Space Use of African Wild Dogs in Relation to Other Large

Home , Iberian lynx, N11 (South Africa), Spotted hyena

SPACE USE OF AFRICAN WILD DOGS IN RELATION TO OTHER LARGE

CARNIVORES IN HLUHLUWE-IMFOLOZI PARK, SOUTH AFRICA

Angela M. Darnell

A Thesis

Presented to

The Faculty of Humboldt State University

In Partial Fulfillment

Of the Requirements for the Degree

Master of Science

In Natural Resources: Wildlife

May, 2012

ABSTRACT

Space Use of African Wild Dogs in Relation to Other Large Carnivores in Hluhluwe- iMfolozi Park, South Africa

Angela M. Darnell

Interaction between species through competition is one of the principle processes shaping the structure of ecological communities. Competition can have significant effects on the behavior, distribution, and ultimately the population dynamics of species, particularly when the animals utilize similar resources. Extensive diet overlap between large African carnivores is associated with high levels of competition which is particularly evident in the interactions between lions (Panthera leo), spotted hyenas

(Crocuta crocuta) and African wild dogs (Lycaon pictus). Using GPS data collected from sightings of large carnivores in Hluhluwe-iMfolozi Park, South Africa, I tested the hypothesis that wild dogs’ space utilization was influenced by other large carnivores. I found that wild dogs had different responses to their two main competitors. They avoided lions, particularly during denning seasons, through a combination of spatial and temporal avoidance. However, the dogs did not alter space use relative to hyenas, likely because pack sizes were large enough to defend their kills adequately without the necessary effort of avoidance. Information from this study is important for managing current carnivore populations, especially as reintroductions and translocations are essential tools used for the survival of endangered African wild dogs.

iii

ACKNOWLEDGMENTS

I would like to thank my advisor, Dr. Micaela Szykman Gunther, for her knowledge and guidance of my project. I would like to thank her and Jan A. Graf for letting me use their data for this project. I would like to thank my committee members

Drs. Richard Golightly and Howard Stauffer for their expertise. I would like to thank Dr.

Dave Druce and the other staff of Hluhluwe-iMfolozi Park (HiP) as well as Ezemvelo

KZN Wildlife for allowing me to spend time in the park gaining additional information.

Funding for this project was provided by Humboldt State University Sponsored Programs

Foundation and the Institute for Wildlife Studies scholarship. My time in South Africa would not have been the same without my housemate (and often chauffer) Leanne Van der Weyde. I would also like to thank Zama Zwane for driving me around the park and for his invaluable knowledge of the wild dogs in HiP. I would also like to thank Dr.

Robert VanKirk for his statistical guidance. I am grateful to my mother, Darlene Darnell, for reading my drafts more times than she could count. I would like to thank my father,

Rodger Darnell, for always supporting me and challenging me to aim high. Finally, I would like to thank my lab-mates (particularly Hilary Cosby for editing all my drafts), fellow graduate students, friends and family for their love and support throughout my graduate school experience.

TABLE OF CONTENTS

ABSTRACT……………………………………………..…………………….…iii

ACKNOWLEDGMENTS……………………………………………………..…iv

TABLE OF CONTENTS…………………………………………………………v

LIST OF TABLES………………………………………………………….……vi

LIST OF FIGURES………………………………………………………….…..vii

LIST OF APPENDICES…………………………………………………...……vii

INTRODUCTION………………………………………………………………...1

METHODS………………………………………………………………………..7

Study Area………………………………………………………………...7

Data Collection…………………………………………………………..11

Spatial Interactions without Temporal Aspect………………………..….12

Spatial Interactions with Temporal Aspect…………………………..…..14

RESULTS………………………………………………………………………..20

Spatial Interacts without Temporal Aspect………………………………20

Spatial Interactions with Temporal Aspect………………………………28

DISCUSSION……………………………………………………………………34

LITERATURE CITED…………………………………………………………..42

LIST OF TABLES

Table Page

1 Notation descriptions used in text for calculating spatial and temporal interactions between African wild dogs, lions, and spotted hyenas in Hluhluwe-iMfolozi Park, South Africa, 2002-2004…..…………………,……..17

2 Comparisons of associations by co-occurrence for African wild dogs (α) and lions (β) in Hluhluwe-iMfolozi Park, South Africa, 2002-2004……….…….….29

3 Comparisons of association by co-occurrence for African wild dogs (α) and spotted hyenas (β) in Hluhluwe-iMfolozi Park, South Africa, 2003-2004..…....30

LIST OF FIGURES

Figure Page

1 Study area showing location of Hluhluwe-iMfolozi Park in KwaZulu- Natal Province with inset showing location of province within South Africa during study in 2002-2006….…………………………………………….…...... 8

2 Map of Hluhluwe-iMfolozi Park showing vegetation, major rivers and roads during study in 2002-2006…………………………………………...... 10

3 Average distance (in km) between neighboring non-overlapping African wild dog and lion core use areas (± SE) during denning (n=19) and not denning (n=10) seasons in Hluhluwe-iMfolozi Park, South Africa, 2002-2004...21

4 Mean percentage overlap (± SE) of home ranges of African wild dogs with lions (n=18) and spotted hyenas (n=10) between seasons in Hluhluwe-iMfolozi Park, South Africa, 2002-2004………………………………………...……...... 22

5 Home ranges of African wild dogs, lions, and spotted hyenas during the a) denning season, b) post-denning season, and c) non-denning season in 2004Hluhluwe-iMfolozi Park, South Africa……………………………….....….23

6 Mean percentage of overlap (±SE) of core use areas of African wild dogs with lions (n=18) and spotted hyenas (n=10) between seasons in Hluhluwe- iMfolozi Park, South Africa, 2002-2004………………………...………...…….25

7 Core use areas of African wild dogs, lions and spotted hyenas during the a) denning season, b) post-denning season, and c) non-denning season in 2004 in Hluhluwe-iMfolozi Park, South Africa…………...…….……………………26

8 Mean percentage volume of intersection of home ranges (± SE) of African wild dogs with lions (n=9) and spotted hyenas (n=5) between seasons using intensity of use (3-dimensional overlap) in Hluhluwe-iMfolozi Park, South Africa, 2002- 2004...…………………………………………………………………………….27

vii

LIST OF APPENDICES

Appendix Page

A Interaction within overlapping home ranges and core use areas for African wild dogs (A) and lions (B), showing spatial attraction or avoidance by each and interaction between groups (ixn) as well as deviation of odds from random in Hluhluwe-iMfolozi Park, South Africa, 2002-2004………………………………………….…...…54

B Interaction within overlapping home ranges and core use areas for African wild dogs (A) and spotted hyenas (B), showing spatial attraction or avoidance by each and interaction between groups (ixn) as well as deviation of odds from random in Hluhluwe-iMfolozi Park, South Africa, 2002-2004………………………………………………....55

viii

INTRODUCTION

Interaction between species through competition is one of the principle processes shaping the structure of ecological communities (Case and Gilpin 1974). Competition can have significant effects on the behavior, distribution, and ultimately the population dynamics of species (Case and Gilpin 1974, Chesson and Rosenzweig 1991, Creel et al.

