Please find here a sample of the research that has been, or is being, undertaken as part of the African Lion Rehabilitation & Release into the Wild Programme. 

Investigating the impacts of captive origin, time and vegetation on the daily activity of African lion prides

Evaluation of activity budgets provides an indication of whether captive history has influenced the daily behaviour of animals within an ex situ reintroduction program. We conducted the first study to compare activity budgets of prides of captive-origin African lions (Panthera leo) to a wild pride investigated under the same methodologies. Behavioural data were collected via direct observations of individual lions. The vegetation type in which each pride was located was recorded at the beginning and end of each observation session. Behaviours were analysed via linear mixed models, using restricted maximum-likelihood analysis. Age, sex, origin, time observed and vegetation type were fitted as fixed factors to assess the main effects of significant interactions, while average temperature was fitted as a covariate. Resting and alert behaviour were found to vary at sex and age levels, with expected peaks and lows coinciding with observation session times. Captive-origin prides showed a decrease in resting and increase in alert and movement behaviours post-1700 hours, while this behavioural change was not observed for the wild prides. Males of the wild prides were observed to rest more than captive-origin counterparts,while this variation was not observed for females across origins. Vegetation was found to influence behaviour, with cubs being more alert and active in riverbed vegetation,and adults and sub-adults in shrubland. Overall, all prides were observed to exhibit behaviours at natural levels. This study provides a comprehensive assessment of the daily activity of lions critical to the pre-release evaluation of prides within an ex situ reintroduction program.

Dunston EJ, Abell J, Doyle RE, Kirk J, Hilley VB, Forsytth A, Jenkins E, McAllister D, Freire R (2017) Investigating the impacts of captive origin, time and vegetation on the daily activity of African lion prides.  Journal of Ethology DOI 10.1007/s10164-017-0508-x.  Read the full text here.

A Social Network Analysis of Social Cohesion in a Constructed Pride: Implications for Ex Situ Reintroduction of the African Lion (Panthera leo)

Animal conservation practices include the grouping of captive related and unrelated individuals to form a social structure which is characteristic of that species in the wild. In response to the rapid decline of wild African lion (Panthera leo) populations, an array of conservational strategies have been adopted. Ex situ reintroduction of the African lion requires the construction of socially cohesive pride structures prior to wild release. This pilot study adopted a social network theory approach to quantitatively assess a captive pride’s social structure and the relationships between individuals within them. Group composition (who is present in a group) and social interaction data (social licking, greeting, play) was observed and recorded to assess social cohesion within a released semi-wild pride. UCINET and SOCPROG software was utilised to represent and analyse these social networks. Results indicate that the pride is socially cohesive, does not exhibit random associations, and the role of socially influential keystone individuals is important for maintaining social bondedness within a lion pride. These results are potentially informative for the structure of lion prides, in captivity and in the wild, and could have implications for captive and wild-founder reintroductions.

Abell J, Kirzinger MWB, Gordon Y, Kirk J, Kokeŝ R, et al. (2013) A Social Network Analysis of Social Cohesion in a Constructed Pride: Implications for Ex Situ Reintroduction of the African Lion (Panthera leo). PLoS ONE 8(12): e82541. doi:10.1371/journal.pone.0082541. Read this article here.

An Assessment of African Lion (Panthera leo) Sociality Via Social Network Analysis: Pre-release Monitoring for an Ex Situ Reintroduction Program

