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University of Southampton
16th August 2016
FACULTY OF NATURAL AND ENVIRONMENTAL SCIENCES
CENTRE FOR BIOLOGICAL SCIENCES
RESOURCE PARTITIONING BETWEEN SPOTTED
HYENAS (CROCUTA CROCUTA) AND LIONS
(PANTHERA LEO)
Arjun Dheer
A dissertation submitted in partial fulfilment of the requirements for the degree of
M.Res. Wildlife Conservation.
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As the nominated University supervisor of this M.Res. project by Arjun Dheer, I confirm that I
have had the opportunity to comment on earlier drafts of the report prior to submission of the
dissertation for consideration of the award of M.Res. Wildlife Conservation.
Signed…………………………………..
UoS Supervisor’s name: Prof. C. Patrick Doncaster
As the nominated Marwell Wildlife supervisor of this M.Res. project by Arjun Dheer, I confirm
that I have had the opportunity to comment on earlier drafts of the report prior to submission of
the dissertation for consideration of the award of M.Res. Wildlife Conservation.
Signed……………………………………
MW Supervisor’s name: Dr. Zeke Davidson
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Abstract
The negative impact of anthropogenic activities on wildlife has led to protected areas being set
aside to prevent human-wildlife conflict. These protected areas are often small and fenced in
order to meet the needs of expanding human communities and to conserve wildlife. This creates
challenges for the management of wide-ranging animals such as large carnivores, especially
those that compete with one another for limited resources. This study focused on resource
partitioning between GPS-GSM collared spotted hyenas (hereafter referred to as hyenas) and
lions in Lewa Wildlife Conservancy (LWC) and Borana Conservancy (BC), Kenya. Scat
analysis revealed that hyenas and lions show a high degree of dietary overlap, though hyenas
have broader diets and feed on livestock species, which lions completely avoid. Spatially, hyenas
show stronger intraspecific avoidance and more exclusive territorial behaviour than do lions.
Hyenas and lions have a high degree of spatial overlap, though lions may influence den site
selection in hyenas. Both species are heavily nocturnal and crepuscular, though hyenas tend to
travel significantly further at night than do lions. Hyenas and lions both display mixed results in
dynamic spatiotemporal interactions, with 40% of hyena-lion pairs showing attraction and 70%
showing simultaneous use of overlapping areas. All but one inter-clan hyena pairs show strong
avoidance, though lion pairs were not as mutually repulsive. This shows that hyenas and lions
may use one another as sources of food and that scavenging and kleptoparasitism likely play a
role in their dynamic. The hyena population is suggested to be growing and healthy, though the
lion population is of concern due to lower density, isolation, and low recruitment. Further
investigation into human-carnivore conflict within surrounding communities, long-term
demographic and behavioural trends of all members of the large carnivore guild, and the
potential development of a dispersal-based metapopulation management scheme will allow for
the continued persistence of large carnivores in the Lewa-Borana Landscape (LBL) and their
coexistence with human communities.
Target journal: African Journal of Ecology
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Table of Contents
1. Introduction ........................................................................................................................... 9
1.1 Background ............................................................................................................................ 9
1.2 The value of large carnivores ................................................................................................ 9
1.3 Resource partitioning between carnivores ........................................................................... 10
1.4 Carnivores in small reserves ................................................................................................ 11
1.5 LWC and BC ....................................................................................................................... 11
2. Aims and objectives ............................................................................................................ 13
3. Materials and methods ......................................................................................................... 14
3.1 Study area ............................................................................................................................ 14
3.1.1 Location ........................................................................................................................ 14
3.1.2 Climate and biodiversity ............................................................................................... 14
3.1.3 Communities and human influence .............................................................................. 15
3.2 Data collection ..................................................................................................................... 18
3.3 Capture and collaring........................................................................................................... 18
3.3.1 Hyenas .......................................................................................................................... 18
3.3.2 Lions ............................................................................................................................. 21
3.4 Dietary partitioning ............................................................................................................. 22
3.4.1 Prey selection ................................................................................................................ 23
3.4.2 Dietary overlap ............................................................................................................. 23
3.4.3 Niche breadth ................................................................................................................ 24
3.5 Spatial partitioning .............................................................................................................. 24
3.5.1 Overlap ......................................................................................................................... 25
3.5.2 Den activity .................................................................................................................. 25
3.5.3 Den buffers ................................................................................................................... 26
3.5.4 Community land use ..................................................................................................... 26
3.6 Temporal partitioning .......................................................................................................... 26
3.6.1 Activity budgets ............................................................................................................ 26
3.6.2 Dynamic analysis .......................................................................................................... 27
3.7 Demographics ...................................................................................................................... 28
3.7.1 Hyenas .......................................................................................................................... 28
3.7.2 Lions ............................................................................................................................. 29
4. Results ................................................................................................................................. 30
4.1 Dietary partitioning ............................................................................................................. 30
4.1.1 Raw proportions ........................................................................................................... 30
4.1.2 Relative proportional contribution................................................................................ 30
4.1.3 Jacobs’ index ................................................................................................................ 32
4.1.4 Pianka’s index .............................................................................................................. 33
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4.1.5 Levins’ index ................................................................................................................ 33
4.2 Spatial partitioning .............................................................................................................. 34
4.2.1 Home range and core sizes ........................................................................................... 34
4.2.2 Overlap ......................................................................................................................... 37
4.2.3 Den activity .................................................................................................................. 41
4.2.4 Den buffers ................................................................................................................... 42
4.2.5 Community land use ..................................................................................................... 45
4.3 Temporal partitioning .......................................................................................................... 46
4.3.1 Activity budgets ............................................................................................................ 46
4.3.2 Dynamic analysis .......................................................................................................... 48
4.4 Demographics ...................................................................................................................... 52
4.4.1 Identified individuals .................................................................................................... 52
5. Discussion ............................................................................................................................ 54
5.1 Collaring .............................................................................................................................. 54
5.2 Dietary partitioning ............................................................................................................. 55
5.3 Spatial partitioning .............................................................................................................. 56
5.4 Temporal partitioning .......................................................................................................... 57
5.5 Demographics ...................................................................................................................... 58
5.6 Conclusions and management recommendations ................................................................ 59
Acknowledgments ................................................................................................................... 62
References ............................................................................................................................... 64
Appendix A ............................................................................................................................. 74
Appendix B .............................................................................................................................. 75
Appendix C .............................................................................................................................. 76
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List of figures
Figure 3.1 Large scale map of LBL within Kenya………………………………………………..16
Figure 3.2 Small scale map of LBL and points of interest………………………………………...17
Figure 3.3 Leg-hold snare trap set for hyenas…………………………..………….......................19
Figure 3.4 CH01 recovering post-immobilisation………………………………………………..20
Figure 3.5 DL01, the male lion in this study……………………………………………………...21
Figure 3.6 Examples of hyena and lion scats in the field………………………………………….23
Figure 3.7 Example of unique left and right spot patterns on BH01……………………………....29
Figure 4.1 Relative proportional contribution of prey items to hyena and lion diets……………...31
Figure 4.2 Jacobs’ index values………………………………………………………….……….33
Figure 4.3 Mean home range and core sizes……………………………………………………...34
Figure 4.4 Map of hyena and lion cores…………………………………………………..…........35
Figure 4.5 Map of hyena and lion home ranges………………………………………..………....36
Figure 4.6 Map of hyena home ranges and cores based on clan membership………………...…...39
Figure 4.7 Map of lion home ranges and cores based on pride membership……………………...40
Figure 4.8 Den distances to intraspecific and interspecific home ranges and cores…………........41
Figure 4.9 Den buffers in relation to hyena home ranges and cores…………………...…….........43
Figure 4.10 Den buffers in relation to lion home ranges and cores………………………...……...44
Figure 4.11 Community use by hyenas and lions……………………………………………..…..45
Figure 4.12 Proportional hyena and lion activity budgets by time windows…………………...…46
Figure 4.13 24-hour chart of hyena and lion mean distance travelled…………………..…….......47
Appendix A Trapping sites for hyenas and lions…………………………………………….…...74
Appendix C Collar deployment progression………………………………………….…….........76
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List of tables
Table 3.1 Collared hyenas by sex and clan membership………………………………………...22
Table 3.2 Collared lions by sex and pride membership…………...…………………………......22
Table 4.1 Raw frequencies of prey hairs in hyena and lion scats……………………………......30
Table 4.2 Relative proportional contribution……………………………………………….........31
Table 4.3 Jacobs’ selection index and standard error values……………………………….........32
Table 4.4 Levins’ index of niche breadth……………………………...………………………...33
Table 4.5 Average percent overlap based on intraspecific and interspecific relationships……...37
Table 4.6 Mann-Whitney U test results for overlap……………………………………………..38
Table 4.7 Average minimum distances based on den activity…………………………………...41
Table 4.8 Frequency of occurrence of hyena den buffers to cores and home ranges…………....42
Table 4.9 Average distances travelled at four separate time windows……………...…………...46
Table 4.10 Doncaster’s test results for hyena-lion pairs…………………………………………49
Table 4.11 Doncaster’s test results for hyena pairs……………………………………………...50
Table 4.12 Doncaster’s test result for a lion pair…………………………………………...........50
Table 4.13 Minta’s test result for interspecific and intraspecific pairs…………………………..51
Table 4.14 Demographics of identified individuals by sex and age class……………………….53
Table B.1 Sample collaring data sheet………………………………………...………………....75
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List of abbreviations
BC Borana Conservancy
CSV Comma-Separated Values
GIS Geographic Information System
GPS Global Positioning System
GR Game Reserve
IUCN International Union for Conservation of Nature
KWS Kenya Wildlife Service
LWC Lewa Wildlife Conservancy
NRT Northern Rangelands Trust
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1. Introduction
1.1 Background
Global expansion of human populations has led to wildlife populations becoming
increasingly confined to protected areas due to habitat loss. Loss of habitat in turn has resulted in
significant biodiversity loss and increased extinction rates across taxa (Brooks et al. 2002). Over
the past 500 years, it is estimated that human-driven extinction rates exceed those in the fossil
record several hundred times (Dirzo and Raven 2003). Biodiversity is a driver of ecosystem
change, resulting in reduction of natural resources available for human use. Loss of biodiversity
in turn limits ecosystem services and function and results in less productive ecological
communities (Cardinale et al. 2012).