2001). Interference competition in particular can play a major role in determining the abundance and distribution of competing species, and its effects usually have different impacts on the species involved (Case and Gilpin 1974, Linnell and Strand 2000, Creel et al. 2001). When animals utilize similar resources, larger competitors may exclude smaller competitors (Johnson et al. 1996).

Competition between carnivores has been considered a key ecological factor affecting carnivore species within the same guild (Caro and Stoner 2003). Past studies have focused on effects of carnivores on prey or vice versa, while ignoring the significant effects carnivores can have on each other (Fuller and Kat 1990, Fuller and Sievert 2001,

Owen-Smith and Mills 2008). Carnivores of the same guild may compete for similar prey resources, often resulting in smaller species either being excluded from or actively avoiding areas with higher densities of the larger competitor when in direct competition

(Johnson et al. 1996, Linnell and Strand 2000, Creel et al. 2001). In Nepal, leopards

(Panthera pardus) avoided habitats where tiger (Panthera tigris) densities were high

(McDougal 1988). Mills and Mills (1982) found a significant pattern of avoidance of spotted hyenas (Crocuta crocuta) by the smaller brown hyenas (Hyaena brunnea).

Studies have also suggested that gray wolves (Canis lupus) displace and exclude coyotes

(Canis latrans) from preferred habitat (Ballard et al. 2003). These studies illustrate the widespread pattern of avoidance and exclusion of smaller carnivores with less competitive advantage due to interference competition.

Displacement by larger carnivores can result in serious effects on smaller species

(Creel et al. 2001, Hunter and Caro 2008). Smaller competitors can be excluded from certain habitats with high prey density, reducing hunting opportunities (Creel et al. 2001,

Caro and Stoner 2003). The amount of prey available may decrease when the density of larger species is high, and rates of food stealing may increase, forcing smaller species to travel long distances to forage (Caro and Stoner 2003). These situations result in smaller species being forced to increase the time and energy expended to obtain sufficient food.

Increased time spent hunting limits the energy available for other essential behaviors, particularly reproduction. In addition, animals can be excluded from preferred breeding areas. Arctic foxes (Vulpes lagopus) reduced use of high quality dens during the breeding season when red foxes were near (Tannerfeldt et al 2002). Thus, interference competition can negatively affect fitness of smaller species with less competitive advantage.

Interference competition among carnivores is particularly hazardous to both carnivores due to their behavioral and morphological adaptations for killing (Creel et al.

2001). This is evidence of an extreme type of interference competition that has appeared in carnivores: intraguild predation (Palomares and Caro 1999, Creel et al. 2001, Donadio and Buskirk 2006). Competition due to overlap of diet between carnivores is one of the main precipitants for interspecific killing, with relative body size as the mediating factor

(Palomares and Caro 1999, Donadio and Buskirk 2006). Wolves commonly kill coyotes

(Ballard et al. 2003), and coyotes often kill kit foxes (Vulpes macrotis), yet the larger competitors rarely consume the carcasses (Nelson et al. 2007), suggesting that this type of predation is a product of competition rather than actual dietary needs. Rates of predation by larger species can be high enough to limit the size of populations by decreasing growth rates (Palomares and Caro 1999, Linnel and Strand 2000, Creel et al.

2001). Lions (Panthera leo) can significantly affect the population size of cheetahs

(Acinonyx jubatus) (Durant 1998); Laurenson (1995) found that over two-thirds of cheetah cub mortality was due to predation by lions.

Overall, interspecific competition between intraguild carnivores can confine spatial distributions, restrict habitat use, reduce prey encounter rates and food intake, and increase mortality of competitors (Creel et al. 2001). Smaller carnivores have developed two main ways of dealing with such competition: spatial and temporal avoidance. Using resources at different places or times allows smaller species to avoid direct interactions with their larger competitors.

Extensive diet overlap between large African carnivores is associated with high levels of competition (Mills and Biggs 1993, Owen-Smith and Mills 2008). This is particularly evident in the interactions between lions, spotted hyenas (hereafter referred to as hyenas) and African wild dogs (Lycaon pictus). Wild dogs are consistently found at lower population densities than any other sympatric carnivore (lions, hyenas, cheetahs, and leopards) (Creel et al. 2004). Interference competition from larger carnivores may be

affecting African wild dog movements, provoking active avoidance spatially and (or) temporally in the areas in which they range.

Wild dogs have distinctive social behaviors that make them an obligate cooperative species. Not only do they hunt more successfully in packs, but packs must have a minimum number of members to successfully reproduce (Courchamp et al. 2000,

Courchamp and Macdonald 2001). Courchamp and Macdonald (2001) suggested there is a minimum pack size for wild dogs. Below this critical threshold, the pack experiences the effect of inverse density dependence, which can lead to extinction of the population

(Somers et al. 2008). Within a pack usually only the alpha male and female reproduce, although subordinate males and females may breed on occasion (Estes 1991, Spiering et al. 2010). Other pack members may aid either through nursing or ‘baby-sitting’ when the pack is hunting as well as providing nourishment through food regurgitation for the pups and the dominant female (Estes and Goddard 1967). Breeding occurs once per year, with nearly equal-length periods for denning (pups restricted to den) and post-denning (pups out of den but too young to travel with the pack on hunts), while the remainder of the year the pack is more mobile and traveling all together.

Wild dogs are excellent hunters and predominantly prey on moderate-sized ungulates that weigh 2-10 times more than their average weight (Estes and Goddard

1967, Fuller and Kat 1990, Estes 1991). As social hunters, wild dogs have a higher hunting success than other large carnivores, and consequently, each pack member has a higher foraging success (Creel and Creel 1995), an important fact considering their high metabolic rate. Wild dogs’ daily energy expenditure is 5-6 times their basal metabolic

rate (Pole et al. 2003). Large pack size not only allows wild dogs to hunt more efficiently, but also allows them to prey on species that will be more energetically profitable (both in terms of species and size) as well as enhance defense of their kills from scavengers (Estes and Goddard 1967, Fanshawe and FitzGibbon 1993). All of the previously stated benefits decrease the quantity of required hunts, which reduces the pack’s energetic costs and risk

(Creel and Creel 1995, Gorman et al. 1998), increasing overall fitness (Gusset and

Macdonald 2010).

Wild dogs, once widespread across sub-Saharan Africa, are now endangered.

Throughout their range, wild dog populations are significantly declining and the species is in danger of extinction (Creel and Creel 1996, Fanshawe et al. 1997, Vucetich and

Creel 1999, Creel et al. 2004). Currently, six countries hold potentially viable populations of the species (Woodroffe et al. 2004). Reasons suggested for the species’ decline such as habitat fragmentation, persecution by humans, and disease affect all large carnivores in sub-Saharan Africa, yet wild dogs in particular continue to decline (Creel et al. 2001).

Because large carnivores are mostly confined to protected areas, they are forced to interact more frequently than they might have historically, which increases the effects of interference competition (Creel et al. 2001). This is especially true in smaller parks that are significantly separated from other populations, such as occurs in highly fragmented areas of South Africa.

