Bachelor thesis

Ex situ lion conservation Behavioural responses to playbacks of competitors

with focus on sex and age differences

Author: Emma Sopelsa Hall

Supervisor: Anders Forsman and

Emma Dunston Examiner: Per Larsson

Academic term: Spring 2017

Subject: Biology

Level: First level, Bachelor

Course code: 2BI01E Nr: 2017:Bi11

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Abstract Due to increasing habitat loss, human-lion conflict, poaching and other reasons, African

lion (Panthera leo) populations have suffered a drastic decline. The African Lion and

Environmental Research Trust (ALERT) is working to stop this pattern and is the first

organization with an ex-situ conservation project for lions. Before releasing lions raised

by captive-bred adults, they must first be ensured to behave properly to make sure they

have the highest chance of survival. One challenge in the wild is encountering and

competition with unknown conspecifics. By conducting playback of unfamiliar lion

roars, the behaviours of lions under this ex-situ reintroduction program were tested and

compared with observations from earlier studies of wild lions. Social interactions were

also collected and a social network analysis was done to give information about the

social structure in the pride. This in turn was compared with boldness scores, calculated

from behavioural responses in the playback experiments. Lastly, I searched for

associations between age and sex with both boldness and social interactions.

The studied pride consisted of 12 lions. The lions were more vigilant when a playback

consisted of numerous lions vocalizing, but playing more than three lions seemed to

make them loose interest, suggesting either habituation or false information. One adult

female and the alpha-male were most bold, followed by five sub-adults. Boldness did

not vary according to sex or age differences, but the social network analysis showed that

some social interactions were more dominated by one sex or age group. These

behaviours were in agreement with comparisons of wild prides.

This study showed that captive-bred lions have developed natural social behaviours.

Based on the behavioural responses observed by the captive-origin lions to the

playbacks of unfamiliar lions, it is unclear whether these lions would appropriately

respond when encountered with unfamiliar conspecifics in the wild post-release.

Keywords Animal behaviour, Ex-situ conservation, Lions, Panthera leo, Playbacks,

Reintroduction, Social network analysis

Acknowledgement First, I would like to acknowledge the African Lion & Environmental Research Trust

for giving me this invaluable opportunity letting me conduct research on their lions. I

would also like to thank Professor Anders Forsman for all the support and advice from

the beginning till the end. Thank you Dr Emma Dunston for teaching and supporting me

both in the field and after during my writing.

Thanks to the faculty of health- and life science at Linnaeus University for the

acceptance of a scholarship which helped me a lot financially in conducting this project.

A last, but not least, big thanks to my friends and family who have been a huge support

and who have been pushing me to help me conduct my own research on lions, a dream

that now has come true.

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Contents

1 Introduction _________________________________________________________ 1 1.1 Reintroductions ___________________________________________________ 1 1.2 African Lion & Environmental Research Trust (ALERT) __________________ 2 1.3 African lions (Panthera leo) _________________________________________ 2 1.4 Aims and hypotheses ______________________________________________ 4

2 Materials and method _________________________________________________ 4 2.1 Study site _______________________________________________________ 4 2.2 Playback experiment_______________________________________________ 5 2.3 Social interactions_________________________________________________ 6 2.4 Statistical analysis_________________________________________________ 6

3 Result ______________________________________________________________ 7 3.1 Playbacks _______________________________________________________ 7 3.2 Social interactions________________________________________________ 10

4 Discussion __________________________________________________________ 11 4.1 Conclusion _____________________________________________________ 15

References ___________________________________________________________ 16

5 Appendices __________________________________________________________ I 5.1 Appendix A - The lions _____________________________________________ I 5.2 Appendix B – Boldness scores _______________________________________ II 5.3 Appendix C – Result for out- and in-degree values ______________________ III 5.4 Appendix D – Result for betweenness centrality values __________________ IV

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1 Introduction

The African lion (Panthera leo) population is declining at an alarming rate. Over the

past 21 years, between 1993 and 2014, the wild population has reduced by

approximately 43% (Bauer et al. 2016). Today the population is estimated to be

between 23,000 (Bauer & Van Der Merwe, 2004) and 32,000 (Riggio et al. 2013), but

likely to be closer to 20,000 (Bauer et al. 2016). One of the major reasons for the

decline is the habitat loss due to the growing human population (Bauer et al. 2015).

Trophy hunting also contributes to the population decline. Because of the poaching of

herbivores, which are food sources for lions, prey populations are declining across

Africa (Craigie et al. 2010). This in turn has led to food depletion for lions,

subsequently causing a decline in lion populations. The last major reason for decline is

the human-lion conflict that has arisen because of the threat lions pose to livestock and

human life. To protect livestock or as an act of revenge, local communities persecute

and kill lions (Schulette et al. 2013).

1.1 Reintroductions

Due to the continuous population decline, both ex-situ and in-situ projects are working

to prevent species from going extinct. Ex-situ management of animals, is a process

aimed to protect them from threats by keeping them outside their natural habitat (IUCN,

2014). This could be done either by translocating wild animals from one population to

another, or by using animals from captive-origin populations (ALERT, 2017).

