Studying personality in captive animals may enable the development of individual-based management decisions, which may improve animal welfare. Asiatic lions at London Zoo represent an opportunity to research an understudied species’ response to new environments since they have experienced social and physical changes, such as new enclosures and increased social interaction with humans. This project aimed to investigate the role of personality in behavioral responses to these changes. Lion personality questionnaires completed by keepers and direct focal animal observations were used to create personality profiles. Time budgets and enclosure use were determined and compared between control nights and event nights and between the lions’ previous enclosure and their new one. The results showed a lack of difference in time budget and enclosure use between control and social event nights, and the spread of participation index values revealed that the lions use their enclosures unevenly. Personality profiles identified various traits that could assist with individual-based management decisions. As the first study to assess Asiatic lions personality, this research contributes to the creation of consistent and valid methodology for evaluating captive animal personality that may improve husbandry and welfare protocols for individual lions, leading to the improved health and success of the species.
Animal personality research has continually developed since Pavlov’s studies on dogs [
Previous studies have demonstrated that animal personality is measureable, has a degree of cross-species comparability, and can be assessed in various species, including mammals, fish, and insects [
Practical applications of personality research in captive animals include husbandry, training, and breeding programs. Personality assessment facilitates more individual-based management, which may help to maximize the welfare and overall success of a captive collection [
Prior to introducing an animal to a captive collection, personality assessment can help to determine how the new individual will affect group dynamics [
Animal personality is evaluated using various methods, including coding personality traits based on observations of natural behavior or a specific test (e.g., novel object test), and trait rating completed by keepers [
Because of possible discordance, it is important to test additional tools to validate personality profiles. The novel object test, which evaluates an animal’s response to new objects (e.g., traffic cone), is commonly used for this purpose [
The Asiatic lions
The Asiatic lion is a lion subspecies that resides in Gujarat, India, and is listed as Endangered by IUCN [
The social and physical changes experienced by the lions guided the development of this study, which aims to evaluate the role of lion personality in their behavioral responses to new environments. This study hypothesizes that personality traits identified from keeper questionnaires and observation data create reliable profiles that associate with individual lion behavioral responses to new physical and social environments. Therefore, because of individual personality variation, this study also hypothesizes that these new environments will alter individual time budgets and enclosure use. To test these hypotheses, previously collected behavioral data (i.e., time budget and enclosure use) from Whipsnade Zoo were compared with data from their new enclosure at London Zoo. These data were also compared between control nights and Sunset Safaris. Considering this behavioral data, personality profiles were constructed to determine if certain traits are associated with individual lion responses to new environments. A sociogram was constructed to determine if the relationships between the lions are impacted by their individual personalities.
This study can be considered a case study that may be used to improve the management of these four individuals. Furthermore, this research has wider implications for management of the species, in terms of husbandry, enclosure design, health, welfare, and breeding program success. As of December 2015, there were approximately 359 Asiatic lions in captivity (Srivastav, 2016, pers. comm.). Therefore, a study on four animals can provide essential captive lion behavior and personality data, which can be applied in other collections around the world.
The Asiatic lion pride at London Zoo consists of three females and one male (Table
Members of the Asiatic lion pride at London Zoo.