As competition can cause a decline in the population size of smaller carnivores, this study has significant implications for wild dogs. Fully understanding the mechanisms causing a species’ decline is crucial for devising actions to solve the problem (to decrease

or reverse the decline) (Linnell and Strand 2000). Additionally, management of wild dog populations should be grounded in a thorough understanding of local threats (Woodroffe et al. 2007). To gain a better understanding of the potential threats to wild dog persistence in South Africa, I utilized a dataset previously collected in Hluhluwe- iMfolozi Park (HiP) in northern KwaZulu-Natal Province. The information was collected by researchers of the Hluhluwe-iMfolozi Predator Project in order to monitor the demographics of the carnivores in the park (Trinkle et al. 2008, Spiering et al. 2010).

African wild dogs were the priority of the project, followed secondarily by lions, and then hyenas, cheetahs, and leopards. The purpose of this project was to continually track locations of carnivores, monitor reproduction, and observe behavior. Researchers collected the above information concurrently on all wild dog packs and lion prides in the park.

Using the Global Positioning System (GPS) data collected from sightings of large carnivores in HiP, I tested the hypothesis that African wild dog space utilization is affected by other large carnivores. As lion populations in HiP are generally clustered and hyenas are more evenly distributed on the landscape (Maddock et al. 1996, Graf et al.

2009), I predicted that wild dogs’ space use would differ relative to their two main competitors. If wild dogs were adjusting their space utilization in response to sympatric carnivores, then evidence of spatial and (or) temporal avoidance would be apparent in their distribution and movements.

METHODS

Study Area

Hluhluwe-iMfolozi Park (Figure 1) is located between 28º00’ and 28º26’S and

31º43’ and 32º09’E in the northern KwaZulu-Natal Province, Republic of South Africa.

The park is approximately 960 km2 and is enclosed by an electrified fence. HiP is about

300 km south of Kruger National Park, which contains the closest historic population of wild dogs. HiP contains many roads, including a regularly used tarmac road which bisects the park. The landscape contains numerous hills and valleys ranging from 60 m to

750 m above sea level, with flat regions restricted to the areas adjacent to the four major rivers in the park: the Hluhluwe river in the north, which has three major tributaries (the

Manzibomvu, Mansiya and Nzsmani Rivers), the Nyalazi River in the south, and the

Black and White iMfolozi Rivers also in the south.

The subtropical climate of the park has unimodal rainfall peaking in summer from

November to February. The winter is usually very dry. Rainfall is the highest in the hills of north-western Hluhluwe and lower in the iMfolozi Game Reserve, ranging between

500-1,100 mm (Berkeley and Linklater 2010). Average temperatures are warm to hot, ranging from 13-35˚ C (Greyling and Huntley 1984).

The park is primarily savannah thornveld, within two veld types (Whateley and

Porter 1983, Acocks 1988): the Lowveld subcategory of the Tropical Bush and Savanna types and Zululand Thornveld, a subcategory of the Coastal Tropical Forest types.

Coastal forest communities are limited to the hills, particularly in Hluhluwe, while

Figure 1. Study area showing location of Hluhluwe-iMfolozi Park in KwaZulu-Natal Province with inset showing location of province within South Africa during study in 2002-2006.

woodlands are found in the river valleys and on the rocky and sandy hillsides (Figure 2).

Much of the park is dominated by shrubland Acacia spp., specifically A. karroo, A. nilotica, A. burkei, A. gerrardii, A. nigrescens, A. tortilis, and A. caffra. True grassland communities are mostly absent from the park.

The heterogeneous environment of HiP supports a large and diverse prey base, from warthogs (Phacochoerus africanus) to greater kudu (Tragelaphus strepsiceros), and as a result, a wide variety of both small and large predators, from jackals (Canis mesomelas) to lions. The carnivores in the park which belong to the same feeding guild as the wild dog are spotted hyenas, lions, leopards and cheetahs. Wild dogs in the park feed largely on nyala (Tragelaphus angasi) and impala (Aepyceros melampus). Despite the electrified fence, wild dogs can find ways to leave the park dispersing into the densely populated areas surrounding HiP.

The wild dog population in Hluhluwe-iMfolozi Park is a product of several reintroductions beginning in 1980-1981 (Andreka et al. 1999, Maddock 1999) with the most recent being in 2006. This population has undergone large fluctuations and is extremely susceptible to stochastic demographic or environmental events (Maddock

1999), making additional information about the population extremely valuable.

Figure 2. Map of Hluhluwe-iMfolozi Park showing vegetation, major rivers and roads during study in 2002-2006.

Data Collection

The large carnivores of HiP were monitored regularly from January 2002 through

December 2006 by researchers of the Hluhluwe-iMfolozi Predator Project. Individual wild dogs (at least 2 per pack in 6 or fewer packs) and lions (at least 1 per pride in 12 or fewer prides) were radio-collared (with transmitters by Sirtrack, Inc., New Zealand and

African Wildlife Tracking, South Africa) by HiP management staff for monitoring purposes. Several researchers and assistants monitored locations, movements and behaviors of all wild dog packs and all lion prides on a daily or weekly basis. One hyena was collared in 2005 and monitored until December 2006. Radio telemetry was conducted from a vehicle (using Telonics and Communication Specialist receivers and 2- and 3-element ‘H’ and yagi antennas). The order of priority of target species being endangered wild dogs, secondarily on lions, then opportunistically on hyenas. Visual sightings of all large carnivores were also recorded on an opportunistic basis: that is, locations for wild dog, lion, hyena, leopard and cheetah that were observed without the use of telemetry were recorded. Date, time and GPS location of the animals were recorded for all large carnivore sightings, as were number of animals observed, and their age, sex and behavior.

Because researchers were primarily interested in population health, demography and behavior (hunting, reproduction, etc.), approximately 90% of radio-tracking efforts resulted in a visual sighting of individuals and thus were accurate to within a few meters.

This obviously biased locations to more open areas and areas close to roads, precluding

the ability to assess habitat utilization or preferences for these species. However, as the park is heavily vegetated, the carnivores in HiP do use the roads preferentially (M. S.

Gunther, 2012, personal communication). In some cases (about 10%), locations of animals were determined via triangulation from distances of no more than 2 km.

In the summer of 2011, while in Hluhluwe-iMfolozi Park, I added to the current data set by checking all of the record logs and identifying sightings with a recorded location but lacking coordinates (approximately 300 locations). I found these locations in the field and identified coordinates for each to ensure the data set was accurate and complete. I then reviewed the entire dataset and removed all non-independent points (any points of the same pack, pride or clan sub-group that were within 12 hours). I used GPS coordinates of all independent points to create maps using ArcGIS (v 10.0,

Environmental Systems Research Institute, Redlands, California) to determine the spatial habits of the carnivores. The data were separated by year and because wild dogs exhibit distinct behavioral changes between seasons (Harris et al. 1990) also into three seasons of equal length: denning (May-Aug), post-denning (Sept-Dec), and non-denning (Jan-Apr).

Data for all of the following analyses were separated by pack, pride, or clan sub-group, because these carnivore groups move as a unit. Statistical tests were considered significant at alpha of 0.05 (Zar 1999).

