WLNP managers and share our data with them. We also make our findings available to the Alberta Division of Fish and Wildlife. Managers incorporate our findings into local (WLNP) management plans for the coming year. They give us feedback on our methods and sampling strategy in order for our work to best support theirs. We consider this an ideal partnership between management and science. Furthermore, our science benefits significantly from feedback and advice from managers.

International Impacts: Our study has bearing on the Crown of the Continent Ecosystem, which straddles the US/Canada border. We collaborate with international conservation organizations who work in this ecosystem, such as the Wildlife Conservation Society, Yellowstone to Yukon, and others, by sharing data. Our findings will have impact on land management in the US as well, within Glacier National Park (GNP) and northern Rocky Mountain US Forest Service (USFS) management units. Our work to support restoration of natural processes is aligned with the Parks Canada Charter (Parks Canada 2011). The PI makes presentations in the US Congress (House Natural Resources Committee) and works with government agency leaders, who occasionally visit us afield. Our work in WLNP also has bearing on First Nations. Kainai tribal leaders, including elders, have joined us afield on numerous occasions to collaborate with and mentor us and share their TEK. WLNP is a site of high international conservation valence. Forming the centerpiece of the much larger transboundary Crown of the Continent Ecosystem, WLNP’s unique physiographic mountain setting, which includes the mountain-prairie interface, has resulted in the evolution of plant communities and ecological complexes that occur nowhere else in the world. In 1932 UNESCO, a specialized branch of the United Nations, combined WLNP (Alberta, Canada) with GNP (Montana, US) to form the world's first International Peace Park. This peace park is also a UNESCO World Heritage site, defined as a landmark site with cultural and/or natural significance so exceptional that it is of common importance for present and future generations of all humanity (UNESCO 2015). It also has UNESCO International Biosphere Reserve designation. UNESCO selected biosphere reserves to foster integration of people and nature for sustainable development, knowledge sharing, poverty reduction, human well- being improvements, respect for cultural values, and to improve society’s ability to cope with global change. As such, any research taking place in WLNP has international impact. In recognition of our long- term ecological research that is helping WLNP and the Kainai First Nation restore the fescue prairie, in November 2017 the Ecohealth Global Network, a Society for Ecological Restoration (SER) and IUCN initiative in fulfillment of the United Nation’s Bonn challenge, selected ours as one of 7 global model restoration ecology projects. Given the high proportion of Earthwatch participants from other nations besides the US and Canada, our project has international impact in terms of raising awareness of conservation and ecological challenges, and also of how stakeholder partnerships can advance conservation globally. Many of the things we are learning (e.g., the use of natural processes such as fire, herbivory, and predation to restore resilience to ecosystems) are relevant in other nations (e.g., using fire in Australia to restore grasslands and reduce shrub and tree invasion onto grasslands). To date Earthwatch

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participants from the following nations have joined us: Australia, Canada, England, France, Germany, Italy, Japan, New Zealand, Scotland, Spain, and the US.

