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Polfus, J. L., M. Manseau, D. Simmons, M. Neyelle, W. Bayha, F. Andrew, L. Andrew, C. F. C. Klütsch, K. Rice, and P. Wilson. 2016.
Łeghágots'enetę (learning together): the importance of indigenous perspectives in the identification of biological variation. Ecology
and Society 21(2):18. http://dx.doi.org/10.5751/ES-08284-210218
Research
Łeghágots'enetę (learning together): the importance of indigenous
perspectives in the identification of biological variation
Jean L. Polfus 1, Micheline Manseau 1,2, Deborah Simmons 3,4, Michael Neyelle 3,5, Walter Bayha 6, Frederick Andrew 3, Leon Andrew 3,
Cornelya F. C. Klütsch 7, Keren Rice 8 and Paul Wilson 7
ABSTRACT. Using multiple knowledge sources to interpret patterns of biodiversity can generate the comprehensive species
characterizations that are required for effective conservation strategies. Caribou (Rangifer tarandus) display substantial intraspecific
variation across their distribution and in the Sahtú Region of the Northwest Territories, Canada, three caribou types, each with a
different conservation status, co-occur. Caribou are essential to the economies, culture, and livelihoods of northern indigenous peoples.
Indigenous communities across the north are insisting that caribou research be community-driven and collaborative. In response to
questions that arose through dialogue with five Sahtú Dene and Métis communities, we jointly developed a research approach to
understand caribou differentiation and population structure. Our goal was to examine caribou variation through analysis of population
genetics and an exploration of the relationships Dene and Métis people establish with animals within bioculturally diverse systems. To
cultivate a research environment that supported łeghágots'enetę “learning together” we collaborated with Ɂehdzo Got'ı̨nę (Renewable
Resources Councils), elders, and an advisory group. Dene knowledge and categorization systems include a comprehensive understanding
of the origin, behaviors, dynamic interactions, and spatial structure of caribou. Dene people classify tǫdzı “boreal woodland caribou”
based on unique behaviors, habitat preferences, and morphology that differ from ɂekwę́ “barren-ground” or shúhta ɂepę́ “mountain”
caribou. Similarly, genetic analysis of material (microsatellites and mitochondrial DNA) from caribou fecal pellets, collected in
collaboration with community members during the winter, provided additional evidence for population differentiation that
corresponded to the caribou types recognized by Dene people and produced insights into the evolutionary histories that contribute to
the various forms. We developed culturally respectful and relevant descriptions of caribou variation through partnerships that respect
the lives and experiences of people that depend on the land. By prioritizing mutual learning, researchers can broaden their understanding
of biodiversity and establish a common language for collaboration.
Key Words: aboriginal; biocultural diversity; biodiversity; caribou; collaborative research; ecology; First Nation; genetic variation;
indigenous communities; population genetics; population structure; Rangifer tarandus; resource management; social-ecological systems;
traditional knowledge
INTRODUCTION
Patterns of biological variation are a result of the replication of
DNA, the potential for DNA mutations and environmental
structure that prevents the complete overlap of groups of
organisms (Hey 2001). Significant scientific effort has been
allocated toward determining where, within the space of genetic
and environmental variation, units emerge that merit
identification (Padial et al. 2010). Although the articulation of
biological categories is a universal human predisposition (Berlin
1973, Atran 1990), the content of named categories reflects a
dynamic exchange between morphological, utilitarian, ecological,
and perceptual factors, all of which are adapted by different
cultures to a particular time and place (Nazarea 2006, Newmaster
et al. 2006). The species concept, which has undergone numerous
iterations and has been, and continues to be, actively debated by
systematists, taxonomists, biologists, and naturalists, illustrates
the complexity of assigning objects to categories (Hey 2001).
Recently, researchers have begun to acknowledge that many, if
not most, species do not have distinct, easy-to-recognize
boundaries (Hey 2006, Mallet 2008) and that species may be best
described as “poorly differentiated way-stations in a continuous
hierarchy of biodiversity” (Mallet 2005:229). However, in order
to identify groups that justify protection, such as species or
subspecies, scientists and managers require not only a firm
understanding of recombination, genetic drift, selection, and
gene flow, but also a critical examination of human perception
and how people connect with and define their world.
Biodiversity can be categorized in many ways as a result of the
inherent complexity and interconnected evolutionary history of
life, e.g., introgressive hybridization, horizontal gene transfer,
lateral exchange, reticulate evolution, etc. (Mallet 2005, Arnold
and Fogarty 2009). Current species taxonomies reflect only one
possible grouping structure out of many alternatives (Atran 1990,
Lakoff 1990, Newmaster et al. 2006, Padial et al. 2010). Critics
of the hierarchical nature of science-based classification systems
cite a lack of flexibility necessary to respond to a world that
includes ambiguous boundaries (Hey 2006). The subjectivity
inherent in species categories can appear inconsequential when
two species or populations are clearly distinct, but in situations
involving closely related taxa, intraspecific variation, or
geographic overlap, it becomes problematic (Mace 2004). The
influence of hybridization and introgression among subspecies,
populations, and species can be difficult because the conservation
1Natural Resources Institute, University of Manitoba, Winnipeg, Manitoba, Canada, 2Office of the Chief Ecosystem Scientist, Parks Canada,
Gatineau, Québec, Canada, 3Ɂehdzo Got'ı̨nę Gots'ę́ Nákedı (Sahtú Renewable Resources Board), Tulı́t'a, Northwest Territories, Canada, 4Aboriginal
Studies, University of Toronto, Toronto, Ontario, Canada, 5Délı̨nę Ɂehdzo Got'ı̨nę (Renewable Resource Council), Délı̨nę, Northwest Territories,
Canada, 6Délı̨nę Land Corporation, Délı̨nę, Northwest Territories, Canada, 7Biology Department, Trent University, Peterborough, Ontario,
Canada, 8Department of Linguistics, University of Toronto, Toronto, Ontario, Canada
Ecology and Society 21(2): 18
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of hybrids, which often display a continuum of genotypes, does
not fit a discontinuous species-based conservation model
(Fitzpatrick et al. 2015). Further, hybridization often necessitates
a subjective decision about the “authenticity” of a certain
genotype over another and poses a challenge to endangered
species policies, e.g., the hybridization of introduced Mallards
(Anas platyrhynchos) with several species of threatened ducks in
New Zealand to the point where all previously considered endemic
ducks may be of hybrid origin (Rhymer and Simberloff 1996,
Fitzpatrick et al. 2015).
The species Rangifer tarandus, known as caribou in North
America and reindeer in Scandinavia and Russia, are highly vagile
and occur across an extremely diverse spectrum of habitats where
they have adapted many different behavioral responses to a range
of ecological conditions. Substantial variation in caribou
behavior, morphology, life history traits, interactions with other
species, movement, diet, and social structures have made attempts
to systematically organize and characterize the species
challenging. In 1961, Banfield classified R. tarandus in North
America into four divisions based on morphological
comparisons: barren-ground caribou (R. t. groenlandicus and R.
t. granti); woodland (R. t. caribou); Peary caribou (R. t. pearyi);
and the extinct Dawson’s caribou that once occurred on the Haida
Gwaii islands of British Columbia (R. t. dawsoni). A range of
subjective subdivisions within Banfield’s designations have since
been applied based on numerous criteria, e.g., calving strategies,
ecotype designations, seasonal distributions, etc., resulting in a
complicated milieu of inconsistently applied naming conventions
that diverge across jurisdictional boundaries. The recent attempt
by the Committee on the Status of Endangered Wildlife in
Canada to define conservation units for caribou, specifically
designatable units (DUs, discrete and significant biological units
that capture irreplaceable components of intraspecific
biodiversity), found that consistent methods and criteria for
organizing the variation inherent to the species are not currently
available (COSEWIC 2011). Effective categories are needed
because units that ignore underlying ecological relationships or
misinterpret population structure lead to confusion when
implementing recovery plans and conservation policies (Crandall
2009).
For example, in the Sahtú region of the Northwest Territories,
Canada, there is considerable overlap among different caribou
herds, groups, and types that exhibit unique life histories and have
acquired different conservation statuses (Fig. 1). In the northern
portion of the region, large herds of barren-ground caribou
migrate between the open tundra and the boreal forest in response
to seasonal pulses of resources and predation pressure (Vors and
Boyce 2009, Nagy et al. 2011). In the Mackenzie Mountains,
caribou display much smaller scale seasonal migrations between
valley bottoms and alpine plateaus (Gullickson and Manseau
2000, Polfus et al. 2011, Letts et al. 2012). Throughout the boreal
forest, boreal woodland caribou exhibit sedentary behavior and
occur in small groups of ~5—15 individuals (Stuart-Smith et al.
1997, O’Brien et al. 2006, Brown et al. 2007, Courtois et al. 2007).
Data on the boundaries and degree of differentiation between
these caribou types is currently limited. Understanding how the
caribou types are structured in the Sahtú region has legal
implications because boreal woodland caribou are listed as
threatened by the federal Species at Risk Act (SARA) and the
territorial Species at Risk (NWT) Act and thus warrant specific
protection and recovery measures (Environment Canada 2012,
Species at Risk Committee 2012). The development of a national
recovery strategy and subsequent action plans for boreal
woodland caribou conservation has been delayed, in part, because
of the complex intraspecific variation that characterizes the
species. To date most range boundaries include a poor
understanding of long-term caribou movements, gene flow, and
genetic divergence. Current research examining zones of contact
and introgression among caribou subspecies aims to help define
evolutionarily significant conservation, management, and
population units (Weckworth et al. 2012, Colson et al. 2014, Røed
et al. 2014, Klütsch et al. 2016); however, alternative sources of
knowledge are rarely considered in the context of caribou ecology
and conservation (but see O’Flaherty et al. 2008, Mager 2012,
Polfus et al. 2014).