Examples of earlier successful ex-situ reintroductions of captive bred animals into the

wild include amongst others the prairie dogs (Cynomys ludovicianus) (Sheir & Owings,

2006), the American red wolf (Canis rufus) (Philips & Parker, 1988), golden lion

tamarins (Leontopithecus rosalia) (Beck et al. 1991), Arabian oryx (Oryx leucoryx)

(Stanley Price, 1989) and New Zealand robins (Petroica australis) (Armstrong et al.

2002).

In reintroduction projects, animals can either be reintroduced through soft or hard

release. A soft release strategy means that the animals will go through a pre- or post-

release training to develop and express instinct and natural behaviour, also getting used

to their new environment before being released (Kleiman, 1989). A hard-release does

not include these conditionings and often leads to a failure of carnivore reintroductions

(Abell et al. 2013). One of the reasons of failure from earlier ex-situ attempts with other

species is that the released animals have not been sufficiently adapted to the wild

environment (Watters & Meehan, 2007; Jule et al. 2008). Kleiman (1989) mentions

important behavioural aspects that should be developed to improve survival when

reintroduced. Two of these include being able to avoid predators and interact accurately

with conspecifics.

There are no earlier attempts of ex situ reintroduction programs for lions. Although,

there are commercial enterprises with lions, where lions are being cuddled and walked

with by tourists to later on being released in an area for canned hunting (Big Cat

Rescue, 2013). These kinds of organizations, mainly situated in South Africa, have

nothing to do with conservation and are only money-oriented (Big Cat Rescue, 2013;

Abell et al. 2013). This kind of practice is one of the reasons why serious attempts of ex

situ reintroduction programmes have been declined (Abell & Youldon, 2013). Also,

these projects are very expensive to run and some have seen them as an attempt of

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failure, not spending the money on conservation of lions in the wild (Hunter et al.

2013).

Thanks to the earlier ex situ reintroduction programmes with other species it has been

easier to identify the main factors for success (Abell et al. 2013). This study was

conducted in the first ex situ reintroduction program for lions, with the aim to release

lions raised by captive-origin ones. I will look at captive-bred and their semi-wild born

sub-adult lions’ reactions when playing unfamiliar lion roars and compare their

behaviours with those reported in previous studies of wild lions’ reactions.

1.2 African Lion & Environmental Research Trust (ALERT)

ALERT is working against the extinction of lions by implementing a model of

conservation in Zimbabwe and Zambia (ALERT, 2017). One facet of program is the

staged ex-situ reintroduction soft-release program of lions. The African Lion

Rehabilitation & Release into the Wild Program is a three-staged program, aiming to

reintroduce lions from a captive-origin into the wild. Firstly, the rehabilitation stage

consists of captive-raised cubs between the age of six weeks and 18 months being

walked in a private game reserve or national park. This encourages the development of

natural behaviours by letting the cubs to interact with their environment, hunt prey

species and do social bonding and play with their siblings. From the age of six months,

tourists are allowed to join trained lion handlers on these walks, raising awareness and

financing the project (ALERT, 2017). At 18 months old, it is no longer safe to take the

lions for walks and are therefore retired and moved to the next phase of the

rehabilitation phase, the Night Encounter program, where the lions can further practice

improving their hunting skills.

Stage two is the release phase, where lions are released as a pride into large fenced wild

area. Here, the purpose is for the lions to form a socially stable and self-sustaining pride

where all human contact is terminated. Within this stage, they can live a normal life and

hunt different prey species, but no other predators or conspecifics are present. In the

Dambwa pride, all individuals are radio-collared to allow for observations of their

movements (ALERT, 2017). During this stage, prides have had cubs which are never

exposed to human contact. As these cubs are raised under a natural environment, they

are subjects for release into national parks and protected game reserves (reintroduction

phase) in the future once of appropriate (ALERT, 2017). To ensure the lions born into

the release phase are well prepared for survival in the wild, long-term monitoring and

assessment of behaviours must first be done to ensure that the lions exhibit natural

behaviours.

1.3 African lions (Panthera leo)

A lion pride consists of three to six related female adults (though it can range up to 18),

their offspring and a coalition of immigrant males (1–4) (Packer et al. 1988; Packer et

al. 1991a). Lionesses are likely to stay in the same pride their whole lives, participating

in cooperative hunting, territory protection and cub raising (Schaller, 1972). Males,

however, leave their pride at young age, around two to three years old, to compete for a

new pride and access to females (Bygott et al. 1979). Because of this, when faced with

intrudes, females and males compete for different resources; females mainly for their

cubs and territory, while males for their females and the risk of being evicted from their

pride (Schaller, 1972). Lions are classified to be cubs up until the age of two, sub-adults

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between the ages of two and four years, and from the age of four they are classified as

adults (Schaller, 1972).

Social mammals, like lions, need to cooperate to defend recourses. When they

encounter competitors, they should be able to consider the number of individuals that

they would be able to interact with and at the same time assess the value of the resource

(McComb et al. 1994; Maynard Smith, 1982). Packer et al. (1990) suggests that the

main reason for lion sociality is because of the threat of conspecifics. Based on studies

on wild prides in Serengeti National Park, lions can distinguish pride mates from

strangers and also determine whether the pride is outnumbered by the intruders or not.

Also, they seem to adopt a more careful approach when being outnumbered (McComb

et al. 1994; Grinnell et al. 1995).