MoS 4.2 Actions or activities that enhance natural and/or social capital: By restoring fire and documenting its impacts in this system, we are gathering data that will be used by the Kainai First Nation and Parks Canada to restore fire regimes on other lands of high historical and cultural value, such as the Blood Timber Limit. Our research is directly and tangibly increasing cultural heritage. We are including Kainai Community Fellows in our field research (4 in 2016, 21 in 2017, and 34 in 2018, Fig. 9-a), and are giving presentations and co-leading field trips for the Kainai First Nation, to teach them how to apply these ecological restoration methods on their lands and, as importantly, for us to learn from them. Since 2016, we have partnered with the Kainai First Nation in our research both inside and outside the park, to incorporate TEK. Kainai elder Mike Bruised Head joined the first Kainai Community Fellows in May and took us to archeological sites in WLNP to discuss Blackfoot heritage, Blackfoot science, and how TEK can inform current land management practices and restoration efforts. Midway through the 10-day Kainai High School Community Fellows’ residency, the tribe and Earthwatch co- hosted a sweat lodge, pipe ceremony, and traditional bison feast, to commemorate our collaboration (Fig. 9-b). WLNP manager Dennis Madsen participated, as did former WLNP ecologist Rob Watt, and tribal elders Peter Weasel Moccasin and Alvine Mountain Horse. Such collaboration has been recommended by experts on adaptive management and TEK (Berkes 2000; Dunn 2017). Specifically, in June 2017 we began data collection on the Blood Timber Limit, with unanimous approval of this work from the Tribal Council and Kainai Ecosystem Protection Association (KEPA). We continued this work in 2018. As a key part of this collaboration, in June 2017 and 2018, we co-led tribal field trips with Kansie Fox and elder Mike Bruised Head, as part of the KEPA Summit. In WLNP we demonstrated aspen survey methods and then visited the Belly River, to mark the first aspen transect on the Blood Timber Limit. Kainai Community Fellows fielded in May, July, and September. These Fellows presented about their experiences on this project at the KEPA Summit. After the Summit and throughout the summer, Kainai school groups, from primary school students to high school students, joined us afield in WLNP and the Blood Timber Limit, as did tribal elders. Community Fellows consisted of tribal members who were teachers, promising youth interested in natural resources ecology, and impactful individuals. Our work with the Kainai First Nation will take place over four field seasons (2017-2020). In recognition of this partnership, in May 2017 the PI was given a Blackfoot name, Mistaksii Kaka'too'sakii (Mountain Star Woman), in a naming ceremony (Fig. 9-c). Our project has significant local impacts on social capital. Kainai First Nation Community Conservation Fellows receive natural resources ecological sampling and ecological restoration training. In 2017 and 2018, each year we employed four Kainai field technicians. In 2019 and beyond, we will continue to provide field technician jobs for the Kainai First Nation. We feed our field crew grass-fed bison meat from the Kainai Reserve and partner with local ranchers, such as the Russell family, to educate and create awareness of the ecology of this system, improve coexistence with carnivores, and do outreach for prescribed burning. Beyond the local, we give many talks nationally and internationally,

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using this project to demonstrate exemplary federal, tribal, provincial, and local partnerships in ecological restoration and TEK.

a)

b) c) c)

Figure 9. Kainai First Nation collaboration, a) Kainai First Nation Community Conservation Fellows in the field in WLNP with EW volunteers; b) Kainai sweat lodge ceremony; c) Kainai naming ceremony.

5. ENHANCING NATURAL AND SOCIO-CULTURAL CAPITAL

MoS 5.1 Conservation of Taxa: In addition to elk, aspen, and wolves, we also are collecting data on grizzly bears, cougars, and wolverines (Gulo gulo), 54 shrub species, and 30 grass species (native and non-native, invasive grasses). Native grass species are of high conservation value, as collectively they form fescue grassland, one of

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the most endangered habitat types in North America (Achuff 2005; Lesica 2012). Non-native, invasive species are of crucial conservation concern, because they compete with and threaten native grasses. WLNP is using our data to help control non-native grass.

Grassland Restoration: We are working with WLNP to restore fescue grassland, a highly at-risk habitat type that provides valuable natural and cultural capital. By returning fire to this fire-adapted plant community, WLNP intends to increase the extent of and the biodiversity in this system, provided that invasive weeds are managed, as fire can also stimulate weed growth. WLNP fire treatments encompass a total of 3,500 ha of this habitat type. Our study, in which we are monitoring the effectiveness of these fires at restoring fescue grassland, provides useful information to managers. Ecological restoration without monitoring within an adaptive-management strategy often fails, as does restoration that does not fully consider the community-level implications of such work (Palmer, Ambrose, and Poss 1997; Young, Chase, and Huddelson 2001; Palmer et al. 2005). Our research is ensuring that WLNP ecological restoration efforts are maximally supported by science. We deliver our data, reports, and analyses annually to the park, which optimizes the flow of information per best adaptive-management practices.