Fig. 1. The Sahtú region of the Northwest Territories, Canada,
includes the overlapping ranges of three types of caribou
(Rangifer tarandus): tǫdzı (boreal woodland caribou; striped
green), ɂekwę́ (barren-ground caribou; blue), and shúhta ɂepę́
(northern mountain caribou; stippled orange). Small black dots
represent locations of caribou fecal, tissue, and blood strip
samples collected in the Sahtú Region and Nahanni National
Park Reserve, Northwest Territories, Canada.
Challenges to the classification of Rangifer present an ideal
opportunity to use multiple knowledge sources to develop a more
thorough and complete understanding of caribou population
organization and variation (Crandall 2009). Knowledge that
arises from indigenous people’s ecological relationships is often
referred to as traditional knowledge (TK) and encodes ways of
knowing and describing environmental diversity (Hunn 2006).
Although significant scholarly work has defined and critiqued the
mechanisms, functions, cultural significance, and cognitive basis
of biological classifications (Berlin 1973, Atran 1990, Lakoff
1990, Ingold 2000, Newmaster et al. 2007), there remains space
for a more practical consideration of the substantive knowledge
indigenous people hold about their environments in the context
of conservation (Fraser et al. 2006). Traditional knowledge is a
product of a dynamic process of individual engagement with
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ecosystems that reflects people’s capacity to respond to a
constantly changing environment (Berkes 2012). Traditional
knowledge is neither static nor antiquated. Processes and
institutions guide the production and legitimation of TK as part
of a living practice that is constantly updated and renewed (Ingold
2000, Davidson-Hunt 2006). For the purpose of this paper we
make the distinction between science and TK. We find it useful
not to conflate the two knowledge traditions because the
procedures that make up science arose from their own particular
social-institutional history and bringing science and TK together
requires substantive interpretive and heuristic procedures (Scott
2011). Thus, TK has the potential to provide robust descriptions
of species variation that can add value to our understanding of
coupled human and natural systems (Fraser et al. 2006, Liu et al.
2007).
Language is one medium by which TK is transmitted and
expressed and in the Sahtú it is crucial to the interpretation,
organization, and articulation of biodiversity (Lakoff 1990,
Basso 1996, Hey 2001, Evans 2012). Knowledge holders often
need to speak their own language (rather than English) to
accurately describe complex components of TK. Examining
multiple language systems in parallel allows for descriptions that
have the potential to reach beyond one dominant biological
classification structure (Davidson-Hunt et al. 2005, Stronen et al.
2014). Languages can express meaning in different ways. This is
partially due to the fact that what must be obligatorily expressed
in one language need not be obligatorily expressed in another, and
thus the structure of the language can implicitly and explicitly
affect how speakers engage with the world and influence memory,
perception, and categorization (Markman and Hutchinson 1984,
Harrison 2007, Deutscher 2010, Boroditsky 2011). Indigenous
languages can provide refined and multifaceted descriptions of
biodiversity (Hale et al. 1992, Newmaster et al. 2007), alternative
ways of examining and relating to nonhuman animals (Ingold
2011, Miller and Davidson-Hunt 2013), and insight into the
underlying processes that create biological structure and drive
patterns of biodiversity (Ragupathy et al. 2009, Gavin et al. 2015).
To achieve effective conservation outcomes there is a need to
explicitly explore, not only variation itself, but the biocultural
forces that shape variation and the relationships people establish
within evolutionary systems (Gavin et al. 2015). The concept of
biocultural diversity emphasizes the reciprocal relationships and
overlapping realms of cultural, biological, and linguistic diversity
and the many compelling similarities between languages and
species as essential units of culture and nature (Loh and Harmon
2014, Gavin et al. 2015). Recent investigations into the link
between biological and cultural diversity have generated
important discourse concerning the vital role of TK, language,
and diverse knowledge systems in conservation and
environmental management (Davidson-Hunt et al. 2012, Gavin
et al. 2015).
Land claim settlements across the Northwest Territories have
introduced new institutions and governance structures that have
the potential to reframe policies influencing lands and resources.
In the Sahtú region, Dene and Métis representatives from local
Ɂehdzo Got'ı̨nę (Renewable Resources Councils; RRCs) recently
passed a resolution that called for a renewed commitment to adopt
TK and the laws of the Dene people as the guiding principles for
caribou research and management. To support the ambitious
goals set forth by the communities, we collaboratively developed
a research approach to explore questions about caribou variation
and differentiation using both traditional and scientific
knowledge. We focused on łeghágots'enetę “learning together”
and acknowledged the complex nature of caribou as part of a
dynamic bioculturally diverse system. Our ultimate goal was to
support the practices that enhance people’s continued
relationships with caribou and promote socially and culturally
appropriate solutions to the complex challenges facing caribou
conservation. In this article, we discuss the potential for
population genetics and TK to deepen our understanding of
caribou variation and the robust relationships that people
maintain with the species.
METHODS
Study area
The Sahtú region surrounds Great Bear Lake and encompasses
280,238 km² of central Northwest Territories (NWT), Canada
(Fig. 1), an area larger than the United Kingdom with a
population of just over 2300 people (Statistics Canada 2012). The
current regional boundaries were defined by the Sahtú Dene and
Métis Comprehensive Land Claim Agreement that concluded in
1993. Dene people have lived in the region for thousands of years
and share a common Sahtú Dene or Athapaskan/North Slavey
cultural and linguistic history (Helm et al. 2000). There are
currently five communities in the region: Délı̨nę, Tulı́t'a, Norman
Wells (Tłegǫ́hłı̨), Fort Good Hope (Rádelı̨ Kǫ́ę́), and Colville
Lake (K'áhbamı̨́ Túé). Until the establishment of local
government administrations and day schools during the post-
WWII period, Dene and Métis peoples led a nomadic existence
in a seasonal harvesting cycle (Abel 2005). Despite the shift to a
more sedentary way of life in the communities, Sahtú Dene and
Métis maintain strong cultural and social-ecological relationships
with the land and wildlife (Andrews et al. 2012a,b, McMillan and
Parlee 2013, Harnum et al. 2014).
There is considerable variation within the Dene language that is
spoken across the region, with varieties differing primarily by
sounds and vocabulary (Rice 1989). Defining linguistic subgroups
of Athapaskan languages is challenging because of historical
intergroup communication and overlapping distributions that
result in borders that in many ways resemble the ambiguous
boundaries of species classifications (Krauss and Golla 1981,
Helm et al. 2000). Language in the Sahtú region varies based on
specific family roots and community social linguistic units. The
main dialect groups are flexible and differ from each other more
or less based on family groups and historical relationships: 1.
K'áálǫ (Willow Lake), Dǝoga (Mackenzie River), and Shúhta
(Mountain) spoken in Tulı́t'a; 2. Sahtú (Bearlake) spoken in
Délı̨nę; 3. K'áhsho (Hare) and Dala spoken in Fort Good Hope
and Colville Lake, respectively (Harnum et al. 2014). Fluency in
Dene language varies greatly across generations and communities.
Elders retain the highest rates of fluency while young people are
less likely to speak Dene as their first language. In the Sahtú, an
estimated 1000 people are able to converse in the language. In
2014, Délı̨nę reported the highest percent of indigenous people
over the age of 15 who could speak Dene (78%; NWT Bureau of
Statistics 2014).
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The subarctic landscape of the Sahtú is diverse and encompasses
four major ecozones: southern arctic, taiga plains, taiga shield,
and taiga cordillera. At treeline, north of Great Bear Lake, tall
shrub tundra ecosystems of dwarf birch (Betula glandulosa) and
willow (Salix spp.) transition into boreal forest dominated by
conifers: black spruce (Picea mariana), tamarack (Larix laricina),
and white spruce (P. glauca). Deciduous stands of trembling aspen
(Populus tremuloides), paper birch (Betula papyrifera), mountain
alder (Alnus tenuifolia), and willow are found in drier and warmer
sites. Lichens, mosses, dwarf birch, cotton-grass (Eriophorum
spp.), Arctic white heather (Cassiope tetragona), Labrador tea
(Ledum groenandicum), and multiple Dryas and Vaccinium species
make up the ground cover. Dynamic fire cycles establish and
control energy flows in the boreal forest. The region contains
zones of continuous, extensive discontinuous and intermediate
discontinuous permafrost (Heginbottom 2000). The climate is
typified by long, cold winters and short, warm summers.
Precipitation is low and restricted by a rain-shadow in the
Mackenzie Valley, which generates milder climates than those to
the east and west (Dyke 2000). Mean temperature in Norman
Wells is -27.4°C in January and 16.7°C in July (Kokelj 2001).
The Dehcho (Mackenzie River) that flows into the Beaufort Sea
dominates the hydrology of the region. Its watershed is the largest
in Canada and covers approximately 1.7 million km² (Kokelj
2001). Sahtú Deh (Great Bear River), a major sub-basin, flows
from an outlet in Great Bear Lake near Délı̨nę over low relief
landscape and enters Dehcho at Tulı́t'a. West of Dehcho, the
Mackenzie and Selwyn Mountains form the northernmost
extension of the Rocky Mountain range and extend along the
Yukon/NWT border from British Columbia to the Peel River
plateau. Climatic zones vary according to the elevation gradient
that ranges from 2972 m (highest peak) to foothills between 200
—800 m. Major tributaries flowing from the mountains into
Dehcho include Begáádǝ́ (Keele), Nǫ́gha Chılı̨ne (Carcajou), and
Fahfá Nı̨lı̨né (Mountain). The Norman Range and Franklin
Mountains, which parallel the east side of Dehcho from Fort
Good Hope to Wrigley, form a series of steep bedrock ridges and
plateaus with elevations of ~1000 m (Morgan and Anderson
2013). The region has extensive karst formations including
prominent sinkholes, caves, dry valleys, and gorges (Ford 2008).