Spatial Interactions without Temporal Aspect

I assessed the spatial interactions without a temporal aspect (static interactions) of the carnivores, which measured species interaction during a time interval of interest, using home ranges and core use areas, when least 50 locations within a season were

available (Seaman and Millspaugh 1999, Garton et al. 2001, Kernohan et al. 2001). Home ranges and core use areas were determined for each wild dog, lion and hyena group using a bivariate normal fixed-kernel estimator in Geospatial Modeling Environment (v 0.5) with smoothing factors calculated using a diagonal plug-in in R statistical software (v

2.14.1). From the kernel density layer, I used Geospatial Modeling Environment to obtain isopleth polygons: 95% for home range and 50% for core use areas.

When the home range and core use areas overlapped, I determined the mean

percentage of overlap as: ( ) (Minta 1992).

I used 2-sample t-tests (using paired t-tests when appropriate) to test for differences between percentage of overlap of home ranges and core use areas between species (wild dog versus lion and wild dog versus hyena) and ANOVA to test for differences between seasons.

I also determined 3-dimensional overlap in space use which takes into account a third dimension: intensity of use in an area. I used the kernel density raster layers, which reflected peaks of use within a home range, to obtain a volume of intersection:

∬ ̂ ̂

where UD 1 and 2 are the utilization distributions (the kernel density layers) for each species (Kernohan et al. 2001, Fieberg and Kochanny 2005). The volume of intersection

(3-dimensional overlap) measures the degree of overlap in shape and location of two utilization distributions. This index ranges from no overlap (0) to complete overlap (1). I

used 2-sample t-tests and ANOVA to test for differences between species and seasons for

2- and 3- dimensional overlap in home range and core use areas. I also used a paired t-test to test for a difference between 2- and 3- dimensional overlap between species.

For non-overlapping core use areas, I determined the average distance separating each species using centroids (the central, most heavily used point) of each core use area, for neighboring groups (packs, prides, or clan sub-groups).

Spatial Interactions with Temporal Aspect

When there was any overlap between carnivore home ranges, I analyzed dynamic spatial and temporal interactions. In contrast to the previous static interactions, dynamic interaction analyses incorporated the temporal aspect of the association between the species (Kernohan et al. 2001). Based on guidelines from Kernohan et al. (2001), I calculated the distance between simultaneous locations (defined as less than 12 hours) of two groups (i.e., pack, pride or clan sub-group) and compared the distances to what would be expected at random. I calculated observed distances (DO) as:

∑ √

where for n pairs of locations for each group (i.e., each pack), x1and x2 and y1 and y2 are the universal transverse mercator (UTM) coordinates (pair of points) for species 1 and 2, respectively. The expected distances (DE) for all recorded observations were calculated as:

∑ ∑ √

I combined all interactions by for each species group (each pack, pride and clan sub- group) and compared differences between observed (DO) and expected distances (DE) using a Wilcoxon signed-rank test (Zar 1999, Gorman et al. 2006). If there was a statistically significant difference between the observed and expected distances, I concluded that the species were expressing either attraction or avoidance. If the interaction was negative (observed distance > expected distance), avoidance was taking place, and if the interaction was positive (observed distance < expected distance), attraction was taking place.

I used methods recommended by Minta (1992) to further analyze spatial and temporal interactions between carnivore species. I tested the null hypothesis that for each group of species (i.e., pack, pride), α and β (see Table 1 for notation descriptions), one species moved randomly, using the overlap area independent of the other (Minta 1992). I tested the above hypothesis when the two species groups had any overlap in home range or core use area and where there were at least 30 independent points for each group within that overlap area. Locations for each group that had overlapping home ranges or core use areas within a season (den, post-den, non-den) were placed into one of the following categories: (1) both groups of species were absent from the overlapped area

(n11); (2) only species group α was present in the overlapped area (n21); (3) only species group β was present in the overlapped area (n12); (4) both groups of species were present in the overlapped area (n22). The above frequency statistics can be summarized as:

Table 1. Notation descriptions used in text for calculating spatial and temporal interactions between African wild dogs, lions, and spotted hyenas in Hluhluwe- iMfolozi Park, South Africa, 2002-2004.

Notation Description α species "A" β species "B" Observed frequencies n11 Both A and B simultaneously present in overlap area n12 Only B present in overlap area n21 Only A present in overlap area n22 Both A and B simultaneously absent in overlap area n+1 Sum of observed simultaneous sightings of A and B, with A present in the

overlap area (n11 + n21) n+2 Sum of observed simultaneous sightings of A and B, with A absent from

the overlap area (n12 + n22) n1+ Sum of observed simultaneous sightings of A and B, with B present in the

overlap area (n11 + n12) n2+ Sum of observed simultaneous sightings of A and B absent from the

overlap area (n21 + n22) Expected frequencies

Both A and B simultaneously present in overlap area p11

Only B present in overlap area p12

Only A present in overlap area ) p21

Both A and B absent in overlap area ( ) p22 p+1 Sum of expected simultaneous sightings of A and B, with A present in the

overlap area (p11 + p21) p+2 Sum of expected simultaneous sightings of A and B, with A absent from

the overlap area (p12 + p22) p1+ Sum of expected simultaneous sightings of A and B, with B present in the

overlap area (p11 + p12) p2+ Sum of expected simultaneous sightings of A and B absent from the

overlap area (p21 + p22)

 | Present Absent Present | n11 n12 | n1+ Absent | n21 n22 | n2+ ------n+1 n+2

Observations of species groups were considered ‘simultaneous’ if they were within 12 hours of each other. Expected frequencies of presence and absence in overlap areas were calculated using areas as recommended by Minta (1992), versus a contingency matrix proportion of frequencies. I then totaled the observed frequencies of presence and absence for each group and determined the expected frequencies using the proportion of overlapped area between the two species in relation to total home range area.

Presence and absence in overlap areas

I analyzed presence and absence of groups by calculating a comparison of association of co-occurrence using a cross-product ratio for each of the four categories

(n11, n12, n21, and n22) of observed and expected frequencies as follows:

̅ ̅

̅ ̅ with p as the probability and

as the estimate of ω. (Note that ω and ‘w’ can also be expressed as odds ratios). To compare the species’ frequency of synchronous (simultaneous) to asynchronous (one group present only) presence or absence within the overlapping area, I calculated the natural log of ‘w’ to obtain L. Thus L is the log of an odds ratio with a chi-square

distribution, with L = 0 indicating independence (or a lack of association between the two groups of species) (Minta 1992). The standard error of L was calculated as:

( )

with the tests statistic, df = 1:

Interactions within overlap areas

I compared expected and observed frequencies for each species to develop an

2 overall chi-square (χ tot). If the species were using overlapping areas independently, then

2 would be small. I partitioned the χ tot into three parts: spatial coefficient of species α, spatial coefficient of species β, and temporal interaction (ixn) coefficient (Mace and

Waller 1997). The effect was the combination of a spatial or temporal coefficient ( ̅

2 and ̅ or Lixn respectively) and a χ probability value. These values were defined as:

Spatial coefficient of species α; df = 1:

̅ ̅

Spatial coefficient of species β; df = 1:

̅ ̅

Temporal interaction (ixn) coefficient; df = 1:

The spatial coefficients indicated the species’ group (pack, pride, or clan sub-group) spatial association within the overlapping area of the home range and revealed attraction, avoidance, or random movement within the area. The temporal interaction effect indicated the group’s temporal association within the overlapping area of the home range.