In response, protected areas are set aside in the face of burgeoning human populations for
conservation purposes and to meet the needs of expanding human communities. There is an
increasing trend worldwide for these protected areas to be fenced to limit human-wildlife conflict
(Packer et al. 2013). However, meta-analyses of protected areas reveal that relying only on large
protected areas can have shortcomings in their effort to conserve biodiversity due to lack of
funding, research on their wildlife populations, and administrative disorganisation (Mora and
Sale 2011). A more holistic approach that includes connected smaller reserves with human
cohabitation and direct oversight and management is a more plausible scenario for the future
(Wackernagel et al. 2002).
This presents challenges for the management of wide-ranging animals that inherently
exist at low densities, such as large carnivores, especially in these small reserves close to human
habitation (Treves and Karanth 2003). Increased close management of such protected areas that
can support viable populations of these predators appears to be the most viable conservation
strategy for the future (Davies-Mostert et al. 2015) and can allow for the coexistence of human
and carnivore populations.
1.2 The value of large carnivores
Apex predators exert top-down effects on lower trophic levels and allow for the proper
functioning of natural systems. They regulate prey populations and in turn promote healthy
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biomass of producers, which form the basis of a food web (Miller et al. 2001). The presence of
large carnivores is an indicator of the ecological integrity of an ecosystem (Schmitz et al. 2000).
They serve as conservation umbrellas for species occupying lower trophic levels and provide
economic benefits due to their appeal to tourists (Linnell et al. 2000). While controversial,
regulated trophy hunting of large carnivores in southern Africa has also provided economic
benefits for local economies, including conservation charities (Lindsey et al. 2006).
However, large carnivore populations worldwide have declined as human populations
have grown (Weber and Rabinowitz 1996). As they occupy the top trophic levels of their
ecosystems, they are particularly sensitive to anthropogenic influences (Woodroffe and Ginsberg
1998). Understanding the ecological needs of large carnivores is therefore necessary to apply
appropriate management policies for the benefit of protected areas. Behavioural studies,
including analyses of diet and spatiotemporal activity, as well as demographic studies for
population management, allow for the development of such policies. East Africa remains one of
the last strongholds for large carnivores left on Earth, though there is a general population
decline in all species of large carnivore in the region (Packer et al. 2013, Schuette et al. 2013).
1.3 Resource partitioning between carnivores
Research on large carnivore ecology in East Africa has been ongoing since the middle
et al. 2008). Understanding competition and niche separation between century (Dalerum th20
carnivore species is needed to determine the roles that different members of the guild have. This
in turn will allow for management decisions to be made at the species level, especially when
study species are of conservation concern (Kamler et al. 2015). Carnivores that occupy similar
niches coexist through resource partitioning (Kamler et al. 2012, Vieira and Port 2006). For
example, in South Africa, bat-eared foxes, cape foxes, and black-backed jackals display dietary
and spatiotemporal resource partitioning (Kamler et al. 2012). In Hwange National Park,
Zimbabwe, hyenas altered their foraging behaviour and prey selection in response to an increase
in lion population (Periquet et al. 2015). In northern Botswana, cheetahs and African wild dogs
display temporal and spatial avoidance of hyenas and lions, which is hypothesised to facilitate
coexistence (Cozzi et al. 2012). In Matusadona National Park, Zimbabwe, lions and hyenas again
displayed dietary partitioning (Purchase 2004). A previous literature analysis reveals that hyenas
generally have a larger dietary niche breadth than lions, which is unsurprising, considering their
opportunistic feeding strategy (Periquet et al. 2014).
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1.4 Carnivores in small reserves
The behavioural ecology of large carnivores with limited ranging ability living in
proximity to humans is poorly understood (Yirga et al. 2012). The studies listed above have
taken place in large, unfenced areas and provide information on carnivore ecology within more
pristine habitats but do not address the pressure that humans are applying to wildlife today.
Given the trend of human population growth and habitat destruction, humans and carnivores are
increasingly coming into conflict (Treves and Karanth 2003). Small reserve management with
respect to large carnivores is therefore moving to the forefront of wildlife conservation. By
understanding the behaviour of carnivores in such areas, policies can be implemented to curb
conflict and allow for improved coexistence in the future as human populations expand. Such
policies can include metapopulation management, culling, reintroductions, and translocations
(Akçakaya et al. 2006).
1.5 LWC and BC
Lewa Wildlife Conservancy (LWC) and Borana Conservancy (BC) in Kenya serve as
exemplars of small, fenced, managed protected areas. At LWC and BC, lions and spotted hyenas
are the two dominant predators in terms of numerical abundance and behavioural interactions.
Research on lions in LBL has been ongoing for several years. However, there is a major gap in
knowledge on the influence that hyenas have on prey and competitors alike in the study area.
Hyenas display remarkable behavioural plasticity over much of their range; they can adopt a
variety of feeding strategies that range from almost exclusive scavenging, as in peri-urban areas
in northern Ethiopia (Yirga et al. 2012) to hunting up to 95% of the time, as in the Ngorongoro
Crater, Tanzania (Kruuk 1972). Furthermore, while the presence of the fission-fusion clan
system is ubiquitous across their range in sub-Saharan Africa, clan size can vary from under 10
in the Kalahari (Mills 1989) to over 70 in the prey-rich grasslands of East Africa (Kruuk 1972).
Territoriality shows a similar pattern: loosely defined territories in the Kalahari and Namib
while territory sizes in the Ngorongoro Crater are generally 2deserts average over 1,000 km
hyenas are the –s contributed to their success (Gittleman 1989). This flexibility ha2 below 40 km
most abundant large carnivores in Africa today despite being heavily persecuted and lacking the
charisma, scientific research interest, and public appeal of canids and felids (Glickman 1995).
Preliminary estimates have shown that the hyena population is between 38-78 individuals
at LWC, while the lion population is known to be 17. Lions are therefore under carrying capacity
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(28) through all four abundance measures used in a recent demographic study (Preston 2015)
whereas hyenas may be past the estimated threshold (50). In neighbouring BC, carrying
capacities for lions and hyenas are estimated at 22 and 36, respectively, while the lion population
is 17 and estimated at 31-57 for hyenas (Preston 2015). Ongoing lion monitoring at the
conservancies has revealed that the lion population between the two conservancies is 34, but the
actual number of hyenas remains unknown. Hyena dietary and spatiotemporal tendencies are
also unknown.
This study addresses the lack of knowledge on large carnivore behavioural ecology and
resource partitioning in small, fenced reserves. It also gathers foundational information on hyena
demography and space use in LBL. Finally, it will provide management recommendations for
large carnivores in small, fenced reserves in close proximity to human communities and evaluate
the translatability of such recommendations to a wider audience. In turn, these recommendations
will allow for improved policy making and mitigation of human-carnivore conflict in East Africa
and beyond.
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2. Aims and objectives
The aim of this project is to better understand the behavioural ecology of hyenas and lions in the
Lewa-Borana Landscape (LBL) and to determine the role of small reserves in large carnivore
conservation. The major outcome is to inform management decisions and policies at LWC and
BC that develop new solutions to conservation challenges in small, fenced reserves. This
overarching aim is met through an investigation of resource partitioning between hyenas and
lions at dietary, spatial, and temporal levels and data collection on their demography.
At the dietary level, the objective is to determine prey selection in hyenas and lions, dietary
overlap between the species, and niche breadth for both species. A compilation of dietary studies
across several countries revealed a 58.6% actual prey species overlap and a 68.8% preferred prey
species overlap between the two (Hayward 2006). This is exceptionally high. It is unusual that
two apex predators can occupy such similar niches and still coexist. This study will explore
whether there is similar extensive dietary overlap or some degree of partitioning.
At the spatial level, the objective is to investigate geographic segregation between hyenas and
lions in the study area at the home range and core levels and determine how lion presence affects
denning behaviour in spotted hyenas. Denning behaviour in hyenas is affected by lion proximity
and presence in Kenya and Zimbabwe (Kolowski and Holekamp 2009, Periquet et al. 2014).
Temporally, both species are considered nocturnal (Kiffner et al. 2008). This project will
determine if there is partitioning in the activity times of spotted hyenas and lions. If there is, it
may explain their ability to coexist while competing for limited resources (Hayward and
Hayward, 2006). Previous studies have demonstrated temporal avoidance amongst large
carnivores, especially subordinate species within the large carnivore guild such as African wild
dogs and cheetahs (Cozzi et al. 2012).
Foundational knowledge on hyena and lion demography within the study area will be gathered.
This information will be gleaned as part of the above three objectives. Furthermore, this
information will set the groundwork for future studies on this species in the area and further
demographic research on the large carnivore guild.
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3. Materials and methods
3.1 Study area
3.1.1 Location
This study was carried out in Lewa Wildlife Conservancy (LWC) and Borana Conservancy (BC)
in central Kenya, straddling the border of Meru and Laikipia counties (00 06′ to 00 17′ North,
370 21′ to 370 32′ East) (Figure 3.1). The total area of this site is 375 km2 (LWC = 250 km2; BC
= 125 km2). The conservancies have fences along their external boundaries, but feature
migratory gaps to allow for animal movement in and out of the conservancies (Dupuis-
Desormeaux et al. 2015).
3.1.2 Climate and biodiversity
The climate in LBL is classified as a tropical savanna climate (Aw) under the Koppen Climate
Classification system (Peel et al. 2007). LBL experiences two wet seasons, the long rains in
March to May, and the short rains from October to December, although periods of drought are
relatively common (Lewa Wildlife Conservancy 2014).
Four main habitat types occur in LWC and BC. Based on the dominant plant species, they
include forest (dominated by Juniperus-Olea forest), plains (dominated by Pennisetum grasses
and Acacia trees), hills and rocky outcrops (dominated by Acacia, Commiphora and Grewia),
and riverine habitat (Mwololo 2011). LBL is situated between montane forests to the south that
give rise to Mount Kenya and lowland grasslands to the north.