When wild lions are intruded by competitors, individuals will exhibit different

behaviours. Captive bred animals have been tested on their ability to respond to

competitors using playbacks of oral communications. For example, McComb et al.

(1994) played single lions roaring and groups of three lions roaring in choruses to

examine if lions in the focal pride would change their behaviours according to the

number of lions in their own and the competitive group. The lions approached the

loudspeaker more slowly when playing three intruders, also glancing back at their

lagging companions and having a tendency of doing more pauses when approaching.

Another response is roaring. Females do this if other pride lionesses are dispersed from

the pride upon hearing the playback to recruit them for help (McComb et al. 1994).

Males, however, do not take this into account as they instead seem to roar irrespective

of the presence of other males (Grinnell et al. 1995).

Heinsohn and Packer (1995) studied the territorial responses of females to other females

and showed that there are lagers and leaders in a pride. The laggards avoid the cost of

fighting, staying behind their companions, while those that lead the pride take much

bigger risks since most territorial fights often lead to death or severe injuries (Schaller,

1972; Packer et al. 1990; Packer et al. 1988). Female laggards could be termed either

“conditional cooperators”, approaching when their help would contribute to higher

chance of winning, “conditional gards”, lagging most when most needed and

“unconditional laggards”, whom would not change their behaviour no matter the odds

of winning a contest (Heinson & Packer, 1995). Behavioural responses of lionesses do

not seem to be related to age and body size when it comes to territorial intrusion

(Heinsohn & Packer, 1995). Neither do female lions show any apparent dominance

hierarchy (Packer & Pusey, 1985). All this suggest that the fighting ability does not

have to be related to the individual differences (Heinsohn & Packer, 1995).

Male laggards can also exist in a pride, but compared to females, male laggards do not

change the grade of lagging when the odds of winning a contest with intruders change

(Grinnell et al. 1995). However, lagging males seemed to approach more slowly, or not

at all, in thicker cover (Grinnell et al. 1995).

Previous work indicate that female lions show an increased probability of approaching

loudspeakers playing unfamiliar lion roars if sub-adults are present in the pride

(McComb et al. 1994). This could either be because of the bigger need to protect their

territory or because of the extra help for defense is present thanks to the younger lions

(McComb et al. 1994). Since my pride consists of six sub-adults, I would expect the

adult females to be defensive when playing the recordings.

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1.4 Aims and hypotheses

The method used by McComb et al. (1994) was applied to study the behavioural

reactions of a pride under the release phase of ALERT’s ex-situ program, intending to

determine whether they would respond in a manner similar to wild lions. I set out to

examine whether and how behavioural responses to vocal playbacks depend on the

number of signaling and receiving lions. I predict that when the risk of losing would be

greater than the chance of winning, the lions should not approach the loudspeaker. This

would be when being outnumbered by the lions being played. When the lions in the

pride outnumber the lions being played they should approach the loudspeaker, but more

cautiously the larger the group of the lions being played are. When playing only one

lion roar, I would expect the lions to approach the loudspeaker.

In this study, I also examined whether there are any behavioural differences depending

on the sex of the lions being played and the sex of the lions in the pride. Males would

risk losing their pride to an invading male, while also losing their entire lifetime

reproductive success (Grinnell et al. 1995). I therefore hypothesize that the males in this

study would approach the loudspeaker and become more defensive when playing male

roars. I further predicted that their behavioural responses will not be as dependent of the

number of females and younger lions present in the pride; instead they will be more

dependent on the number of defenders (Grinnell et al. 1995). Females, however, are

expected to adjust their decision to approach according to the composition of their

group (McComb et al. 1994).

If the lions exhibited similar behaviours of those observed for wild lions, the work of

ex-situ reintroduction could indicate that lions with a captive-origin are capable of

developing a suite of natural behaviours, which could increase the chance of surviving

in the wild, knowing how to approach and avoid competitors. This could show that

playback experiments might be a successful method to use in other ex-situ projects.

Being a social animal, lions, just like humans, prefer some individuals more than others.

A social network analysis can provide information of the social structure in the pride

(Farine & Whitehead, 2015) and inform how lions are connected to each other in the

network and in what way. By looking at different social interactions, relationship

between boldness and sociality can be investigated and again also comparing sex and

age differences. My hypotheses are that some lions are more connected to each other

than to others and that there will be some differences in interactions, especially between

the sub-adults and adults due to the age difference.

A positive result would be one step forward towards a release of the lions into the wild,

since this study will provide information on a captive-origin pride’s behaviour prior to

release when it comes to both behavioural responses and social behaviour. This is a

contribution to the work of preventing the decline of African lion populations.

2 Materials and method

2.1 Study site

One of ALERT’s research sites is situated in the Dambwa Forest, Zambia (17°48’S,

25°51’N) (Figure 1), which is a government owned land leased by ALERT. The lions

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that were studied are fenced within an area of 707 acres and can hunt prey species like

puku (Kobus vardonii), impala (Aepyceros melampus) and duikers (Cephalophinae).

The pride consists of twelve lions, six adults and six sub-adults. Of the adults, one male

and five females, were released into the release site in 2011, and in 2013 the first litter

of cubs were born (two females and one male). In 2014 another litter of cubs were born

(one female and two males) in the site to another female (ALERT, 2017) (Appendix A).