This showed concurrent versus single group occupation of the overlap area in relation to overall home range use, revealing if temporal attraction or avoidance was evident. If there was a significant interaction, I determined if the species were exhibiting spatial attraction or avoidance, temporal attraction or avoidance, or random movement. If coefficients approached zero, it implied random use of the overlap zone (Mace and

Waller 1997).

RESULTS

Spatial Interactions without Temporal Aspect

There were 1,647 total independent wild dog pack locations for three packs, with

42%, 31%, and 27% from denning, post-denning, and non-denning seasons respectively;

1,466 independent lion pride locations for 12 prides with 39%, 33%, and 28% from each season, and 428 independent hyena clan sub-group locations with 43%, 38%, and 19% from each season. There were sufficient locations (at least 50 within a season) to analyze space use within nine seasons between wild dogs and lions and five seasons between wild dogs and hyenas. The average number of locations used to create home ranges for each group was: wild dog packs: 111.8 ± 20.8, lion prides: 189.1 ± 45.1, and hyena sub- groups: 70.0 ± 8.3. The number of locations used to determine home range was not correlated with the size of the home range (r33 = 0.229)

Wild dog packs remained an average of 16.6 ± 2.1 km away from neighboring lion prides during the denning season, whereas packs only maintained an average distance of 6.7 ± 1.6 km from lion prides during the other times of year (collectively ‘not denning’), (t26 = 3.76, p = 0.001; Figure 3). In contrast, the average distance from wild dogs to hyenas during the denning season (2.0 ± 1.9 km) was not significantly different from ‘not denning’ (2.5 ± 6.0 km; p = 0.894).

Overlap in home ranges was significantly lower during denning than non-denning season for wild dogs and lions (F17 = 6.85, p = 0.008; Figures 4, 5), but not for wild dogs and hyenas (p = 0.887). Overlap in core use areas did not differ significantly between

Figure 3. Average distance (in km) between neighboring non-overlapping African wild dog and lion core use areas (± SE) during denning (n=19) and not denning (n=10) seasons in Hluhluwe-iMfolozi Park, South Africa, 2002-2004. (Note: open circle at the top of “Denning” indicates the top value of the range of distances).

100 With lions

90 With hyenas 80 70 60 50 40 30

20 Overlap of home ranges (%) ranges home of Overlap 10 0 Denning Post-denning Non-denning

Figure 4. Mean percentage overlap (± SE) of home ranges of wild dogs with lions (n=18) and spotted hyenas (n=10) between seasons in Hluhluwe-iMfolozi Park, South Africa, 2002-2004.

a b ) )

a c ) c

)

c Figure 5. Home ranges of African wild dogs, lions, and spotted hyenas during ) the a) denning season, b) post-denning season, and c) non-denning season in 2004 in Hluhluwe-iMfolozi Park, South Africa.

seasons for any species (p = 0.635 with lions, p = 0.745 with hyenas; Figures 6, 7).

Overlap in core use areas was significantly less than overlap in home ranges for wild dogs and lions (t18 = -7.86, p < 0.001; compare Figure 5 to Figure 7), but there was no significant difference for wild dogs and hyenas (p = 0.052). Core use areas of wild dogs overlapped significantly more with hyenas than with lions (t7 = -3.34, p = 0.016; Figures

6, 7).

When taking into account intensity of use in an area, the volume of intersection of home ranges (3-dimensional overlap) between wild dogs and lions was significantly less than the mean (2-dimensional) overlap (t8 = -4.96, p = 0.001; Figure 8). However, there was no difference for wild dogs and hyenas (p = 0.223). There was no significant difference between the overlap with intensity of use (3-dimensional) of core use areas and home ranges of wild dogs with lions (p = 0.690). However the overlap for core use areas of wild dogs and hyenas was significantly higher than for home ranges (t3 = -3.95, p

= 0.029).

The 3-dimensional overlap of wild dog home ranges with lions was significantly less than the mean (2-dimensional) overlap for all seasons: denning (t6 = 4.89, p = 0.001), post-denning (t5 = 2.31, p = 0.035), and non-denning (t2 = 3.03, p = 0.047). However, there was no significant difference between seasons of mean (2-dimensional) and 3- dimensional overlap in home ranges of wild dogs with hyenas (p = 0.959, 0.750, and

0.797 for denning, post-denning, and non-denning, respectively). There was also significantly less 3-dimensional overlap of wild dogs with lions during the denning

100 With lions

80 With hyenas 70 60 50 40 30

Overlap of core use core of (%) Overlap 20 10 0 Denning Post-denning Non-denning

Figure 6. Mean percentage of overlap (±SE) of core use areas of wild dogs with lions (n=18) and spotted hyenas (n=10) between seasons in Hluhluwe-iMfolozi Park, South Africa, 2002-2004.

a b )

c )

Figure 7. Core use areas of African wild dogs, lions, and spotted hyenas during the a) denning season, b) post-denning season, and c) non-denning season in 2004 in Hluhluwe-iMfolozi Park, South Africa.

100 With lions

80 With hyenas 70 60 50 40 30

Volume of intersection (%) intersectionof Volume 20

10 0 Denning Post-denning Non-denning

Figure 8. Mean percentage volume of intersection of home ranges (± SE) of wild dogs with lions (n=9) and spotted hyenas (n=5) between seasons using intensity of use (3- dimensional overlap) in Hluluwe-iMfolozi Park, South Africa, 2002-2004.

season compared to ‘not denning’ (post-denning and non-denning seasons combined) (t5

= -2.26, p = 0.037), but this pattern did not hold with hyenas (p = 0.989).

Spatial Interactions with Temporal Aspect

There was no significant difference between observed and expected distances between wild dogs and lions over all seasons (p = 0.620), but wild dogs were significantly further from lions than expected when comparing denning to not denning seasons (t7 = 2.04, p = 0.04). Hyenas were significantly closer to wild dogs than expected overall (Ws < 0.001, p = 0.011), but there was no significant difference between seasons

(p = 0.549).

Presence and absence in overlap areas

In the comparison of association by co-occurrence for wild dogs and lions, wild dogs exhibited avoidance (either spatially or temporally) 100% of the time, as indicated by the differential use of overlap areas (Table 2). Lions were dominant to wild dogs in overlapping home range areas 44% of the time, suggesting spatial avoidance by wild dogs. Wild dogs exhibited temporal avoidance of lions the other 56% of the time. In core use areas, wild dogs avoided lions spatially 33% of the time and temporally 67% of the time. Comparisons of co-occurrence with hyenas indicated that wild dogs never avoided hyenas (Table 3). Within overlapping home range areas, wild dogs were dominant to hyenas 60% of the time (indicating spatial avoidance by hyenas). There was temporal avoidance by hyenas 20% of the time, and the remaining 20% of the time there was no significant relationship, indicating random movement within the overlap area with regard

to each other. Hyenas exhibited 40% spatial avoidance, 20% temporal avoidance and

40% non-significant random movement within overlapping core use areas.

