The Effects of Drought on Diets of Apex Predators in the South African Lowveld Inferred by Fecal Hair Analysis" (2016)

Western Kentucky University TopSCHOLAR® Honors College Capstone Experience/Thesis Honors College at WKU Projects

9-1-2016 The ffecE ts of Drought on Diets of Apex Predators in the South African Lowveld Inferred by Fecal Hair Analysis Shelby Wade Western Kentucky University, [email protected]

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Recommended Citation Wade, Shelby, "The Effects of Drought on Diets of Apex Predators in the South African Lowveld Inferred by Fecal Hair Analysis" (2016). Honors College Capstone Experience/Thesis Projects. Paper 669. http://digitalcommons.wku.edu/stu_hon_theses/669

This Thesis is brought to you for free and open access by TopSCHOLAR®. It has been accepted for inclusion in Honors College Capstone Experience/ Thesis Projects by an authorized administrator of TopSCHOLAR®. For more information, please contact [email protected]. THE EFFECTS OF DROUGHT ON DIETS OF APEX PREDATORS IN THE SOUTH

AFRICAN LOWVELD INFERRED BY FECAL HAIR ANALYSIS

A Capstone Experience/Thesis Project

Presented in Partial Fulfillment of the Requirements for

the Degree Bachelor of Science with

Honors College Graduate Distinction at Western Kentucky University

By:

Shelby K. Wade

*****

Western Kentucky University

2017

CE/T Committee:

Dr. Michael Stokes, Advisor Approved by

Dr. John Andersland ______Advisor Dr. Karen Furgal, DPT Department of Biology

ABSTRACT

To properly manage offtake quotas and conservation efforts, Balule Nature

Reserve (South Africa) instituted a study in 2014 to determine prey species selection by megapredators. In 2015, Balule Nature Reserve received about 190 mm less rainfall between the months of January and June than in 2014 (116 mm less than average). This study compares the diets of lions and hyaenas between 2014 and 2015. Prey species consumed were determined by fecal analysis, and results were compared to prey availability. Sixteen, 1 km2 plots were chosen from the 400 km2 Reserve. Between June and August 2015, we walked three, 1 km transects in each plot, collecting 87 fecal samples. Hairs from each sample were selected for microscopic analysis via multiple subsampling methods and the hairs identified. As a reference, we developed a pictorial atlas of hairs from 17 mammalian game species showing cuticular scale and medullary patterns visible with scanning electron and light microscopes. Diet composition and

Jacob’s Selectivity Index were calculated for both predator species. Impala were less represented as prey than expected, and zebra, nyala, bushbuck, and duiker were more represented as prey than expected relative to their populations.

Keywords: Apex Predators, South Africa, Balule Nature Reserve, Diet Composition, Fecal Analysis, Pictorial Atlas

ACKNOWLEDGEMENTS

We thank all of the volunteers at Transfrontier Africa for their encouragement and support. We also thank Limpopo Taxidermy, the Louisville Zoo, and the Thomas M.

Baker Collection for supplying hairs used for this project. Dr. John Andersland directs the electron microscopy facility and ensured best results. Funding for the project was provided by the Honors College and FUSE program at WKU.

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VITA

December 12, 1994 ...... Born—Louisville, Kentucky

May 25, 2013 ...... Central Hardin High School, Cecilia, Kentucky

April 2014 ...... Employee at All Creatures Animal Hospital

Summer 2015 ...... Study Abroad in South Africa

PUBLICATIONS

None

FIELDS OF STUDY

Major Field: Biochemistry

TABLE OF CONTENTS

Page

Abstract……………………………………………………………………………………ii

Acknowledgments………………………………………………………………………..iii

Vita………………………………………………………………………………………..iv

List of Figures…………………………………………………………………………….vi

List of Tables……………………………………………………………………………..xi

Chapters:

1. An SEM image reference guide to hairs of seventeen species of large African

mammals…………………………………………………………………………..1

2. Diets of two apex predators in a period of drought in the Balule Nature Reserve,

Hoedspruit, South Africa………………………………………………………...33

Bibliography……………………………………………………………………………..68

LIST OF FIGURES

Figure Page

1.1 SEM image of impala (Aepyceros melampus) guard hair………………………...7

1.2 SEM image of impala (Aepyceros melampus) guard hair...………………………8

1.3 SEM image of blue wildebeest (Connochaetes taurinus) guard hair……………..9

1.4 SEM image of waterbuck (Kobus ellipsiprymmus) guard hair…………………..10

1.5 SEM image of klipspringer (Oreotragus oreotragus) guard hair………………..11

1.6 SEM image of gemsbok (Oryx gazella) guard hair……………….……………..12

1.7 SEM image of steenbok (Raphicerus campestris) guard hair...…………………13

1.8 SEM image of common duiker (Sylvicapra grimmia) guard hair...……………..14

1.9 SEM image of African buffalo (Syncerus caffer) guard hair…………………….15

1.10 SEM image of male nyala (Tragelaphus angasii) guard hair...………………….16

1.11 SEM image of male nyala (Tragelaphus angasii) bristle hair…...……………....17

1.12 SEM image of bushbuck (Tragelaphus scriptus) guard hair...…………………..18

1.13 SEM image of greater kudu (Tragelaphus strepsiceros) guard hair……………..19

1.14 SEM image of giraffe (Giraffa camelopardalis) guard hair……………………..20

1.15 SEM image of giraffe (Giraffa camelopardalis) bristle hair…………………….21

1.16 SEM image of warthog (Phacochoerus africanus) guard hair...………………...22

1.17 SEM image of lion (Panthera leo) guard hair...…………………………………23

1.18 SEM image of lion (Panthera leo) bristle hair...………………………………...24

1.19 SEM image of leopard (Panthera pardus) light guard hair...…………………....25

1.20 SEM image of leopard (Panthera pardus) dark guard hair...……………………26

1.21 SEM image of spotted hyaena (Crocuta crocuta) guard hair...………………….27

1.22 SEM image of Burchell’s zebra (Equus burchellii) guard hair...………………..28

1.23 SEM image of Burchell’s zebra (Equus burchellii) bristle hair...……………….29

2.1 Map of Balule Nature Reserve and corresponding management districts……….36

2.2 Plots sampled in 2014……………………………………………………………37

2.3 Plots sampled in 2015……………………………………………………….…...37

2.4 Grid used for subsampling method………………………………………………38

2.5 Petri dish with 18 points, used for hair selection………………………………...39

2.6 Distribution of all 87 predator scat samples collected in 2015…………………..43

2.7 Bar graph comparing proportions of species found in lion fecal samples from

2015, aerial game counts, and lion fecal samples from 2014…………....………46

2.8 Bar graph comparing proportions of species found in hyaena fecal samples from

2015, aerial game counts, and hyaena fecal samples from 2014…………….…..46

2.9 Bar graph demonstrating Jacob's Index values for all species in lion and hyaena

diets in 2014………………………………………………………………...……47

2.10 Bar graph demonstrating Jacob's Index values for all species in lion and hyaena

diets in 2015……………………………………………………………………...47

2.11 Brightfield image of impala (Aepyceros melampus) guard hair medulla

pattern……………………………………………………………………….…...49

2.12 Cast of impala (Aepyceros melampus) guard hair scale pattern…………………49

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2.13 Brightfield image of blue wildebeest (Connochaetes taurinus) proximal region of