At the time of the study, only the adult lions were collared with telemetry collars.

Figure 1. The release site is situated in the Dambwa Forest, southern Zambia, close to the boarder of Zimbabwe (Google Earth, 2017).

2.2 Playback experiment

A BOSE SLIII speaker was used for the playback and was connected through Bluetooth

to an iPhone where the mp3 files of the playbacks were saved. Playbacks consisted of

unfamiliar lion vocalisations which differed in terms of numbers of lions and genders

(one male, three males, three females, five, eight, thirteen and eighteen lions of mixed

genders). One to two playbacks were done per week with at least five days apart,

minimizing the risk of lions becoming habituated to playbacks. The speaker was located

outside the fence and hidden in high grass, at a minimum distance of 200 meters from

the lions. The playbacks were either conducted in the morning between 0800-1100 h or

in the afternoon between 1500-1600 h when the temperature was not too high. All

reactions were filmed with a Nikon Coolpix L330 until all lions had stopped paying

attention to the recording or for a total of 30 minutes.

To quantify the boldness of each individual, they were given positive or negative scores

ranging between -5 and 5 depending on their reaction of the playback (Appendix B). A

high score indicated a bolder response, while a low score indicated a more fearful

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response. The total final score for each lion, when summarized the boldness for every

treatment, was thereafter used for the comparison of sex and age differences. Also, for

every minute spent active paying attention to the playback, each lion received a score.

As with the boldness scores, these values were summarized and analyzed to see the

effect of number of intruders, and how the sex and age of the lions differed in reactions.

2.3 Social interactions

The social interactions were recorded at all occurrences and included all grooming,

greet, play and aggression, earlier classified by Schaller (1972). For each interaction, the

initiator and receiver and whether it was accepted or rejected was recorded. If the

behaviour was repeated or another one was initiated within one minute after the first

one, it was not included in the analysis to avoid pseudo replication.

2.4 Statistical analysis

Eleven treatments were conducted, whereby two of them were replications of three

males and thirteen lions. The total boldness and time score across all treatments for each

lion was calculated to compare and analyze age and sex differences for social degree

values, boldness and time scores. Since the behavioural response data was not normally

distributed, the non-parametric tests Spearman’s rank correlation and two-sample

Wilcoxon were performed using the statistical program R, Version 2.3-2 (Fox &

Bouchet-Valat, 2017). To see the effect of number of intruders on boldness and time

reactive to playback, the total boldness and time score for each treatment was

summarized separately and analyzed using Spearman’s rank correlation (Fox &

Bouchet-Valat, 2017).

A total of 104 social interactions were collected during eight weeks. All data were

compiled into weighted directed matrices for all social interactions and the program

Ucinet, Version 6.543, was thereafter used to do a Social Network Analysis (Borgatti et

al. 2002). Density, degree and betweenness centrality analyses were done for all social

interactions (groom, greet, play and aggression) and also for the total interactions (all

social) to see which individuals that were dominating each network in the pride. Density

values indicate how connected individuals are within the pride. A value of 1 indicates

that all individuals in the network are connected to each other, while a value of 0 means

no connection (Wasserman & Faust, 1994). The degree measures the activity level of an

individual (Vital & Martins, 2009), where the in-degree score shows the level of

receiving from other individuals, while the out-degree score represents the level of

initiating (Wey et al. 2008). Betweenness centrality values indicate how individuals

control or mediate relations between other individuals that are indirectly connected

(Knoke & Yang, 2008). The higher the value of betweenness centrality is for an

individual, the more it acts as a connector and mediator of interactions between

unconnected lions within the network (Knoke & Yang, 2008). To test whether social

interactions differed between males and females, a two-sample Wilcoxon test was

conducted using R (Fox & Bouchet-Valat, 2017).

Sociograms for all social networks were generated using the program Netdraw, Version

2.147 (Borgatti, 2002). Sociograms provided an illustration of social networks,

providing a clear indication of who interacts with whom and to what level. Each lion is

represented by a node, which differs depending upon the sex (node shape) and age

(node size) of the individual. Arrows connect individuals, showing the direction of the

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interactions while line thickness showing the number of interactions between lions, with

a thicker line indicating more interactions (Hanneman & Riddle, 2005).

3 Result

3.1 Playbacks

The playback experiment showed that adult female Rusha was the boldest individual in

the pride (Figure 2), with a score of fourteen. None of the other adult females

approached the loudspeaker, regardless of the sex or number of intruders on the

playback. These other adult females were also observed to have shorter response times

to playbacks, resulting in these individuals having the lowest scores, while sub-adult

RS1 spent the most time being reactive, followed by her brother RS2 and mother Rusha

(Figure 3).

The alpha-male, Zulu, was observed to only approach when playing males (one male

and three males), but was more likely to be unresponsive to all other playbacks. During

the second playback of three males, five lions were absent from the pride, including the

oldest male sub-adult RS2. This was also the only time when Zulu started roaring when

approaching the location of the speaker and after. All sub-adults except female RS3

approached as a response for some of the playbacks. For the lions that approached, they

only paused on the way towards the speakers when playing three lions.

Figure 2. Total boldness score for each lion across the eleven playbacks.