Despite its small geographic area, from a conservation perspective, LBL is home to strategically
important populations of threatened species such as Grevy’s zebra (Equus grevyi), black
rhinoceros (Diceros bicornis), and white rhinoceros (Ceratotherium simum) (Low et al. 2009).
Apart from dense herbivore populations, LWC and BC contain the full suite of species from the
East African large carnivore guild: lions (Panthera leo), spotted hyenas (Crocuta crocuta),
leopards (Panthera pardus), cheetahs (Acinonyx jubatus), striped hyenas (Hyaena hyaena), and
African wild dogs (Lycaon pictus).
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3.1.3 Communities and human influence
Human communities reside permanently both in and around LBL (Figure 3.2) and therefore
come into close contact with wildlife. Intensive agriculture and pastoralism are primary sources
of income for local communities (Nyaligu and Weeks 2013). Herds of cattle, sheep, and goat are
grazed in controlled plots within the conservancies and graze freely within community lands.
This controlled grazing within the conservancies is carried out to control low-nutrition grass
species such as Pennisetum stramineum and P. mezianum (Lewa Wildlife Conservancy 2006)
and in turn benefit grazing herbivores with improved grassland biodiversity. Several
photographic tourism lodges also operate within LWC and BC and are a primary source of
employment and economic activity in the area (Gaitho 2014).
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Figure 3.1 – Location of LBL within Kenya.
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Figure 3.2 – Map of LBL with points of interest.
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3.2 Data collection
The bulk of data collection took place between February and May 2016, but scat data and prey
count data from 2014, 2015, and 2016 were used to assess long-term prey selection. Dietary
separation was determined using scat analysis based on a reference collection of hairs
(Mukherjee et al. 1994). Dietary partitioning focused on prey species relative contribution, prey
selection, dietary overlap, and niche breadth.
Spatial separation was determined through point and area overlap of core and home ranges for
the two species as well as den site selection by hyenas (Kamler et al. 2013). Data from collars
were downloaded as CSV (Comma-Separated Values) files and exported to ArcMap 10.4 for
analysis.
Temporal separation was determined through a comparison of activity budgets and analysis of
concurrent spatial avoidance between sympatric and non-sympatric pairs (Kitchen et al. 1999,
Doncaster 1990, Minta 1992).
Demographic data were collected through camera trapping at known hyena den sites, baited
traps, and opportunistic photographing of encountered individuals.
The confidence interval considered for analyses was 95%; therefore, p<0.05 demonstrated
significance. All statistical analyses were performed in R statistical software (R Core Team
2014) and ArcMap 10.4 (ESRI 2014).
3.3 Capture and collaring
3.3.1 Hyenas
A total of seven adult hyenas were collared with GPS-GSM collars within the study area,
representing five distinct clans (Table 3.1). Collars were manufactured by Savannah Tracking, a
Kenya-based telemetry and radio-tracking company.
Adults were targeted because hyenas’ necks grow throughout their lives; collaring a young hyena
can put the animal in discomfort as it grows. These collars were deployed between 23/02/2016
and 27/03/2016 (Appendix C).
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Free darting of hyenas at LBL was not possible due to their skittish behaviour. Hyenas were
captured using leg snares (n=1; Figure 3.3) and cage traps (n=6; Figure 3.4) with bait. Cages
were built by LWC’s workshop in advance of the trapping effort and were made of steel mesh,
measuring 3 m x 1.5 m x 1 m. Once baited and set, cage floors were padded with plywood
boards and thick, dead grass. Cages were also kept under bushes or trees to provide shade in the
event of a hyena being successfully trapped. Bait was purchased from local markets or acquired
opportunistically from carcasses. Prey species used for bait included cattle, goat, camel, eland,
and sheep. Bait was dragged roughly 100 meters from intended capture sites to form scent trails.
As hyenas are olfactory foragers (Woodmansee et al. 1991), this technique proved to be
essential. Trapping sites were either within 500 m of communal dens or in areas where rangers
frequently sighted hyenas. As hyenas were collared, the collaring effort was expanded outwards
to increase the probability of collaring conspecifics from different clans (Appendix A).
Figure 3.3 – Leg-hold snare trap for hyenas.
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In accordance with Kenya Wildlife Service (KWS) policy, veterinarians were present to dart and
treat the study animals. Hyenas were immobilized with a combination of ketamine hydrochloride
and medetomidine hydrochloride, collared, and checked for body condition and dimensions
(Appendix B). They were then reversed with atipamezole. Hyenas were recumbent in an average
of 12.1 minutes (range: 8.0-30.0) and recovered in an average of 17.5 minutes (range: 6.0-30.0).
One mortality occurred during the study period. SH01’s collar was found along with her almost
entirely consumed carcass near the western boundary of her clan’s territory. The thoroughness of
her consumption and the presence of three large hyena scats and hyena spoor in the vicinity of
her carcass make it likely that she was killed and cannibalised by conspecifics from the
neighbouring Borana clan. As she had previously been re-collared, a significant amount of
potential data was lost, resulting in there being less than 14 days’ worth of data from her collar.
Her collar was re-deployed on SH02, a clan member whose data was used in this study.
Therefore, SH01’s data have been excluded from all analyses in this report.
Figure 3.4 – CH01 recovers after immobilisation and collaring following capture with a baited cage trap in LWC.
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3.3.2 Lions
Long-term information on lion ecology has been collected by the research department at LWC
for several years and lions have been the focal species for previous Master’s-level projects at the
field site. They are also a primary tourist target at LWC and BC. Accordingly, lions in the area
are considered well-habituated. Free-darting was therefore used.
A total of three GPS-GSM collars were deployed on adult lions within the study area,
representing three distinct prides (Table 3.2). These collars were deployed between 26/02/2016
and 29/04/2016 (Appendix C).
Lions were recumbent in an average of 8.7 minutes (range: 11.0-15.0) and recovered in an
average of 40.3 minutes (range: 12.0–66.0). No mortalities occurred for collared lions during the
study period. However, one lion collar, deployed on SL01, stopped working on 03/05/3016 and a
new collar was deployed on the same lioness on 18/05/2016. Analysis of her data includes
information before and after her re-collaring but excludes the hiatus period.
Figure 3.5 – DL01, the male lion in this study, with unique right and left whisker spot patterns.
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Lion ID Sex Pride
DL01 Male Dalma
SL01 Female Sarah
WL01 Female Western
3.4 Dietary partitioning
Determination of dietary separation was based on scat analysis (Mukherjee et al. 1994). Scats were
identified (Figure 3.6) and collected in the field at kill sites, rest sites, hyena latrines, and
opportunistically. One scat was collected per site to prevent pseudo-replication from occurring.
Scat was placed in a plastic bag, dried, and treated with a mixture of 150 ml 75% ethanol solution
and 350 ml boiled water. 20 hairs were extracted per scat, mounted, and dried overnight. Scats
were then examined under microscope using analysis of hair roots. Each hair was compared to
reference collections present at LWC for species identification. Frequencies of different prey items
were recorded and tabulated for each scat sample. 118 hyena scats and 87 lion scats were used for
this analysis.
Hyena ID Sex Clan
BH01 Male Borana
CH01 Male Charlie
NH01 Female Nala
NH02 Male Nala
SH02 Female Shamba
UH01 Male Utalii
SH01 Female Shamba
Table 3.1 – Collared hyenas by sex and clan membership.
Table 3.2 – Collared lions by sex and pride membership.
23
Raw and relative (# F.O. species n / # occurrences for all species) contributions were calculated
for further analysis of selection, overlap, and breadth. Relative contribution of prey species
standardises the data to add up to 100% and is useful for determining how much a given prey
item contributes to a predator’s diet (Lyngdoh et al. 2014).
3.4.1 Prey selection
Jacobs’ selection index (Jacobs 1974) was used to assess prey selection:
r is the proportion of kills or scat samples of a given prey species;
p is the proportional availability of the given species.
The value can range from -1.0 to 1.0 depending on the degree of avoidance or selection,
respectively. This method uses abundance data on annual game counts from the LWC Research
Department from years 2014, 2015, and 2016 for scats from these respective years. Livestock
counts were not available so these have been excluded from the analysis.
3.4.2 Dietary overlap
Pianka’s index (Pianka 1974) was used to assess dietary overlap:
Figure 3.6 – Examples of hyena (L) and lion (R) scats identified in the field. The white coloration, chalky texture, and bulky size of hyena scat is diagnostic.
24
Pij is the relative proportion of prey species i in predator species j scat samples;
Pij is the relative proportion of prey species i in predator species k scat samples.
This method results in a standardized output from 0.0 (no overlap) to 1.0 (complete overlap).
3.4.3 Niche breadth
Levins’ measure of niche breadth (MacArthur and Levins 1967, Wallace Jr 1981) was used to
determine the diversity of their diets:
Y is the total number of individuals sampled;
Nj is the number of individuals using prey species j.
The raw Levins’ measure was then further standardised to a 0.0-1.0 scale:
n is the number of possible prey species.
3.5 Spatial partitioning
Analysis of spatial segregation included multiple spatial and mapping techniques to compare
geographic behaviour in the species. All analyses were completed in ArcMap 10.4. Areas of
25
interest, such as hyena dens, were marked via a GPS device and uploaded to ArcMap base layer
maps.
Collars were set to collect one fix every hour over a 24-hour period and report every six hours at
03:00, 09:00, 15:00, and 21:00. A total of 15,912 hyena fixes (range: 2,460–3,025) and 7,041
lion fixes (range: 1,619–3,227) were collected and used for analysis during the study period.
Overall, deployed GPS-GSM collars reported fixes 92.1% of the time (range: 79.0%-97.0%).
Hyena collars averaged 90.4% report success whereas lion collars averaged 96.2% report
success.
3.5.1 Overlap
Home ranges and cores were mapped out for the different species. Cores use the 50% Kernel
Density Estimate (KDE) method and home ranges use the 95% KDE. Home ranges and cores
were determined on ArcMap using the Home Range Tools extension (Rodgers et al. 2015) and
sizes (km2) were compared for the two species using the Mann-Whitney U test. They were then
analysed as follows:
(i) Core point and area overlap percentage;
(ii) Home range point and area overlap percentage;
(iii) Core-home range area overlap percentage.
