Introduction
Human land uses and development are altering landscapes worldwide and
restricting the movements of wide-ranging species such as large
carnivores (Crooks et al. 2011; Ripple et al. 2014). One
outcome of the expanding human footprint is that carnivores and people
are increasingly overlapping with one another. Several tools have been
employed in human-dominated landscapes to mitigate carnivore
interactions with people, including fencing protected areas or other
barriers and policies to separate humans from wildlife (e.g., McInturff et al. 2020). Depending on the relative scale and permeability of
these physical structures, such efforts often result in new patterns of
wildlife movement and navigation (e.g., Wilkinson et al. 2021b).
On coexistence frontiers , where human infrastructure and activity
have either recently appeared or are rapidly increasing, carnivores must
learn to traverse novel landscapes with novel risks. In such regions, it
is critical to identify predictors of landscape navigation for
carnivores and determine whether and how these species can adaptively
traverse these increasingly human-dominated landscapes (Sanjayan &
Crooks 2005).
Understanding carnivore landscape navigation within coexistence
frontiers requires social-ecological approaches rather than only
considering ecological factors (Lute et al. 2020; O’Neal
Campbell, 2014). In general, wildlife sharing space with people must
navigate three main elements present on the landscape, namely ecological
factors, human infrastructure, and human acceptance, that determine social-ecological landscape permeability (e.g., Ghoddousi et al. 2021). This guiding framework is critical because while
certain carnivore species may be able to persist in human-altered
habitats (see Athreya et al. 2013; Breck et al. 2019;
Chapron et al. 2014; Devens et al. 2019), human
intolerance may be a strong enough limiting factor that it can override
adaptability for carnivore populations or survival of individuals
navigating developing landscapes (Moss, Alldredge & Pauli 2016). Thus,
human perceptions are likely to be an important factor in determining
how carnivores navigate landscapes (Behr et al. 2017). On
coexistence frontiers, people may not have developed tolerance or
acceptance for species which they have not encountered before or which
they are now encountering more frequently (e.g., Lute & Carter 2020).
In this case, intolerance may serve as a proxy for people employing
nonlethal hazing or deterrents or as an indicator for tendencies to
conduct illicit persecutory actions toward wildlife (Benson et
al. 2023; Ditmer et al. 2022; Manfredo et al. 2021), such
as poisoning (see Ogada 2014) or habitat destruction (see Ripple et al. 2014). In these social-ecological spaces, people also
interact with, respond to, and alter ecological features and processes
that influence carnivore movement at different scales, such as
vegetation availability (e.g., Bateman & Fleming 2012; Suraci et
al. 2020) and climatic season (e.g., through changing livestock
movements; Schuette et al. 2013).
Spatial and ecological scales are important for contextualizing human
coexistence with carnivores (Carter & Linnell 2016). For instance,
anthropogenic development may have community-level effects by pushing
some species into limited remaining natural habitats (Parsons et
al. 2018), yet at a fine scale, carnivores can exhibit adaptations to
anthropogenic development – or even be synanthropic—on an individual
level (i.e., Moss et al. 2016; Nisi et al. 2021; Wang et al. 2015; Suraci, Nickel & Wilmers 2020). Through these
scale-dependent processes, human-dominated landscapes near protected
areas can result in a source-sink dynamic, whereby carnivore populations
from protected areas that venture into more densely human-populated
regions are more likely to die through anthropogenic causes (Lamb et al. 2020). However, individuals may succeed in these spaces by
taking advantage of anthropogenic resources at finer scales, depending
on human tolerance (i.e., Moss et al. 2016). Thus, for large
carnivores, which are often highly mobile, social-ecological landscape
permeability across scales is essential to maintain populations.
Carnivore species living in both human-dominated environments and
protected environments may avoid anthropogenic features such as roads
and fences (e.g., Baker & Leberg 2018; McInturff et al. 2020;
Young et al. 2019) or change their activity patterns to adjust
for human presence (e.g., Gaynor et al. 2018). Yet carnivores may
also either avoid or be attracted to human infrastructure at different
scales (Poessel et al. 2014), underpinned by the density of the
infrastructure (Morales-Gonzalez et al. 2020; Xu et al .
2023), the infrastructure’s impact on resource availability (i.e.,
Belton et al. 2016), the species involved (Wilkinson et
al. 2021b), or individual animal characteristics, such as life stage
(Thorsen et al. 2022).
As a widely reviled (Glickman 1995; Macdonald et al. 2022) and
behaviorally plastic apex predator (Holekamp & Dloniak 2010), spotted
hyenas (Crocuta crocuta , hereafter “hyenas”) are a model
species for understanding the nature of carnivore adaptiveness to
human-caused landscape change and negative human perceptions. Hyenas are
generally considered one of the most behaviorally flexible carnivore
species, yet there have been few empirical conclusions regarding the
extent and mechanisms of their adaptiveness to human activities,
infrastructure, and tolerance. Green et al. (2018) found that hyena
populations in Maasai Mara, Kenya increased in an area of anthropogenic
disturbance, possibly linked to increased livestock consumption. In
densely populated areas in Ethiopia where native prey is depleted,
hyenas have become almost entirely dependent on anthropogenic food
(e.g., Yirga et al. 2012). However, other studies have found
negative, neutral, or nuanced responses to people. In one study in
Kenya, hyena activity shifted in response to livestock grazing and other
anthropogenic activities (Kolowski et al. 2007), while a study in
South Africa found that hyena propensity to visit anthropogenic sites
varied depending on season, age, or individual (Belton et al. 2018). Determining whether a species is surviving or thriving in the
presence of humans is complex, as even highly synanthropic species may
be exposed to greater levels of stress, toxicants, and disease while
living in anthropogenic landscapes (Murray et al. 2016; Murray et al. 2019). While movement is thus solely the broadest scale at
which to assess carnivore survival and wellbeing within coexistence
frontiers, understanding how hyenas and other behaviorally plastic
mammalian carnivores navigate anthropogenic landscape change is
fundamental to forecasting the resilience of movements, food webs, and
ecosystems in rapidly developing landscapes.
We sought to provide insight into spotted hyena abilities to navigate
coexistence frontiers by examining the following questions in and around
Lake Nakuru National Park and Soysambu Conservancy, Kenya: 1) How do
spotted hyenas navigate anthropogenic infrastructure, human activity,
tolerance, and ecological factors across scales, seasons, and management
types (i.e., fully protected vs. multi-use), and 2) How many spotted
hyenas cross through the region’s conservation fence, and how does
spotted hyena fence navigation manifest at a fine scale? We predicted
that (1) hyenas would respond differently to ecological factors than
they do to anthropogenic infrastructure and activity, (2) hyena
selection for and against landscape characteristics would differ at
different scales (i.e., fine-scale and landscape scale), (3) hyenas
would exhibit different responses to covariates in the rainy vs. the dry
season, (4) hyenas with dens in the fully protected national park would
be more avoidant of barriers, less avoidant of roads, and more attracted
to verified and perceived livestock predation hotspots outside of the
park than would hyenas with dens in the multi-use conservancy, and (5)
fence crossing would be limited to a few select individuals and hyenas
likely perceive fence navigation to be risky. We employed resource
selection functions (RSFs) and step-selection functions (SSFs) to
determine hyena space use and landscape navigation at the home range and
step scales, respectively (Reinking et al. 2019). We then used
this information to infer whether and how to consider a suite of
social-ecological factors when designing for hyena landscape
permeability, and present the implications of these inferences for
global human-carnivore coexistence.