guard hair medulla pattern…….…………………………………………………50

2.14 Cast of blue wildebeest (Connochaetes taurinus) guard hair scale pattern……...50

2.15 Brightfield image of blue wildebeest (Connochaetes taurinus) distal region of

guard hair medulla pattern….……………………………………………………50

2.16 Brightfield image of waterbuck (Kobus ellipsiprymnus) proximal guard hair

medulla pattern…………………………………………………………………...51

2.17 Cast of waterbuck (Kobus ellipsiprymnus) guard hair scale pattern……………..51

2.18 Cast of waterbuck (Kobus ellipsiprymnus) distal region of guard hair scale

pattern……………………………………………………………………………51

2.19 Brightfield image of klipspringer (Oreotragus oreotragus) guard hair medulla

pattern…………………………………………………………………………....52

2.20 Cast of klipspringer (Oreotragus oreotragus) guard hair scale pattern………….52

2.21 Brightfield image of steenbok (Raphicerus compestris) guard hair medulla

pattern……………………………………………………………………………52

2.22 Cast of steenbok (Raphicerus compestris) guard hair scale pattern……...……...52

2.23 Brightfield image of duiker (Sylvicapra grimmia) guard hair medulla pattern.....53

2.24 Cast of duiker (Sylvicapra grimmia) proximal region of guard hair scale

pattern……………………………………………………………………………53

2.25 Cast of duiker (Sylvicapra grimmia) medial/distal region of guard hair scale

pattern……………………………………………………………………………53

2.26 Brightfield image of African buffalo (Syncerus caffer) proximal guard hair

medulla pattern……………………………………………………………...……54

viii

2.27 Cast of African buffalo (Syncerus caffer) guard hair scale pattern………...……54

2.28 Brightfield image of African buffalo (Syncerus caffer) distal region of guard hair

medulla pattern…………………………………………………………………...54

2.29 Brightfield image of nyala (Tragelaphus angasii) bristle hair medulla pattern....55

2.30 Cast of nyala (Tragelaphus angasii) bristle hair scale pattern………...………...55

2.31 Brightfield image of nyala (Tragelaphus angasii) guard hair medulla pattern.....55

2.32 Cast of nyala (Tragelaphus angasii) guard hair scale pattern………...…………55

2.33 Brightfield image of bushbuck (Tragelaphus scriptus) guard hair medulla

pattern…………………………………………………………………………....56

2.34 Cast of bushbuck (Travelaphus scriptus) guard hair scale pattern………………56

2.35 Brightfield image of greater kudu (Tragelaphus strepsiceros) guard hair medulla

pattern……………………………………………………………………………56

2.36 Cast of greater kudu (Tragelaphus strepsiceros) guard hair scale pattern……....56

2.37 Brightfield image of giraffe (Giraffa camelopardalis) bristle hair medulla

pattern……………………………………………………………………….…...57

2.38 Cast of giraffe (Giraffa camelopardalis) bristle hair scale pattern………………57

2.39 Brightfield image of giraffe (Giraffa camelopardalis) proximal region of guard

hair medulla pattern……………………………………………………………...57

2.40 Cast of giraffe (Giraffa camelopardalis) guard hair scale pattern………………57

2.41 Brightfield image of giraffe (Giraffa camelopardalis) distal region of guard hair

medulla pattern…………………………………………………………………..58

2.42 Brightfield image of lion (Panthera leo) dark bristle hair medulla pattern...……59

2.43 Cast of lion (Panthera leo) bristle scale pattern…………………………………59

2.44 Brightfield image of lion (Panthera leo) light bristle hair medulla pattern...……59

2.45 Cast of lion (Panthera leo) guard hair scale pattern……………………………..59

2.46 Brightfield image of lion (Panthera leo) guard hair medulla pattern...………….60

2.47 Brightfield image of leopard (Panthera pardus) guard hair medulla pattern...….61

2.48 Cast of leopard (Panthera pardus) guard hair scale pattern……………………..61

2.49 Brightfield image of spotted hyaena (Crocuta crocuta) proximal/medial region of

guard hair medulla pattern……………….………………………………………62

2.50 Cast of spotted hyaena (Crocuta crocuta) guard hair scale pattern……………...62

2.51 Brightfield image of spotted hyaena (Crocuta crocuta) distal region of guard hair

medulla pattern…………………………………………………………………...62

2.52 Brightfield image of Burchell’s zebra (Equus burchellii) dark bristle hair medulla

pattern………………………………………………………………….………...63

2.53 Cast of Burchell’s zebra (Equus burchellii) bristle hair scale pattern...…………63

2.54 Brightfield image of Burchell’s zebra (Equus burchellii) white bristle hair

medulla pattern………………………………………………………..…………63

2.55 Cast of Burchell’s zebra (Equus burchellii) guard hair scale pattern………...….63

2.56 Cast of Burchell’s zebra (Equus burchellii) guard hair scale pattern………...….64

LIST OF TABLES

Table Page

1.1 Species whose hairs were imaged with SEM ...... 4

2.1 Species whose hair was gathered for reference and characterized ...... 40

2.2 Number of scats containing hairs of each prey species by region consumed by

lion ...... 44

2.3 Number of scats containing hairs of each prey species by region consumed by

hyaena ...... 44

2.4 G-statistic results for the African lion when compared to aerial game counts and

results from the 2014 study ...... 48

2.5 G-statistic results for the spotted hyaena when compared to aerial game counts

and results from the 2014 study ...... 48

CHAPTER 1

AN SEM IMAGE REFERENCE GUIDE TO HAIRS OF SEVENTEEN SPECIES OF

LARGE AFRICAN MAMMALS

Introduction

Morphological features of mammalian hairs have proven to be useful in a variety of biological studies, including predator diet analyses, conservation management, forensic studies, medical uses, and even archaeological studies (Johnson and Hansen

1979, Rosas-Rosas et al. 2003, Deedrick and Koch 2004, Manfredi et al. 2004, Breuer

2005, Ott et al. 2006, Backwell et al. 2009, Mansilla et al. 2011, Wentworth et al. 2011,

Braczkowski et al. 2012, Mbizah et al. 2012, Verma and Joshi 2012, Davidson et al.