These overlap percentages for interspecific competitors were further compared to intraspecific
overlap between hyena pairs and lion pairs from different social groups – five hyena clans and
three lion prides using the Mann-Whitney U test:
n1 is sample size 1;
n2 is sample size 2;
R1 is the rank of the sample size.
3.5.2 Den activity
26
Active vs. inactive minimum den distances were compared to one another using the Mann-
Whitney U test for lion cores and home ranges, inter-clan hyena cores and home ranges, and
intra-clan hyena cores and home ranges. This was done in order to determine if den sites chosen
by hyenas during the study period were influenced by proximity to intraspecific or interspecific
competitors.
3.5.3 Den buffers
500 m, 1000 m, and 2000 m ring buffers were placed around each hyena den to further examine
spatial relationships between den site selection and intra-clan home ranges and cores, inter-clan
home ranges and cores, and lion home ranges and cores. Comparisons were drawn between the
six active and 10 inactive hyena dens to determine if interspecific or conspecific pressure play a
role in den site choice. A total of 16 den sites were included in this analysis, six of which were
confirmed to be active during the field season and 10 of which were inactive.
3.5.4 Community land use
Use of non-protected community land by hyenas and lions was compared in order to gauge
spatial avoidance of human habitation. Mean percentages of home range and core areas
overlapping with communities were compared between the species using the Mann-Whitney U
test.
3.6 Temporal partitioning
3.6.1 Activity budgets
Activity budgets were developed based on data from the GPS-GSM collars, which give location
fixes every hour. GPS fixes were uploaded to ArcMap and distances measured between each
successive fix. Data from all six hyena collars and all three lion collars were used in this
analysis. Mean distance travelled (meters/hour) was calculated for both species for every one-
hour window.
Mean distance travelled per hour was then consolidated into time windows: dawn (05:00-09:00),
day (09:00-17:00), evening (17:00-21:00), and night (21:00-05:00) in order to determine if there
27
were differences in diurnal, nocturnal, or crepuscular activity patterns. These distances were
compared between species using Welch’s two-sample t test (Kitchen et al. 1999).
3.6.2 Dynamic analysis
Spatiotemporal avoidance of lions by hyenas was analysed by comparing the concurrent mean
distance between hyena-lion sympatric pairs to the distance expected by chance using dynamic
analysis of mutual interaction (Doncaster 1990, Minta 1992). The proportion of simultaneous
fixes that were below a critical distance of 200 m and a time window of 30 minutes was
compared to that which would be expected based on the distribution of distances between all
fixes using the widlifeDI package in R. This produced a contingency table of paired and non-
paired fixes with an associated p-value from a Chi-squared test (Long 2014).
Of the 18 possible hyena-lion pairs, 10 were sympatric and therefore used in this analysis. The
eight pairs with 0% overlap were excluded from the analysis, as their movements with respect to
one another were assumed to be random. Pairs were classified into two groups: those with
significant attraction based on Doncaster’s non-parametric test of interaction (p<0.05) and those
without (p>0.05). Home range overlap percentages between the two groups were then analysed
to determine if the group which displayed significant spatiotemporal attraction had a greater
mean overlap than the group with random movements. This was done through a Mann-Whitney
U test.
Of the 36 possible hyena pairs, five had any degree of home range overlap and could be used for
this analysis, four of which were pairs from different clans. Clan mates NH01 and NH02 were
expected to display attraction within their home ranges whereas inter-clan hyenas were expected
to display avoidance behaviour.
Of the three possible lion pairs, all three were sympatric. However, while WL01 is sympatric
with both SL01 and DL01, there were zero paired events, whether temporally matched or not.
Analysis was only run between SL01 and DL01, as they are sympatric and featured matched
fixes. They were accordingly expected to display avoidance behaviour. Therefore, SL01-DL01
was the only instraspecific lion analysis possible with Doncaster’s test.
Minta’s test for spatial and temporal interaction was also used for dynamic analysis, as it
accounts for differences between shared or overlapping home range areas and non-overlapping
areas with respect to space use (Long 2014, Minta 1992, Darnell 2014). Home ranges for a
28
sympatric pair are divided into areas used by individual A (LAA), individual B (LBB), and both.
The number of fixes in each area are tested against expected values based on the probability of
finding the animal in a given area based on real overlap percentages (Long 2014). LAA or LBB
values greater than 0 indicate attraction to the shared area while negative values indicate
avoidance. The Lixn statistic indicates simultaneity of use; a positive value indicates mutual
attraction to the shared area while a negative value indicates avoidance.
In Minta’s test, the test statistics LAA and LBB are calculated from a 2x2 contingency table, from
which a chi-squared value can be calculated for significance. Minta’s test was run for all 10
hyena-lion sympatric pairs, all five sympatric hyena pairs and the three sympatric lion pairs. This
technique used rgeos, wildlifeDI, and adehabitatHR packages in R.
3.7 Demographics
3.7.1 Hyenas
Prior to the field season, informal interviews with local rangers and scouts provided information
on known hyena dens. Den sites were visited on foot, marked with a GPS device, and camera
traps were placed to monitor activity. A photographic database of individual hyenas based on left
and right side spot patterns was developed from photos at dens and bait sites. If hyenas were
sighted opportunistically, they were photographed and identified.
Hyenas were assigned to a specific clan based on the following criteria:
(i) If the hyena was observed at the clan’s communal den;
(ii) If the hyena was observed with a hyena that has been assigned to a given clan;
(iii) If the hyena was observed within the core area of a collared hyena’s territory.
The database is split into five sections based on clan membership. Age classes were classified
into adult, sub-adult, and cub based on body size and pelage.
29
3.7.2 Lions
As part of its ongoing predator monitoring work, LWC already has an identification database of
the 34 lions within the LBL population. Lions were identified based on whisker spot patterns and
the photos within the database were updated when individuals were sighted.
Figure 3.7 – Example of unique left and right spot patterns on BH01.
30
4. Results
4.1 Dietary partitioning
4.1.1 Raw proportions
For both species, pains zebra was the most frequently counted prey item in scats (n=112, 70 for
hyenas and lions, respectively). For hyenas, impala (n=99) and Grevy’s zebra (n=73) were the
second and third most frequently counted prey species. For lions, Grevy’s zebra (n=60) and
eland (n=50) were second and third most frequent, respectively (Table 4.1). Zero lion scats
contained livestock (cattle, sheep, and goat) hairs while they were counted in 93 hyena scats
(78.8% of hyena scats).
4.1.2 Relative proportional contribution
Based on relative frequency of occurrence, the hyenas’ diet consists primarily of plains zebra
(18.9%), impala (16.7%), and Grevy’s zebra (12.3%) while the lions’ diet consists primarily of
plains zebra (22.5%), Grevy’s zebra (19.3%), and eland (16.1%). Livestock species altogether
contribute to 15.7% of the hyenas’ diet and 0.0% to the lions’ diet (Table 4.2, Figure 4.1).
Hyena Lion
Species # Scats (n=118) Proportion # Scats (n=87) Proportion
Plains zebra
Grevy's zebra
Warthog
Buffalo
Impala
Kudu
Giraffe
Oryx
Waterbuck
Eland
Cattle
Sheep
Goat
112
73
10
53
99
0
57
13
45
39
36
29
28
0.95
0.62
0.09
0.45
0.84
0.00
0.48
0.11
0.38
0.33
0.31
0.25
0.24
70
60
22
25
36
4
34
4
6
50
0
0
0
0.81
0.69
0.25
0.29
0.41
0.05
0.39
0.05
0.07
0.58
0.00
0.00
0.00
Table 4.1 – Raw frequencies of prey hairs in hyena and lion scats.
31
Prey Species Hyena Lion
Plains zebra 0.19 0.23
Grevy's zebra 0.12 0.19
Warthog 0.02 0.07
Buffalo 0.09 0.08
Impala 0.17 0.12
Kudu 0.00 0.01
Giraffe 0.10 0.11
Oryx 0.02 0.01
Waterbuck 0.08 0.02
Eland 0.07 0.16
Cattle 0.06 0.00
Sheep 0.05 0.00
Goat 0.05 0.00
Table 4.2 – Relative proportional contribution.
0.00
0.05
0.10
0.15
0.20
0.25
Pro
port
ion
Hyena Lion
Figure 4.1 – Relative proportional contribution of prey species to hyena and lion diets.
32
4.1.3 Jacobs’ index
Jacobs’ index values reveal selection of five prey species each for both hyenas and lions. Hyenas
select for waterbuck, giraffe, Grevy’s zebra, eland, and warthog, and against kudu, buffalo,
plains zebra, oryx, and impala. Lions select for warthog, eland, Grevy’s zebra, giraffe, and kudu
and against buffalo, oryx, impala, waterbuck, and plains zebra (Table 4.3, Figure 4.2). Of the 10
prey species, hyenas and lions shared selection in eight cases; the only differences were in kudu
(hyena negative, lion positive) and waterbuck (hyena positive, lion negative).
Species Hyena Lion
Plains zebra -0.19 ±.02 -0.19 ±.07
Grevy's zebra 0.38 ±.01 0.51 ±.09
Warthog 0.02 ±.06 0.59 ±.17
Buffalo -0.43 ±.07 -0.55 ±.11
Impala -0.03 ±.01 -0.34 ±.02
Kudu -1.00 0.37 ±.18
Giraffe 0.40 ±.10 0.38 ±.08
Oryx -0.07 ±.09 -0.41 ±.04
Waterbuck 0.48 ±.15 -0.27 ±.03
Eland 0.19 ±.07 0.54 ±.14
Table 4.3 – Jacobs’ index and standard error values.
33
4.1.4 Pianka’s index
Pianka’s index of niche overlap was found to be 0.8852 on the standardised scale, indicating
high dietary overlap between hyenas and lions at LBL and therefore similar dietary niches. This
outcome will be interpreted and compared to findings from other studies in the dietary
partitioning discussion (Section 5.2).
4.1.5 Levins’ index
Levins’ index of niche breadth reveals that hyenas have a broader diet than lions in LBL (Table
4.4), owing to their wider prey base and more uniform distribution of prey selection. Species
consumed by hyenas that were not consumed by lions include cattle, sheep, and goat – all
domesticated livestock. The only species consumed by lions that hyenas did not consume was
kudu, which contributed a small (1%) amount to the lions’ diet.