Ungulates in the region include caribou, moose (Alces alces),
muskox (Ovibos moschatus), mountain goats (Oreamnos
americanus), and Dall sheep (Ovis dalli). The large mammal
predator community consists of grizzly bears (Ursus arctos), black
bears (U. americanus), wolverines (Gulo gulo), wolves (Canis
lupus), and Canada lynx (Lynx canadensis).
Research design
Our community-collaborative research project was developed and
implemented within the current institutional and political
structures of the Sahtú Land Claim and the Mackenzie Valley
Resource Management Act (1998). Under this political structure
the Ɂehdzo Got'ı̨nę Gots'ę́ Nákedı (Sahtú Renewable Resources
Board; SRRB) and the five local RRCs of the Sahtú Region are
responsible for managing renewable resources in the region
including wildlife and habitat. The research project initiated a
collaborative partnership between the SRRB, RRCs, university
researchers, and the NWT Department of Environment and
Natural Resources (ENR). We developed partnerships with key
knowledge holders, elders, and an advisory group with support
from the formal institutional structures. In September 2012, the
Sahtú RRCs passed a joint resolution supporting the adoption
of TK and Dene law as essential components of caribou research.
Our research project is a direct outcome of the resolution and was
designed to support the initiatives proposed by communities in
line with the principals of community-based participatory
research frameworks and methodologies (Appendix 1; Hall 1979,
Ferreira and Gendron 2011).
Our research included community members in all phases of the
research process and created an open and transparent dialogue
between scientific and traditional knowledge (Cruikshank 1981,
Cornwall and Jewkes 1995). We prioritized łeghágots'enetę
“learning together” and knowledge generation in an attempt to
develop a richer, culturally respectful, and relevant understanding
of caribou variation. The principles and protocols governing the
research were covered by a multiyear research license from the
Aurora Research Institute (15217, 15443, and 15597), wildlife
research licenses from ENR (WL500104, WL500307), and a
University Ethics Protocol (J2012:202).
In December 2012, we held research planning meetings in
Norman Wells, Fort Good Hope, Tulı́t'a, and Délı̨nę to discuss
the project and plan for winter field work. Discussions facilitated
the development of research priorities, research questions, and
appropriate methods for the current and future monitoring of
caribou populations. In January 2013, we held a series of RRC
and public meetings to plan for winter sampling, build awareness
for the program, and train community members in sampling
techniques. We developed a Memorandum of Understanding
with each RRC to confirm the governing principles of the
research, budgets, research methods, intellectual property rights,
and administration of the project. Subsequent RRC meetings
were held in each community (including the addition of Colville
Lake) to continue to guide the research during the winters of 2013
and 2014.
In alignment with our approach of łeghágots'enetę “learning
together” we prioritized opportunities that allowed for the
establishment of collaborative relationships between an
interdisciplinary group of community researchers, local experts,
and academic researchers (discussed in more depth in Appendix
1). We promoted ongoing communication with the public through
outreach and relationship building in the communities. Local
experts shared knowledge about caribou histories, movements,
and identities during formal and informal interactions on the
land. The knowledge helped to guide sample collection and
identified concepts and ideas that were discussed in depth at focus
group meetings, with the advisory group, and among coauthors.
All community participants received honoraria for their time.
Focus group meetings
We held focus group sessions (Agar and MacDonald 1995,
Morgan 1996, Berman and Kofinas 2004) to build a
comprehensive understanding of the origin, dynamic
interactions, and spatial structure of caribou in the Sahtú region.
Workshops lasted between one and two days. Three to 10 local
experts, selected in partnership with the local RRCs, participated
in five focus group sessions held in each of the communities (total
39 people) in April of 2013. Focus group participants had
extensive firsthand knowledge of caribou populations in the
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region. Interpreters helped to develop appropriate metaphors to
describe genetic concepts in ways that resonated with community
members. We found that simultaneous interpretation was often
unsuccessful when explaining complex scientific topics. Instead,
we more often used sequential interpretation, which allowed more
time for ideas to be heard and understood (many participants
were bilingual). We also worked with community researchers and
interpreters to build a robust understanding of Dene concepts
and avoid back-translations from English. Focus group meetings
included significant discussions in Dene language.
At meetings we provided an overview of the research process and
described the methods and preliminary results. We documented
information through word maps, flow charts, diagrams,
geographic maps (Google Earth 7.1.5.1557), and note taking. We
digitally audio-recorded meetings with participant consent.
Consistent with the iterative nature of our research process, we
concluded focus group meetings with a discussion of how the
project could be improved. Key sections of the meetings were later
translated and transcribed by local language specialists in English.
Genetic sampling
Community members, researchers, collaborators, and industry
monitors collected caribou fecal pellets noninvasively during the
winters of 2013 and 2014 by gathering frozen pellet piles found
on the snow in plastic bags. In general, hunters and trappers
collected samples while traveling on skidoo trails, winter roads,
seismic lines, and traditional trails during normal on-the-land
activities. The sampling area represented the range of all three
types of caribou in the region. We encouraged community
members to help with sample collection during outreach at public
meetings, through promotional posters, regional newspaper
stories, on local radio, and in Facebook posts. Community
members received a $25 gift card for gas at a local gas station for
each caribou fecal pellet sample they provided. The RRCs and
Norman Wells ENR staff oversaw sample collection, data entry,
and gift card distribution. Collaborations with industry partners
also allowed for targeted sampling by helicopter to locate areas
of caribou activity (tracks, cratering, etc.) during the winter of
2013. In April 2014, we spent an additional three days flying by
helicopter with participants selected in collaboration with the
RRCs of Fort Good Hope, Tulı́t'a, and Délı̨nę to collect scat
samples and fill sampling gaps. We also collected muscle tissue
samples and blood strip samples from hunted animals in
collaboration with a caribou health monitoring study (Brook et
al. 2009). Finally, we included caribou fecal samples collected in
Nahanni National Park Reserve in the southern Mackenzie
Mountains through collaborations with Parks Canada.
Microsatellite genotyping
We followed microsatellite genotyping protocols that had been
established as part of a long-term caribou genetic database to
ensure the production of high quality genetic profiles (Ball 2007,
Ball et al. 2010, Galpern et al. 2012a, Hettinga et al. 2012, Klütsch
et al. 2012). To isolate DNA we swabbed and amplified the
mucosal layer covering the caribou fecal pellets. We genotyped a
panel of nine microsatellite loci (BM848, BM888, MAP2C, RT5,
RT6, RT7, RT9, RT24, and RT30; Bishop et al. 1994, Wilson et
al. 1997, Cronin et al. 2005). In May 2013 swabbing took place in
Norman Wells where we worked with local students and trained
technicians to build capacity in the communities and to continue
to foster collaboration during the research process. Subsequent
genetic analysis took place at Trent University. Extraction
protocol followed Ball (2007) and profiling procedures can be
found in Galpern et al. (2012a) and Klütsch et al. (2016).
Electropherograms were scored by at least two individuals with
GENEMARKER v. 1.9.1 (SoftGenetics, LLC) to determine
allele sizes. Samples included in the final dataset had a minimum
of eight successfully amplified loci. We used AlleleMatch 2.5
(Galpern et al. 2012b) to screen profiles for genotyping errors,
remove duplicate profiles, and identify the number of individual
caribou sampled. Only one sample from each individual caribou
was included in subsequent analysis.
Mitochondrial DNA (mtDNA) sequencing
Genetic analysis of nuclear and mtDNA (mitochondrial DNA),
larger sample sizes, and improved analytical methods have
influenced our understanding of caribou taxonomy and
evolutionary history in North America (McFarlane et al. 2009,
Klütsch et al. 2012, Mager et al. 2013, Yannic et al. 2014).
Specifically, analysis of mtDNA has revealed two distinct
phylogenetic groups of caribou that represent separate northern
and southern glacial refugia during the Pleistocene. Many of the
boreal woodland caribou in the southern Canadian provinces
originated south of the Laurentide ice sheet that covered most of
present-day Canada (North American lineage; NAL), while
barren-ground caribou likely originated in the northern
unglaciated refugium of Beringia and the Canadian high arctic
(Beringian-Eurasian lineage; BEL, McDevitt et al. 2009, Klütsch
et al. 2012) To examine phylogeographic structure we amplified
and sequenced a 429 bp mtDNA control region fragment using
the primers: L15394:5′ - AAT AGC CCC ACT ATC AGC ACC
C - 3′ and H15947:5′ - TAT GGC CCT GAA GTA AGA ACC
AG - 3′ (Flagstad and Røed 2003). Only samples from individual
caribou were sequenced. We followed lab procedures outlined in
Klütsch et al. (2012). We used the program BioEdit 7.2.5 (Hall
1999) to check and align sequences. Mutations were manually
double-checked and all newly identified haplotypes were
resequenced to confirm their identity.
Genetic analysis
We used the program Structure 2.3.4 (Pritchard et al. 2000) to test
for population subdivision and assign individual caribou to
inferred subpopulations (Falush et al. 2003). We performed five
runs with 1,000,000 burn-in iterations and 10,000,000 Markov
chain Monte Carlo (MCMC) repetitions. We varied the number
of K between 1 and 15 under the admixture model with correlated
allele frequencies, and specified no a priori models of
subpopulation structure. We plotted the mean and variance in
likelihood per K using Structure Harvester v0.6.94 (Earl and
vonHoldt 2012). We found the average individual membership
coefficients across the five iterations using the programs
CLUMPP 1.1.2 (Jakobsson and Rosenberg 2007) and
DISTRUCT 1.1 (Rosenberg 2004). We mapped the structure
output by interpolating the average probability assignment score
using the inverse-distance-weighted interpolation in ArcGIS 10.1
(ESRI, Redlands, California, USA) and confining the
interpolation to sampled locations (Twomey et al. 2014).