The wild population of the African lion Panthera leo continues to decline, requiring alternate conservation programs to be considered.  One such programme is ex-situ reintroduction. Prior to release, long-term monitoring and assessment of behavior is required to determine whether prides and coalitions behave naturally and are sufficiently adapted to a wild environment. Social Network Analysis can be used to provide insight into how the pride as a whole and individuals within it, function. Our study was conducted upon two captive-origin prides who are part of an ex-situ reintroduction program, and one wild pride of African lion. Social interactions were collected at all occurrence for each pride and categorised into greet, social grooming, play and aggression.  Betweenness centrality showed that offspring in each pride were central to the play network, while degree indicated that adults received (indegree) the greatest number of overall social interactions, and the adult males of each pride were least likely to initiate (outdegree) any interactions. Through the assessment of individual centrality and degree values, a social keystone adult female was identified for each pride. Social network results indicated that the two captive-origin prides had formed cohesive social units and possessed relationships and behaviors comparable with the wild pride for the studied behaviors. This study provided the first SNA comparison between captive-bred origin and a wild pride of lions, providing valuable information on individual and pride sociality, critical for determining the success of prides within an ex-situ reintroduction program.  

Dunston EJ, Abell J, Doyle RE, Kirk J, Hilley VB, Forsyth A, Jenkins E, Freire R (2016) An assessment of African lion Panthera leo sociality via social network analysis: pre-release monitoring for an ex-situ reintroduction program.  Current Zoology OnlineFirst.  Download this article here.

Re-introduction of the African lion from a captive origin: Zambia & Zimbabwe
In March 2016 the IUCN SSC Re-introduction Specialist Group published the 2016 edition in its Global Re-introduction Perspectives series.  ALERT's contribution is a case study that summarises the aims of the African Lion & Rehabilitation & Release into the Wild Programme, and reports on the success and failures of this ongoing conservation project.
Youldon D, Abell J, Monks N (2016) Re-introduction of the African lion from a captive origin: Zambia & Zimbabwe.  In: Global Re-introduction perspectives: 2016.  Case Studies From Around the Globe (ed. Soorae, PS).  IUCN SSC Re-introduction Specialist Group and Abu Dhabi, UAE: Environment Agency - Abu Dhabi. xiv + 276 pp (pp153 - 156).  Download the book from here.

Exploring African lion (Panthera leo) behavioural phenotypes: individual differences and correlations between sociality, boldness and behaviour

Increasing our understanding of personality, at an individual and group level, is crucial to the pre-release assessment of social species within ex situ reintroduction programs. We conducted the first exploration into the personality of a captive-origin pride of African lions (Panthera leo), assessing behavioural variations and consistencies in daily activity, social and hunting behaviour, and boldness. Data were collected via direct observations, while a species-specific protocol for testing boldness, using playbacks, was developed. Differences in sex, age and session time for the activity budget were evaluated using Pearson correlations and repeated-measures ANOVA, while social interactions were analysed using social network analysis. Spearman’s correlations were conducted to assess for associations between boldness scores, activity and sociality. The two boldness tests provided a range of scores per lion, indicating that the test was effective. Correlations and variations in individual behaviour indicated that adults and sub-adults have specific roles within pride behaviour. Correlations between boldness and activity and social behaviours provided information on the role of individuals, allowing investigation into the behaviour of a dominant and a social keystone. Our study indicates that evaluating various aspects of behaviour in conjunction with boldness has the potential to assist the pre-release assessment of a pride within an ex situ reintroduction program.

Dunston EJ, Abell J, Doyle RE, Evershed M, Freire R (2016) Exploring African lion (Panthera leo) behavioural phenotypes: individual differences and correlations between sociality, boldness and behaviour. Journal of Ethology doi:10.1007/s10164-016-0473-9

Behavioural correlates between daily activity and sociality in wild and captive origin African lions
Study of behavioural correlations within and across populations has long been of interest to ethologists. An exploration of behavioural correlations between sociality and behaviour of African lions (Panthera leo) was undertaken to examine if this approach is better able to reveal important aspects of lion behaviour not easily discernible by looking at these behaviours separately. Resting behaviour and received play interactions were correlated in two captive-origin prides and one wild pride, attributable to the involvement of cubs and sub-adults.
Direct and exploratory movement was negatively correlated with groom centrality in two of the three prides, due to adults engaging in high levels of both of these activities. Exploration of these behavioural correlations highlighted the differences between age-groups in activity and sociality, facilitating the understanding of the complex behaviour and interactions of lions. In addition, the finding of similar behavioural correlations between captive-origin and the wild prides provides confidence in the suitability if captive-origin candidates for ex-situ release. This is imperative to ensure the success of sub-groups and prides under an ex-situ reintroduction program.
Dunston EJ, Abell J, Freire R (2016) Behavioural correlates between daily activity and sociality in wild and captive origin African lions.  Communicative & Integrative Biology, DOI: 10.1080/19420889.2016.1208874