Species # Spp. Consumed Levins’ index Std. Levins’ index
Hyena 12 8.72 0.59
Lion 10 6.61 0.43
-1
-0.8
-0.6
-0.4
-0.2
0
0.2
0.4
0.6
0.8
1
Hyena Lion
Figure 4.2 – Jacobs’ index values based on game count data from 2014-2016.
Table 4.4 – Levins’ index of niche breadth, raw and standardised.
34
4.2 Spatial partitioning
4.2.1 Home range and core sizes
Lions had larger home ranges (U = 0, p<0.05) and cores (U = 0, p<0.05) than hyenas in all cases.
Home ranges for hyenas averaged 44.58 km2 (S.E.: ±5.81 km2, range: 37.00 km2–68.39 km2,
n=6) and for lions averaged 122.63 km2 (S.E.: ±24.46 km2, range: 76.34 km2–159.46 km2, n=3).
Cores for hyenas averaged only 9.87 km2 (S.E.: ±1.27 km2, range: 6.61 km2–14.83 km2, n=6) and
for lions averaged 40.64 km2 (S.E.: ±14.54 km2, range: 20.95 km2–69.02 km2, n=3) (Figure 4.3).
Figure 4.3 – Average home range and core sizes for hyenas and lions at LBL.
-10
10
30
50
70
90
110
130
150
Core Home Range
Mean a
rea (
km
2)
Hyena Lion
35
Figure 4.4 – Cores for collared hyenas and lions at LBL.
36
Figure 4.5 – Home ranges for collared hyenas and lions at LBL.
37
4.2.2 Overlap
Point and area percent overlap of hyena (HH, n=28), hyena-lion (HL, n=18), lion (LL, n=3), and
lion-hyena (LH, n=18) cores and home ranges were calculated and averaged to reveal greater
intraspecific avoidance than interspecific avoidance, particularly with respect to hyenas (Table
4.5, Table 4.6). Core-home range area overlap confirmed this trend. Intraspecific hyena
relationships revealed 0.00% overlap of cores and cores-home ranges, indicating strong inter-
clan avoidance. The NH01-NH02 relationship was excluded from this analysis, as they were in
the same clan.
Relationship PC PHR AC AHR CHR
HH 0.03 ± 0.00 0.89 ± 0.09 0.00 ± 0.00 2.04 ± 0.25 0.00 ± 0.00
LL 3.67 ± 1.09 33.18 ± 10.88 0.26 ± 0.00 14.65 ± 10.58 14.39 ± 11.27
HL 15.06 ± 2.75 80.82 ± 8.83 11.38 ± 0.28 70.27 ± 6.59 79.23 ± 9.26
LH 8.17 ± 0.48 67.90 ± 3.47 5.53 ± 0.53 51.09 ± 2.18 56.79 ± 4.11
Table 4.5 – Mean percent overlap of difference intraspecific and interspecific relationships based on point-core (PC), point-home range (PHR), area-core (AC), area-home range (AHR), and core-home range
(CHR) overlap.
38
Hyenas displayed significant intraspecific avoidance in comparison to interspecific avoidance
across all five spatial relationships (PC: U=170.5, p<0.05; PHR: U=150.5, p<0.05; AC: U=182,
p<0.01; AHR: U=144, p<0.01; CHR: U=140, p<0.01) (Table 4.6). Lions did not display
differences in overlap across any metric. Hyena-lion and lion-hyena overlap also did not reveal a
significant difference in interspecific avoidance from either species’ perspective.
U p-value
PC
HH-HL 170.5 0.01
LL-LH 58.5 0.77
HL-LH 155.5 0.83
PHR
HH-HL 150.5 0.02
LL-LH 61 0.66
HL-LH 189 0.38
AC
HH-HL 182 <0.01
LL-LH 57 0.84
HL-LH 157.5 0.87
AHR
HH-HL 144 <0.01
LL-LH 70 0.29
HL-LH 202 0.19
CHR
HH-HL 140 <0.01
LL-LH 58.5 0.77
HL-LH 183 0.47
Table 4.6 – Results of Mann-Whitney U Test.
39
Figure 4.6 - Home ranges and cores of collared individual hyenas, differentiated by clan association.
40
Figure 4.7 - Home ranges and cores of collared individual lions, differentiated by pride association.
41
4.2.3 Den activity
Average distance to active (n=6) vs. inactive (n=10) hyena dens was not significantly different
across any relationship (Table 4.7). Active dens were predictably closer to intra-clan cores (HC)
and home ranges (HHR), but surprisingly, also inter-clan intraspecific cores (OHC) and home
ranges (OHHR).
0
1000
2000
3000
4000
5000
6000
7000
8000
9000
HC HHR LC LHR OHC OHHR
Dis
tance (
m)
Active Inactive
Active Inactive U p-value
HC 705.5 ± 313.7 1514.5 ± 596.8 37 0.80
HHR 164.1 ± 153.5 297.6 ± 150.8 34.5 0.34
LC 1866.3 ± 771.9 564.9 ± 295.8 46.5 0.08
LHR 210.5 ± 174.5 142.0 ± 134.7 36 0.38
OHC 5471.5 ± 692.3 7363.6 ± 664.6 17 0.18
OHHR 3891.2 ± 652.9 4683.4 ± 585.3 20 0.31
Figure 4.8 - Average minimum distances (m) of active dens vs. inactive dens from intra-clan cores and home ranges, lion cores and home ranges, and inter-clan cores and
home ranges.
Table 4.7 – Average minimum distances (m) of active dens vs. inactive dens from intra-clan cores and home ranges, lion
cores and home ranges, and inter-clan cores and home ranges.
42
4.2.4 Den buffers
Placing 500 m, 1000 m, and 2000 m buffers around the hyena dens revealed strong inter-clan
avoidance, confirming the territoriality of hyenas at LBL (Table 4.8, Figure 4.9). Only one
confirmed den site was located within 2000 m of an inter-clan hyena’s home range; under all
other measures, there was no overlap between inter-clan cores or home ranges and hyena dens,
whether active or inactive.
Predictably, intra-clan dens were located within cores and home ranges of their respective
collared hyenas, regardless of den status (active or inactive). All 10 inactive dens were also
within the 2000 m buffer of an intra-clan hyena’s core, indicating the possibility of little seasonal
change in home ranges and relatively consistent spatial behaviour.
Given the large home ranges and cores of lions (Figure 4.10), it is also unsurprising that the
majority of hyena dens are located within lion home ranges. However, only 2/6 active hyena
dens were located within 500 m of lion cores, indicating possible lion avoidance.
500 m 1000 m 2000 m
Active Inactive Active Inactive Active Inactive
HC 4/6 6/10 5/6 6/10 6/6 8/10
HHR 5/6 9/10 6/6 10/10 6/6 10/10
LC 2/6 7/10 2/6 8/10 4/6 9/10
LHR 5/6 9/10 5/6 9/10 6/6 10/10
OHC 0/6 0/10 0/6 0/10 0/6 0/10
OHHR 0/6 0/10 0/6 0/10 1/6 0/10
Table 4.8 – Frequencies of occurrence of hyena dens within 500 m,
1000 m, and 2000 m buffers to intraspecific and interspecific cores
and home ranges.
43
Figure 4.9 – Hyena dens with buffers overlaid with hyena cores and home ranges.
44
Figure 4.10 – Hyena dens with buffers overlaid with lion cores and home ranges.
45
4.2.5 Community land use
Mean home range-community overlap for hyenas was 15.9% (S.E.: ±1.6%, range: 9.2% - 20.4%,
n=6) and for lions was 28.3% (S.E.: ±8.9%, range: 7.3% - 44.4%, n=3). Mean core-community
overlap for hyenas was 4.9% (S.E.: ±3.7%, range: 0.0% - 25.0%, n=6) and for lions was 6.5%
(S.E.: ±5.3%, range: 0.0% - 19.4%, n=3). Lions and hyenas did not differ significantly with
regards to home range (U=12, p=0.55) or core (U=10, p=0.90) overlap with communities lands,
indicating that both species utilise the human communities similarly.
Figure 4.11 - Mean proportion of home range and core areas located within human communities for hyenas and lions.
0
0.05
0.1
0.15
0.2
0.25
0.3
0.35
0.4
Home range Core
Pro
port
ion
Hyena Lion
46
4.3 Temporal partitioning
4.3.1 Activity budgets
Hyenas and lions displayed similar activity patterns, though hyenas travelled significantly further
at night (t=11.37, p<0.01, df=9.47) (Table 4.9). Both species displayed nocturnal tendencies,
with night (hyena: 39%, lion: 41%) contributing the greatest proportion to mean daily distance
travelled followed by evening (hyena: 31%, lion 28%), dawn (hyena: 27%, lion 26%), and day
(hyena: 3%, lion 5%) (Figure 4.12; Table 4.9). Hyenas reach maximum activity between 20:00 –
21:00 (1,020.5 m/hr) whereas lions peak at 03:00 – 04:00 (659.1 m/hr) (Figure 4.13).
Window Hyena Lion T p-value df
Dawn 582.8 ± 155.3 346.8 ± 102.0 0.81 0.46 5.18
Day 54.8 ± 8.7 68.7 ± 8.7 -1.13 0.28 13.99
Evening 677.4 ± 201.8 373.7 ± 105.8 1.29 0.26 4.53
Night 857.5 ± 10.4 555.8 ± 24.4 11.37 <0.01 9.47
Dawn Day Evening Night Dawn Day Evening Night
Table 4.9 – Mean distances travelled (m/hr) at five separate time windows by hyenas and lions at LBL.
Figure 4.12 – Activity budgets of hyenas (L) and lions (R) at LBL based on distance travelled at separate time windows as a proportion of total 24h distance travelled.
47
0
200
400
600
800
1000
1200
0-1 1-2 2-3 3-4 4-5 5-6 6-7 7-8 8-9 9-10 10-11 11-12 12-13 13-14 14-15 15-16 16-17 17-18 18-19 19-20 20-21 21-22 22-23 23-0
Dis
tance t
ravelle
d (
m/h
r)
Hyena Lion
Figure 4.13 - Average distance travelled each hour by hyenas and lions at LBL, with standard errors.