We assessed genetic diversity by calculating genetic indices of the
structure informed populations with GenAlEx 6.501 (Peakall and
Smouse 2006, 2012) and HP-Rare 1.1 (Kalinowski 2005). We
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tested for significant deviations from Hardy-Weinberg
equilibrium (HWE) and linkage disequilibrium (LD) per locus
and population using GenePop 4.2 (Rousset 2008). We used
SPAGeDi 1.5 (Hardy and Vekemans 2002) to test the
microsatellite pairwise differentiation with FST and RST and used
ARLEQUIN 3.5.2.2 (Excoffier and Lischer 2010) to test the
mtDNA pairwise differentiation (ΦST) and analysis of molecular
variance (AMOVA; see Klütsch et al. 2016 for further details).
Collaborative analysis
We analyzed information shared during focus group sessions
using a combination of thematic analysis and modified grounded
theory methods to identify important concepts and themes
(Glaser and Strauss 1967, Bernard 2002, Berman and Kofinas
2004). We used the program NVivo (QSR International Pty Ltd.
Version 10) to code meeting notes, transcripts, and other TK
reports and publications. Hierarchical categories emerged
through the process of coding and were developed into potential
themes as the ideas became more concrete through repeated
identification. To support a collaborative production of
knowledge we facilitated a process for coanalysis of the TK and
genetic data. With the help of RRCs, individuals were selected
from all Sahtú communities for their expertise and interest in
participating in an advisory group. The advisory group’s role was
to guide the project, ensure that Dene knowledge was properly
and respectfully interpreted, and provide the TK context needed
to help accurately interpret the genetic data. We discussed TK
themes, language, and genetic results in two separate 3-day
meetings to clarify and develop important concepts (Fig. A1.1
and A1.2). Our first advisory group meeting, held in June 2014,
included seven participants who helped plan and select additional
elders to participate in a follow-up meeting. The meeting was
transcribed and coded. The second meeting was held in February
2015 and included 11 participants.
We employed what we call a language-based methodology by
working to elucidate conceptual Dene TK needed to ground the
concepts and themes. Elders often requested to speak only in Dene
language when discussing TK. Walter Bayha uses a translation
of the teachings of his Shúhtagot'ı̨nę grandfather, Joseph Bayha,
to explain this affinity: “Our history is written on the land. The
language comes from the land.” Dene knowledge holders stressed
the importance of respecting dialect differences and were very
careful to avoid making assumptions about how speakers of other
dialects would express a word or concept. The advisory group
agreed that in general it is best to defer to Dene words in a dialect
with a direct relationship to the caribou population being
discussed. In this paper, we generally use Shúhta (S) dialect when
referring to caribou in the mountains and Sahtú/Délı̨nę (D)
dialect with respect to barren-ground caribou terminology. We
also include K'áhsho (K) words where possible (Fig. 2).
The advisory group focused on key concepts and ideas from
previous focus group meetings, TK literature, and publications.
We used visual facilitation techniques to guide the advisors to
expand on important topics and explain the genetic data. We paid
special attention to clarifying Dene TK and descriptions of the
types of caribou found in the Sahtú region as well as presenting
our preliminary interpretation of the genetic data to help assess
how appropriate our inferences were with respect to TK of
caribou ecology in the region. Beyond the two formal meetings,
several of the advisory group members have continued to guide
the research, interpret genetic data, work on the details of Dene
language translations, review drafts of manuscripts, copresent the
research in schools and during public presentations, and coauthor
this manuscript.
Fig. 2. Word descriptions and definitions in the Shúhta (S),
Sahtú/Délı̨nę (D), and K'áhsho (K) dialects of the North
Slavey language in the Sahtú region of the Northwest
Territories, Canada.
RESULTS
Traditional knowledge
During focus group meetings elders continually expressed their
personal relationship with caribou as being crucial to
understanding caribou knowledge. One example of this
association was described by Alfred Taniton as bedélé t'á núzhǫ
(D) that translates to “we grew up with their blood” (Fig. 2).
Alfred Taniton said, “we were raised with the blood from the
caribou. In the past, the people have always survived because of
the blood of the animals.” The intimate interaction between
human and nonhuman animals highlights how many indigenous
people recognize the importance of their relationships with other
beings on a daily basis. The concept behé ts'enézhǫ (D) “we grew
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up with them,” or as Walter Bayha translated, “we are people with
them,” further illustrates how Dene people relate to caribou as
unique entities, capable of intelligence, identity, perception, self-
awareness, rationality, and intentionality.
When Dene people relate to nonhuman animals autonomously
they follow important Dene laws regarding bets'erı̨hchá (D)
“respect” and łegháts'eredı (D) “we give to each other.” Dene
people recognize that individual animals have unique perspectives
that allow them to gain knowledge and intelligence in distinctive
ways (Legat 2012). Elders repeatedly state that caribou are their
own bosses and cannot (and should not) be controlled by people.
Rather, Dene laws provide guidance for mutual respect and honor
that require that Dene take care of caribou so that caribou will
reciprocally provide for them. Gordon Yakeleya stated the
following:
It’s very important that we look after the animals, we
have to have respect for them. There’s a reason why they
do what they do. They want to survive like we want to
survive. It’s the same thing. That’s what my mom and dad
always said: “Animals are like human beings.” They do
everything for a reason, just like we do. Like we go to
store, they get food for the whole winter. They raise their
young ones and teach their young ones. We do the same
thing.
Fundamental to the Dene relationship with caribou is a profound
knowledge of caribou morphology, behavior, and habitat
preferences. Dene people identify three main types of caribou in
the region: shúhta ɂepę́ (S) “mountain caribou,” ɂekwę́/ɂedǝ (D/
K) “barren-ground caribou,” and tǫdzı “boreal woodland
caribou” (Fig. 2). Participants describe tǫdzı as bekwı́ dezene (D)
“darker colored/having a dark head,” larger and heavier than
ɂekwę́. Shúhta ɂepę́ are identified by their large size and close
association with the mountains. The caribou types can also be
distinguished based on their tracks (size, shape, and the encoded
behavior) and their general location. Though the ranges of tǫdzı
and ɂekwę́/ɂedǝ (D/K) overlap in many areas during the winter,
knowledgeable hunters and elders are able to distinguish between
the types. For example, Gabe Kochon of Fort Good Hope
described a situation where he once saw a very large male tǫdzı
(that was a dark color) in the center of a group of female barren-
ground caribou during rut many years ago. Hunters have even
reported being able to distinguish types by the taste of the meat.
Interestingly, some Shúhtagot'ı̨nę elders describe a fourth type of
caribou called tęnatł'ǝa (S) “the fast runners” that live in the
Mackenzie Mountains, migrate long distances, and are identified
by particular morphological markings.
Although there are dialect differences in the words used to refer
to barren-ground caribou, the classificatory term tǫdzı “boreal
woodland caribou” is shared across the region and mirrors the
caribou population’s distribution. The word tǫdzı is also found
in the Tɬı̨chǫ language of central NWT and the second part of
this word, dzı, is commonly associated with Athapaskan caribou
words and is found in many related languages including Tɬı̨chǫ,
Dene Su̜ɬıné, and languages of the Yukon, Alaska, and British
Columbia. Further, in Délı̨nę the word ɂekwę́wa (D) can be used
when discussing different types of caribou and translates to “the
real (or the original/prototypical) caribou,” which links the
language with the histories of the caribou. Gabe Kochon of Fort
Good Hope also explained through translation that the “regular”
caribou have always been in the north (living in the barren-land
north of Fort Good Hope) whereas tǫdzı probably had a different
origin.
Knowledge of differences in behavior between different caribou
types is essential to successful hunts. Dene hunters describe how
tǫdzı react much more strongly to the presence of humans than
ɂekwę́ or shúhta ɂepę́. Focus group participants explained that to
successfully hunt tǫdzı it is necessary to anticipate the animal’s
behavior. They used the Dene phrase goecha fehtǝ (S) to describe
a situation in which a tǫdzı will loop back on his or her own trail
so he/she can rest (lie down) in a sheltered area downwind from
his/her path and thus be alerted to the scent of potential predators
that might be following his/her tracks. The hunter must react by
predicting the caribou’s behavior and looping around behind to
ensure that the animal can’t smell the hunter before they can take
a shot. In Dene, this can be described as goecha gots'anele (S) “to
hunt from downwind.” This behavior is also described for moose,
but not ɂekwę́ or shúhta ɂepę́. Interestingly, a similar word is found
in the place-name Gocha Túé (D) that was first translated in the
1860s by French Oblate missionary Father Émile-Fortuné Petitot
as Shelter Lake (and which he renamed Lac Ste-Thérèse/Lac
Sainte-Therese; Petitot 1893). Walter Bayha was able to use
Petitot’s translation to uncover the obscured meaning of the
place-name through the word “gocha” and the sheltered
snowdriftless characteristics of the lake where he spent time when
he was young.
Monitoring caribou population fluctuations has been imperative
to Dene survival for millennia (Beaulieu 2012). Over time,
significant migrations and range-shifts have occurred between
caribou groups. Dene descriptions of large-scale caribou
movements help explain current caribou distribution, temporal
patterns, and can be used to predict future movements.