0

2

4

6

8

10

12

14

16

Zulu

Kel

a

Kw

andi

Ley

a

Lom

a

Rush

a

RS

1

RS

2

RS

3

LE

1

LE

2

LE

3

Bold

ness

scores

Lions

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Figure 3. Average number of minutes individual lions spent being reactive to playbacks across the eleven playbacks.

The lions showed a higher probability of approaching when playing three males, but

when playing more than three lions (five, eight, thirteen and eighteen) the likelihood of

approaching dropped (figure 4).

Figure 4. Total boldness scores plotted against number of intruders. The figure shows boxplots, which describes the total boldness scores for each treatment with different number of lion roars being played to the pride. The median lines at 0 are due to many scores of zero for each playback. The playback of one female, one male, three females, three males 1 and thirteen lions show unfilled circles, outliers, describing values that stand out from the others in the dataset of each treatment.

There was no significant correlation between boldness and age (rs = -0.25, p = 0.43).

Also, a two-sample Wilcoxon test showed no relationship between boldness and sex (W

(9), p = 0.25). Figure 5 shows the distribution of boldness scores with age as the

independent variable. When testing for boldness scores and number of intruders no

significant correlation was found (rs = -0.59, p = 0.074). Time reactive decreased

0.0

1.0

2.0

3.0

4.0

5.0

6.0

7.0

Zulu

Kel

a

Kw

andi

Ley

a

Lom

a

Rush

a

RS

1

RS

2

RS

3

LE

1

LE

2

LE

3

Tim

e a

cti

ve (

min

)

Lions

A B C D E F G H I J

01

23

45

67

Intruders

Boldn

ess

Eigh

teen

Thirt

een

2

Thirt

een

1

Eigh

t

Five

Thre

e m

ales 2

male

s 2

Thre

e m

ales 1

Thre

e fe

male

s

One

male

One

fem

ale

One

fem

ale

One

mal

e

Thr

ee f

emal

es

Thr

ee m

ales

1

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ee m

ales

2

Five

Eig

ht

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Thi

rtee

n 2

Eig

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Intruders

Bol

dn

ess

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significantly with increasing number of intruders (rs = -0.722, p = 0.018). A boxplot was

created to visualize the distribution of time scores for each treatment (Figure 6). The

probability of the lion to approach was independent of sex (rs = -0.40, p = 0.199) and

age (W (10.5), p = 0.395).

Figure 5. Boldness scores plotted against the age of the lions. Squares represent males and circles represent females. Individuals with the same age and boldness score are seen as one, but larger, shape. The triangle represents two individuals, one female and one male of the same age.

Figure 6. The time each lion spent reactive (minutes) for each playback treatment. All lions spent different amount of time paying attention to the playback and received a score for every minute. The black lines represent the median time for each treatment. Maximum and minimum time can also be read. The playback of five, eight and eighteen lions have outliers, again having values standing out from the other values because of some individuals being much

more active than others.

0

2

4

6

8

10

12

14

16

0 1 2 3 4 5 6 7 8 9 10

Boldness

Age

A B C D E F G H I J

05

1015

2025

30

Intruders

Tim

e.ac

tive

Eigh

teen

Thirt

een

2

Thirt

een

1

Eigh

t

Five

Thre

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ales

2

mal

es 2

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ales

1

Thre

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mal

es

One

mal

e

One

fem

ale

Tim

e ac

tive

(min

)

Intruders

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3.2 Social interactions

The density values for greet, groom, aggression, play and all social were 0.30, 0.11,

0.04, 0.08, and 0.47, respectively. For all social interactions, the lion to initiate most

interactions was Rusha, with an out-degree value of 1.73, while initiate least for all

social were Zulu and Kwandi (0.18). The individual receiving most interactions for all

social was Zulu (in-degree value 2.09) and those that received the least attention were

the sub-adults RS3 and LE1 (0.36) (Appendix C). Rusha was also the lion with the

highest betweenness centrality value of 8.08 for all social interactions and the lions with

the lowest value was sub-adult LE3 (Appendix D). There were no significant

correlations between boldness and any of the social interactions. The closest

relationship found was between boldness and in-degree of all social interactions (rs =

0.541, p = 0.069), suggesting that bolder individuals were likely to receive more social

interactions. There was a significant positive correlation between grooming indegree

and outdegree (rs = 0.88, p < 0.001), indicating that individuals who initiate grooming

are also those who receive the interaction. Initiating grooming behaviour was

overrepresented by females (W (29), p = 0.026). Males received more aggression than

females (W (5.5), p = 0.041). There was no significant difference for which sex initiated

the most aggressive interactions. A significant negative correlation was found between

age and play (rs = -0.66, p = 0.021), indicating that the younger lions were still more

playful than the adults. It was also the younger lions that received the most aggression

(rs = -0.63, p = 0.028).

The lions most central in the all social network were the ones that initiated most greets

(rs = 0.66, p = 0.021). Play betweenness centrality values were negatively correlated

with being aggressive (rs = -0.59, p = 0.044) and positively associated with play out-

degree (rs = 0.67, p = 0.018), which indicates that the lions most central to the play

network were also the most playful individuals (the sub-adults). The lions most central

in the groom network greeted the most (rs = 0.61, p = 0.037) and received most

grooming (rs = 0.75, p = 0.005). Lastly, greet betweenness centrality values increased

with greet in-degree (rs = 0.71, p = 0.01) and decreased with play out-degree (rs = -0.69,

p = 0.013), suggesting that the lions most central in greet received most greetings but

initiated play least of all.