48
4.3.2 Dynamic analysis
Dynamic analysis of mutual interactions using Doncaster’s test on 10 hyena-lion pairs revealed
attraction in four cases and indifference in six cases (p-values = <0.01–1.00; Table 4.10). Total
paired fixes between sympatric hyena-lion pairs ranged from 1,537–2,868, with paired fixes that
fell within the 200 m threshold ranging from 0–14 and those above the 200 m threshold ranging
from 1,561–2,855.
Three of the four inter-clan hyena pairs showed no attraction (all p-values = 1.00). Surprisingly,
NH01-SH02 showed attraction, though this result was based on just one paired event, which is
attributable to a territorial dispute or kill site. To test the validity of these results, the interactions
between NH01 and NH02 were also analysed, revealing strong attraction between the two
(p<0.01) and underscoring the difference between intra-clan and inter-clan relationships (Table
4.11). The only lion pair analysed using Doncaster’s test revealed no attraction (p=0.43; Table
4.12).
Home range overlap between hyena-lion pairs that expressed attraction and those that expressed
indifference did not differ (U=15.5, p=0.51), indicating no relationship between overlap and
attraction.
49
Pair Below Above Totals
BH01-WL01
Paired 3 1561 1564
Non-Paired 1277 2443255 2444532
Totals 1280 2444816 2446096
p-value 0.06
CH01-DL01
Paired 0 2428 2428
Non-Paired 144 5892612 5892756
Totals 144 5895040 5895184
p-value 1.00
CH01-WL01
Paired 0 1330 1330
Non-Paired 141 1767429 1767570
Totals 141 1768759 1768900
p-value 1.00
NH01-DL01
Paired 14 2651 2665
Non-Paired 5242 7094318 7099560
Totals 5256 7096969 7102225
p-value <0.01
NH02-DL01
Paired 5 2591 2596
Non-Paired 5492 6731128 6736620
Totals 5497 6733719 6739216
p-value 0.10
SH02-DL01
Paired 3 2380 2383
Non-Paired 1177 5675129 5676306
Totals 1180 5677509 5678689
p-value <0.01
SH02-SL01
Paired 14 1897 1911
Non-Paired 1114 3648896 3650010
Totals 1128 3650793 3651921
p-value <0.01
SH02-WL01
Paired 0 1537 1537
Non-Paired 77 2360755 2360832
Totals 77 2362292 2362369
p-value 1.00
UH01-DL01
Paired 13 2855 2868
Non-Paired 9407 8213149 8222556
Totals 9420 8216004 8225424
p-value <0.01
UH01-SL01
Paired 2 2264 2266
Non-Paired 1026 5131464 5132490
Totals 1028 5133728 5134756
p-value 0.12
Table 4.10 – Table of contingency values for all 10 hyena-lion pairs, including Chi-squared p values.
50
Pair Below Above Totals
BH01-SH02
Paired 0 2268 2268
Non-Paired 143 5141413 5141556
Totals 143 5143681 5143824
p-value 1.00
CH01-NH02
Paired 0 2121 2121
Non-Paired 15 4496505 4496520
Totals 15 4498626 4498641
p-value 1.00
CH01-NH01
Paired 0 2210 2210
Non-Paired 17 4881873 4881890
Totals 17 4884083 4884100
p-value 1.00
NH01-NH02
Paired 98 2344 2442
Non-Paired 61819 5899103 5960922
Totals 61917 5901447 5963364
p-value <0.01
NH01-SH02
Paired 1 2238 2239
Non-Paired 33 5010849 5010882
Totals 34 5013087 5013121
p-value <0.01
Pair Below Above Totals
DL01-SL01
Paired 3 2480 2483
Non-Paired 3774 6159032 6162806
Totals 3777 6161512 6165289
p-value 0.43
Table 4.11 – Table of contingency values for all five hyena pairs, including Chi-squared p values.
Table 4.12 – Table of contingency values for DL01-SL01.
51
Minta’s test revealed hyenas were attracted to shared spaces with lions 40% of the time (LAA), of
which 75% were significant, and shared spaces with other inter-clan hyenas 0% of the time. The
intra-clan pair NH01-NH02 revealed significant attraction, as expected. Lions were attracted to
shared spaces with hyenas 40% of the time (LBB), of which 50% were significant, and shared
spaces with inter-pride lions 33% of the time, of which 50% were significant. In terms of
simultaneity of use (Lixn), hyena-lion pairs displayed attraction 70% of the time, of which 85.7%
were significant, inter-clan hyena pairs displayed attraction 0% of the time, and inter-pride lion
pairs displayed attraction 67% of the time, none of which were significant (Table 4.13).
Pair LAA pLAA LBB pLBB Lixn pLixn
Hyena-lion
BH01-WL01 1.85 <0.01 0.32 <0.01 0.51 <0.05
CH01-DL01 -0.42 <0.01 -1.06 <0.01 0.94 <0.01
CH01-WL01 -0.81 <0.01 0.25 0.21 -0.60 0.18
NH01-DL01 0.89 <0.01 -0.25 <0.01 -0.03 0.74
NH02-DL01 0.17 0.41 -0.29 <0.01 0.37 <0.01
SH02-DL01 -0.08 <0.01 -0.32 <0.01 -0.13 0.16
SH02-SL01 -0.12 <0.01 0.05 0.80 0.14 <0.01
SH02-WL01 -1.06 <0.01 -0.44 <0.01 0.05 0.65
UH01-DL01 1.04 <0.01 0.83 <0.01 0.31 <0.01
UH01-SL01 -0.44 <0.01 -0.48 <0.01 0.29 <0.01
Hyena-hyena
BH01-SH02 -1.61 <0.01 -1.43 <0.01 -0.98 <0.01
CH01-NH01 -∞ <0.01 -∞ <0.01 -∞ 0.84
CH01-NH02 -∞ 0.36 -∞ 0.44 -∞ 0.99
NH01-NH02 0.71 <0.01 0.34 <0.01 0.13 0.09
NH01-SH02 -1.23 <0.01 -0.07 0.70 -2.68 <0.01
Lion-lion
DL01-WL01 0.03 0.96 -∞ <0.01 -0.03 0.99
DL01-SL01 0.25 <0.05 -0.76 <0.01 0.06 0.27
SL01-WL01 -1.65 <0.01 -0.68 0.02 0.35 0.75
Table 4.13 – Test statistics and p-values for Minta’s spatial interaction test for hyena-lion, hyena-hyena, and lion-lion pairs.
52
4.4 Demographics
4.4.1 Identified individuals
A total of 89 hyenas were identified during the course of this study, with a range of 3-35 between
the clans (Table 4.14A). This number is undoubtedly incomplete as it relied on opportunistic
encounters with hyenas and camera trap photos from the dens. For the Shamba and Borana clans,
photographs were lost due to camera trap malfunctioning and partially recovered.
However, this result provides a minimum density of 0.24 hyenas/km2 and falls within the
estimates provided by a demographic study in 2015 (Preston 2015). Continued monitoring of
dens and development of the photographic database will complete the picture. However, this
effort has provided a baseline knowledge of hyena clan structure and membership within LBL.
Sex ratios were difficult to determine for the hyenas at LBL, leaving 67 of the 89 individuals
unidentified. Given the morphological similarities between male and female hyenas (East 2001),
females could only be positively identified if they were visibly lactating suckling cubs. Sexes of
collared individuals were determined while hyenas were immobilised.
Identification based on pride association of lions (Table 4.14B) confirmed that there are 34 lions
in LBL, providing a density of 0.09 lions/km2. Male coalitions were grouped with their resident
prides for the purposes of this study. As of July 2016, cubs were only present within one of
prides, Western, and none of the cubs had been identified yet by sex. The hyena to lion ratio
based on identified individuals is therefore, at minimum, 2.62:1.
53
A
B
Clan
Adults Sub-adults Cubs Male / Female / Unknown Totals
Borana 3 0 0 1 / 0 / 2 3
Charlie 16 3 5 1 / 3 / 20 24
Nala 21 9 5 1 / 8 / 26 35
Shamba 3 2 4 0 / 2 / 7 9
Utalii 10 2 6 1 / 5 / 12 18
Totals 53 16 20 4 / 18 / 67 89
Pride Adults Sub-adults Cubs Male / Female / Unknown Totals
Dalma 3 0 0 2 / 1 / 0 3
Sarah 3 2 0 0 / 5 / 0 5
Western 10 0 8 5 / 5 / 8 18
Mole 0 6 0 4 / 2 / 0 6
Linda 2 0 0 0 / 2 / 0 2
Totals 18 8 8 11 / 15 / 8 34
Table 4.14 - Individually identified hyenas (A) and lions (B) at LBL based on clan and pride membership, respectively.
54
5. Discussion
5.1 Collaring
While this study focused on interactions between hyenas and lions, it did not explore sex
differences in spatiotemporal patterns due to small sample size. Hyenas are matriarchal and
females tend to spend more time at den sites than males, particularly if they have cubs (Boydston
et al. 2003). Furthermore, each member of a hyena clan has a social rank. High ranking females
have smaller home ranges and cores and a higher proportion of fixes at the communal dens than
do low ranking females (Boydston et al. 2003). They also breed more frequently and display
higher fitness (Holekamp et al. 1996, Watts and Holekamp 2009). Finally, high-ranking females
display bolder behaviour with lions than do low-ranking females and males, including in
disputes over food (Shaw 2012). These factors affect spatiotemporal interactions with lions.
Because this study included the first collaring effort of hyenas and the first study to focus on this
species in LBL, hyenas were nervous. This made direct observations impossible, resulting in the
inability to differentiate individuals based on rank or sex – the latter only being possible with
collared individuals or lactating females. Therefore, differences may have occurred in
spatiotemporal patterns since four of the six hyenas in this study are male, and social ranks are
unknown for all individuals.
For lions, males tend to have larger home ranges and unless they are resident pride males,
display highly variable movement patterns when dispersing (Loveridge et al. 2009).