Uncommon movements of barren-ground caribou herds that
winter around Great Bear Lake are often recalled and discussed
because they can influence caribou health and hunting methods.
For example, Gordon Yakeleya remembers how in the winter of
1988 barren-ground caribou migrated all the way down to K'áálǫ
Tué (Willow Lake) near Tulı́t'a and displaced resident tǫdzı. In
Fort Good Hope, elders speak about a large herd of caribou that
crossed the Dehcho and headed into the foothills of the mountains
many years ago. Gabe Kochon said that they never saw the entire
herd return or migrate back across the river. He related the
following in Dene language:
There was a lot of them, I have witnessed the caribou
crossing ... ice, even though it was broken up, there was
lots of them... Many years ago, the caribou crossed to
the other side.... They have been gone a long time and
people are saying that they have become lots again and
they have been using that area for calving ... This is
according to the elders and they also say that they will
never disappear.
Microsatellites
We collected caribou scat samples from the Sahtú region and
Nahanni National Park Reserve with the cooperation of ~100
community members and project collaborators including grade
school and high school students, hunters, trappers, environmental
monitors, researchers, and industry partners. We obtained 1036
caribou fecal samples, 96 caribou tissue samples, and 16 caribou
blood strips from localities across the Sahtú region and Nahanni
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Table 1. Genetic diversity estimates, averaged across nine microsatellite loci, for the three major caribou (Rangifer tarandus) groups
identified by structure analysis: number of samples (N), number of alleles (NA), allelic richness (AR), private allelic richness (ARP),
observed (HO) and expected (HE) heterozygosity, FIS estimates, and standard errors (SE) for each estimate.
Group N NASE ARARP HOSE HESE FIS
Barren-ground 123 15.1 1.20 14.97 2.50 0.84 0.013 0.87 0.010 0.035
Boreal woodland 171 12.0 1.43 11.26 1.05 0.79 0.023 0.79 0.023 0.005
Mountain 260 17.1 2.93 14.38 2.29 0.82 0.014 0.84 0.011 0.031
National Park Reserve (Fig. 1). A subset of these samples were
analyzed and 996 were successfully amplified at > 8 loci and
included in the analysis. We identified 555 individual animals
(47% female, 41% male, and 12% unknown gender). Missing data
in the microsatellite dataset was low (3.43%). Average number of
alleles per locus was 20 across all individuals (Table A2.1). We did
not find evidence for systematic deviations from HWE in specific
populations (4/27 cases were significant after Bonferroni
correction) or LD (no cases significant).
Structure analysis identified K = 2 as the highest level of
substructure (using the ∆K criterion) that corresponded to a
boreal woodland group and a mountain/barren-ground group
(Fig. A2.1). Further finer-scale structure of K = 3 corresponded
with an additional split between mountain and barren-ground
(Fig. 3, Fig. A2.2). These partitions were largely concordant with
the TK on caribou types in the region, supporting an ecological
foundation for three inferred groups that represent clusters of (1)
barren-ground caribou; (2) boreal woodland caribou; and (3)
mountain caribou (Fig. 3). The barren-ground population had
the highest levels of allelic diversity and heterozygosity (Table 1).
Pairwise comparisons between groups (FST and RST) indicated
low levels of differentiation, though the boreal woodland group
was the most differentiated from the other two types (Table A2.2,
A2.3).
mtDNA
We identified 69 mtDNA control region haplotypes from 337
individual caribou (Table 2, Fig. A2.3). We fit the mtDNA data
into the well-resolved phylogeny of NAL and BEL (see Klütsch
et al. 2012, 2016). Unlike the nuclear markers (microsatellites)
that showed intraspecific divisions between types at the regional
scale, the phylogenetic mtDNA analysis revealed that caribou of
the Sahtú belong predominantly to the BEL (96.7% Beringian).
However, very few haplotypes (n = 12) were found in more than
one of the three clusters identified by structure (Fig. A2.3). Most
haplotypes were nonoverlapping, signifying long-standing
diversification among the types. We identified only 3 NAL
haplotypes (in 11 caribou) in the study area, most belonging to
haplotype 50 (n = 9 boreal woodland caribou) in the Sahtú region
(Fig. A2.3). This was an especially surprising result and suggests
that boreal woodland caribou in the northern extent of their range
are distinct from more southern boreal woodland caribou that
generally belong to the NAL (Klütsch et al. 2012). Pairwise
comparisons using the mtDNA data (ΦST) were low but significant
(Table A2.4) and showed the strongest differentiation between
boreal woodland and mountain, and barren-ground and
mountain, which may suggest that the mountain group have been
historically isolated. The AMOVA of the three groups revealed
that ~14% of the mtDNA genetic variation was found among
populations (Table A2.5).
Fig. 3. We analyzed microsatellite data from caribou (Rangifer
tarandus) genetic samples collected in the Sahtú region and
Nahanni National Park Reserve of the Northwest Territories,
Canada from 2012 to 2014. We used structure software to
assign individual caribou to inferred genetic clusters. We found
support for K = 3 populations (shown in bottom bar) that
coincided with clusters of (1) barren-ground (blue), (2) boreal
woodland (green), and (3) mountain (red). Vertical colored bars
indicate the probability that an individual belongs to a certain
group. We mapped the structure output using the inverse-
distance-weighted interpolation in ArcGIS and constrained the
interpolation to the sampled locations.
Caribou spatial diversity
Collection sites were distributed across the range of all types of
caribou in the Sahtú region (Fig. 3) and were focused on
traditional hunting areas in the mountains (along the Begáádǝ́
“Keele” and Nǫ́gha Chılı̨ne “Carcajou” rivers, Tets'ehxe “Drum
Lake,” and “Canol Lake”), Nahanni National Park Reserve, the
boreal forest in the Mackenzie Valley, and the winter-ranges of
the Bluenose West (area surrounding K'áhbamı̨́ Túé “Colville
Lake,” Nılı̨n Túé “Lac Belot” and Tashı́n Túé “Lac Des Bois”)
and Bluenose East (Ɂehdaı̨la “Caribou Point” and around Délı̨nę)
barren-ground caribou herds.
Strikingly, the spatial distribution of the boreal woodland genetic
cluster encompassed the known range of boreal woodland
caribou and was restricted to the boreal forest of the Mackenzie
valley (Fig. 3). Although the geographic cohesion was strong there
was overlap with the two other clusters that demonstrates at least
some level of intergradation between the types. It is well known
that barren-ground caribou herds often overlap in distribution
with the much less numerous boreal woodland caribou in the
winter. However, as described above, knowledgeable hunters are
able to distinguish between the types and this was also
demonstrated by the genetic results. In the winter of 2013, Wilbert
Kochon, a Colville Lake hunter, killed three caribou, which he
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Table 2. Haplotype (mtDNA) genetic diversity for the three major caribou (Rangifer tarandus) groups identified by structure analysis:
number of individual caribou samples analyzed for mtDNA (N), number of samples assigned to the North American and Beringian-
Eurasian haplogroup lineages (NALN and BELN, respectively), number of haplotypes in the NAL (NALH) and BEL (BELH)
haplogroups, nucleotide diversity (π), and gene diversity (GD) with standard deviations (SD).
Group N NALNBELNNALHBELHπ SD GD SD
Barren-ground 93 1 92 1 37 0.0170 0.0090 0.9385 0.0165
Boreal woodland 110 9 99 1 12 0.0192 0.0100 0.844 0.0195
Mountain 134 1 133 1 31 0.0213 0.0110 0.9174 0.0144
identified as tǫdzı, in an area where overlap with barren-ground
animals was clearly occurring (based on tracks and reported
sightings). Later, genetic analysis of tissue samples from the three
caribou indicated they clustered with the boreal woodland group
(average probability of assignment was 0.942 to the boreal
woodland cluster). Interestingly, the few samples collected along
the Mackenzie River near Nahanni National Park Reserve also
clustered most strongly with the boreal woodland group (Fig. 3).
The spatial boundaries between the barren-ground and mountain
clusters were less distinct, however the two groups generally occur
in the vicinity of Great Bear Lake and within the Mackenzie
Mountains, respectively (Fig. 3). Admixture is apparent
throughout the Mackenzie Mountains as well as in the winter-
ranges of the barren-ground herds. Interestingly, genetic analysis
of the nine individual caribou sampled in the foothills of the
mountains across from Fort Good Hope revealed that the group
was genetically more similar to the barren-ground cluster than
the mountain cluster. This coincides with the event described by
Gabe Kochon about the historic movement of a large group of
barren-ground caribou that crossed the Dehcho (Fig. A2.4).
DISCUSSION
In this study, we used a participatory approach to examine the
biological variation of caribou populations of the Sahtú region
of the Northwest Territories. Our community-collaborative
research engaged local indigenous experts in all stages of the
project and generated results that united Dene TK and population
ecology. The participatory framework and iterative methods
generated space for the refinement of collaborative research
questions and allowed for a rigorous knowledge coproduction
process. Our results provide evidence for genetic, linguistic,
historical, phenotypic, and behavioral differentiation among the
caribou types in the region. By recognizing the lived experience
and TK of indigenous people we were able to develop more
profound understanding of caribou ecology through which we
were able to more accurately interpret the population genetic
results. The genetic subpopulation structure corresponded to
caribou types that are recognized and distinguished by Sahtú
Dene and Métis people through their language. Detailed
descriptions of tǫdzı “boreal woodland caribou,” ɂekwę́ (D)
“barren-ground caribou,” and shúhta ɂepé (S) “mountain
caribou” denote quantifiable characteristics that categorize
caribou in the region.