Reintroduction of social carnivores: effect of composition on cooperation and hunting success in lion hunting groups

Reintroduction of endangered species is important in conservation biology. An additional aspect in social carnivores is forming a social bond to increase their fitness. Lions form social prides to hunt cooperatively on larger prey and become more efficient hunters. This study was within a reintroduction project in Zambia, where captive-bred lions learn to hunt and form a social pride in a four stage program. The main question was what the effect is of mixing lion hunting groups on their cooperative behaviour and which composition is the most effective. Composition of groups was changed, by putting lions together that did not hunt together before. The results show no difference in cooperation among different group compositions. However cooperation decreases when hunting groups change every encounter. Another result is the lightest lion starts the stalk, while the heaviest lion makes the kill. Just like wild lions they are able to cooperate by a division of labour based on body size. But towards the end of the study, the lions decided to hunt more solitary instead of cooperatively, meaning the experimental setup had a negative influence on lion’s cooperative behaviour. Lions do not need to be together from young age, to cooperate during hunting. They still cooperate in new group compositions. This study adds to the knowledge how to successful introduce social carnivores into the wild. Often the causes of reintroduction failure are unknown, which underlines the importance of researching the used methods.

Jansen S (2011) Reintroduction of Social Carnivores:  Effect of composition on cooperation and hunting success in lion hunting groups.  Master’s thesis Biology / Animal Ecology.  Supervisor: Dr. Pim van Hooft, Resource Ecology Group, Wageningen University.

Read the full article here:

Influence of social upbringing on the activity pattern of captive lion (Panthera leo) cubs: benefits of behaviour enrichment

The influence of social upbringing on the activity pattern of lion (Panthera leo) cubs was investigated at three sites. In this study, stimulus objects such as sticks, grass, fresh dung (elephant Loxondota africana, zebra Equus quagga, impala Aepyceros melampus, duiker Sylvicapra grimmia, kudu Tragelaphus strepsiceros, giraffe Giraffa camelopardalis and wildebeest Connochaetes taurinus) and cardboard boxes, were utilized in an enrichment program aimed at encouraging active behaviours of captive lion cubs in two sites of the African Lion Rehabilitation & Release into the Wild Program. Lion cubs at Chipangali were not behaviourally enriched. Activity patterns were recorded for 10 days at each site. We recorded moving, resting, playing, grooming, visual exploration and display of hunting instincts. We found that behavioural enrichment enhanced the active behaviours of captive lion cubs. Orphan-raised cubs spent more time moving, playing and displaying hunting instincts than mother-raised cubs, but the time spent grooming was similar across areas and suggests that grooming is not influenced by enrichment. Mother-raised cubs spent more time engaged in visual exploration than orphan-raised cubs and this could be a behaviour acquired from mothers or a result of confidence to explore because of their presence. Activity patterns were different among time treatments across our three study sites. Based on these findings, we suggest that lion cubs raised in captivity could benefit from behavioural enrichment to encourage active behaviours essential for eventual reintroduction into the wild.

