Social-ecological landscape navigation and fine-scale
movement decisions
Hyenas in this region are selecting for different factors at the
landscape scale than at a fine scale. Differences in resource selection
at different scales were particularly apparent for infrastructure and
human experience characteristics, and less apparent for environmental
characteristics, selection for which largely remained the same across
RSF and SSF results. On a fine scale, hyena navigation appears to be
more heavily influenced by roads, fences, and human experiences and
acceptance than by environmental characteristics such as NDVI and
proximity to water. These scale-dependent affects likely stem from a
combination of factors, with fine-scale movements dictated by hyena
behavioral flexibility, and landscape scale selection dictated by
broader resource availability. While there are changes in the magnitude
of effect, at both broad and fine scales hyenas in this region generally
select for vegetation greenness, lakes, and roads, and against rivers,
boundaries, slopes, and high elevation. In arid environments, spotted
hyena hydration is mostly derived from prey rather than water sources
(Green et al. 1984), and hyenas can also survive for a week or
more without water (Holekamp & Dloniak 2010); thus, selection against
rivers could reflect the aridity of the region. Additionally, though the
two major lakes in the region are highly alkaline, hyenas have been
observed hunting flamingos, wallowing, and using the lakesides’ heavy
vegetation as refuge during the heat of the day (authors’ observations).
Thus, even undrinkable water sources can serve as critical habitat for
hyenas (e.g., Matsumoto-Oda 2021).
At the landscape scale, hyenas are selecting against participatory
mapped livestock predation areas and for areas of participatory mapped
risk, while the opposite holds true for fine scale selection. The
landscape-scale selection for and against these human acceptance
covariates may indicate that hyenas are broadly selecting for areas in
which they may face hazing or be poisoned (i.e., perceived risks from
and low tolerance for hyenas), and against areas that people use for
livestock grazing (i.e., participatory mapped livestock predation).
People are also likely perceiving the highest risks from hyenas in
places that constitute suitable hyena habitat, which supports hyena
landscape-scale selection for these areas. In densely populated areas
where tolerance is high or there are policies against wildlife killing,
carnivore populations may thrive (Athreya et al. 2013;
Gebresenbet et al. 2018), yet in areas where tolerance is low
(such as in our study site), carnivore populations can be negatively
affected by retaliation and other practices (Ripple et al. 2014).
Hyenas in this study site may also be avoiding participatory mapped
livestock predation areas—which are likely most often used for
grazing—as locations with consistent, predictable risks (i.e., through
deterrents and higher levels of human activities). This reflects
findings from previous studies in regions with less development, in
which spotted hyenas reduced their activities in response to livestock
grazing (i.e., in the Maasai Mara, Kolowski et al. 2007).
Our results also showed seasonal differences in landscape-scale and fine
scale hyena selection for environmental and anthropogenic
characteristics. At the landscape scale, in the dry season, hyenas
exhibited an increase in selection for vegetation greenness, boundaries,
and participatory mapped livestock predation. The latter two support our
findings that hyenas may be expanding their ranges and traveling outside
of the protected areas during the dry season. When resources are scarce,
animals living alongside people may be more likely to choose
anthropogenic food sources (Johnson et al. 2015), and hyena
predation on livestock or scavenging of livestock and other
anthropogenic food sources may be increasing during dry seasons. The
boundary navigation result at the landscape level also supports these
hyenas’ tendency toward dry season range expansion. Meanwhile, at a fine
scale, the dry season showed a minor increase in hyena navigation toward
roads, with minimal seasonal differences in selection for environmental
and human experience characteristics. While species in other studies
have been known to use roads for easier traversal of the landscape
(Abrahms et al. 2015; Hill et al. 2020), hyenas in this
study area may also use roads in the dry season for dust bathing and
access to artificial water points, particularly in the conservancy.
When looking at variation in movement for hyenas living in different
management types across both seasons, at the landscape scale roads were
more likely to be avoided by Soysambu hyenas than LNNP hyenas, despite
human use of roads being extremely low in the conservancy at night. One
reason for this could be that despite being active at night, hyenas in
Soysambu associate roads with the abundance of human activities that
occur during the day on the roads within the conservancy, while in LNNP,
there is only one activity happening on roads: tourism. Vehicle speeds
in the national park are also heavily regulated. Hyena avoidance of
roads in Soysambu stands in contrast to research that has found that
animals may select for human infrastructure at night for resources or
ease of movement while avoiding it during the day when humans are more
active (e.g., Toverud 2019). However, hyenas are generally more skittish
in the conservancy than in the national park, possibly due to historical
hyena bounties (K. Combes, pers. comm.) before the conservancy’s
designation as a wildlife habitat. The anthropogenic activity signature
on Soysambu’s roads may thus have a strong enough effect on the spotted
hyena’s human-caused “landscape of fear” (Smith et al. 2017;
Suraci et al. 2019) as to contribute to nighttime avoidance of
roads that are devoid of human activity.
SSFs revealed that hyenas with dens in LNNP are also selecting against
boundaries to a much stronger degree than hyenas with dens in Soysambu.
However, fine scale selection against vegetation greenness and toward
verified livestock predation locations outside of the park, as well as
known fence-crossing behaviors by LNNP hyenas (Wilkinson et al. 2021b), point to a lack of sufficient resources or space in the national
park.