Name | Age | Sex | Relationship |
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Rubi | 7 | F | Full siblings |
Heidi | 5 | F | Full siblings |
Indi | 5 | F | Full siblings |
Bhanu | 6 | M | Unrelated |
Data collection took place from May 31 to July 19, 2016. Focal animal behavioral observations using continuous sampling were completed to record the state and event behaviors at one minute intervals for each animal [
Each 60-minute observation period was divided such that 15 minutes were spent observing each animal. An observation session ended if the focal animal spent five consecutive minutes out of the observer’s sight (e.g., indoor). Total observation time summed between observation periods was approximately 87 hours. Included in each observation period were recordings of weather (i.e., sunny, cloudy, or rainy), temperature (
The behaviors recorded followed a standardized felid ethogram compiled by Stanton et al. [
Behavioral classes used to create time budgets. Individual behaviors come from the full ethogram, included in Table
Class | Behaviors included |
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Inactive | Lie, sit, stand, stretch, stare |
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Locomotion | Walk, run, stalk, chase, climb, crouch |
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Stereotypic | Pace |
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Reproductive | Mount, sniff anogenital region, lordosis |
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Maintenance | Defecate, urinate, self-groom, scratch |
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Marking | Spray, scratch object |
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Vocalizations | Growl, grunt, roar, cough |
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Feeding | Eat, drink |
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Exploratory | Any interaction with objects, sniff, flehmen, dig |
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Interactions | Allogroom conspecific, bite conspecific, play with conspecific, chase conspecific, stalk conspecific, swat conspecific, head/body rub conspecific, tail up, band on glass |
The London Zoo enclosure was divided into 27 zones to distinguish areas that may be used for different purposes. Twenty-one zones were located in the females’ section of the enclosure and six in the male’s section. The Whipsnade Zoo enclosure consisted of eight zones. These zones were assigned so that an animal’s specific location could be recorded during each observation, which was used to determine each lion’s enclosure use for each observation period. Maps of the London and Whipsnade Zoo enclosures and zone descriptions are available in Figures
The spread of participation index (SPI) was calculated to determine evenness of enclosure use. SPI was developed as described by Plowman [
London Zoo time budgets and enclosure use were compared to Whipsnade Zoo data, which was collected using the same methodologies in 2015. The data were also compared between Sunset Safaris and control nights. A sociogram was constructed showing the strength of relationships between individuals using time spent in proximity of another lion (i.e., at body-length or nearer). This was completed by calculating Association Index (AI) values for each relationship, as used by Schaller [
Personality profiles were compiled using questionnaires completed by seven London and Whipsnade Zoo keepers in 2015. The methodology for these questionnaires was adapted from Chadwick’s research on cheetah personality [
Behaviors recorded during observations were coded similar to time budgets such that classes could be compared to some of the traits on the personality questionnaire (Table
Personality trait classes consisting of behaviors from full ethogram, included in Table
Class | Behaviors included |
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Active | When an animal is exhibiting any observable behavior other than staring |
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Aggressive to conspecific | Bite conspecific, swat conspecific |
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Curious | Play with object, pounce on object, stalk object, swat object, bite object, dig, sniff, flehmen |
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Eccentric | Pacing |
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Friendly to conspecific | Allogroom conspecific, head/body rub conspecific, play with conspecific, tail up |
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Playful | Chase conspecific, play with conspecific/object, stalk conspecific/object, pounce on conspecific/object |
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Solitary | Time spent alone (i.e., greater than one body length away from conspecific) |
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Vocal | Growl, grunt, roar, cough |
Data analysis was completed using Microsoft Excel 2013 and IBM Statistical Package for the Social Sciences. Due to a small sample size, most tests for statistical significance were deemed inappropriate and therefore analysis focuses on descriptive statistics. Interrater reliability was calculated for the personality questionnaires using intraclass correlation ICC
Bhanu spent little time in his outdoor enclosure during the study because he was still adapting to the enclosure, which totaled to only a few minutes of observation data. Therefore, he was not included in data analysis.
The females’ time budgets were calculated for each observation period. These were also combined to create overall time budgets for each observation period and in total for all observations. The charts, including data values, are displayed in Figures
Rubi’s time budget for each observation period and overall at London Zoo. Data values are included to show exact percentages of time for each behavior class.
Heidi’s time budget for each observation period and overall at London Zoo. Data values are included to show exact percentages of time for each behavior class.
Indi’s time budget for each observation period and overall at London Zoo. Data values are included to show exact percentages of time for each behavior class.
Overall time budgets for each observation period and a complete time budget for all observations at London Zoo. Data values are included to show exact percentages of time for each behavior class.