Ncube S, Ndagurwa HGT (2010) Influence of Social Upbringing on the Activity Pattern of Captive Lion (Panthera leo) cubs:  Benefits of behaviour enrichment.  Current Zoology 56 (4). Undergraduate thesis in Forest Resources and Wildlife Management.  Supervisor Prof. Peter Mundy, Department of Forest Resources and Wildlife Management, Faculty of Applied Sciences, National University of Science & Technology, Bulawayo, Zimbabwe

Read the full article here, or download as a (pdf)

The possible benefit of human presence and interaction upon play in lion cubs (Panthera leo) and differences in play types amongst different cub age groups

Four categories of play behaviour exhibited by captive bred African lion (Panthera leo) cubs were investigated: social, object, locomotory and predatory. The aim of this study was to establish the effect of human presence and absence upon play and determine the benefits of human presence upon juvenile development. This study also investigated the differences between play types and frequencies amongst three groups of cubs of different ages. Data was collected over four months within the African Lion Reintroduction & Release into the Wild Program in the Mosi‐Oa‐Tunya Natonal Park in Zambia. Observations were conducted under three conditions, human presence during walks, human presence during enclosure time and no human presence during enclosure time.  The three groups of cubs were aged (at the beginning of the study) 17 months, 10 months and 5 months. This study found that younger cubs exhibited play behaviours thought to develop neuromuscular and adult skills (object and locomotory play) more often than older cubs, which exhibited social play more often comparatively to other play behaviours. This study also found all three groups of cubs exhibited play far more frequently during human presence than absence. This is thought to be due to the mimicry of other pride members people create when present upon walks and when in enclosures. Based on these findings this study recommends the use of human‐wildlife relationships with cubs within carefully managed and controlled ex‐situ reintroduction programmes to help promote play behaviour and thus improve development of skills required for self‐sufficiency during adulthood.

This paper is currently being prepared for submission to an academic journal.

Kokes HR (2010)  The possible benefit of human presence and interaction upon play in lion cubs (Panthera leo) and differences in play types amongst different cub age groups.  Master’s thesis in Animal Behaviour.  Supervisor Dr. Barry Stevens-Wood, Behavioural and Environmental Biology Department of Biological Sciences, Manchester Metropolitan University

Behavioural sequencing and character development in lion cubs

This study aims to identify character traits within lions and then work out at what age they can be reliably assessed. Studies have observed that when vocalisations of non-resident prides are played back to the resident pride not all lions approach the “intruders”. Interestingly, when these tests are repeated over time observers found that the approaching lions will always be the same individuals and approach in the same order. Observations in the wild have shown that when territorial fights break out between resident prides some individuals will only join in when the pride is outnumbered; otherwise they refrain from fighting despite the advantage it would give their own pride. These behaviours are consistent over time. This may indicate that it is possible to identify individuals that are likely to either be highly territorial, or be non-confrontational.  There is little evidence of formal roles within prides but it has also been observed that lions repeatedly take the same roles during hunts.

This study aims to link character traits to roles within a pride with the goal of identifying lions that will be suitable for pride releases together in the African Lion Reintroduction into the Wild Program with the highest chance of success. Lions that can be introduced to each other at an early age have a much stronger chance of socially bonding. This is observed in the wild as well as in captivity. Lion introductions with adults can be extremely dangerous and result in fatalities, much as territorial disputes occur in the wild.  As such, being able to identify the development of different character traits in lions at a young age will assist in forming release prides earlier with advantages to social bonding, whilst also ensuring that the pride consists of a natural balance of character types.

Clifforde L (ongoing study) Behavioural Sequencing and Character Development in Lion Cubs.  MPhil thesis.  Supervisor Dr. Sasha Dall, Department of Biosciences, Exeter University

A critical analysis of equity in nature based tourism

The purpose of this research is to explore the concept of equity in the context of nature based tourism (NBT), using a case study of ALERT & Lion Encounter in Livingstone, Zambia.  Specifically, the various conceptualizations of equity are identified and their influence on NBT expectations, decision-making, and outcomes are analyzed. 

Critical theory is discussed as a way to analyse the fundamental traditions (ethics, distributive justice and procedural justice) that shape how equity is conceptualized, and systems thinking is presented to better understand equity with NBT.  Since NBT benefit distribution demonstrates a link between decision-making power, benefit distribution, and equity, NBT governance is also addressed.