2013, Taru and Backwell 2013, Taru and Backwell 2014). Hair is made mostly of keratin, which causes it to be very durable, as keratin is more resistant to decay and digestive alteration than other mammalian tissues such as dentin or bone (Deedrick and Koch 2004,

Backwell et al. 2009, Mansilla et al. 2011, Taru and Backwell 2013). Under ideal conditions, keratin may last hundreds or even thousands of years without undergoing significant changes, and is therefore useful when it comes to species identification from historical samples (Backwell et al. 2009, Mansilla et al. 2011, Taru and Backwell 2013).

Short (1978) and Keogh (1983) found that there are no recognizable differences between

1 otherwise identical hairs from fresh kills, from museum specimens, or from digested materials.

Mammalian hair shafts consist of three main concentric layers: the cuticle, the cortex, and the medulla (Deedrick and Koch 2004). The cuticle is the superficial layer of the hair. It consists of scales which typically overlap each other and form cuticular scale patterns. The cortex is the intermediate layer of the hair. It typically contains pigment granules, which give a hair its color, and sometimes contains ovoid bodies, which are large, pigmented, oval-shaped structures. The medulla forms the central core of the hair.

It is made of dead, collapsed cells; the number present can vary greatly. In some hairs, the medulla may be absent.

Longitudinally, hair also consists of three regions: proximal (the end nearest the root or follicle), medial (the middle region), and distal (the free end). Scale patterns can change down the length of the hair, so images should be taken at all three regions for every species (Seiler 2010). This is very important, especially when analyzing hairs from scat, because hairs will often be fragmented, so scale patterns from only one region of the hair may be available. Cuticular scales always point from the proximal end of the hair to the distal region, and hair shafts tend to decrease in diameter towards the distal end

(Deedrick and Koch 2004).

Previous catalogs have used several terms to describe the different common cuticular scale patterns. These terms include imbricate, mosaic, chevron, coronal, and petal-like or spinous (Homan and Genoways 1978, Deedrick and Koch 2004, Keogh

1983). Imbricate is described as flattened, overlapping scales with margins close together

(Deedrick and Koch 2004). Mosaic is described as “being composed of a number of

2 units” (Keogh 1983). Chevron is a waved pattern in which the trough, crest, or both of the scales resemble a “V” shape (Keogh 1983). Coronal involves only one or two scales surrounding the hair shaft and bearing the resemblance of a stack of paper cups (Homan and Genoways 1978, Deedrick and Koch 2004, Keogh 1983). Petal-like, or spinous scales protrude from the hair shaft. They resemble overlapping flower petals and may be diamond or triangular in shape (Deedrick and Koch 2004, Keogh 1983).

The distances between two adjacent scale margins are often used as descriptors and diagnostic markers in addition to the shape of the scales. The three most common words used to describe the distances are close, near, and distant (Keogh 1983). These terms are only relative, and are not quantified, so it is less precise to describe a hair in this way, but can help in cases where the identity of the species is uncertain. Lastly, the scale margins themselves can be described as smooth, crenate, or rippled (Keogh 1983).

In several studies, cuticular scale patterns of hairs were used to aid in the process of species identification (Johnson and Hansen 1979, Rosas-Rosas et al. 2003, Manfredi et al. 2004, Breuer 2005, Ott et al. 2006, Wentworth et al. 2011, Braczkowski et al. 2012,

Mbizah et al. 2012, Davidson et al. 2013). This is often accomplished by creating a scale imprint by mounting a hair on a slide using clear nail polish or a gelatin and allowing it to dry (Homan and Genoways 1978, Keogh 1983, Mizutani 1999, Ott et al. 2006,

Wentworth et al. 2011, Mbizah 2012) . Once dry, the hair is removed and a mold of the scale pattern is left. This mold is then observed under a light microscope and compared to a reference catalog of known hair scale patterns.

Using a scanning electron microscope (SEM) allows for a more detailed analysis of cuticular scale patterns. Using the SEM, scales can be observed directly, instead of

3 through use of imprints or molds. The SEM provides higher resolution images of the cuticle scale patterns than light microscopes (Sahajpal and Goyal 2009). This gives a much clearer picture of the scale patterns than light microscopy methods, revealing different scale shapes, scale margin patterns, and accurate distances between scales.

It has been argued that the scale patterns are not enough to accurately identify mammalian species to a taxonomic level finer than genus (Short 1978, Taru and

Backwell 2014). The purpose of this catalog is not to address this point, but rather to provide a clear reference system for those who are looking to use scale patterns to aid in the identification of African game species from hair samples.

Species Studied

This catalog was created in conjunction with another study, in which the feces of apex predators within the Balule Nature Reserve of the Greater Kruger National Park region of South Africa were examined to determine which prey species were consumed.

Therefore, only the larger, more prevalent species found within that area are included.

We photographed guard hairs or bristle hairs from 17 species (Table I). Classifications follow Wilson and Reeder (2005):

Scientific Name Common Name

Aepyceros melampus—Lichtenstein 1812 impala

Connochaetes taurinus—Burchell 1824 blue wildebeest

Kobus ellipsiprymnus—Ogilby 1833 waterbuck

Oreotragus oreotragus—Zimmerman 1783 klipspringer

Oryx gazella—Linnaeus 1758 gemsbok

Raphicerus campestris—Thunberg 1811 steenbok

Sylvicapra grimmia—Linnaeus 1758 bush duiker

Syncerus caffer—Sparrman 1779 African buffalo

Tragelaphus angasii—Angas 1849 nyala

Tragelaphus scriptus—Pallas 1766 bushbuck

Tragelaphus strepsiceros—Pallas 1766 greater kudu

Giraffa camelopardalis—Linnaeus 1758 giraffe

Phacochoerus africanus—Gmelin 1788 warthog

Panthera leo—Linnaeus 1758 lion

Panthera pardus—Linnaeus 1758 leopard

Crocuta crocuta—Erxleben 1777 spotted hyaena

Equus burchellii—Gray 1824 Burchell’s zebra

Table 1 Species whose hairs were photographed with SEM

Materials and Methods

Most hairs were collected from specimens in a taxidermy office located in

Phalaborwa, Limpopo Province, South Africa. Lion hairs and warthog hairs were collected from the Louisville Zoo, located in Louisville, Kentucky, USA. Bushbuck, gemsbok, and leopard hairs were donated by the Thomas M. Baker collection of Bowling

Green, Kentucky, USA. Guard hairs were collected from the dorsoscapular, scapular,

5 chest, or axillary region of each animal, as was done in previous studies (Homan and

Genoways 1978, Short 1978, Keogh 1983, Hess et al. 1985, Seiler 2010). If bristle hairs were present, they were collected as well. The bristle hairs were always taken from the mane of the species.