The sociograms for each social interaction type are presented in figure 7, where the

visualization of all social interactions (1) together indicate that the strongest connection

was between Rusha and Zulu. There were also strong linkages between Rusha and

Leya, LE1 and Zulu, and finally between LE3 and Zulu. The lion with least connections

within the network was RS3, only seen interacting with five out of eleven lions.

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Figure 7. Sociograms illustrating the social relationships between lions in the pride for each social network; all social (1), aggression (2), groom (3), greet (4) and play (5). Squares represent males and circles represent females. Size is in proportion to the age of the lion, with larger the node, the older the lion.

4 Discussion Rusha was the boldest individual in this pride and also had most social interactions in

the network. At the same time, sub-adult RS3 was the lion with the lowest boldness

score and had least interactions. Pike et al. (2008) explored the effect of social

12

interactions in an animal network in three-spinned sticklebacks (Gasterosteus

aculeatus), but had the opposite result. The boldest individuals were those having fewer

interactions, while the shy individuals had more, but with a small number of group

members. Even though no significant correlation could be found between boldness and

degree in the present study of lions, it was still a close relationship between being bold

and receiving interactions. A conclusion can therefore not be made that with a larger

sample size and more data, a significant correlation between these variables would still

not be found.

Interestingly, RS1 and RS2 spent the most time being reactive to the playback, followed

by their mother Rusha. Even though they were not consistent in boldness throughout the

playbacks, they both approached when conducted the first playback of three males.

These three pride members had both the highest degree of receiving interactions, after

Zulu, and also spent most time reactive to the playbacks. It is therefore possible that

these sub-adults will become more like their mother as they become adults, being bold

and protective when they have been released into the wild.

It was expected that Zulu would receive the most greets from the other lions, since he is

the alpha-male and greeting him is a way of showing respect (Schaller, 1972). However,

I found that initiation of grooming was female dominated in the pride. Schaller (1972)

observed that grooming is a way of strengthening the social bond between adult females

within the pride. Since males are not supposed to stay in the same pride their whole

lives, as females do, it might not be as important for them than for the females. Female

grooming behaviour can also be seen in several different primates (Henzi & Barrett,

1999). The reason why age was not correlated with grooming behaviour might be

because the sub-adults are on the verge of becoming adults. The sub-adults were still

more playful than the adults, which is expected since the younger the lions the more

playful they are in the wild (Schaller, 1972). The lions in the pride were most aggressive

towards the male sub-adults, but also to female RS1. It was not very surprising given

that the sub-adults received most aggression, since the adults do this to discipline the

younger pride members.

The focus on sex and age differences led me to the social network analysis, which is a

good way of evaluating the social structure of a pride, while also providing important

information when preparing for an ex-situ reintroduction. Before releasing the sub-

adults, an individual’s role in a group must be known to help predict their success post

release. These analyses can, and should, be done over a longer period of time to see if

the structure changes and if so, establish a reason for these changes. By having a study

conducted over time, each individual’s social role can be assessed for information of

how important it is socially for the pride’s stability (McCowan et al. 2008; McCowan et

al. 2011). Overall, sex and age did not affect how central an individual was in mediating

and controlling relationships within the pride, and except for RS3 and LE3, the other

sub-adults appeared to contribute to keeping the network together. This in turn might

suggest that when reaching adulthood and post-release these lions will hopefully be

capable of establishing a socially cohesive pride. RS3 and LE3, however, might not be

as cohesive to their siblings when released, being candidates to become nomads. This

can of course not be concluded, especially not from this short time of study, but from

the look of their low betweenness values they might not stay together as close as the

other lions after release.

13

Different studies suggest various explanations for variation in behavioural traits. These

range from genetics (Van Oers et al. 2005) to life-history stage, individuals’ experiences

(Sih et al. 2004; Réale et al. 2007; Dall et al. 2004) and also on maternal effects (Stamps

& Groothuis, 2010). Yet, experience in life can make a consistent behaviour non-stable.

Bell and Sih (2007) found a higher correlation between boldness and aggression in the

threespined stickleback (Gasterosteus aculeatus) when have been exposed to predators.

An association between being bold and aggressive has been reported in more species

(Dochtermann & Jenkins, 2007; Duckworth & Badyaev, 2007; Johnson, & Sih, 2005;

Kortet & Hedrick, 2007; Moretz et al. 2007). However, in the house cricket (Acheta

domesticus), for example, no correlation could be found between boldness and being

aggressive (Wilson et al. 2009). This was also the case in this study of lions, where no

correlation could be found between boldness and aggressiveness. Having grown up in a

safe environment with no threats around them, the lions, especially the bolder ones,

might become more aggressive when encountered with unfamiliar conspecifics and

competitors, like hyenas, in the wild.