Ecosystem Services: As observed elsewhere, these prescribed burns are increasing nutrient cycling and biodiversity (Berger et al. 2001; Bailey and Witham 2002; Fontaine and Kennedy 2012; North, Collins, and Stephens 2012). In this fire-adapted system, our data are linking fire to intensive sprouting and growth of aspen and native grass. These dynamics improve habitat for some wildlife species and taxa, including elk, bears, wolves, songbirds, and insects (Hobbs et al. 1981; Bailey and Witham 2002). In 2016 and 2017 we observed intensive bear use of burned portions of the Eskerine Complex. The bears were foraging on forbs and shrubs. For example, in July 2016, during a two-hour period, we encountered 8 bears at ranges from 50 m to 500 m. They appeared to have a good body condition (per body condition score metrics). Such foraging responses by bears and other wildlife have been observed commonly in early-seral communities created by disturbance such as fire (Swanson et al. 2010) and are linked to an increase in ecological resiliency (Hobbs and Huenneke 1992; Rogers, Eisenberg, and St. Clair 2013).

Are you enhancing, restoring, or maintaining populations of any species of conservation significance as part of your project? For each species include where possible:

1. Scientific and common names 2. Significance of the species 3. Impact on the species e.g. range increased, population size increased, improving population structure, maintaining/enhancing genetic diversity

All of the native grass species listed in Appendix A have high conservation value, as described above. Our work complements and contributes to WLNP’s fire policy, wildlife management policy, and invasive species control practices. Appendix B provides names of the shrubs we have identified and are

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studying, with notes on their nutritional value to hooved herbivores. Additionally, our data contributes to conservation of wolves and grizzly bears, which are highly sensitive species. Grizzly bears are listed as threatened in Alberta under the Species at Risk Act and as threatened in the US under the Endangered Species Act. While wolves are not a T&E species in Canada, they are considered of high conservation value due to their impact on whole food webs via trophic cascades. The prescribed burns we are studying directly affect conservation of endangered species of songbirds, such as Le Conte’s sparrow (Ammodramus leconteii), which is found in shortgrass prairie, and which we detected in our study site when doing songbird point counts in 2007 and 2008. Finally, our work has direct positive bearing on bison conservation.

MoS 5.2 Conservation of Habitats – Are you enhancing, restoring, or maintaining habitats as a result of your project? Yes, see below:

1. Habitat being restored: Fescue prairie, a highly at-risk habitat, provides valuable natural and cultural capital and which historically contained fire, wolves, and bison. Mixed-grass prairie, considered essential bison habitat, consists of native short grasses and tall grasses, and occurs on Kainai land (Fig. 10).

Figure 10. 2017 and 2018 Kainai Community Fellow Monroe Fox in a patch of intact mixed-grass prairie on Kainai land that is the site of a buffalo jump, and where a bison reintroduction is planned • Photo by Cristina Eisenberg

2. Treatment: Recovering wolves and returning fire to this fire-adapted system, which should increase the extent of and the biodiversity in this system, provided that invasive weeds are managed, as fire can also stimulate weed growth; gathering baseline data about fire, grass, and elk diet that can inform bison

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conservation. We are collaborating with the Iinnii Initiative to prepare for the bison reintroduction that is currently underway, which would significantly contribute to grassland restoration.

3. Area of habitat enhanced: A total of 5,500 hectares are part of this study.

4. Source of baseline data: Parks Canada has baseline information on the extent of the fescue prairie; the PIs have baseline information on the extent of aspen cover and on the integrity of shortgrass fescue prairie. Parks Canada and the PIs have baseline data on wildlife.