Hair shafts were soaked in 91% isopropyl alcohol for about one hour for cleaning, and then soaked in distilled water for 3-5 minutes to rinse. After air-drying, they were prepared for observation under a scanning electron microscope. They were mounted onto stubs using very smooth tabs (Cat #16084-20, Ted Pella) and sputter-coated with gold- palladium alloy (60% gold, 40% palladium) for three minutes. The stub was then turned

180° and again sputter-coated with the gold palladium alloy. Whole hairs were mounted; in some cases long hairs had to be mounted in sections to fit on the stubs.

Samples were observed using a JEOL JSM-6510LV scanning electron microscope at 10 kV and 500x magnification (unless otherwise specified) and photographed in three different locations—the proximal, medial, and distal regions. These settings gave the best resolution with the least amount of contrast on the edges of hairs. The same magnification was used for most hairs to give a reference for size differences among hairs and among different regions of the same hair. If more detail was needed, or if the hair was too big for the frame, the magnification was altered.

Results

Figure 1 SEM image of impala (Aepyceros melampus) guard hair

Figure 2 SEM image of impala (Aepyceros melampus) guard hair

Figure 3 SEM image of blue wildebeest (Connochaetes taurinus) guard hair

Figure 4 SEM image of waterbuck (Kobus ellipsiprymnus) guard hair

Figure 5 SEM image of klipspringer (Oreotragus oreotragus) guard hair

Figure 6 SEM image of gemsbok (Oryx gazella) guard hair

Figure 7 SEM image of steenbok (Raphicerus campestris) guard hair

Figure 8 SEM image of common duiker (Sylvicapra grimmia) guard hair

Figure 9 SEM image of African buffalo (Syncerus caffer) guard hair

Figure 10 SEM image of male nyala (Tragelaphus angasii) guard hair

Figure 11 SEM image of male nyala (Tragelaphus angasii) bristle hair

Figure 12 SEM image of bushbuck (Tragelaphus scriptus) guard hair

Figure 13 SEM image of greater kudu (Tragelaphus strepsiceros) guard hair

Figure 14 SEM image of giraffe (Giraffa camelopardalis) guard hair

Figure 15 SEM image of giraffe (Giraffa camelopardalis) bristle hair

Figure 16 SEM image of warthog (Phacochoerus africanus) guard hair

Figure 17 SEM image of lion (Panthera leo) guard hair

Figure 18 SEM image of lion (Panthera leo) bristle hair

Figure 19 SEM image of leopard (Panthera pardus) light guard hair

Figure 20 SEM image of leopard (Panthera pardus) dark guard hair

Figure 21 SEM image of spotted hyaena (Crocuta crocuta) guard hair

Figure 22 SEM image of Burchell's zebra (Equus burchellii) guard hair

Figure 23 SEM image of Burchell's zebra (Equus burchellii) bristle hair

Discussion

We have presented the cuticular scale patterns of hair from 17 mammalian species commonly found in the South African lowveld. In most cases, the process of Seiler

(2010) was followed, and the three regions of hair shafts (proximal, medial, and distal) were photographed to display changes in the scale pattern along the length of the hair.

Our samples of male nyala’s bristle hair, warthog’s guard hair, and Burchell’s zebra’s guard hair, had damage to the hair shaft that destroyed the cuticle. In these cases, certain regions of the hair could not be photographed. Seiler (2010) also noted damaged, unrecognizable cuticle patterns in a number of her hair samples. She attributed this damage to museum storage. Keogh (1983), however, records that hairs collected from specimens in museum storage do not show a significant difference from those that are collected from live specimens.

Most samples were collected from the dorsoscapular area of each animal. Lion guard hairs were collected both from lion feces gathered by the Louisville Zoo in

Louisville, Kentucky, USA, and from the axillary region of a taxidermic specimen from the Thomas M. Baker collection of Bowling Green, Kentucky, USA. Hairs from the axillary region had no cuticular scale patterns. It is uncertain if this is natural or caused by damage from preparing the specimen for taxidermy. Only guard hairs collected from the fecal sample were photographed. These hairs were swallowed during grooming.

Because these guard hairs were collected from feces, it is unsure from which part of the body these hairs originated. Also, leopard hairs were collected from the lower left flank, and gemsbok and bushbuck guard hairs were collected from the sternal region.

As noted in similar studies, a limiting factor of this study was the lack of hairs collected from different body regions of each species. No hairs were collected from the posterior, ventral, or cranial regions of the animals. There are also no scale patterns representing hairs from juveniles; all hairs came from adults.

It is important to note that impala has two different types of scale patterns. The first scale pattern (Figure I) appears to be streaked, with a raised surface running down the center of the hair shaft. The second (Figure II) appears to be imbricate overall, with smooth, yet irregularly waved scales. Both are unique to the impala species and make this species easily identifiable. Klipspringer hair is unmistakable, with several scales along the entire length of the hair in a mosaic, petal-like pattern. Steenbok hair is also unique, with scales appearing mosaic proximally and slowly transitioning into a smooth, coronal shape. Duiker and lion guard hairs are both coronal throughout the length of the shaft, and medullary characteristics are necessary to distinguish between the two. Giraffe guard hairs can be recognized by their regularly waved scales. Bristle hairs of all species—as well as guard hairs for buffalo, warthog, wildebeest, and waterbuck—can be distinguished by their large size, as well as the close proximity and very rippled appearance of scale margins. Guard hairs of greater kudu, bushbuck, leopard, hyaena, and zebra are all characterized by close, rippled scale margins, as well, but the hair shaft tends to be thinner than in the species previously listed. We did also observe hairs from each species at 10 kV and 10,000x magnification to search for distinguishing characteristics at the extremely microscopic level. However, no additional distinctive features were found to discriminate among different species. To produce a fully diagnostic key, the scale

31 patterns found in this study should be studied along with medullary characteristics, color patterns, dimensions, and other diagnostic features of known reference hairs.

CHAPTER 2

DIETS OF TWO APEX PREDATORS IN A PERIOD OF DROUGHT THROUGHOUT

THE BALULE NATURE RESERVE, HOEDSPRUIT, SOUTH AFRICA

Introduction

Population control is a very important aspect of conservation. One way to manage the populations of species is to set hunting quotas for each of the various species within a certain area. Hunting quotas are largely determined based upon the abundance of species, as well as the hunting patterns of nonhuman predators. In the Balule Nature Reserve of

South Africa, there are five apex predators which often pass through the area. These are lion (Panthera leo), spotted hyaena (Crocuta crocuta), leopard (Panthera pardus), cheetah (Acinonyx jubatus), and wild dog (Lycaon pictus). This study seeks to determine the diet compositions of two of these apex predators in Balule Nature Reserve, South

Africa: the lion and spotted hyaena.