Time budgets for each female for Whipsnade Zoo in 2015. Data values are included to show exact percentages of time for each behavior class.
Similar to time budgets, the females’ enclosure use was calculated for each observation period and for all observations (Table
Enclosure use values at London Zoo for each female and overall for each observation period, and in total for all observations.
Daytime | Control nights | Sunset Safari |
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Rubi | Heidi | Indi |
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Rubi | Heidi | Indi |
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Rubi | Heidi | Indi |
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Zone 1 | 50.5% | 45.5% | 57.5% |
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0.0% | 1.3% | 2.6% |
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6.3% | 1.4% | 0.8% |
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Zone 2 | 9.2% | 9.8% | 23.4% |
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6.7% | 0.0% | 0.0% |
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6.6% | 0.0% | 0.0% |
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Zone 3 | 0.6% | 1.3% | 0.7% |
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0.0% | 0.4% | 0.0% |
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0.0% | 0.0% | 0.0% |
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Zone 4 | 0.2% | 6.7% | 0.4% |
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0.9% | 0.4% | 1.3% |
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1% | 0.0% | 0.0% |
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Zone 5 | 0.6% | 0.4% | 0.9% |
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0.0% | 0.0% | 0.0% |
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5.2% | 0.3% | 0.0% |
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Zone 6 | 1.7% | 1.6% | 0.0% |
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0.0% | 2.1% | 0.0% |
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1.7% | 0.7% | 0.0% |
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Zone 11 | 0.0% | 0.0% | 0.0% |
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0.0% | 0.0% | 0.0% |
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0.7% | 0.0% | 0.0% |
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Zone 12 | 4.1% | 3.8% | 2.9% |
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13.4% | 8.1% | 0.4% |
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27.3% | 0.0% | 0.0% |
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Zone 13 | 14.4% | 14.4% | 5.6% |
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57.6% | 51.3% | 66.8% |
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31.8% | 61.6% | 72% |
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Zone 14 | 2.2% | 0.2% | 0.0% |
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0.0% | 0.0% | 0.0% |
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0.3% | 0.0% | 0.0% |
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Zone 15 | 11.6% | 4.4% | 7.6% |
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0.0% | 0.0% | 0.0% |
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9.1% | 5.1% | 0.0% |
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Zone 16 | 0.0% | 1.8% | 0.0% |
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0.0% | 0.9% | 0.0% |
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0.3% | 5.1% | 0.0% |
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Zone 17 | 0.0% | 0.0% | 0.0% |
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0.0% | 0.0% | 0.0% |
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0.0% | 0.0% | 0.0% |
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Zone 18 | 0.7% | 2.2% | 0.2% |
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0.0% | 3.4% | 0.4% |
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0.3% | 3.4% | 0.8% |
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Zone 19 | 1.1% | 2.5% | 0.0% |
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0.0% | 1.7% | 2.6% |
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5.9% | 2.1% | 2.4% |
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Zone 20 | 0.4% | 0.4% | 0.0% |
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0.0% | 1.3% | 0.9% |
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0.3% | 2.7% | 3.1% |
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Zone 21 | 1.1% | 1.8% | 0.2% |
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0.0% | 3.8% | 2.2% |
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2.4% | 5.1% | 6.3% |
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Zone 22 | 0.2% | 0.7% | 0.0% |
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0.0% | 0.4% | 2.2% |
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0.3% | 5.1% | 0.8% |
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Zone 23 | 0.0% | 0.2% | 0.0% |
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0.0% | 0.0% | 0.4% |
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0.0% | 0.7% | 0.4% |
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Zone 24 | 1.1% | 0.2% | 0.0% |
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0.4% | 0.4% | 0.4% |
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0.0% | 1.4% | 0.0% |
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Zone 25 | 0.4% | 1.1% | 0.7% |
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21.0% | 24.4% | 19.8 |
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0.0% | 5.1% | 7.5% |
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General enclosure use for each observation period and for all observations.