MoS 5.3 Conservation of Ecosystem Services: Are you enhancing, restoring, or maintaining ecosystem services as a result of your project? Please state the ecosystem service affected and the impact your project has had, with supporting numbers or data if possible. These prescribed burns are increasing the cycling of nutrients in this system and increasing biodiversity (e.g., grasses, shrubs, birds).

a)

Figure 11. a) Kainai High School Community Fellows and project staff at sweat lodge ceremony and pipe offering to honor fellows; b) Kainai technician teaching Kainai High School Community Fellows how to drum, while a dinner of stew made from the meat of a bison produced by tribal elder Dan Fox is being prepared on our project.b)

MoS 5.4 Conservation of Cultural Heritage: Are you enhancing, restoring, or maintaining intangible or tangible cultural heritage as a result of your project? Please state the heritage component affected and the impact your project has had, with supporting numbers or data if possible. Yes, by restoring fire and documenting its impacts in this system, we are gathering data that will be used by the Kainai First Nation and Parks Canada to restore fire regimes on other lands of historical and high cultural import, such as the Blood Timber Limit. Our research is directly and tangibly increasing cultural heritage, because we are examining the role of two species of high cultural and spiritual significance to the Kainai—bison and wolves. We are incorporating Kainai Community Fellows in our field research, and the PI is giving presentations and leading field trips for the Kainai First Nation, to teach them how to apply these ecological restoration methods on their lands. Considering the ecological role in this system of wolves and bison, species of high cultural and spiritual value to the Kainai, directly

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contributes to conserving cultural heritage. We also acknowledge and include Kainai ceremonies, which are part of TEK observations and cultural heritage (Fig. 11 a-b).

MoS 5.5 Impacting Local Livelihoods: Are you enhancing, restoring, or maintaining livelihoods in the local community as a result of your project? “Livelihood assets” include persons benefitting from economically applicable training, local employment for duration of project, and community assets such as clean water, access to resources, development of a community trust or setting up a museum, for example. In 2018, through the Kainai First Nation Community Fellows Program, we trained Kainai community members in ecological sampling and hired 4 Kainai field technicians. We began to develop a graduate fellowship to support a Kainai master’s student working on our project to do a master’s thesis focusing on TEK and ecological restoration. We locally purchase much of the food we use to feed Earthwatch participants, fellows, and staff, supporting the local economy. For example, annually we buy 200 pounds of locally produced grass-fed bison and elk and moose meat hunted by Kainai tribal members on tribal land.

Acknowledgements, Funding and Appendices Project funding: Earthwatch is working towards improving long-term sustainability of its projects and we see that sourcing funding from multiple sources is a key component towards this. Provide a brief statement relating to other funding sources (or funding sources you anticipate receiving from or plan to apply to) and the relative (or anticipated) contribution of Earthwatch support for project success. Table 3 (above) provides details of our research partnerships. We have received considerable funding from these sources over the years. We expect to continue to receive support from Parks Canada in the form of housing for our research team (PIs, technicians, and volunteers), funding for laboratory analyses, and logistical support. This support is secured in part through our Partnership Agreement with WLNP.

ACKNOWLEDGEMENTS In 2018 Earthwatch Institute, WLNP, Michigan Technological University, the Kainai First Nation, Whitefish Community Foundation, and Oregon State University provided financial and logistical support for our project. We thank Scott Murphy and Randall Schwanke for their expert fire management over the years. We gratefully acknowledge the support of WLNP managers and ecologists Dennis Madsen, Barb Johnston, Robert Sissons, Adam Collingwood, Kimberly Pearson, and Ryan Peruniak. Thanks to Rob Watt, Mike Bruised Head, Kansie Fox, Annette Bruised Head, Tanner Broadbent, Heather Pruiksma, Stan Rullman, Stephen Hart, and Steve Eisenberg for logistical help and help afield. Special thanks Elliot Fox, for providing exemplary leadership and guidance as our lead Kainai Technician from 2016-2018, and to our graduate student Chris Anderson and field technicians Justin Bruised Head, Alex Shade, Dustin Fox, Maire Kirkland, Hannah Kirkland, and Andrew Acosta, and intern Kristen Arakawa. We are deeply grateful to the Earthwatch citizen-science participants and Kainai First Nation Community Conservation

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Fellows who joined us for the 2018 field season. Heartfelt thanks to the Russell family for their friendship and support. Thanks to the Kainai First Nation for your wisdom, support, and collaboration.