When hunting, apex predators demonstrate both preference and avoidance of different species. Selective feeding takes place if a predator consumes equally common species at different rates (Jacobs 1974). Preferences in diets occur when species are consumed more than expected based on their relative abundance within an area (Hayward and Kerley 2005, 2008). When a species is avoided as prey by a predator, the species is

33 consumed less than expected based on its relative abundance (Hayward and Kerley 2005,

2008). The diets of apex predators are largely dependent upon the mass of prey, relative abundance, herd size, and evolutionary adaptations that render a species more or less susceptible to predators (Hayward and Kerley 2005, Hayward 2006, Rapson and Bernard

2007, Mbizah 2012, Clements et al 2014).

Lions prefer prey species that fall between the range of 92 kg and 632 kg and avoid species that are < 32 kg and > 632 kg (Clements et al 2014). Both lions and hyaenas have rich, diverse diets, and have a large dietary overlap (Hayward and Kerley

2008). They both fulfill the upper niche of Africa’s predatory guild, and hyaenas are also often seen scavenging at lion kill sites (Hayward and Kerley 2008).

Spotted hyaenas prefer prey species between 91 kg and 139 kg and avoid species

< 15 kg. Spotted hyaenas are the only apex predators that do not have a tendency to avoid prey above a certain mass (Hayward 2006, Clements et al 2014). This is most likely because hyaenas are often scavengers (Breuer 2005, Hayward 2006, Trinkel 2010,

Clements et al 2014), so prey that they do not kill will also appear in their scat. It could also be due to their ability to hunt in both group and solo fashions (Breuer 2005,

Hayward 2006).

On average, the Balule Nature Reserve receives 243.4 mm of rainfall between the months of January and June (Olifants West 2015). In 2014, the area received 317.4 mm of precipitation within the same amount of time (Olifants West 2015). In the first six months of 2015, the Balule Nature Reserve experienced a drought, with only 127.6 mm of rainfall (Olifants West 2015). I compared the diets of lions and spotted hyaenas to a similar study conducted in 2014 to observe changes in diets of these species within a

34 drought period. Prey species have been shown to adjust their behavior by increasing herd size and vigilance while at waterholes during times of drought, when the risk of encountering predators is increased (Valiex et al 2009). Therefore, it can be reasonably hypothesized that the diets of lions and spotted hyaenas will change significantly during this period of drought, due to changes in behavior of both predators and prey. The results of this study will also contribute to a long-term study of predator/prey relationships within the Balule Nature Reserve’s ecosystem.

Study Area

This study took place in Balule Nature Reserve, which is 60 km southwest of

Phalaborwa and 17 km north of Hoedspruit in the Limpopo Province of South Africa

(Olifants 2013). The nature reserve covers 40,000 hectares and makes up the western boundary of the Greater Kruger National Park, as well as the Association of Private

Nature Reserves (APNR) (Olifants 2013). There are nine administrative units within the

Balule Nature Reserve: Grietjie, Mohlabetsi River (MRNR), Mohlabetsi South, Olifants

East (OREC), Olifants North (ON), Olifants South (OS), Olifants West (OW), Parsons, and York. Over the past 30 years, Balule has received an average annual rainfall of a little over 415 mm, which places it within the category of a “semi-arid savannah biome”

(Olifants 2010, page 5; Olifants 2011, page 3). While the landscape and topography within and between the regions of Balule can fluctuate, Tupsuvihniö (Aristida congesta) and Curly Leaf (Eragrostis rigidior) have the highest frequency of occurrence among grass species (Olifants 2014, page 23). Trees dominating the landscape consist of

Knobthorn (Acacia nigrescens), Raisin Bush (Grewia spp.), Red Bushwillow

(Combretum apiculatum), and Corkwood (Commiphora spp.) (Olifants 2014a).

Figure 1 Map of Balule Nature Reserve and corresponding management districts; image from http://www.olifantsreserve.co.za/documents/Balule%20Management%20Regions.jpg

Methodology

Fecal Collection In 2014, the Balule Nature Reserve adopted a systematic method for collecting fecal samples, created by Audrey N. Sohikian (Olifants 2014b, page 46-68), in which 30 plots were chosen within the reserve, and three transects were walked within each plot. This year, 16 of those plots were sampled, each measuring one square kilometer. Three one-kilometer transects were walked in every plot. Transects were

36 separated by 500 meters. Plots were hiked between June 8, 2015, and August 6, 2015.

Anywhere from two to five observers walked along the different transects, scanning the ground for predator feces. Apex predator feces are easily distinguished from other scat by their size, white appearance, and presence of hair. When scat was found, it was placed in a plastic sealable bag, labeled, and the GPS coordinates were recorded. Scats were identified with the assistance of field guides, as well as a scat identification book (Murray

2011).

Figure 2 Plots sampled in 2014 (Olifants 2014b)

Figure 3 Plots sampled in 2015

Fecal Analysis To analyze all fecal samples without bias, it was necessary to develop a sampling method to randomly select hairs for identification. Similar to Eagle and Pelton

(1983), Hellgren (1993), Marker et al (2003), Ott et al (2006), and Martins et al (2011), a grid was created on a wooden board with 24 columns and 12 rows, totaling 288 cells with an area of 9 cm2 each. Fecal samples were crumbled by hand, and evenly spread out along to grid. Using a random number generator (random.org), five cells were chosen, and all hairs were collected from these cells.

Figure 4 Grid used for subsampling method

After collecting hairs from the grid, they were distributed evenly in a petri dish with 25 labeled points on the bottom. The hair closest to each point was chosen for identification. If the sample contained fewer than 25 hairs, all hairs were identified. Hairs were placed in distilled water for one hour and then moved to 91% isopropyl alcohol for a few minutes for cleaning (Sahajpal and Goyal 2009). After completing 24 samples, it was determined that no more than 18 hairs were necessary for identifying all species within a sample.