Table 2. Comparisons of associations by co-occurrence for African wild dogs (α) and lions (β) in Hluhluwe- iMfolozi Park, South Africa, 2002-2004. Measure of Correlation Year Season associationa SEb pc coefficientd Interpretatione 2002 denning home range -1.809 0.496 < 0.001 0.373 β dominant to α: spatial avoidance core use -1.885 0.676 0.003 0.364 β dominant to α: spatial avoidance post- den home range -3.095 0.603 < 0.001 0.628 β dominant to α: spatial avoidance core use -1.792 1.080 < 0.001 0.818 β dominant to α: spatial avoidance non- den home range -4.357 0.911 < 0.001 0.788 temporal avoidance 2003 denning home range -2.640 0.350 < 0.001 0.573 temporal avoidance core use -3.466 0.559 < 0.001 0.696 temporal avoidance post- den home range -1.910 0.513 < 0.001 0.409 β dominant to α: spatial avoidance core use -2.639 0.592 < 0.001 0.548 β dominant to α: spatial avoidance non- den home range -2.688 0.411 < 0.001 0.586 temporal avoidance core use -4.981 1.196 < 0.001 0.819 temporal avoidance 2004 denning home range -3.044 0.594 < 0.001 0.586 temporal avoidance core use -3.989 1.130 < 0.001 0.878 temporal avoidance post- den home range -3.562 1.051 < 0.001 0.630 β dominant to α: spatial avoidance core use -3.312 0.769 < 0.001 0.667 temporal avoidance non- den home range -3.127 0.609 < 0.001 0.541 temporal avoidance core use -2.967 0.874 < 0.001 0.594 temporal avoidance a measure of association between different groups of species b standard error of measure of association c p-value for χ2 test with values < 0.05 indicating measure of association is significant d phi correlation coefficient demonstrates strength of measure of association between different species groups e simplest ad hoc interpretation based on associated cell values (n11, n12, n21, n22)

Table 3. Comparisons of association by co-occurrence for African wild dogs (α) and spotted hyenas (β) in Hluhluwe- iMfolozi Park, South Africa, 2003-2004. Measure of Correlation Year Season associationa SEb pc coefficientd Interpretatione 2003 denning home range -1.643 0.446 < 0.001 0.089 α dominant to β: spatial avoidance core use -1.044 0.508 0.036 0.217 α dominant to β: spatial avoidance post- den home range -0.211 0.391 0.543 - random core use -0.622 0.467 0.181 - random 2004 denning home range -1.991 0.488 < 0.001 0.425 α dominant to β: spatial avoidance core use -1.727 0.621 0.003 0.363 α dominant to β: spatial avoidance post- den home range -1.284 0.544 0.015 0.261 α dominant to β: spatial avoidance core use -1.509 0.611 0.011 0.358 temporal avoidance non- den home range -4.153 1.080 < 0.001 0.670 temporal avoidance core use -1.386 0.844 0.091 - random a measure of association between different groups of species b standard error of measure of association c p-value for χ2 test with values < 0.05 indicating measure of association is significant d phi correlation coefficient demonstrates strength of measure of association between different species groups e simplest ad hoc interpretation based on associated cell values (n11, n12, n21, n22)

Interactions within overlap areas

The analysis of interactions of wild dogs and lions within overlap areas

(Appendix A) was consistent with the previous results of comparisons of association by co-occurrence. Within home ranges, wild dogs exhibited spatial avoidance (main effects of A) of the overlap area 78% of the time. Twenty-two percent of the time there was random use by wild dogs. Lions (main effects of B) demonstrated spatial attraction to the overlap area 67% of the time, with 22% not significant and one occurrence of avoidance.

Wild dogs exhibited spatial avoidance of overlapping core use areas 12% of the time, spatial attraction 38% (with the other 50% non-significant). Lions showed 75% attraction to the overlap area and 25% random use. All of the temporal interaction analyses were negative indicating solitary use was greater than simultaneous use. For home range overlap, I found 56% spatial avoidance and 44% temporal avoidance of lions by wild dogs. In core use areas, I found 12% spatial avoidance and 88% temporal avoidance of lions by wild dogs in the overlap area.

The analysis of interactions with hyenas within overlap areas (Appendix B) also supported the comparisons of association by co-occurrence results. Within home ranges, wild dogs exhibited spatial attraction to the overlap areas 60% of the time, with 20% non- significant random use and one occurrence of spatial avoidance. Hyenas demonstrated spatial avoidance 80% of the time with one occurrence of spatial attraction. Within core use areas, wild dogs exhibited 60% spatial attraction and 40% non-significant random use, while hyenas showed 100% spatial avoidance. I found 80% of the interactions analyses significant and of these all were negative (solitary use > simultaneous use).

Within both home range and core use areas, hyenas exhibited 75% spatial avoidance of the overlap area and 25% temporal avoidance.

DISCUSSION

My results support the hypothesis that African wild dogs utilize space differently relative to their two main competitors in Hluhluwe-iMfolozi Park. Overall, wild dogs remained further from lions than from hyenas, and their core use areas overlapped significantly more with hyenas than with lions (Figures 3, 6, 7). Also, overlap in space use (when taking into account the 3rd dimension intensity of use) between wild dogs and lions was significantly less than mean (2-dimensional) overlap, while there was no such difference between wild dogs and hyenas (compare Figures 4 and 8). The above results are consistent with Webster et al. (2011) who found that wild dogs actively avoid lions more than they avoid hyenas. In addition, other studies have shown that cheetahs adjust their behavior more often in response to lions than to hyenas (Laurenson 1994, Durant

1998).

Competition with its two main competitors, lions and hyenas, can be a major hindrance to wild dog populations (Vucetich and Creel 1999). One of the main causes of natural mortality in wild dogs is intraguild predation, most often by lions (Woodroffe et al. 2004, 2007). Therefore, it is not surprising that wild dog densities are consistently low in areas where lion densities are high (Creel and Creel 1996; Creel and Creel 2002). The significant overlap of home ranges between wild dogs and lions in HiP is likely due to the relatively small size of the park as well as the fact that the park is fenced. Carnivore densities in HiP are relatively high (Trinkel et al. 2008, Graf et al. 2009), and there is limited space. However, it appears possible for wild dogs to avoid lions through

adjustments in core space use. Wild dogs spent a majority of their time in areas free of lions, as was evidenced by the significantly lower overlap of core use areas compared to home ranges (compare Figures 5 and 7). Additionally, the analysis of spatial overlap including intensity of use also indicated that peaks of space use in the home ranges of wild dogs differed significantly from lions when compared to mean overlap (compare

Figures 4 and 8). Wild dogs of Pilanesberg National Park, South Africa responded to lions in a similar manner avoiding areas where lion activity was high (van Dyk and

Slotow 2003).

Wild dogs utilized both spatial and temporal avoidance of lions within overlap areas (Table 2, Appendix A). Wild dogs demonstrated temporal avoidance considerably more than spatial avoidance in core use areas, while using spatial and temporal avoidance strategies approximately equally within their larger home ranges. Although my study did not have the information necessary to incorporate habitat characteristics into the analyses, past studies have suggested that wild dogs accomplish spatial segregation through exploitation of different habitats (Mills and Gorman 1997, Mills and Funston 2003). This is consistent with studies of similar species that indicate habitat heterogeneity can create refuges that allow smaller competitors to persist (Palomares et al. 1996; Fedriani et al.