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Appendix A

All grass species detected in WLNP and the Blood Timber Limit 2015 – 2018. Terms Used: Status: N = native; NN = non-native; Indic Spp. = key species for which the park encourages monitoring. P = focal prairie grass species (identified by Agriculture Alberta as native grass species that characterize fescue prairie); I = invasive weeds that are part of the park’s weed-control program.

Scientific Name Common Name Code Origin Grazing Response Forage Value Achnatherum nelsonii needlegrass, Columbia ACH NEL N (form. Stipa nelsonii) Achnatherum occidentale needlegrass, Western ACHN OCC N Increaser Good (form. Stipa occidentalis) Achnatherum richardsonii needlegrass, Richardson’s ACHN RIC N Decreaser Good (form. Stipa richardsonii) Agropyron repens quack grass AGRO REP NN, I Decreaser Good Agrostis scabra ticklegrass AGRO SCA N Invader Good Alopecurus aequalis foxtail, shortawn ALOP AEQ N Decreaser Good Alopecurus magellanicus foxtail, alpine meadow ALOP MAG N N/A N/A Bromus carinatus brome, mountain BROM CAR N Decreaser Good Bromus ciliates brome, fringed BROM CIL N Decreaser Good Bromus inermis brome, smooth BROM INE NN, I Invader Good Calamagrostis montanensis plains reedgrass CALA MON N Dactylis glomerata orchard grass DACT GLO NN Increaser Good Danthonia intermedia oatgrass, timber DANT INT N Increaser Good Danthonia parryi oatgrass, Parry’s DANT PAR N, P Increaser Good Elymus dasystachyum wheatgrass, northern ELYM DAS N Elymus glaucus Wildrye, blue ELYM GLA N Increaser Fair Elymus trachycaulus subsp. subsecundus wheatgrass, slender ELYM TRA N Decreaser Good (form. Agropyron trachycaulum) Festuca campestris fescue, foothills rough FEST CAM N, P Decreaser Good Festuca idahoensis fescue, Idaho FEST IDA N, P Increaser Good Festuca occidentalis fescue, Western FEST OCC N Increaser Good Festuca ovina fescue, sheep FEST OVI NN, I Increaser Good Hesperostipa spartea porcupinegrass HESP SPA N (form. Stipa spartea) Koeleria macrantha Junegrass KOEL MAC N Increaser Good Leymus triticoides wildrye, beardless LEYM TRI N N/A N/A (form. Elymus triticoides) Nassella virdula needlegrass, green NASS VIR N Decreaser Good (form. Stipa viridula) Psathyrostachys juncea wildrye, Russian PSAT JUN NN (form. Elymus juncea) Pascopyrum smithii wheatgrass, Western PASC SMI N Decreaser Good (form. Agropyron smithii) Phleum pretense Timothy grass PHLE PRA NN, I Invader Good Poa alpine bluegrass, alpine POA ALP N N/A N/A Poa pratensis bluegrass, Kentucky POA PRA NN, I Invader Good Poa secunda bluegrass, Sandberg POA SEC N Increaser Good (form. Poa ampla) Pseudoroegneria spicata wheatgrass, bluebunch PSEU SPI N, P Decreaser Good (form. Agropyron spicatum) Schendonorus pratensis fescue, meadow SCHE PRA NN, I Invader Good

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Appendix B

Shrubs detected by Eisenberg et al. in WLNP and on the Blood Timber Limit, 2015 – 2018.