Figure 5 Petri dish with 18 points, used for hair selection

After cleaning, hairs were placed on a microscope slide with a drop of distilled water, and a cover slip was placed on top. They were viewed under a WILD M20-90715 microscope to observe the medulla and cortex. Cuticular scale imprints were made for each hair by spreading a thin layer of Henkel NB Multi-Purpose Cement (part #1589347) on a microscope slide and pressing the unknown hair overtop. The glue was allowed to dry, hairs were pulled off using a pair of thin-point tweezers, and the scale impressions were viewed under the microscope. Medullary characteristics and scale patterns were compared to reference materials (Keogh 1983, Backwell et al 2009, Seiler 2010, Taru and

Backwell 2013, Taru and Backwell 2014). A catalog of reference hairs from Limpopo

Taxidermy, the Louisville Zoo (Kentucky, USA), and the Thomas M. Baker Collection

(Kentucky, USA) were also used. These microscopic characteristics, along with some macroscopic characteristics, were used to identify hairs to the species level when possible. If more than one item from the same species was found in a scat sample, it was considered to be from the same individual animal.

Species Studied Only hairs from game species in the Balule Nature Reserve were identified. Classifications follow Wilson and Reeder (2005):

Scientific Name Common Name

Aepyceros melampus—Lichtenstein 1812 impala

Connochaetes taurinus—Burchell 1824 blue wildebeest

Kobus ellipsiprymnus—Ogilby 1833 waterbuck

Oreotragus oreotragus—Zimmerman 1783 klipspringer

Raphicerus campestris—Thunberg 1811 steenbok

Sylvicapra grimmia—Linnaeus 1758 bush duiker

Syncerus caffer—Sparrman 1779 African buffalo

Tragelaphus angasii—Angas 1849 nyala

Tragelaphus scriptus—Pallas 1766 bushbuck

Tragelaphus strepsiceros—Pallas 1766 greater kudu

Giraffa camelopardalis—Linnaeus 1758 giraffe

Phacochoerus africanus—Gmelin 1788 warthog

Panthera leo—Linnaeus 1758 lion

Panthera pardus—Linnaeus 1758 leopard

Crocuta crocuta—Erxleben 1777 spotted hyaena

Equus burchellii—Gray 1824 Burchell’s zebra

Table 1 Species whose hair was gathered for reference and characterized

Statistical Analysis Several different tests were used to analyze the diet compositions of both apex predators. First, the proportions for all species found in scat samples were determined by dividing the number of occurrences of one species found in all samples by the total occurrences of all species. These proportions were compared with the species availability determined by aerial game counts, as well as with the results from the study completed in 2014 (Olifants 2014b, page 46-68), using the Jacob’s Index (Jacobs 1974) and the g-test goodness-of-fit (Rohlf and Sokal 1981, Sokal and Rohlf 1981).

The Jacob’s Index was applied to each species found in scat samples to determine selected and non-selected prey species of both apex predators. The Jacob’s Index is as follows:

푟 − 푝 퐷 = 푟 + 푝 − 2푟푝 where r is equal to the proportion of a species found in scat samples, and p is equal to the proportion of a species found in game counts (Jacobs 1974). D values range from -1 to

+1, with values less than zero indicating a species was consumed less than expected based on its relative abundance, zero indicating prey was consumed equal to its relative abundance, and values greater than zero indicating prey was consumed more than expected based on its relative abundance.

The g-tests determined whether there were any significant differences in the total diet composition of either predator when comparing it to both the game counts and the results from 2014. The formula is as follows:

푂푖 푔 = 2 ∑ 푂푖 × ln 퐸푖

41 where Oi represents the number of a certain prey species observed in all samples, and Ei represents the number of times that same species would be expected to appear in the same number of samples based on the proportions from aerial game counts or 2014’s observations (Rohlf and Sokal 1981, Sokal and Rohlf 1981). This test was conducted twice for each predator—once to compare with the expected results coming from aerial game counts, and once to compare with the expected results coming from 2014’s results.

If any species was not detected in the fecal samples, they were omitted from the g-test. Therefore, when comparing the proportions of prey species found in lion scat in

2014 to proportions of prey species found in lion scat in 2015, steenbok, warthog, and wildebeest were omitted from the g-test equation. Wildebeest was not detected in any of the fecal samples found in 2014, and steenbok and warthog were not detected in any of the fecal samples tested in 2015. Because steenbok and warthog were not detected in any of the fecal samples in 2015, they were also omitted from the g-test when comparing the proportions of prey species found in lion scat in 2015 to the proportions of prey species found during aerial game counts. Nyala was not detected in hyaena scat in 2014, so it was omitted from the g-test when comparing proportions of prey species found in hyaena scat in 2014 to the proportions of prey species found in hyaena scat in 2015.

Results

In total, 87 fecal samples were collected between June 8, 2015, and August 6,

2015. Of the 87 samples collected, 68 scats contained hair that was able to be identified.

Hairs in exceptionally old, deteriorated samples were damaged past the point of

42 identification. Therefore, it should be noted that it is not of use to collect scat samples which have been exposed to much weathering and degradation (Breuer 2005, Yihune and

Bekele 2014).

Most scat samples were collected from the regions of Olifants South, Olifants

West, Jejane, and York. No scat samples were found within transects in the regions of

Parsons, Olifants North, and Greitjie. While some scat samples were spotted in these plots, they were not located within transects, and thus could not be collected. It would be wise to allow opportunistic scat collection while walking between different transects, to allow more opportunity for scat collection.

Figure 6 Distribution of all 87 predator scat samples collected in 2015

Of the 68 scat samples that could be identified, 34 scat samples were identified as hyaena scat and 34 were identified as lion scat. Tables 2 and 3 display the number of

43 scats containing hairs of each prey species consumed by each predator, and in which regions the fecal samples containing those prey hairs were located.

Lion prey consumption across all regions of Balule Nature Reserve

Jejane MRNR OS OW OREC York

bushbuck 1 0 0 0 0 0

duiker 0 0 1 0 0 2

giraffe 2 0 1 0 0 0

impala 4 2 3 1 0 11

kudu 0 1 0 1 1 2

nyala 1 1 1 0 0 1

steenbok 0 0 0 0 0 1

warthog 0 0 0 0 0 0

waterbuck 0 0 0 0 0 1

wildebeest 1 0 0 0 0 0

zebra 0 0 0 0 0 8

Table 2 Number of scats containing hairs of each prey species by region consumed by lion

Hyaena prey consumption across all regions of Balule Nature Reserve

Jejane MRNR OS OW OREC York

bushbuck 0 3 0 0 0 0

duiker 1 1 0 0 0 0

giraffe 0 2 0 0 0 0

impala 4 8 0 2 0 6

kudu 1 2 0 0 0 3

nyala 0 1 1 0 0 2

steenbok 0 0 1 0 0 0

warthog 0 0 1 0 0 0

waterbuck 3 0 0 1 0 3

wildebeest 0 1 0 2 0 0

zebra 1 0 0 0 0 9

Table 3 Number of scats containing hairs of each prey species by region consumed by hyaena

When comparing proportions of species consumed found in lion scat, impala and zebra made up the greatest proportions. Steenbok and warthog were totally absent. These proportions were compared with the proportions of species found during aerial game counts, as well as the results found in the 2014 study.