Dene concepts reflect ecological processes and relationships that
bring the complexity of dynamic biocultural systems to light. The
consistency of the word tǫdzı across all Sahtú region dialects as
well as the Tɬı̨chǫ region may suggest stability of the boreal
woodland caribou phenotype in the region (though more research
is needed to understand the contrasting pattern of variability in
barren-ground nomenclature). Interestingly, we found substantial
genetic differentiation between tǫdzı and other caribou in the
microsatellite genetic structure (Fig. 3). This is surprising because
in other areas of North America where overlap among caribou
types occurs, such clear delineation is not observed (Boulet et al.
2007, Klütsch et al. 2016, Pond et al. 2016). Thus, even in the face
of extensive overlap and known mixing with barren-ground and
mountain caribou populations (described by local Dene people),
there are likely important adaptive traits that are necessary to
retaining the behavioral and genetic characteristics of tǫdzı.
Mechanisms that can produce intraspecific population structure
across continuous habitats include isolation by distance,
divergence with barriers, drift after expansion, and local
evolutionary adaptation (Puckett et al. 2015). In British
Columbia, strong differentiation between wolf populations,
which are capable of large dispersing movements, has been
attributed to evolutionary adaptation to different ecological
conditions (Stronen et al. 2014).
It is likely that the differentiation between tǫdzı and other types
of caribou in the region is due in part to ecological divergence.
Dene knowledge of the association between tǫdzı and the boreal
forest is ubiquitous across the Sahtú region and was recorded as
far back as the 1860s when Petitot documented Dene language
and culture during his travels as a missionary (Moir 1998).
Petitot’s writings include descriptions of woodland caribou as
loners that lived in the forests (Petitot 1893). Knowledge about
differences in behavior among the types of caribou that occurred
in different habitats was crucial to Dene survival. Historically,
some of the most renowned Dene hunters were those who could
successfully hunt and kill the large and widely dispersed tǫdzı.
The more closely people associate with nonhuman animals, the
more intimate and detailed the knowledge becomes (Brightman
2002). As Fred Sangris, a Yellowknives Dene, said, “We learn by
being in the field, by being with ekwǫ́ [barren-ground caribou] all
the time” (Sangris 2012:77). The historic caribou movements only
observed by people with intimate knowledge of the environment
also played an important role in refining our questions and
methods. Gabe Kochon’s detailed knowledge of historic caribou
movements was crucial to our decision to collect samples from
that specific region of the mountains. The information provided
by the TK also allowed us to accurately interpret the genetic
patterns that otherwise would have been difficult to understand,
and supported the historic occupancy of distinct caribou groups
in the region.
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Likewise, Alfred Taniton’s description of growing up with the
blood of the animals demonstrates how Dene people’s survival
and ways of life are linked with wildlife. Historically, people
traveled across the land to hunt caribou for essential food,
clothing, and tools and these practices are part of the expression
of their identity. The depth of Dene people’s relationship with
caribou is revealed within Dene language. Dene language includes
numerous descriptors that help facilitate communication and
improve hunting success. Language can also provide clues to the
histories of the caribou through descriptions like ɂekwę́wa “the
original or prototypical caribou” that is used in Délı̨nę to refer to
barren-ground caribou and inform alternative ways of classifying
the relationships between caribou types. Although it is difficult
to determine the exact time-frame reflected in the language, it is
well established that Dene people have a substantial history with
caribou, and barren-ground caribou in particular have been a
crucial and relatively consistent resource for at least the last 8000
years (Gordon 2003, 2005, Andrews et al. 2012b, Beaulieu 2012).
The rich vocabulary of Dene caribou words points to the ease
with which TK holders are able to describe complex behaviors
and actions in Dene language that are difficult and unwieldy to
depict in English. The use of words like goecha gots'anele in the
context of hunting are understood by a relatively small portion
of community members and require significant discussion in
Dene language with elders, knowledgeable hunters, and
interpreters to capture the detailed nuances in English.
Conducting work in both Dene language and English was a
consistent challenge. However, the dialogue and back and forth
that was required to refine terms and concepts allowed us to come
to a common understanding and identify deeper ecological
connections that might not be apparent on the surface. As Walter
Bayha pointed out, “If I didn’t speak the language I wouldn’t be
able to make these connections.” Thus, the examples we provide
show that Dene language is deeply adapted to the environment
in which it evolved and that a focus on Dene language, facilitated
by the distinct disciplinary backgrounds of our research team, is
one of the most important contributions of the collaborative work
(see Appendix 1 for more information).
The complex nature of caribou substructure is further revealed
by the differences between genetic marker types. Examining
multiple markers concurrently can provide information about
how phenotypic differences may be contributing to historical
isolation and present patterns of gene flow (Wood et al. 2014). In
the case of tǫdzı in the Sahtú, the mtDNA haplogroups do not
coincide geographically with the microsatellite markers. Similar
divergence among markers has been shown in shovel-nosed
snakes (Wood et al. 2014) and chipmunks (Good et al. 2008, Hird
and Sullivan 2009). Surprisingly, results from the pairwise
mtDNA analysis indicated that boreal woodland/tǫdzı and
barren-ground appear to be more closely related than either are
to the mountain group (Table A2.4). Recent research suggests that
mtDNA can introgress quickly, even at low levels of gene flow,
while other loci remain resistant to introgression (Chan and Levin
2005, Hird and Sullivan 2009). Thus, one possible explanation for
the phylogenetic pattern present in tǫdzı is potential historic
introgression with northern Beringian lineage animals.
Alternatively, the boreal woodland caribou/tǫdzı phenotype may
be an independent derivation from the BEL with little or no
contribution from southern evolved boreal woodland caribou
that carry NAL haplotypes. Analysis of competing evolutionary
models will help identify and date divergence events and historical
introgression between populations that have contributed to the
current spatial genetic variation (e.g., Klütsch et al. 2016).
Variation below the species level is an important component of
biodiversity because it provides the genetic variation required for
incipient speciation and local genetic adaptation (Wood et al.
2014, Mee et al. 2015, Hamilton and Miller 2016). Cryptic
intraspecific diversity, as is displayed between caribou ecotypes
and subspecies (Pond et al. 2016), can be especially contentious
because it is not always clear how to best identify, delimit, or
maintain genetic lineage diversity (Mace 2004, Wood et al. 2014,
Fitzpatrick et al. 2015). Further, from a purely scientific
perspective, there can exist multiple valid interpretations because
rules for finding discontinuity in genetic or spatial ecological data
are at some level arbitrary. Combining multiple knowledge
systems can help to provide complementary criteria for
designating distinct units for conservation. We found this was the
case in our study and used Dene TK to help us interpret the genetic
statistical output (choosing K = 3 as the most biologically relevant
inference that was supported by both the statistical analysis and
TK). In doing so we use multiple knowledge sources to guide the
translation of data to understanding. Thus, through a pluralistic
approach we were able to demonstrate the ways that linguistic,
TK, and genetic patterns corroborate each other and allowed us
to identify criteria that can be used to identify and differentiate
between groups of animals for biodiversity conservation.
Preserving evolutionarily significant diversity in caribou that is
identified through the analysis of multiple genetic markers and
TK is essential because caribou populations in the southern
portions of their range face extirpation (Hebblewhite et al. 2010,
Johnson et al. 2015). Further, recent research on loggerhead sea
turtles (Caretta caretta) has shown that populations at the margins
of the species range can be important reservoirs of genetic
diversity and “contribute disproportionally to the adaptive
potential and future viability of the population” (Stiebens et al.
2013:8). Thus, the genetic differentiation of tǫdzı and the rich TK
on the unique attributes of the type provide evidence for their
prioritization as an irreplaceable component of Canada’s
biodiversity. However, sustainable conservation strategies must
find ways to maintain not simply the categorical entities (like
subspecies) but rather the dynamic relations among peoples and
species as the basis of bioculturally diverse systems.
The importance of supporting social-ecological relationships and
processes is gaining momentum in conservation science (Gavin et
al. 2015). Proponents of this viewpoint maintain that
conservation priorities should be not be defined in relation to
discontinuous species, but rather directed toward the protection
of essential processes that create adaptive potential and sustain
biological variation (Bowen 1999, Crandall et al. 2000, Moritz
2002, Eizaguirre and Baltazar-Soares 2014). This viewpoint
acknowledges the subjectivity of species categories and highlights
the importance of conserving the dynamic nature of functioning
ecosystems. Strategies for identifying units for conservation that
integrate multiple biological criteria, acknowledge the dynamic
nature of intraspecific diversity, respond flexibly to specific
circumstances, and adapt to differing situations are needed to
cultivate evolutionary potential in a changing environment
(Fraser and Bernatchez 2001).
Ecology and Society 21(2): 18
http://www.ecologyandsociety.org/vol21/iss2/art18/
Similarly, ethnoecological explorations of the intrinsically
adaptive nature of categorization systems place emphasis not on
categorical entities (contents of categories such as species) but on
defining elements of an ecosystem in relation to the other elements
that surround them in time and space (Ingold 2011). As a
consequence, more attention is given to an entity’s function, its
role in the larger spatial and temporal environment, rather than
its intrinsic qualities that are devoid of context. Identifying
important connections among ecosystem components allows the
unpredictable emergent properties of a system to become
apparent (Berkes et al. 2003, Ingold 2011) and can lead to
improved conservation planning (Alcorn 1993, Fraser et al. 2006).
For example, research suggests that TK classification systems can,
in some cases, identify more taxa than science-based systems
(Newmaster et al. 2007, Ragupathy et al. 2009) or be especially
suited to identifying intraspecific diversity (Fraser et al. 2006). In
the Sahtú, the description of a distinct group of caribou in the
mountains known as tęnatł'ǝa warrants further study because they
may harbor unique genetic diversity and could play an important
role in intraspecific dynamics. Thus, the analysis of genetic
variation in conjunction with the relationships indigenous people
maintain with species has the potential to reveal complex patterns
that would likely not be apparent when evaluated separately.