Furthermore, home range size varies across different environments and also according to pride
size and environmental variables such as mean annual precipitation and prey density (Schaller
1972, Funston and Mills 2006). DL01, the male lion in this study, is considered resident within
his pride and therefore represents a third distinct social group. However, being male, his data
may exhibit differences from WL01’s or SL01’s due to sex, both of whom are females.
For the two hyena clan mates NH01 and NH02, analysis was conducted for both separately and
they were treated as independent when looking at interspecific and intraspecific spatiotemporal
trends. NH01 is a female, and as expected, her core area was smaller than NH02’s, a male (6.61
km2 versus 14.83 km2).
Accordingly, despite these factors that may affect spatiotemporal behaviour in these species, the
similarities and differences that exist must be looked at through a species-specific lens as
55
opposed to a sex-based one. Ideally, a study can be conducted where all hyenas share the same
social rank and are the same sex and all lions are from the same-sized prides and are the same
sex, along with other demographic factors such as age. However, this is impossible in a
previously completely unstudied population.
5.2 Dietary partitioning
This study revealed high dietary overlap (Ô = 0.8852) between the study species, which
corroborates the findings of a meta-analysis of dietary partitioning between hyenas and lions
(Periquet et al. 2014). This overlap is indicative both of their similar niches and also a possible
scavenging or kleptoparasitic relationship between the two. In the Ngorongoro Crater, lions are
noted to scavenge from hyenas more frequently than hyenas do from lions (Kruuk 1972) and
lions do not surrender carcasses to hyenas until they have eaten their fill (Höner et al. 2002).
Hyenas are also much less successful at supplanting lions from kills when a male is around
(Kissui and Packer 2004). Given the small size of LBL and the high spatial overlap between
hyenas and lions, there is undoubtedly direct interaction between the species for kills.
As hyenas in this population have a broader diet (�̂� = 0.59 versus 0.43), they are utilising a prey
base outside of the lions’. One clear difference is that hyenas feed on livestock species in LBL,
while no livestock hairs were found in lion scat. This may be a mechanism for the hyenas to
avoid or mitigate competition for wild prey species. Therefore, it is possible that hyenas are
utilizing livestock species as a small but important alternative to wild ungulate prey (15.7% of
the hyenas’ diet). This has strong implications from a human-wildlife conflict perspective, as
local pastoralists rely on livestock for their livelihoods.
Jacobs’ index reveals that although plains zebra and impala are important contributors to both
species’ diets, they do not necessarily selectively feed on them. This suggests an abundance-
related tendency to focus on plains zebra and impala – they may simply hunt them because there
are more of them. The negative values for both plains zebra should not be regarded as blatant
avoidance but rather opportunistic, random feeding. Hyenas and lions both select against buffalo,
which supports previous dietary findings for lions in LBL (Pratt 2014). It is also surprising, as
the buffalo tends to be among the most favoured prey item for lions across populations (Hayward
and Kerley 2005, Davidson et al. 2013). However, different lion populations are known to
develop different cultures with prey selection; over time, the tendency to avoid buffaloes at LBL
may change (Power and Compion 2009).
56
5.3 Spatial partitioning
There are spatial parallels between the LBL hyena population and that of the Ngorongoro Crater,
Tanzania, which has been studied since the early 1970s. Hyenas display strong inter-clan spatial
exclusivity, reminiscent of the Ngorongoro population (Kruuk 1972). The Crater floor is 260
km2 and the ecosystem also features human communities both in and around the conservation
area. Average home range size at LBL (44.58 km2) is small when compared to other populations
of Crocuta, but larger than in areas with extremely high hyena density (≥1.00 individuals/km2)
such as the Ngorongoro Crater and Amboseli National Park, where home ranges are closely
defended and are generally 20–30 km2 (Holekamp and Dloniak 2011). The death of SH01 may
point towards such aggressive territoriality; in populations with tightly packed home ranges,
conspecific intruders are sometimes viciously attacked and killed (Smale et al. 1997). This sort
of territorial behaviour is in stark contrast to hyena populations in the Serengeti (Hofer and East
1995), where females travel far outside their core areas for several days due to the wildebeest
migration before returning to suckle cubs.
Lion home ranges are also noted to vary across landscapes (Gittleman and Harvey 1982). In
Nairobi National Park, Kenya, lion home ranges are between 25–51 km2, while in the lower
density Etosha National Park, Namibia, home ranges can be over 2,000 km2 (Stander 1991). The
home ranges at LBL can therefore be described as moderate in size, despite the small area of the
study site and intense human pressure. Lions do not display the same degree of territoriality that
hyenas do in this population, perhaps due to a lower population density and therefore less
intraspecific competition for prey. Lions are also inherently different from hyenas in that they do
not have communal dens; once cubs are mobile, they will accompany the pride as it moves
(McComb et al. 1993). Adult females, unless recently whelped, can freely move with their pride
members and cubs without needing to return to a den site for suckling and social rituals as adult
female hyenas do.
Despite the limited space available in LBL for these predators to coexist, there does appear to be
avoidance in core areas between intraspecific and interspecific competitors alike. Hyenas may
prefer core areas that minimise their chances of encountering lions, but still may use them as a
food source, therefore allowing for some degree of home range overlap. Lions may in turn also
utilise hyenas as a food source and are noted to respond favourably to recorded calls of hyenas
squabbling over kills (Kiffner 2008).
57
Two of the hyenas dens found during this study were located outside the conservancy
boundaries, and one was confirmed as being active. This den was attributed to the Borana clan,
as BH01’s fixes have indicated visits to this den site. While the majority (84.1%) of hyena home
range area in this study were located within LWC and BC, there is still clearly use of community
lands, as reflected in the scat data. Hyena clans are also noted, anecdotally, to reside permanently
within the ranches and community lands surrounding LBL, further confirming their ability to
reside in close proximity to humans (Yirga et al. 2011, Yirga et al. 2012, Yirga et al. 2013).
While lions also display utilisation of human settlements, no livestock hairs were found in their
scats. This appears to be a contradictory finding. This discrepancy may be attributed to the
chance that livestock hairs may not have been counted in the lion scats; livestock hairs may be
counted in the future with increased sample size. There are anecdotal reports of lions preying on
cattle in LWC in 2016 (Z. Davidson, Conservation Biologist, Marwell Wildlife, pers. comm.).
Hyenas that are based in the conservancies may be limited in how far they can venture into the
communities by other clans, as evidenced by the low spatial overlap between clans in this
population. Lions do not face such an intraspecific barrier.
5.4 Temporal partitioning
Both hyenas and lions at LBL have similar activity patterns, apart from hyenas being
significantly more active at night. As hyenas prey on domestic livestock from the communities,
it is possible that this heavily nocturnal behaviour is a response to human presence. Analysis of
hyena temporal trends in the Masai Mara Game Reserve (GR) reveals that hyenas in close
contact to human habitation tend to be more nocturnal than hyenas towards the centre of the GR
(Pangle and Holekamp 2010). All the hyena clans in LBL have home range boundaries that
overlap with human communities, which directly puts them at risk of retaliatory persecution due
to livestock losses. Overall, both species appear to be largely nocturnal and crepuscular, which is
to be expected, especially given the high human impact in LBL as a compounding factor.
Because 40% of the interspecific cases of spatiotemporal avoidance actually revealed attraction
by hyenas and lions using both Doncaster’s and Minta’s tests, a likely factor playing a role in
these relationships is prey. It is plausible that hyenas are scavenging from lions at LBL, or vice
versa. In Minta’s test, mutual simultaneity of use revealed 70% of the hyena-lion cases also
shows a surprisingly high level of direct interactions. All hyena pairs revealed avoidance apart
from the NH01-NH02 and the NH01-SH02 pairings. The former is unsurprising due to the pair
belonging to the Nala clan; the latter is based on just one paired encounter which could be
attributed to a dispute over a carcass or territorial conflict. Both NH01 and SH02 are large, adult
58
females, which makes them likely participants in inter-clan conflicts for space or prey (Shaw
2012). Lions showed less intraspecific avoidance than hyenas, with 67% of lion pairs showing
simultaneous use and 33% showing attraction. This points towards the exclusivity of hyena clans
in this landscape, and to a lesser degree, lion prides. It also reveals the high degree of
interspecific sharing of space and time, for the acquisition of prey.
Direct encounters between hyenas and lions often result in hyena mortality. Considering the
frequency of intraguild predation by lions on hyenas and the limitation of hyena fitness by lions,
the benefit of food is a strong driver for such bold behaviour around lions by hyenas (Watts and
Holekamp 2008). In the Masai Mara, prior to human expansion towards the GR in the 1990s,
lions were the leading cause of adult hyena mortality but have since been supplanted by humans
for hyenas with home ranges bordering human communities (Pangle and Holekamp 2010).
Coming into close proximity with lions is a potentially lethal decision for hyenas to make.
Therefore, the potential of acquiring food through such encounters must outweigh the risk of
mortality; the same can be said for hyena-human contact.
5.5 Demographics
The 89 hyenas identified so far in the LBL population is an incomplete figure, but represents a
density of 0.24 hyenas/km2, which is higher than areas such as the Okavango Delta (Cozzi et al.
2013), Kruger National Park, Etosha National Park, and Central Kalahari Game Reserve
(Holekamp and Dloniak 2011). Only three hyenas were identified from the Borana clan, which is
a definite underestimate. Anecdotal reports from researchers who have worked in LBL as well as
rangers suggest that hyenas are frequently seen and heard in BC (S. Gilisho, Predator Monitoring
Officer, LWC, pers. comm.). Furthermore, sightings of groups of 47, 41, 20, 16, and 10 adult
hyenas together at different locations within LBL since 2015 (S. Gilisho pers. comm.) point
toward a robust population. It is likely that the LBL hyena population is growing, given the
identification of 36 individual hyenas below breeding age in this study. Furthermore, the
presence of permanent hyena clans outside of LWC and BC is a sign that there is adequate gene
flow in and out of the conservancies, given the migratory gaps. The Mount Kenya elephant
corridor also provides for landscape connectivity between LBL and Mount Kenya National Park
(Nyaligu and Weeks 2013), and hyenas are known to use it. There is limited information,
however, on hyena demography in central Kenya. In Laikipia, an estimated 71 hyenas are killed
annually by humans on ranches, not counting road kill (Frank 1998), and hyenas are declining
outside of protected areas across Africa (Woodroffe 2000). Therefore, while there are some
59
encouraging signs on the stability and growth potential of the hyena population within LBL,
there are still gaps in knowledge that need to be filled.