As previously mentioned, experiences in life can change an earlier consistent behaviour

(Bell & Sih, 2007). The future release into the wild of the six sub-adults means a big

change for these individuals, with a completely new environment and challenges to

face. The reactions shown by the sub-adults do not necessarily indicate that, when in the

wild, they would exhibit the same level of boldness. It is also difficult to say who would

have a higher chance of survival, since being a bold individual can both be beneficial

and a disadvantage. In a study of captive-bred swift foxes (Vulpes velox), the

individuals with the highest boldness score when captive were those with the lowest

survival after release (Bremner-Harrison et al. 2004). The reason for this is probably

because of the lower fear of approaching a novel object, or that bolder individuals are

less likely to avoid intrusion of territorial conspecifics, predators and anthropogenic

stimuli (Bremner-Harrison et al. 2004). On the other hand, boldness could be species

specific since Sinn et al. (2013) showed that reintroduced Tasmanian devils that

survived were 3.5 times bolder than those that did not. Being bold has also shown to

increase the reproductive success in other animals, especially in males (Smith &

Blumstein, 2008), coming to the conclusion that boldness is also a “trade-off” in fitness

across different situations (Smith & Blumstein, 2008).

All adult female lions in this study, except Rusha, had a total boldness score of zero.

Either the others were just not interested, or whether they might have known that Rusha

is the one who would cover for them is unknown. These females could in that case be

termed laggards, since those are the individuals not sharing equally in defending the

pride as Rusha, a leader, would (Breed & Moore, 2011). Consistent behaviours make

individuals predictable, which can be beneficial in a social group (Wolf et al. 2011).

The response of an individual can be influenced of its group member’s earlier behaviour

in a specific situation, and so, the laggards might act as they do because they know that

their group members will take the lead and face the threat. In this pride, Rusha’s

behaviour might was predictable to the other females, which could be one explanation

for their lagging behaviour.

An earlier study did not find any relationship of body size and territorial responses of

female lions (Heinson & Packer, 1995) and no measures were taken on the pride in

Zambia to investigate this. However, Rusha is by far the largest female in body size and

height, while Kela and Kwandi with the lowest boldness scores, are the smallest

14

lionesses. With a longer period of study and height measures of the lions, this would be

interesting to see whether body size is associated with boldness.

When conducted the second playback of three males, the oldest male sub-adult, RS2,

was absent and Zulu started roaring. Since three incoming males would be the biggest

threat for males in a pride, it is possible that roaring in response to this playback was for

calling RS2 to join for defense. Male lions often form coalitions that consist of either

kinship or non-relatives (Packer et al. 1991b). The purpose of forming a coalition is for

cooperation of keeping the access to females increasing their reproductive success and

helping each other to protect their pride from incoming males to take over (Bygott et al.

1979). As RS2 gets older, it would be interesting to see whether he and Zulu would

actually form an apparent coalition.

The non-significant difference in age for reactions to the playbacks indicates that the

younger lions are not so dependent of their mother anymore as they are reaching

adulthood. Heinsohn et al. (1996) studied juvenile lions’ reactions to playbacks of

unfamiliar lion roars and the developmental responses between the age of eight and 42

months old. The result showed that when no adults were present, it was a 70% chance

of approaching by the age of 37-42 months. Females showed a higher probability of

responding with age, while males did not, but were seen most often in the back of the

group when responding. The sub-adults in the Dambwa pride were never seen without

adults when conducting the playbacks, since the pride usually stayed all together, and

neither was there any difference between the sexes. Again, having almost turned four

years old and considering the other juveniles were three years old, the sub-adults should

be in the age of being bold enough to approach. The lack of time and replications also

made it difficult to find any consistencies in boldness, making it impossible to come to a

conclusion regarding boldness differences between the individuals.

Because the pride studied here usually stayed together, it was not possible to examine

the effect of group size, which has been done for wild prides (McComb et al. 1994).

Because of the bigger reactions when five individuals were not present (if this had to do

with this or the fact that three males were played), a longer period of study might have

allowed for looking at the changes of responses depending on the size of the pride.

McGregor (1992) proposed some of the problems with conducting playbacks to

animals. The first problem is about the risk of habituation. When animals are exposed to

the same sounds, and even different sounds but at the same or similar situations, they do

tend to ignore them in the end or the reactions will not be as strong as during the first

playbacks. The second and also more serious problem, is that the vocalizations being

played might not mean what they actually were thought to. This study was designed to

play territorial lion roars, but even though this was the search criteria when looking for

the recordings, they might not have been perceived aggressive for the pride. The more

playbacks that were done, the more the lions seemed to pay less attention to the

playback, which could have been because of habituation. In nine out of ten playbacks,

the lions stayed at the same location close to the fence, making it difficult to change the

place to put the loudspeaker. This might have made the playbacks even less trustworthy

towards the end of the experiment. Also, staying close to the fence might affected their

response in the way that they would feel more secure being protected behind the fence.

Neither sex nor age had any effect on boldness and time active to the playbacks, though

the higher number of intruders being played increased the time active to it. But there

15

was only an increase until playing more than three lions and the following increase in

number gave an unstable mean value of time scores. The low response of the lions when

playing eight, thirteen and eighteen lions may be due to the possible confusion of the

number of lions being played. As it is known about the lions’ ability to count the

number of roaring intruders, they might also hear when it does not sound like a real

pride. Earlier studies have only focused on playing maximum three lions at the same

time (McComb et al. 1994; Grinnell et al. 1995; Heinsohn & Packer, 1995; Heinsohn et

al. 1996). However, one of the aims in this study was to investigate their reactions when

being outnumbered. The hypothesis was that they were not going to approach when

playing more lions than the number present in the pride, which was true in this case, but

looking at the time active to the playback it seemed as it was more a question of

interest.