Weekly enclosure use values for London Zoo daytime observations to demonstrate how the females’ enclosure use changed throughout the study. Weeks 4 and 5 are combined as there were fewer observation sessions in Week 5.
Week 1 | Week 2 | Week 3 | Weeks 4-5 | |
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Zone 1 | 82.2% | 52.5% | 30.2% | 54.1% |
Zone 2 | 13.5% | 19.2% | 15.9% | 7.9% |
Zone 3 | 1.2% | 0.9% | 1.1% | 0.4% |
Zone 4 | 0.4% | 1.1% | 2.2% | 5.3% |
Zone 5 | 0.0% | 0.4% | 0.2% | 1.4% |
Zone 6 | 0.0% | 0.2% | 3.4% | 0.4% |
Zone 11 | 0.0% | 0.0% | 0.0% | 0.0% |
Zone 12 | 0.0% | 0.0% | 0.2% | 12.1% |
Zone 13 | 1.5% | 21.6% | 10.5% | 7.7% |
Zone 14 | 0.0% | 0.0% | 0.4% | 2.2% |
Zone 15 | 1.2% | 0.2% | 27.7% | 0.0% |
Zone 16 | 0.0% | 0.0% | 0.0% | 2.0% |
Zone 17 | 0.0% | 0.0% | 0.0% | 0.0% |
Zone 18 | 0.0% | 1.5% | 1.6% | 0.6% |
Zone 19 | 0.0% | 0.7% | 2.7% | 1.2% |
Zone 20 | 0.0% | 0.0% | 0.7% | 0.2% |
Zone 21 | 0.0% | 0.9% | 0.9% | 1.8% |
Zone 22 | 0.0% | 0.0% | 0.9% | 0.2% |
Zone 23 | 0.0% | 0.0% | 0.0% | 0.6% |
Zone 24 | 0.0% | 0.2% | 0.7% | 0.6% |
Zone 25 | 0.0% | 0.7% | 0.7% | 1.2% |
General weekly enclosure use (daytime observations) to compare how the females’ enclosure use changed throughout the study.
Enclosure use values for each female and overall at Whipsnade Zoo in 2015.
Rubi | Heidi | Indi | Overall | |
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Zone 1 | 52.7% | 76.7% | 54.5% | 61.5% |
Zone 2 | 2.1% | 0.9% | 4.2% | 2.5% |
Zone 3 | 2.5% | 1.9% | 1.5% | 1.9% |
Zone 4 | 0.0% | 0.9% | 0.3% | 0.4% |
Zone 5 | 38.9% | 14.5% | 39.0% | 30.6% |
Zone 6 | 0.4% | 0.0% | 0.0% | 0.1% |
Zone 7 | 3.5% | 5.0% | 0.6% | 3.0% |
Zone 8 | 0.0% | 0.0% | 0.0% | 0.0% |
SPI values for each female for Whipsnade Zoo and London Zoo observations.
Whipsnade Zoo | Daytime | Control nights | Sunset Safari |
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Rubi | 0.69 | 0.78 | 0.87 | 0.70 |
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Heidi | 0.70 | 0.69 | 0.77 | 0.69 |
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Indi | 0.69 | 0.84 | 0.81 | 0.78 |
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Decibel levels were averaged for each observation period and are displayed in Table
Maximum, minimum, and average decibel levels for each observation period and overall.
Max | Min | Average level | |
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Daytime | 85.9 | 37.7 | 63.2 |
Control nights | 78.2 | 32.9 | 56.4 |
Sunset Safari | 86.2 | 48.2 | 62.9 |
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Although sociograms are generally used for larger groups of animals, one is provided here for both Whipsnade Zoo and London Zoo to allow for visualization of the AI values and the strength of the relationships between the lions (Figure
Sociograms displaying the Association Index values for the relationship between the lions at Whipsnade Zoo (a) and London Zoo (b).