Scientific name COMMON NAME Family CODE PALATABILITY Acer glabrum Rocky Mountain maple Aceraceae ACER GLA 1 Alnus incana mountain alder Betulaceae ALNU INC 1 Alnus viridis green alder Betulaceae ALNU VIR 3 Amelanchier alnifolia Saskatoon; serviceberry AMEL ALN 1 Artemisia tridentata big sagebrush Asteraceae ARTE TRI 3 Arctostaphylos uva-ursi kinnickinnick Ericaceae ARCT UVA 3 Betula glandulosa dwarf birch Betulaceae BETU GLA 2 Betula nana dwarf birch Betulaceae BETU NAN 2 Betula occidentalis water birch Betulaceae BETU OCC 2 Betula pumila dwarf birch Betulaceae BETU PUM 2 Ceanothus velutinus snowbrush Rhamnaceae CEAN VEL 2 Clematis occidentalis virgin’s bower Ranunculaceae CLEM OC Cornus sericea red-osier dogwood Cornaceae CORN SER 1 Crataegus douglasii black hawthorn Rosaceae CRAT DOU 3 Crataegus chrysocarpa round-leaved hawthorn Rosaceae CRAT CHR 3 Dasiphora fruticosa shrubby cinquefoil Rosaceae DASI FRU 3 Elaeagnus commutata wolf-willow Eleagnaceae ELEA COM 3 Juniperus communis ground-juniper Cupressaceae JUNI COM 3 Juniperus horizontalis creeping juniper Cupressaceae JUNI HOR 3 Lonicera ciliosa twinberry Caprifoliaceae LONI CIS 2 Lonicera dioica Western honeysuckle Caprifoliaceae LONI DIO 3 Lonicera involucrata bracted honeysuckle; black twinberry Caprifoliaceae LONI INV 1 Lonicera utahensis red twinberry Caprifoliaceae LONI UTA 2 Mahonia repens Oregon grape Berberidaceae MAHO REP 3 Menziesia ferruginea false azalea Ericaceae MENZ FER Pachistima myrsinites mountain lover Celastraceae PACH MYR Prunus pensylvanica pin cherry Rosaceae PRUN PEN 2 Prunus virginiana choke cherry Rosaceae PRUN VIR 2 Rosa acicularis prickly rose Rosaceae ROSA ACI 1 Rosa arkansana prairie rose Rosaceae ROSA ARK 1 Rosa woodsii wild rose Rosaceae ROSA WOO 1 Ribes hudsonianum northern black currant Grossulariaceae RIBE HUD 2 Ribes inerme whitestem gooseberry Grossulariaceae RIBE INE 3 Ribes lacustre bristly black gooseberry Grossulariaceae RIBE LAC 3 Ribes oxyacanthoides northern gooseberry Grossulariaceae RIBE OXY 3 Ribes viscosissimum sticky currant Grossulariaceae RIBE VIS 3 Rhamnus alnifolia alder-leaved buckthorn Rhamnaceae RHAM ALN 2 Rubus acaulis wild raspberry Rosaceae RUBU ACA 1 Rubus idaeus wild raspberry Rosaceae RUBU IDA 1 Rubus parviflorus thimbleberry Rosaceae RUBU PAR 2 Rubus pubescens dwarf red blackberry Rosaceae RUBU PUB Salix bebbiana beaked willow Salicaceae SALI BEB 1 Salix discolor pussy willow Salicaceae SALI DIS 1 Salix exigua sandbar willow Salicaceae SALI EXI 1 Salix planifolia flat leafed willow Salicaceae SALI PLA 1 Sambucus racemosa elderberry Caprifoliaceae SAMB RAC 1 Shepherdia canadensis Canada buffaloberry Eleagnaceae SHEP CAN 1 scopulina western mountain ash Rosaceae SORB SCO 3 Spiraea betulifolia birchleaf spirea Rosaceae SPIR BET 3 Symphoricarpos albus snowberry Caprifoliaceae SYMP ALB 1 Symphoricarpos occidentalis snowberry Caprifoliaceae SYMP OCC 1 Vaccinium membranaceum mountain huckleberry Ericaceae VACC MEM 2 Vaccinium scoparium huckleberry Ericaceae VACC SCO 2 Viburnum edule low-bush cranberry Caprifoliaceae VIBU EDU 2

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