Proportions of prey species found in lion fecal samples in 2014 and 2015 and in aerial game counts 1

0.5

0 1 2 3 4 5 6 7 8 9 10 11

Proportion of species found in lion fecal samples in 2014 Proportion of Species Consumed Species of Proportion Proportion of species found in lion fecal samples in 2015 Proportion of species found during aerial game counts

Figure 7 Bar graph comparing proportions of species found in lion fecal samples in 2014, 2015, and aerial game counts

Proportions of prey species found in hyaena fecal samples were also compared to proportions found during aerial game counts and results from the study conducted in

2014. Similar to the lion, impala and zebra also made up the largest proportion of species found in hyaena diets for 2015.

Proportions of prey species found in hyaena fecal samples in 2014 and 2015 and in aerial game counts 1 0.8 0.6 0.4 0.2 0 1 2 3 4 5 6 7 8 9 10 11

Proportion of species found in hyaena fecal samples in 2014

Proportion of Species Consumed Species of Proportion Proportion of species found in hyaena fecal samples in 2015 Proportion of species found during aerial game counts

Figure 8 Bar graph comparing proportions of species found in hyaena fecal samples in 2014, 2015, and in aerial game counts

These proportions were used when calculating the Jacob’s Index for each species consumed by each predator. The Jacob’s Index is shown here for both 2014 and 2015 for an easy comparison between studies.

Comparison of Jacob's Index Values Between Lion and Hyaena in 2014 1

0.5

-0.5

-1

Axis Title Jacob's Index Values for All Species All for Values Index Jacob's Lion Hyaena

Figure 9 Jacob's Index values for all species in lion and hyaena diets in 2014

Comparison of Jacob's Index Values Between Lion and Hyaena in 2015 1 0.8 0.6 0.4 0.2 0 -0.2 -0.4 -0.6 -0.8 -1

Jacob's Index Values for All Species All for Values Index Jacob's Lion Hyaena

Figure 10 Jacob's Index values for all species in lion and hyaena diets in 2015

These proportions were also used to calculate g-values to compare proportions of different prey species in the scat of each predator species to both aerial game counts and the results from the 2014 study. The results are shown in Tables 4 and 5.

Lion compared with… G-statistic P-value Significance?

Game Counts g = 56.477 p < 0.001 Significantly different

Year 2014 g = 26.946 p < 0.001 Significantly different

Table 4 G-statistic results for the African lion when compared to aerial game counts and

results from the 2014 study

Hyaena compared with… G-statistic P-value Significance?

Game Counts g = 73.395 p < 0.001 Significantly different

Year 2014 g = 55.655 p < 0.001 Significantly different

Table 5 G-statistic results for the spotted hyaena when compared to aerial game counts

and results from the 2014 study

Light Microscopy Images

Images were taken using a light microscope. If the appearance of the hair shaft or scale pattern changed along the length of the hair, multiple images are displayed and labeled according to the region of the hair shown in the image.

Medulla Scale Patterns

impala (Aepyceros melampus) impala (Aepyceros melampus)

Figure 11 Brightfield image of impala Figure 12 Cast of impala (Aepyceros (Aepyceros melampus) guard hair medulla melampus) guard hair scale pattern pattern

49 blue wildebeest (Connochaetes taurinus) blue wildebeest (Connochaetes taurinus)

Figure 13 Brightfield image of blue Figure 14 Cast of blue wildebeest wildebeest (Connochaetes taurinus) (Connochaetes taurinus) guard hair scale proximal region of guard hair medulla pattern pattern

Figure 15 Brightfield image of blue wildebeest (Connochaetes taurinus) distal region of guard hair medulla pattern

50 waterbuck (Kobus ellipsiprymnus) waterbuck (Kobus ellipsiprymnus)

Figure 146 Brightfield image of Figure 17 Cast of waterbuck (Kobus waterbuck (Kobus ellipsiprymnus) ellipsiprymnus) guard hair scale pattern proximal region of guard hair medulla pattern

Figure 18 Brightfield image of waterbuck (Kobus ellipsiprymnus) distal region of guard hair medulla pattern

klipspringer (Oreotragus oreotragus) klipspringer (Oreotragus oreotragus)

Figure 19 Brightfield image of Figure 20 Cast of klipspringer klipspringer (Oreotragus oreotragus) (Oreotragus oreotragus) guard hair scale guard hair medulla pattern pattern

steenbok (Raphicerus campestris) steenbok (Raphicerus campestris)

Figure 21 Brightfield image of steenbok Figure 22 Cast of steenbok (Raphicerus (Raphicerus campestris) guard hair campestris) guard hair scale pattern medulla pattern

duiker (Sylvicapra grimmia) duiker (Sylvicapra grimmia)

Figure 23 Brightfield image of duiker Figure 24 Cast of duiker (Sylvicapra (Sylvicapra grimmia) guard hair medulla grimmia) proximal region of guard hair pattern scale pattern

Figure 25 Cast of duiker (Sylvicapra grimmia) medial/distal region of guard hair scale pattern

African buffalo (Syncerus caffer) African buffalo (Syncerus caffer)

Figure 26 Brightfield image of African Figure 27 Cast of African buffalo buffalo (Syncerus caffer) proximal region (Syncerus caffer) guard hair scale pattern of guard hair medulla pattern

Figure 28 Brightfield image of African buffalo (Syncerus caffer) distal region of guard hair medulla pattern

nyala (Tragelaphus angasii) nyala (Tragelaphus angasii)

Figure 29 Brightfield image of nyala Figure 30 Cast of nyala (Tragelaphus (Tragelaphus angasii) bristle hair medulla angasii) bristle hair scale pattern pattern

Figure 31 Brightfield image of nyala Figure 32 Cast of nyala (Tragelaphus (Tragelaphus angasii) guard hair medulla angasii) guard hair scale pattern pattern

bushbuck (Tragelaphus scriptus) bushbuck (Tragelaphus scriptus)

Figure 33 Brightfield image of bushbuck Figure 34 Cast of bushbuck (Tragelaphus (Tragelaphus scriptus) guard hair medulla scriptus) guard hair scale pattern pattern

greater kudu (Tragelaphus strepsiceros) greater kudu (Tragelaphus strepsiceros)

Figure 35 Brightfield image of greater Figure 36 Cast of greater kudu kudu (Tragelaphus strepsiceros) guard (Tragelaphus strepsiceros) guard hair hair medulla pattern scale pattern

giraffe (Giraffa camelopardalis) giraffe (Giraffa camelopardalis)