The lion population described in this study, as it stands at 34, is considered accurate. A density
of .09 lions/km2 is moderate and healthy in large ecosystems such as Selous in Tanzania (Creel
and Creel 1997). However, unlike hyenas, lions are not known to reside permanently in the
community areas immediately surrounding LBL and there is limited gene flow in and out of this
population. Inbreeding between siblings has been confirmed in at least one case (M. Mwololo,
Research Officer 1, LWC, pers. comm.) and is of concern. The lion population in the
Ngorongoro Crater has also been noted to have high rates of inbreeding due to the closed nature
and small area of the Crater, which in turn has costly fitness consequences and places such
populations at risk of local extinction (Riggio et al. 2013). Small populations are inherently
susceptible to stochastic variations (Lande 1993). Just eight cubs were identified this year, from
only the Western pride. One encouraging sign is that lions are known to reside in ranches to the
west of BC in Laikipia (Frank et al. 2005), which may allow for appropriate management
measures to be taken to encourage dispersal. There may be potential for dispersal to and from
ranches further west in Laikipia, but the presence of inbreeding within this population points to
limited connectivity.
5.6 Conclusions and management recommendations
Overall, hyenas display stronger intraspecific than interspecific avoidance in spatiotemporal
trends. Both species share similar diets and activity patterns, though lions have larger home
ranges and cores. Nonetheless, there are some differences. Hyenas travel further at night than do
lions. Hyenas feed on livestock whereas lions do not. Hyenas have small, exclusive home ranges,
whereas lions have large, slightly overlapping ones. There are known hyena clans within the
communities, but no lion prides. The hyena population overall appears to be healthy, with a high
number of cubs and sub-adults, but the lion population may be suffering from inbreeding, a
limited ability to disperse, and low cub recruitment.
A human-hyena conflict study would be useful in determining local attitudes towards hyenas, the
impact that hyenas are having on livestock, and human economic loss to livestock predation.
This can provide for the development of solutions to mitigate losses, such as compensation
schemes or the development of corrals to protect livestock. Estimates of livestock abundance in
comparison to wild ungulate abundance can provide a clearer picture of whether hyenas are
preferentially feeding on domesticated prey. A comparison of hyena clans inside versus outside
60
the conservancies would shed light on the impact of human communities on hyena spatial and
dietary ecology, as well as differences in stress hormones such as cortisol. This information can
provide for policies that allow for humans and carnivores to coexist. Deploying GPS collars on
hyenas from as many clans as possible both inside and outside the conservancies would be ideal
in this scenario to maximise sample sizes.
Expanding monitoring efforts to other members of the large carnivore guild will provide a more
complete picture of predator ecology in the region. African wild dogs, leopards, cheetahs, and
striped hyenas are all present in LBL. Leopards and cheetahs were both observed in chance
encounters during the field season, and the cheetahs were well-habituated. A leopard and a
striped hyena were also seen on camera trap at hyena baits. It would be useful to gather data on
the complete demography, spatiotemporal behaviour, and dietary ecology of these species not
just in LBL but in surrounding ranches as well. It is also advised that direct observations be used
in this effort. Lions and cheetahs in LBL are already well-habituated to vehicles, but hyenas,
leopards, and wild dogs are not. Habituating these species to vehicles for observational research
can provide accurate hunting versus scavenging data. Ideally, this research can be done long
term to examine seasonal differences in diet or activity and longer term spatial trends.
Culling of predators is strongly advised against based on the findings in this study. Lions and
hyenas exist in a natural balance; lion control leads to massive rebounds in hyena populations
and disrupts this balance. In Amboseli National Park, hyena populations exploded from a density
of 0.13 hyenas/km2 to 1.65 hyenas/km2 between the mid-1990’s and mid-2000’s due to local lion
extinction (Watts and Holekamp 2008). Although they demonstrate the ability to exist in human-
dominated landscapes, hyenas are vulnerable once locally extirpated due to female philopatry
and conservative recolonisation of unclaimed territories, even with intact source populations
nearby (Frank 1998). Furthermore, no rhino hairs were found in any of the hyena or lion scats
analysed in this study. While hyenas have been noted to attack rhino calves in Aberdare National
Park, there were no confirmed successful cases of predation (Sillero-Zubiri and Gottelli, 1991).
The lion population is already low and putting external control on them can have strong negative
outcomes, including the possibility of extinction. Eight lions were poisoned in LBL since 2014
as retaliation for livestock attacks (Z. Davidson pers. comm.). For a small population, this is a
major loss. Lions are classified as Vulnerable by the International Union for Conservation of
Nature (IUCN) (Bauer et al. 2008); preserving this population can be a small step towards
ensuring their continued persistence in Kenya.
Landscape connectivity and dispersal are important for both hyenas and lions. Male lion
dispersal ability is strongly correlated to population connectivity and East African lions now
61
exist in population fragments due to anthropogenic pressure (Dolrenry et al. 2014). While hyenas
in this area appear to enjoy the presence of multiple clans both in and around the conservancies,
the lion population is more isolated. It may be necessary to provide more wildlife corridors
within the study area to allow for genetic exchange between lion populations in surrounding
areas such as Laikipia, Samburu, Meru, and Mount Kenya. An alternative to this is
metapopulation management, which relies on hands-on removal and introduction of individuals
between disconnected areas. Metapopulation management has demonstrated success in South
Africa with African wild dogs (Gusset et al. 2006) and has been suggested as a management
strategy for lions (Dolrenry et al. 2014), cheetahs (Johnson et al. 2010), and leopards (Balme et
al. 2010). Expanding the current predator monitoring effort inside and outside of LBL,
facilitating carnivore dispersal, and determining carnivore impacts on human communities will
allow for management decisions to be made that allow for the coexistence of predators and
humans alike in LBL and beyond.
62
Acknowledgments
I owe many people thanks for their help and support during the course of this project. Many
thanks to Dr. Zeke Davidson and Prof. C. Patrick Doncaster for their helpful guidance and
feedback during this project. Dr. Davidson opened his home to me and was always helpful with
fieldwork-related aspects and the initiation of the project. Prof. Doncaster provided me with
considerable support regarding statistical analysis and the writing of this report. Many thanks to
both for rigorously reviewing my proposal, dissertation drafts, and providing me with detailed
feedback.
I am grateful to Dr. Marc Dupuis-Desormeaux and Prof. Suzanne MacDonald for taking an
interest in LBL’s hyenas and getting this project started. They provided funding for the GPS-
GSM collars and their guidance at the start of the field season as we prepared for the trapping
endeavour was incredibly helpful. I must also give thanks to LWC’s and BC’s management
teams and Northern Rangelands Trust (NRT) for allowing this project to take place – Ian Craig
and Mike Watson took a keen interest in this project and I appreciate their support and
encouragement.
I also would like to thank Dr. Alayne Cotterill for the extremely informative tutorial on trapping
hyenas (and for getting our first hyena) as well as Dr. Laurence Frank for generously allowing
the team to borrow his equipment for leg-hold snares. This was my first ever crash course in
collaring spotted hyenas and it would have been impossible without their support.
Special thanks must also be extended to the research departments at LWC and BC, in particular
Geoffrey Chege, Mary Mwololo, Edwin Kisio, and of course, Saibala Gilisho. Chege, Mary, and
Edwin were exceptionally organised with vehicles, bait, and reporting hyena and lion sightings.
Saibala spearheaded all the work prior to my arrival and very effectively set the foundation for
this research, which made the process much smoother than it otherwise would have been. He
worked extremely long hours with me that went above and beyond my expectations.
I want to thank the drivers as well – Logeme, Sumbere, and Karmushu – for providing me with
transport in challenging field conditions and always assisting during the collaring efforts with a
smile.
63
To all the casuals whose names I never learned or could not remember – I express my gratitude
here. Studying large carnivores requires a lot of manual labour and they were always keen to
lend a helping hand when I had to move traps or baits.
I also greatly appreciate the help provided by Dr. Bernard Rono and Dr. Matthew Mutinda, the
two veterinarians I worked with during the collaring effort. Monitoring the spatiotemporal
aspects of these predators would have been impossible without collaring them, and collaring
them would have been impossible without their immobilisation. Dr. Rono and Dr. Mutinda made
themselves available and kept the study animals safe even in extremely logistically challenging
situations and on short notice, and for that I am very grateful.
Finally, more than anything, I owe thanks to my parents and siblings. Without their support and
encouragement over the years to pursue this field, I would not be where I am today. I understand
why it can be challenging at times to have a son (or brother) who wants nothing more than to
spend time in the remote wilderness with large carnivores. However, they accept and embrace it
with open arms and have taken a genuine interest in my career, and for that I am eternally
grateful.
64
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Appendix A
Figure A.1 – Collaring sites for hyenas and lions between February – April 2016.
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Appendix B
Capture Data Sheet
Date Sex
Time Age (approx.)
ID Collar Freq.
Condition (1-5) Area/ Grid ref.
Skull length Weight
Skull width Mane lower jaw
Nose – tail root (HB) Mane cheek
Tail Mane at crest skull
Chest girth(half x2) Mane b/w shoulders
Shoulder height Mane mid chest
Canine upper length curve Comments:-
Time dart in:
Time to Ataxa:
Time to Recumbency:
Reversal time:
Recovery time:
Drugs:
Immobilisation (Mg/Kg):
Reversal (Mg/Kg):
Smooth Immobilisation?
Top-up Administered?
Smooth Recovery?
Condition mammae:
Blood sample:
Nasal swab:
Hair sample:
Other:
Other:
Canine upper base width
Canine length straight
Canine lower length
Canine lower base width
Wear on teeth
Front paw length
Front paw width
Rear leg (knee - paw)
Base of tail circum.
Table B.1 – Sample collaring data sheet template.
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Appendix C
Figure C.1 – Progression of collar deployment and collar data windows during field season.
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