1999, Campbell 2004, Berger and Gese 2007, Nelson et al. 2007).

My results were also consistent with others with respect to temporal avoidance by wild dogs and similar carnivores. Schoener (1974) suggested that time is one of three critical dimensions of a species’ ecological niche. Thus, temporal partitioning by altering activity patterns to reduce encounters with larger competitors decreased competition

effects and allowed animals to coexist (Johnson et al. 1996, Arjo and Pletscher 1999,

Creel et al. 2001, Kronfeld-Schor and Dayan 2003, Lucherini et al. 2009). Hayward and

Slotow (2009) showed that activity patterns of wild dogs differed significantly from those of lions. The two species had the least overlap in times of activity of any of the five carnivores studied (wild dogs, lions, spotted hyenas, leopards and cheetahs). Saleni et al.

(2007) also found temporal avoidance by wild dogs to avoid interference competition with their major competitors, lions and spotted hyenas.

Wild dogs remained significantly further from lions during the denning season as indicated from the spatial interaction analyses both with and without temporal components (Figure 3). Additionally, home ranges of wild dogs and lions overlapped significantly less during the denning period (Figures 4, 5). Webster et al. (2011) found that wild dog packs with younger pups had greater vigilance and alarm calls in response to lions. It is likely that the packs adjusted their behavior while denning to avoid lions

(i.e., choosing den sites far from lions prides), as almost half of all juvenile wild dog mortality is as a result of lion predation (Woodroffe and Ginsberg 1997). Wild dogs demonstrated temporal avoidance more often than spatial avoidance during the denning season (Table 2, Appendix A). This logical considering the restricted movement during this time as wild dog packs must return to the den after each hunting foray to feed the alpha female and her pups. During the non-denning season, wild dogs did not alter their space use as considerably as both their mean overlap and overlap incorporating intensity of use with lions was similar to their overlap with hyenas (Figures 4, 5, 8). Instances of wild dog attraction to overlap areas where lion densities are high likely corresponded to

areas of high prey density (Mills and Gorman 1997), and thus it may be necessary for wild dogs to spend time in those areas despite the presence of lions, especially when traveling with growing pups.

It has been suggested that conservation of high densities of competing carnivores in small, fenced reserves may not be feasible and may lead to the extinction of the smaller competitor (Creel and Creel 1996, Mills and Gorman 1997). However, it appears that the wild dogs of HiP have been able to adapt to life in a fenced reserve with lions through a combination of spatial and temporal avoidance, adjusting their behavior as necessary based on the season (i.e., through increased use of temporal avoidance during denning when the packs are less mobile). Moehrenschlager et al. (2007) found that kit foxes were able to successfully avoid coyotes within relatively small areas indicating that sometimes smaller competitors may be able to coexist with larger competitors without extensive habitat separation. As the wild dog population in HiP is currently relatively stable (~100 individuals in 8 packs), this study supports the conclusion that smaller wild dogs can coexist with larger lions and hyenas.

Wild dogs in HiP did not appear to alter their space use significantly in relation to hyenas (Figures 4-8). Past studies have found mixed results on the effects of hyenas with some indicating hyenas have a negative impact on wild dogs. This is due to frequent stealing of wild dog kills, a phenomenon termed kleptoparasitism, a largely one-way interaction (Fanshawe and FitzGibbon 1993, Creel and Creel 1996, Gorman et al. 1998,

Carbone et al. 1997, 2005). Saleni et al. (2007) suggested that wild dogs in HiP temporally avoid hyenas. Other studies have found little effect of hyenas on wild dogs

(Mills and Funston 2003, Hayward and Slotow 2009, Webster et al. 2011). The space use results of my study support the latter findings. Dynamic interaction analyses indicated that hyenas were closer than expected to wild dogs and demonstrated no evidence of wild dogs avoiding hyenas either spatially or temporally (Table 3, Appendix

B). This pattern is likely even stronger considering that hyena observations were limited, and there were likely many more hyenas present in areas with wild dogs than were reported.

The lack of avoidance of hyenas by wild dogs is likely due to the relatively large size of wild dog packs in HiP, which can adequately defend their kills against kleptoparasitic hyenas. The average wild dog pack size in the park during the study years was 17 individuals, significantly higher than 5-9 individuals, the average pack sizes of other parks (Creel and Creel 2002). Whateley and Brooks (1978) found the average hyena clan size in HiP to be relatively small: between 9-14 individuals. Hyena numbers have increased in the park through increasing numbers of clans (Graf et al. 2009).

However, feeding groups (sub-groups of clans) remain small (Andreka et al. 1999,

Kruger et al. 1999). Furthermore, the highest density of hyenas is in the northern

(Hluhluwe) section of the park (Graf et al. 2009) where only one wild dog pack resides.

The others reside in the iMfolozi section of the park to the south. Large wild dog packs can better defend their kills and for longer periods of time than smaller packs.

Kleptoparasitism will only negatively affect wild dogs when hyenas take over kills quickly, as wild dogs can fill their stomachs on a kill within minutes (Fanshawe and

FitzGibbon 1993, Carbone et al. 2005). I often observed hyenas arriving at a wild dog

kill and using a whoop call to attract more hyenas. However the hyenas never appropriated the kill, most likely because hyena numbers were never greater than those of the wild dog packs. This is in contrast to a past study in Serengeti National Park,

Tanzania where there were more hyenas than wild dogs at kills almost a third of the time

(Fanshawe and Fitzgibbon 1993).

As it appears that wild dog pack numbers in HiP are large enough to avoid most cases of kleptoparasitism, extra effort to avoid hyenas becomes unnecessary and in some cases would be detrimental due to the high total energy needs of the packs (Gorman et al.

1998). If a larger competitor does not have a significant effect, then it is not crucial to alter behavior to avoid it (Durant 1998, Webster et al. 2011). Fanshawe and FitzGibbon

(1993) found that for wild dogs in the Serengeti, it was likely that competition with hyenas was one of the factors keeping pack numbers high. Courchamp and Macdonald

(2001) concluded the same thing. This could also be the case in HiP as hyena densities are relatively high, despite clan sizes being relatively low (Graf et al. 2009).

The high overlap in space use among these two competitors may also be due to an attraction of hyenas to wild dogs. The seemingly contradictory spatial analysis

(including the temporal component) that indicated that hyenas avoid wild dogs was likely due to the differing number of available locations between wild dogs and hyenas. With only one collared hyena, most hyena sightings were opportunistic. Thus sightings of wild dogs alone in overlap areas were extremely high compared to hyena sightings and likely biased the results. Estes and Goddard (1967) noted that hyenas often remain near a resting wild dog pack for hours waiting for them to begin hunting. It is common for

hyenas to follow wild dogs during hunts and they often arrive only minutes after wild dogs have killed prey (personal observations; Estes and Goddard 1967, Fuller and Kat

1990). It is rare for a wild dog kill to be free of hyenas (Fanshawe and FitzGibbon 1993;

Carbone et al. 2005), and because it is common for hyenas to eventually appropriate the kill (generally once the wild dogs have nearly finished feeding), it is logical that they are often found near wild dog packs (Fanshawe and FitzGibbon 1993). Furthermore, this pattern could be indirectly caused by both wild dogs’ and hyenas’ avoidance of lions, as lions are also a significant source of hyena mortality (Kruuk 1972, Trikel and Kastberger

2005).