4.1 Conclusion

This study does not only provide information about a captive-origin lion pride, but also

about lions’ social behaviours. The lions in the Dambwa pride did not really behave as

expected when it came to playback experiments compared to wild lions. Although, the

playbacks gave information on who was the boldest and least bold individual in the

pride. These findings were then compared with their social behaviour, where bold

individuals were more likely to receive social interactions from other lions and the

opposite. Being more central in a specific network was associated with other social

behaviours, and some social interactions were more dominated by a certain sex or age.

Social network analysis was a good method of exploring the current social structure in

the pride. There was evidence of some individuals who preferred each other more than

others and some were better in keeping the pride together socially than others. This

should be followed up until the day of release to see if any changes occur and establish

whether the individuals to be reintroduced are well-prepared. The differences in

sociality associated with age and sex were mostly in agreement with what has

previously been reported based on studies of behaviours of wild lions, indicating that

captive-bred lions still display natural behaviours, which is of great importance when

preparing for a release.

16

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I

5 Appendices

5.1 Appendix A - The lions

Name Sex Birth

Zulu M 8th July 2008

Kela F 3rd May 2008

Kwandi F 3rd May 2008

Leya F 26th May 2008

Loma F 26th May 2008

Rusha F 9th August 2008

RS1 F 21st June 2013

RS2 M 21st June 2013

RS3 F 21st June 2013

LE1 M 30th January 2014

LE2 F 30th January 2014

LE3 M 20th January 2014

II

5.2 Appendix B – Boldness scores

Yes No

Approach > 5 m 1 0

Approach 5-10 m 2 0

Approach 10-20 m 3 0

Approach 20-50 m 4 0

Approach > 50 m 5 0

Going opposite direction -6 0

Retreats after > 5 m -5 0

Retreats after 5-10 m -4 0

Retreats after 10-20 m -3 0

Retreats after 20-50 m -2 0

Retreats after > 50 -1 0

Glance back to pridemembers

-1 0

1-3 pauses -1 0

4-6 pauses -2 0

> 7 pauses -3 0

No pauses 1 0

Roaring 1 0

III

5.3 Appendix C – Result for out- and in-degree values

Table 1. Out-degree values for all interactions. The higher value the more does the individual interact with others.

Out-degree

Aggression Greet Groom Play All social

Zulu 0.00 0.18 0.00 0.00 0.18

Kela 0.18 0.36 0.27 0.00 0.82

Kwandi 0.00 0.09 0.09 0.00 0.18

Leya 0.09 0.36 0.18 0.00 0.64

Loma 0.00 0.64 0.09 0.18 0.91

Rusha 0.09 1.36 0.27 0.00 1.73

RS1 0.00 0.27 0.27 0.46 1.00

RS2 0.09 0.46 0.00 0.09 0.91

RS3 0.00 0.18 0.09 0.27 0.27

LE1 0.00 0.82 0.09 0.09 1.18

LE2 0.00 0.36 0.18 0.18 0.82

LE3 0.00 0.27 0.09 0.18 0.82

Table 2. In-degree values for all social interactions. The higher value, the more interactions does the individual receive from others.

In-degree

Aggression Greet Groom Play All social

Zulu 0.00 1.91 0.00 0.18 2.09

Kela 0.00 0.27 0.27 0.00 0.55

Kwandi 0.00 0.73 0.00 0.00 0.73

Leya 0.00 0.27 0.27 0.18 0.73

Loma 0.00 0.27 0.18 0.00 0.46

Rusha 0.00 0.64 0.27 0.00 0.91

RS1 0.09 0.36 0.27 0.36 1.09

RS2 0.18 0.46 0.00 0.27 0.91

RS3 0.00 0.00 0.00 0.36 0.36

LE1 0.09 0.09 0.18 0.00 0.36

LE2 0.00 0.36 0.18 0.09 0.64

LE3 0.09 0.27 0.00 0.00 0.64

IV

5.4 Appendix D – Result for betweenness centrality values

Table 3. Betweenness centrality values for four different social interactions and all together. Age and sex of the lions are also shown to more easily compare for the differences in social behaviours.

Betweenness

Lions Age Sex Groom Greet Play Aggression All

social

Zulu 9 M 0.00 8.91 10.00 0.00 3.67

Kela 9 F 17.58 4.09 0.00 5.46 2.29

Kwandi 9 F 0.00 8.85 0.00 0.00 1.87

Leya 9 F 12.73 1.42 6.36 0.00 1.17

Loma 9 F 20.61 1.52 8.18 0.00 2.65

Rusha 9 F 5.15 24.91 0.00 0.00 8.08

RS1 3 F 15.15 0.46 11.82 0.00 2.60

RS2 3 M 0.00 1.94 0.00 9.09 4.77

RS3 3 F 0.00 0.00 15.46 0.00 0.57

LE1 4 M 18.18 1.42 0.00 0.00 1.43

LE2 4 F 5.15 2.18 20.91 0.00 6.81

LE3 4 M 0.00 2.49 7.27 0.00 0.46