Personality questionnaires were completed in 2015 by seven keepers who worked with the lions at Whipsnade Zoo or London Zoo (Table
Summary of keepers who completed lion personality questionnaires in 2015.
Keeper | Sex | Experience with these lions | Hours/week with the lions | Average range between ratings |
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1 | M | 6 months | 25+ | 2.5 |
2 | M | 6 years | 8 | 2.0 |
3 | F | 3.5 years | 3 | 1.8 |
4 | M | 5 years | 2 | 1.1 |
5 | M | 3 years | 7 | 2.2 |
6 | M | 6 years | 10 | 1.5 |
7 | M | 6 years | 8 | 1.9 |
Ethogram.
State behaviour | Description |
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Out of sight (OOS) | Beyond one’s range of vision |
Decubitus–dorsal (DD) | Lays down on the dorsum |
Decubitus–lateral (LD) | Lays down laterally |
Decubitus–lateral–legs raised (DLLR) | Lays down laterally, one back leg raised |
Decubitus–sternal (SD) | Lays down on the sternum |
Sternal–sphynx (SPH) | Lays down on the sternum, back legs parallel and orientated forward |
Sternal–lunula (LUN) | Lays down on the sternum, legs put to one side |
Ears forward (EF) | Ears oriented forward |
Ears backwards (EB) | Ears oriented backward |
Facing conspecific (FC) | Stares at another animal of the same species |
Facing observer (FO) | Stares at the observer |
Facing public (FP) | Stares at the public |
Proximity to conspecific–body length (BL) | Within one body length of other animal |
Proximity to conspecific–far (F) | More than one body length away from the other animal |
Proximity to conspecific–contact (C) | In body contact with conspecific |
Sitting (SIT) | Upright position, all four feet on ground, front legs straight, back legs folded |
Standing (STA) | Stands with all four legs extended, paws on the ground, immobile |
Event behaviour | Description |
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Allogroom (AG X) x is the animal | Licks the fur of a conspecific |
Allogroomed (AGD b X) | Has the fur licked by a conspecific |
Bare teeth (BAT X a) a for active | Animal opens its mouth and pulls the lips back, exposing its teeth |
Receiving bare teeth (BAT X p) p for passive | Is on the receiving end of bared teeth |
Bite (BT X) | Mouth closes on object or conspecific |
Bitten (BT b X) | Is bitten by conspecific |
Belly up (B UP) | Animal lies on its back with throat and belly exposed to the opponent |
Belly up defensive posture (B UP DP) | Animal lies on its back with bared teeth, all four paws up with claws unsheathed |
Chase (CH X) | Runs after conspecific or other being/object |
Chased (CHD b X) | Pursued by conspecific |
Climb up (CU) | Ascends an object or structure |
Climb down (CD) | Descends an object or structure |
Defensive open mouth (DOM X) | Mouth wide open in defensive stance |
Drink (DR) | Lapps up water and swallows |
Defecate (DF) | Relieves colon, releases faeces |
Eat (EAT) | Ingests food by chewing and swallowing |
Eat grass (EAG) | Ingest grass by chewing |
Stretching (STR) | Extend all body and forelegs forward and put the back and tail up |
Fight (F X) | Assaults conspecific |
Assaulted (ASS b X) | Is assaulted by conspecific |
Jump on (JM) | Attack suddenly and forcefully jump on the back of conspecific |
Paw (PW) | Strike with the paw someone else |
Flehmen (FH) | Sniffs, then lift head with open mouth, breath in, eyes almost closed and upper lip curled |
Head butt (HB X) | Briefly pushes/bumps its head against a conspecific’s head |
Head butted (HB b X) | Has is head briefly bumped by a conspecific’s head |
Scratch (SRT) | Damage and mark the surface of by scraping with nails |
Lick object (LO) | Protrudes tongue from the mouth and strokes object with it |
Lick lips (LL) | Protrudes tongue from the mount and lick lips |
Pace (PC) | Repetitive locomotion in a fixed pattern |
Head shake (HSH) | Repetitive move of the head with short and quick movements |
Circling (CIR) | Repetitive locomotion in a circle around |