CONCLUSION
A renewed focus on multidisciplinary conservation frameworks
demonstrates the importance of studying human and natural
systems (social-ecological systems) in tandem (Liu et al. 2007,
Collins et al. 2011, Bodin and Tengö 2012). By exploring
indigenous people’s relationships with caribou, which have been
actualized through language, we developed new insights into the
underlying processes that create structure and drive patterns of
caribou biodiversity. We contend that indigenous languages
provide an obvious place to ground research processes and build
collaborations. Words can be used to strengthen people’s
relationship with local ecosystems and create appropriate and
unifying dialogue. As Frederick Andrew affirmed, “The most
important thing is to talk the old language and honor our
ancestors that went before us.” As a direct outcome of our
research, the SRRB has made the decision to use the word tǫdzı
in all official correspondence relating to boreal woodland caribou.
The process of changing vocabulary has the potential to allow for
the development of common-ground from which new
relationships can move forward (Stevenson 1998). By recognizing
the validity of other knowledge systems it is possible to broaden
the worldview of the listener (Gavin et al. 2015). In doing so the
world “becomes richer as our ability improves to view it from a
variety of angles” (Cruikshank 1981:86).
Through the process of łeghágots'enetę “learning together” we
were able to embrace the synergies that come from the sometimes
intangible process of knowledge expansion and develop
comprehensive descriptions of caribou populations that reflect
biodiversity. Our results point to the importance of assessing
multiple criteria simultaneously when determining population
boundaries and characterizing population structure. We found
clear connections between Dene people’s descriptions of caribou
ecology and other domains of knowledge such as population
genetics where connectivity and delineation of groups are central
themes. Utilizing multiple methods has the potential to strengthen
evidence-based decisions with respect to range mapping as part
of the boreal woodland caribou range and action plans
(Environment Canada 2012) and environmental assessments in
response to potential shale-oil development in the region. At the
national scale, our results provide guidance on the delineation of
DUs for caribou across Canada and suggest practical approaches
toward the inclusion of TK in the development of policies related
to SARA.
Multidisciplinary research broadens the scope of biological
inquiry and recognizes the significant contribution that multiple
knowledge sources provide (Gavin et al. 2015). By exploring
multiple ways of organizing knowledge our research was able to
forge the basis for cross-cultural collaboration. For example, by
investing in cooperation from the onset, our project produced
results that have been acknowledged from different world views,
thus our research outcomes may be more broadly accepted. As
Walter Bayha pointed out, “The future of research in the north
will include more and more cases of science confirming the history
of aboriginal people and thus add to the overall knowledge that
has existed since time immemorial.” Likewise, as demonstrated
by our research, TK also has the potential to inform and improve
scientific methods, processes, and outcomes. Through
collaboration and łeghágots'enetę “learning together” our
research outlines ways to respectfully draw upon indigenous
knowledge and support relationships between people and wildlife.
By working with local communities, combining methods from
different disciplines, and establishing potential for transformative
dialogue, we can generate new insights and assist managers in
confronting the daunting conservation challenges of the future.
Responses to this article can be read online at:
http://www.ecologyandsociety.org/issues/responses.
php/8284
Acknowledgments:
We would like to express heartfelt gratitude to all the individuals
who participated in the project and our community partners, the
Ɂehdzo Got'ı̨nę (Renewable Resources Councils), who made this
work possible: specifically we thank Roger Boniface, Sarena
Kaskamin, Wilfred Jackson, Lawrence Manuel, Patricia Manuel,
Frank T'selele, Harry Harris, Roger Odgaard, Ricky Andrew, David
Menacho, Roderick Yallee, Valerie Yakeleya, Julie Lennie, Dion
Lennie, William Horassi, Joe Bernard, David Etchinelle, Rocky
Norwegian, Alfred Taniton, Leon Modeste, Morris Neyelle, Charlie
Neyelle, Sideny Tutcho, Mitchell Naedzo, Wilbert Kochon, and
Marie Kochon among many others. Special thanks go to the late
Angus Shae, Norm Hodgson, and Doug Urquhart whose knowledge
was profound and whose presence is notably missed. The research
advisory group was crucial to the work and we are indebted to them:
Michael Neyelle, Walter Bayha, Jimmy Dillon, Gordon Yakeleya,
Frederick Andrew, Leon Andrew, Maurice Mendo, Michel Lafferty,
Judy Lafferty, Richard Kochon, Hyacinthe Kochon, Gabe Kochon,
Lucy Jackson, and Camilla Rabisca. The SRRB board and staff
provided consistent encouragement as well as financial, logistical,
and administrative support for this project. Particular thanks are
owed to Catarina Owen, Lori Ann Lennie, Kristen Kodakin, and
Joe Hanlon. We thank ENR staff Richard Popko, Stephanie
Ecology and Society 21(2): 18
http://www.ecologyandsociety.org/vol21/iss2/art18/
Behrens, Heather Sayine-Crawford, Laurel McDonald, Keith
Hickling, Ron Doctor, Leroy Andre, Mabel Tatchinron, James
Hodson, and Jeff Walker who enabled this work. Máhsı cho to
Morris Modeste for enlarging our understanding of Dene language,
generous hospitality, and collecting samples. We thank all those who
helped collect samples especially Jeffery Jackson, who single
handedly submitted over 130 samples and Veronique Kochon, our
youngest caribou scat collector - máhsı! Terrence Mackeinzo,
Cheyenne Menacho, and high school students from the Mackenzie
Mountain School helped with lab work in Norman Wells. Thanks
to the staff of Nahanni National Park Reserve and Saoyú-Ɂehdacho
National Historic Site who collected and transported samples. Rick
Farnell, Susan Kutz, and Anja Carlsson generously provided caribou
samples from their research projects. Essential laboratory
assistance was provided by colleagues at Trent University. We
especially thank Marina Kerr, Jill Lalor, and Bridget Redquest. We
are grateful for discussions with colleagues that contributed to this
work including Iain Davidson-Hunt, Amy Flasko, Laura Hebert,
Sam McFarlane, Pauline Priadka, Paul Galpern, and Sandra
Marken. Jesse Tigner of Explor provided helicopter flight time.
General funding for this research was provided by the SRRB, ENR,
Cumulative Impact Monitoring Program, Environmental Studies
Research Fund, Parks Canada, University of Manitoba, and an
NSERC Strategic Grant held by MM and PW. JLP thanks Claire
Polfus, Morgan Moffitt, Jonaki Bhattacharyya, and Joe Hanlon
for reviewing early versions of the manuscript and the Wilburforce
Foundation for support through the Wilburforce Fellowship in
Conservation Science.
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Appendix 1. Community-collaborative research background.
In northern Canada, landscapes, people and wildlife are inextricably and compellingly
intertwined. Federal control over wildlife and the exclusion of indigenous people in wildlife
management and decision-making for much of the 19th century has generated environmental,
jurisdictional, and political conflicts revolving around natural resources (Sandlos 2011). For
many years science-based wildlife conservation approaches often had negative impacts on the
traditional harvesting practices of indigenous peoples. For example, concerns about declining
populations led to legislative controls by the federal government in the early 1900s, prohibiting
(and in some cases criminalizing) subsistence hunting of caribou and other large mammals and
birds by Dene and Inuit hunters in the Northwest Territories (Sandlos 2011). Understanding the
historical context of unequal power relations is an important part of developing new approaches
to environmental research (Fletcher 2003, Nadasdy 2005, McGregor et al. 2010, Tobias et al.
2013).
In the 1970s researchers working in the north began to acknowledge the importance of
indigenous people’s knowledge and priorities in natural resource research (Cruikshank 1981).
However, early traditional knowledge (TK) research focused mostly on collecting objective and
quantifiable information that could be packaged and accessed within scientific frameworks
(Stevenson 1998). This led to substantial misrepresentation and the appropriation of knowledge
(Nadasdy 2005, Castleden et al. 2012). A more recent shift in the orientation of research
advocates for collaborative processes that serve indigenous interests, provide ownership and
control of research outcomes, and include local people in decision-making processes (Hall 1979,
21:(2).
18.
2
Simpson 1999, Smith 1999, Simpson and Driben 2000). For example, community-based
participatory research frameworks emerged in response to disrespectful and exclusionary
approaches that concentrated research on people rather than with people (Simpson and Driben
2000, Fletcher 2003)
Participatory research (also including community-engaged, community-participatory,
community-based, collaborative, cooperative; Ferreira and Gendron 2011) is intended to include
people as active participants in all phases of the research process to “facilitate a more accurate
and authentic analysis of social reality” (Hall 1979) and have been adapted in the fields of
education (Hall 2005), public health (Christopher et al. 2011, Ferreira and Gendron 2011, Tobias
et al. 2013), social science (Fletcher 2003), resource management (McKinley et al. 2012), and
linguistics (Czaykowska-Higgins 2009) among others. The principles of participatory research
include fostering a co-learning environment, answering relevant community-driven questions,
focusing on co-capacity building and sustainable solutions, sharing decision-making
responsibilities, and above all reflecting critically on the roles and power relations of those
involved in the research process (Cornwall and Jewkes 1995, O'Fallon and Dearry 2002,
Davidson-Hunt and O'Flaherty 2007)
In alignment with the principals of participatory research we brought together an
interdisciplinary team of research partners and co-authors to build a solid foundation across
diverse fields. Our research process was iterative and built on information and questions
developed and refined over time. Significant knowledge exchange and gots'enet “learning
together” between the co-authors and research partners occurred as ideas for the project were
developed, at focus group meetings, during the selection of the field sampling sites, while
3
collecting samples, and on the land during day trips, hunting trips and overnight trips to cabins.