The case study is expected to demonstrate how equity is conceptualized in NBT decision making and benefit distribution but also how equity contributes to broader tourism, conservation and development discussions.

Becarra L (ongoing study) A critical analysis of equity in nature based tourism.  PhD thesis.  Supervisors Prof. Wayne Freimund & Dr Norma Nickerson, Department of Society and Conservation, University of Montana.

Acoustic analysis of lion roars

Part One: Data collection, spectrogram and wave analyses

Felids are one of the most successful carnivore families ever to exist, and within the 35–40 different cat species that exist today several different vocalizations can be found, with different functions, ranging from the well-known purring to the most impressive sound of them all: roaring of lion (Panthera leo) fame. This paper focuses on the impressive lion roaring, and highlights methodological problems associated with the collection of animal vocalizations data.

Eklund R, Peters G, Ananthakrishman G, Mabiza E (2011) An acoustic analysis of lion roars. I: Data collection, spectrogram and waveform analyses.  Proceedings from Fonetik 2011.

Part two: Vocal tract characteristics

This paper makes the first attempt to perform an acoustic-to-articulatory inversion of a lion (Panthera leo) roar. The main problems that one encounters in attempting this, is the fact that little is known about the dimensions of the vocal tract, other than a general range of vocal tract lengths. Precious little is also known about the articulation strategies that are adopted by the lion while roaring. The approach used here is to iterate between possible values of vocal tract lengths and vocal tract configurations. Since there seems to be a distinct articulatory changes during the process of a roar, we find a smooth path that minimizes the error function between a recorded roar and the simulated roar using a variable length articulatory model.

Ananthakrishman G, Eklund R, Peters G, Mabiza E (2011) An acoustic analysis of lion roars. II: Vocal tract characteristics. Proceedings from Fonetik 2011.

Eklund et al. are a group comprising individuals from: the Department of Cognitive Neuroscience, Karolinska Institute; the Centre for Speech Technology, Royal Institute for Technology in Stockholm; the Department of Computer Science, Linköping University in Sweden, and Forschungsinstitut Alexander Koenig, Bonn.

Read the full articles here

Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).

A behaviour study was carried out at Antelope Park in Zimbabwe on 16 captive male breeding Africa lions (Panthera leo) in order to evaluate the effectiveness of three low cost behavioural enrichments and their effects on territorial behaviour. The behaviours that were observed for the study included active locomotion, vocalisation, resting, behaviour enrichment (BE) - feeding related, BE- touch related, social behaviour, territorial behaviour, grooming, aggression and stereotypical behaviour. The study was carried out over an 18 day period producing 108 hours of observations in total. The three different types of enrichments which were tested with the lions in the investigation on a rotational basis included a log covered in zebra (Equus quagga) dung, a Flat White Boer Ford pumpkin (Cucurbita maxima) and a log with Cow blood. 

The hypotheses that were formulated for the behavioural study included: “Lions will interact most with the food related enrichment (cow blood log) more than the other two behavioural enrichments”. “Zebra dung and cow blood behavioural enrichments will receive most interactions throughout the day”. “More territorial behavioural displays will be exhibited with the presence of the Zebra dung log and cow blood log enrichments” and “Zebra dung log and cow blood log will increase activity levels”.

The results from the investigation concluded that the presence of the log with zebra dung enrichment significantly increased behaviours such as active locomotion, vocalisation and territorial behaviour. The statistical tests carried out showed an increase in these behaviours with a significant P-Value of <0.05, indicating a significant difference between data. Both behavioural enrichment interaction categories; BE- feeding related and BE- touch related was also increased with the presence of the log with zebra dung enrichment, leading to the conclusion that this enrichment type was the most successful treatment in this investigation. 


Keel-Stocker M (2014) Evaluating the effectiveness of natural, low cost behavioural enrichments and their effects on territorial behaviour of conservation breeding male African lions (Panthera leo).  Bachelor's degree thesis in Animal Conservation & Biodiversity.  University of Greenwich.