Figure 37 Brightfield image of giraffe Figure 38 Cast of giraffe (Giraffa (Giraffa camelopardalis) bristle hair camelopardalis) bristle hair scale pattern medulla pattern

Figure 39 Brightfield image of giraffe Figure 40 Cast of giraffe (Giraffa (Giraffa camelopardalis) proximal region camelopardalis) guard hair scale pattern of guard hair medulla pattern

giraffe (Giraffa camelopardalis) giraffe (Giraffa camelopardalis)

Figure 41 giraffe (Giraffa camelopardalis) distal region of guard hair medulla pattern

warthog (Phacochoerus africanus) warthog (Phacochoerus africanus)

No image available No image available

lion (Panthera leo) lion (Panthera leo)

Figure 42 Brightfield image of lion Figure 43 Cast of lion (Panthera leo) (Panthera leo) dark bristle hair medulla bristle hair scale pattern pattern

Figure 44 Brightfield image of lion Figure 45 Cast of lion (Panthera leo) (Panthera leo) light bristle hair medulla guard hair scale pattern pattern

lion (Panthera leo) lion (Panthera leo)

Figure 15 Brightfield image of lion (Panthera leo) guard hair medulla pattern

leopard (Panthera pardus) leopard (Panthera pardus)

Figure 47 Brightfield image of leopard Figure 48 Cast of leopard (Panthera (Panthera pardus) guard hair medulla pardus) guard hair scale pattern pattern

spotted hyaena (Crocuta crocuta) spotted hyaena (Crocuta crocuta)

Figure 49 Brightfield image of spotted Figure 50 Cast of spotted hyaena hyaena (Crocuta crocuta) (Crocuta crocuta) guard hair scale pattern proximal/medial region of guard hair medulla pattern

Figure 51 Brightfield image of spotted hyaena (Crocuta crocuta) distal region of guard hair medulla pattern

Burchell’s zebra (Equus burchellii) Burchell’s zebra (Equus burchellii)

Figure 52 Brightfield image of Burchell's Figure 53 Cast of Burchell's zebra (Equus zebra (Equus burchellii) dark bristle hair burchellii) bristle hair scale pattern medulla pattern

Figure 54 Brightfield images of Burchell's Figure 55 Cast of Burchell's zebra (Equus zebra (Equus burchellii) white bristle hair burchellii) guard hair scale pattern medulla pattern

Burchell’s zebra (Equus burchellii) Burchell’s zebra (Equus burchellii)

Figure 16 Brightfield image of Burchell's zebra (Equus burchellii) guard hair medulla pattern

Discussion

Based on hairs found in feces, African lions’ diets on Balule consist of impala

(44%), zebra (19%), kudu (10%), and nyala (8%). Zebra (25%) and impala (13%) constituted the largest fractions of African lions’ diets in the 2014 study as well. (Olifants

2014b). Prey species making up the largest fraction of spotted hyaena diets during this study include impala (33%), zebra (17%), waterbuck (12%), and kudu (10%). In 2014, spotted hyaena diets were composed of zebra (22%), warthog (19%), buffalo (11%), and impala (10%). There is similarity between years 2014 and 2015 of the simple frequencies of prey species that comprised the diets of both apex predators. However, these data are inconclusive unless compared with abundance of prey species within the Reserve, which was the purpose of the Jacob’s Index.

The results of the Jacob’s Index calculations for African lions indicate that lions are consuming nyala, duiker, zebra, and giraffe more than expected. Impala, despite making up the largest portion of prey species found in lion fecal samples, were significantly less represented than expected when compared to their abundance in the

Reserve. Steenbok and warthog were completely absent from any fecal samples deposited by lions. Previous studies have shown lions consuming prey species in the weight range of 32 - 632 kg at a rate greater than or equal to their abundance in the environment

(Clements et al 2014). Nyalas have a weight range between 55 and 126 kg, duikers weigh between 15 and 25 kg, zebras weigh between 300 and 320 kg, and giraffes weigh between 800 and 1200 kg (Kruger 2016). Discrepancies found between this study and the literature could be because we found many samples within close proximity to each other, indicating that many identified species found in separate fecal samples could have originated from the same individual animal.

Jacob’s Index values for spotted hyaena diets indicate that hyaenas consume nyala, bushbuck, zebra, and duiker more frequently than expected based on their abundance in the environment. Impala were also significantly less represented in hyaena diets than expected when compared to aerial game counts. Previous studies have indicated a tendency for spotted hyaenas to consume prey species with a mass of 15 kg and above at rates greater than or equal to their abundance in the environment (Clements et al 2014). This tendency was consistent with this study, as nyala weigh between 55 and

126 kg, bushbuck weigh between 30 and 77 kg, zebra weigh between 300 and 320 kg, and duikers weigh between 15 and 25 kg (Kruger 2016). The degree of dietary overlap

66 between the spotted hyaena and the African lion agrees with results found in previous studies (Hayward and Kerley 2008).

Significant differences were found with both apex predator diets when compared to results from 2014. Jacob’s Index values during both years were similar for bushbuck, duiker, impala, nyala, and zebra, but that is where the similarities end. Results from 2015 indicate both apex predators consume giraffe and wildebeest more than expected relative to their abundance, while in 2014 they received low Jacob’s Index values. In 2014, warthog was consumed more than expected by both apex predators, whereas neither apex predator showed a preference for them during this study. Besides the significant differences found in proportions, both apex predators consumed the same prey species at rates greater than expected during both years. During both studies, lions consumed nyala, duiker, and zebra more than expected. They also consumed waterbuck and impala at rates lower than expected when compared with their abundance. Hyaenas consumed bushbuck, duiker, and zebra more than expcted throughout both studies and consumed impala significantly less than expected during both 2014 and 2015.

Information gathered in this study is used to make educated decisions on hunting quotas for hunters on the Balule Nature Reserve. Animals on the Balule Nature Reserve are managed to maximize profits while maintaining species’ populations at sustainable levels. When creating hunting quotas, it is important to note the hunting patterns of predators, as well as the relative abundance of all species within the Reserve (Caro et al

1998). If a prey species with a lower abundance on the Reserve is consumed at a higher rate by carnivorous predators (i.e. zebra), the hunting quota for that species should be lower. If a prey species with a higher abundance on the Reserve is consumed at a lower

67 rate by carnivorous predators (i.e. impala), the hunting rate for that species should be higher. It is also important to consider the age and sex structure of populations (Caro

2009). Lastly, the management of large nature reserves such as Balule should take into account the effects of illegal hunting on the population when determining offtake hunting quotas (Caro et al 1998).

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