The distinct disciplinary backgrounds of team members, who spoke different first languages,
necessitated significant dialogue to come to common understanding for a project.
The commitment, interest and openness of community research partners in the Saht
region was crucial to the collaborative research process. Michael Neyelle, Walter Bayha,
Frederick Andrew, and Leon Andrew are all native Dene language speakers and have significant
TK experience and knowledge from their personal experiences and their parents and ehts
“grandparents”. They have worked in collaboration with the ehdzo Got' '
(Sahtú Renewable Resources Board; SRRB) and other non-Dene researchers on various research
projects over the years. Their interest in this research project, commitment to helping support the
research, guidance on TK practices, and help with interpretation of the language, and their
leadership positions within the communities allowed for new knowledge to be created and a
common understanding to be reached. A focus on language during the research process was a
means for Dene and non-Dene speakers to explore knowledge and understanding of the
environment in more depth. For example, TK holders were able to unearth older knowledge that
is not used every day. Non-Dene partners were able to explore the ways in which the words we
use and the ideas we express influence the collaborative environment.
The project also included extended place-based research by non-Dene partners (Jean
Polfus and Deborah Simmons live and work in the community of Tul t'a) that allowed for
opportunities to participate in activities on-the-land and in the communities (thus learning was
not restricted to research activities/agendas). Jean Polfus also traveled throughout the
communities in the Sahtú to provide support for the ehdzo Got' n (Renewable Resources
4
Councils), collect caribou fecal samples with community members, participate in hunting
activities, meet with students at local schools and Aurora Colleges and coordinate sampling
efforts. The understanding required to respond appropriately to cultural cues and respectfully
engage in gots'enet “learning together” is on-going, intangible and personal (for all
research partners and co-authors) – but this exploration provides the necessary foundation
needed to produce truly collaborative research.
Over time relationships were fostered that provided space for non-Dene researchers to
learn important lessons regarding hunting traditions, on-the-land safety, and Dene ekw
“caribou laws” required to demonstrate respect for the land and wildlife. Likewise, community
members were also able to benefit from the collaborative relationship through increased contact
with “outsider knowledge” (Caine et al. 2007), including expertise in wildlife biology,
population genetics and linguistics, the chance to be involved in long-term natural resource
management research and planning, and access to other resources that the non-Dene researchers
could more easily acquire. The union of knowledge traditions can only be achieved though
shared experiences, considerable time, and strong local and regional governance (McGregor et
al. 2010). A large amount of knowledge was gained over time and cannot be readily summarized
in a manuscript. The research process was organic and agreement on the interpretation of the
results was gradual, forcing everyone to explore their own knowledge in depth, and in some
cases leading to new questions and additional analysis.
Meeting the demands of academic requirements, funding agencies, and indigenous
communities in the same process is fundamentally challenging and levels of participation,
control and ownership of the research process and products often vary based on the complex
5
constraints on each project (Cornwall and Jewkes 1995, Simpson and Driben 2000, Tondu et al.
2014). Our research benefitted considerably from the partnership with the SRRB. The board is a
land-claim organization responsible for managing renewable resources. The SRRB’s
contributions to this project allowed our research to be firmly grounded in the communities
needs and questions from the onset of the research because the project was built on past-
experiences and related work. The SRRB also facilitated ongoing communication with the public
and local research partners by developing connections between various research agendas and
other co-occurring projects. The opportunity for long-term planning and stability in the research
process (implemented through connections with multiple community-driven projects and long
term institutional research strategies and programs) is an important contribution of collaborative
interdisciplinary research and the SRRB’s involvement was essential to the success of the long
term collaborative project. Thus, we were able to produce research contributions that were
deeper and more robust than could have been achieved by a single, stand-alone academic
research project.
6
Figure A1.1. We discussed traditional knowledge themes, language, and genetic data with a local
group of experts (advisory group, including co-authors) in two separate 3-day meetings to clarify
and develop important concepts and themes related to caribou populations in the Saht region
and Nahanni National Park Reserve of the Northwest Territories, Canada. Advisory group
members June 2014 in Tul t'a, Northwest Territories: Gordon Yakeleya, Frederick Andrew,
Michael Neyelle, Jean Polfus, Walter Bayha, Camilla Rabisca, Deborah Simmons, Michel
Lafferty, and Judy Lafferty.
7
Figure A1.2. Advisory group members February 2015 at Deochah (Bennett Field), Northwest
Territories: Back two rows – Jean Polfus, Gordon Yakeleya, Frederick Andrew, Richard
Kochon, Jimmy Dillon, Walter Bayha, Deborah Simmons, Leon Andrew, Nicole Beaudry
(ethnomusicologist), Michael Neyelle, and Lucy Jackson. Front row – Corrine Andrew (cook),
Gabe Kochon, Maurice Mendo, and Hyacinth Kochon.
8
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10
Tondu, J. M. E., A. M. Balasubramaniam, L. Chavarie, N. Gantner, J. A. Knopp, J. F.
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Appendix 2. Population genetic summary statistics for caribou genetic samples collected in the
Saht region and Nahanni National Park of the Northwest Territories, Canada from 2012 to
2014.
Table 1. Summary of genetic diversity estimates for each microsatellite loci including allelic size
range in base pairs, number of alleles (NA), observed (HO) and expected (HE) heterozygosity, FIS
estimates and standard errors (SE) for caribou genetic data from the Northwest Territories,
Canada. The original references for each loci are provided.
Microsatellite
Locus
Allelic
Range
NA HO SE HE SE FIS Source
BM848 356-386 16 0.819 0.0234 0.864 0.0150 0.053 Bishop et al. 1994
BM888 162-260 51 0.882 0.0041 0.865 0.0071 -0.019 Bishop et al. 1994
Map2C 89-115 16 0.824 0.0113 0.850 0.0232 0.030 Moore et al. 1992
RT5 88-116 15 0.768 0.0283 0.816 0.0373 0.058 McLoughlin et al. 2004†,
Wilson et al. 1997‡
RT6 88-112 14 0.837 0.0122 0.833 0.0245 -0.005 Wilson et al. 1997
RT7 210-232 12 0.746 0.0242 0.772 0.0109 0.033 Wilson et al. 1997
RT9 100-128 15 0.840 0.0257 0.859 0.0077 0.022 Wilson et al. 1997
RT24 205-227 24 0.780 0.0497 0.786 0.0707 0.008 Wilson et al. 1997
RT30
183
-
211
19
0.829
0.0130
0.862
0.0242
0.039
Wilson et al. 1997
†Reverse primer
‡Forward primer
Polfus, J. L., M. Manseau, D. Simmons, M. Neyelle, W. Bayha, F. Andrew, L. Andrew, C. F. C. Klütsch, K. Rice,
and P. Wilson. 2016. eghágots'enet (learning together): the importance of indigenous perspectives in the
identification of biological variation. Ecology and Society 21:(2).
Wilson. 2016. Ł
e
18
Table 2. Pairwise FST values based on microsatellites for the three major groups identified by
structure analysis (below diagonal) and pairwise P values (above diagonal).
FST Barren-
ground
Boreal
woodland Mountain
Barren-ground - 0.0000 0.0000
Boreal woodland 0.040 - 0.0000
Mountain 0.011 0.041 -
Table 3. Pairwise RST values based on microsatellites for the three major groups identified by
structure analysis (below diagonal) and pairwise P values (above diagonal).
RST Barren-
ground
Boreal
woodland
Mountain
Barren-ground - 0.0303 0.0028
Boreal woodland 0.030 - 0.0269
Mountain 0.003 0.027 -
Table 4. ST values based on mtDNA for the three major groups identified by structure
analysis (below diagonal) and pairwise P values (above diagonal).
ST Barren-
ground
Boreal
woodland Mountain
Barren-ground - 0.0000 0.0000
Boreal woodland 0.079 - 0.0000
Mountain 0.138 0.173 -
Table 5. Analysis of molecular variance (AMOVA) based on mtDNA haplotype data for the
three groups identified by structure analysis. FST represents the variance within groups relative to
the total variance.
Source of variation d.f. Variance
components % Variation F P
Among groups 2 0.63 13.9
Within groups 334 3.91 86.1 FST = 0.139 0.0000
Figure 1. Most likely number of population clusters (K = 2) identified by the Evanno method
(Evanno et al. 2005) using Structure Harvester v0.6.94 (Earl and vonHoldt 2012).
Figure 2. Mean likelihood for each K plus standard deviation as retrieved from Structure
Harvester v0.6.94 (Earl and vonHoldt 2012).
Figure 3. Frequency of mtDNA haplotypes for the three major groups identified by structure
analysis: 1) barren-ground (blue), 2) boreal woodland (green), and 3) mountain woodland (red).
Three haplotypes belong to the woodland haplogroup lineage: 50, 522 and 523, all other
haplotypes belong to the Beringian haplogroup lineage.
Figure 4. During a focus group meeting, Gabe Kochon of Fort Good Hope, Northwest
Territories, Canada, described a historic event where a large herd of caribou crossed the Dehcho
(Mackenzie River) and headed into the foothills of the mountains many years ago. We collected
samples from the area identified by the arrow during our 2 April 2014 helicopter survey and
identified 9 individual caribou from the site in subsequent genetic analysis. Structure analysis of
these samples found a high probability of assignment (average 0.73) to the barren-ground
caribou cluster (shown as blue in bottom bar and represented as blue on the map). Genetic
structure analysis identified k=3 clusters of 1) barren-ground (blue) 2) boreal woodland (green)
and 3) mountain (red).
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