The alternative hypothesis to the one presented in my study is that the carnivores in HiP could be distributed according to prey preferences. In other areas, lions have been shown to move according the distribution of their prey (Hopcraft 2002). Although there is high diet overlap between wild dogs, lions and hyenas, there is some diet segregation. That is, lions prefer buffalo (Syncerus caffer; Hayward and Kerley 2005) that the smaller wild dog and hyena rarely hunt. In HiP, wild dog’s preferred prey is medium-sized antelope such as impala and nyala (Kruger et al. 1999), which are seldom taken by lions. Thus, the habitat requirements of the prey combined with the prey preferences of each species could also influence space use of competing carnivores.

Temporal changes in the population densities of lions and spotted hyenas in relation to wild dogs suggest the limiting effects these dominant competitors have on the density of wild dogs. Wildlife managers and conservationists often simply consider exploitative competition when developing management strategies. As the success of

conservation efforts may rely on the interactive role between species, managers should account for competitive relationships between sympatric carnivores when devising management tactics (Johnson et al. 1996, Vucetich and Creel 1999, Creel 2001). My study suggests that taking into account interference competition between species may be equally important. In KwaZulu-Natal Province, where most wild dog packs exist in fenced parks and reserves and individual numbers are generally low and often not self- sustaining, management through reintroductions and relocations is common. Even in

Hluhluwe-iMfolozi Park, where the population is relatively stable at this time, individual introductions are necessary to avoid inbreeding and genetic drift (Spiering et al. 2010,

2011). Thus, the information from my study will be useful particularly in regards to choosing appropriate reintroduction and relocation sites. My study also demonstrates that wild dogs can survive and even thrive in areas with high densities of larger competitors, especially if the habitat is heterogeneous. However, as wild dogs avoided lions in almost all instances, when considering reintroduction and translocation sites, it would likely be better to place them in areas with low lion density as past studies have shown that wild dogs suffer greater mortality when lion densities are high (Creel and Creel 1996,

Woodroffe and Ginsberg 1997).

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PERSONAL COMMUNICATIONS

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Appendix A. Interaction within overlapping home ranges and core use areas for African wild dogs (A) and lions (B), showing spatial attraction or avoidance by each and interaction between groups (ixn) as well as deviation of odds from random in Hluhluwe-iMfolozi Park, South Africa, 2002-2004. Spatial Effects Interaction Effects Odds for each celld a b a b c b Year Season LA:Ā pA ̅ pB Lixn pixn n11 n12 n21 n22 2002 denning home range -0.699 0.001 0.852 < 0.001 -1.000 < 0.001 0.567 1.870 0.438 0.699 core use -1.103 0.002 1.877 < 0.001 -1.519 < 0.001 0.613 4.480 0.414 0.458 post- den home range -0.687 0.002 0.230 0.314 -1.579 < 0.001 0.341 2.078 1.256 0.693 core use 2.205 0.700 4.675 < 0.001 -4.755 < 0.001 0.000 17.857 1.520 0.167 non- den home range -1.639 < 0.001 1.044 < 0.001 -2.711 < 0.001 0.336 9.754 0.709 0.528 2003 denning home range -0.402 0.004 -0.102 0.470 -1.332 < 0.001 0.386 2.012 1.345 0.500 core use 1.446 < 0.001 1.483 < 0.001 -3.363 < 0.001 0.000 3.878 3.053 0.240 post- den home range -0.198 0.343 0.445 0.047 -0.909 < 0.001 0.734 1.586 1.146 0.367 core use 0.322 0.185 0.919 0.004 -1.120 < 0.001 0.847 1.616 1.680 0.228 non- den home range -1.069 < 0.001 0.284 < 0.001 -0.599 < 0.001 0.169 1.746 4.837 3.448 core use 0.760 0.008 0.864 0.003 -1.832 < 0.001 0.615 2.016 2.129 0.048 2004 denning home range -0.074 0.717 1.052 < 0.001 -1.278 < 0.001 0.827 2.442 1.060 0.149 core use 0.934 0.134 2.197 < 0.001 -4.483 < 0.001 0.000 5.569 2.246 0.088 post- den home range -2.355 < 0.001 1.190 < 0.001 -1.956 < 0.001 0.619 1.079 5.035 0.245 core use 0.530 0.050 -0.474 0.092 -1.796 < 0.001 0.471 1.035 3.484 0.279 non- den home range -0.939 0.032 -0.628 0.001 -1.557 < 0.001 0.339 1.683 2.053 0.448 core use 0.129 0.672 0.487 0.113 -1.272 < 0.001 0.718 1.857 1.346 0.179 a spatial main effects of A (wild dogs) or B (lions) indicating attraction (+) or avoidance (-) of overlap area b p-value for χ2 test with values < 0.05 indicating effects were significant c interaction effects d odds indicate departure from expectation of each cell (n11, n12, n21, n22), with values close to 1 indicating use of shared area was as expected at random, values < 1 indicating use less than expected, and > 1 indicating use more than expected

Appendix B. Interaction within overlapping home ranges and core use areas for African wild dogs (A) and spotted hyenas (B), showing spatial attraction or avoidance by each and interaction between groups (ixn) as well as deviation of odds from random in Hluhluwe-iMfolozi Park, South Africa, 2002-2004. Spatial Effects Interaction Effects Odds for each celld a b a b c b Year Season LA:Ā pA ̅ pB Lixn pixn n11 n12 n21 n22 2003 denning home range 0.979 < 0.001 -1.345 < 0.001 -1.192 < 0.001 0.591 0.520 3.283 0.564 core use 0.799 0.001 -0.921 < 0.001 -0.744 0.001 0.678 0.544 2.186 0.620 post- den home range 0.014 0.956 -2.097 < 0.001 -0.111 0.371 0.479 0.533 3.741 4.184 core use 0.322 0.159 -1.726 < 0.001 -0.523 0.002 0.568 0.568 4.217 2.269 2004 denning home range -0.558 0.011 -2.054 < 0.001 -0.506 0.004 0.223 1.273 4.313 2.530 core use 1.611 < 0.001 -2.764 < 0.001 -1.814 < 0.001 0.363 0.271 10.256 1.352 post- den home range 0.590 0.018 -2.446 < 0.001 -1.162 < 0.001 0.533 0.516 1.342 2.224 core use 0.393 0.163 -0.738 0.008 -0.919 0.001 0.645 0.848 2.857 0.833 non- den home range 1.234 < 0.001 0.975 < 0.001 -0.764 < 0.001 1.745 1.733 2.081 0.032 core use 2.178 < 0.001 -1.668 < 0.001 -1.730 < 0.001 1.096 0.614 0.290 9.357 a spatial main effects of A (wild dogs) or B (hyenas) indicating attraction (+) or avoidance (-) of overlap area b p-value for χ2 test with values < 0.05 indicating effects were significant c interaction effects d odds indicate departure from expectation of each cell (n11, n12, n21, n22), with values close to 1 indicating use of shared area was as expected at random, values < 1 indicating use less than expected, and > 1 indicating use more than expected