Twitch (TW) | Moving with a sudden, quick and short movements as reaction to something/someone |
Move backwards (MB b X) | Moving backwards with ears backwards and head down as reaction to someone |
Play object (PLO) | Interacts with objects |
Play with conspecific (PL X a) | Initiates interaction with conspecific in a nonharmful manner (chasing, jumping, wrestling, etc.) and gets no response |
Play with conspecific and is reciprocated (PL X) | Initiates interaction with conspecific in a nonharmful manner (chasing, jumping, wrestling, etc.) and gets some response |
Played by conspecific (PL X p) | Passive receiver of conspecific play |
Roll (RO) | Lying on the ground, the animal rotates its body from side to side; during the roll, the back is rubbed against ground, the belly is exposed and all paws are in the air |
Rub–Body (RB) | Rubs body on conspecific or object |
Rub–Head (RH) | Rubs head on conspecific or object |
Rubbed (RBD) | Rubbed by a conspecific |
Self-groom (SG) | Licks own fur |
Sniff (SNF) | Smells by inhaling air through the nose |
Spray (SP) | Stands with tail raised vertically and releases a jet of urine backwards against a vertical surface or object. |
Stalk (STL) | Usually slow, forward locomotion with back and head slightly lowered and eyes focused on the stalked individual/object |
Stare (STR) | Looks fixedly to something/someone |
Tail up (TU) | Tail is held vertically, in a upright position |
Tail slash (TS) | Standing or moving with tail bent over body, slashing |
Tail tip (TT) | Prolonged, repeated movement of tip of the tail |
Tail twitch (T TW) | A rapid flick of the tail in either a side to side or up to down motion |
Urinate (U) | Releases urine, standing or squatting |
Vocalization | Produces sounds or calls with is mouth/throat |
Vocalization–chuff (CHF) | Cat expels jets of air through the nose creating a low-intensity, soft, pulsed sound, described as being similar to the snorting of a horse |
Vocalization–grunt/cough (GRT) | Short, throaty call, characterized by the deep contraction and expansion of the diaphragm |
Vocalization–growl (GRL) | A low-pitched, throaty, rumbling noise produced while the mouth is closed |
Vocalization–hiss (HS) | A drawn-out, low-intensity hissing sound produced by rapid expulsion of air from the cat’s mouth, usually during exhalation |
Vocalization–roar (RO) | Long, throaty, high intensity call |
Vocalization–syndetic call (SC) | Amiable call with the purpose of gather or appease conspecifics |
Walk (WK) | Forward locomotion at a slow gait |
Run (RU) | Forward locomotion at a quick gait |
Warning bite (W BT X) | Snap teeth in response to an unwelcomed closing individual |
Yawn (YN) | The mouth is opened widely, the head tips back, lips are pulled back so that the teeth are exposed |
Look around (LOA) | Turn one’s eyes toward something or in some direction in order to see |
Crouch (CR) | Bend close to the ground or stoop low for lay down |
Crouch for other lion (CR X) | Stoop low and lays down on the sternum with ears backwards, head down or open mouth for submit to someone |
Dive in (DIN) | Plunge into water and stay in the water |
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Mount (MT) | Moves on top of conspecific in the attempt to copulate |
Nape bite (N BT) | The male performs an inhibited nape bite, where he will place his mouth on or around the back of the female’s neck at the moment of, or just after, ejaculation, but is unlikely to actually bite down |
Being mounted (BM) | Is mounted by other lion |
Sniff anogenital (SNA) | Smells the anogenital region of conspecific |
Zone descriptions of London zoo.
Zone | Features | Approx. % of total section area |
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1 | Back right corner of enclosure; next to entrance to indoor area; borders raised walkway along back wall | 4.85 |
2 | Front right corner; contains chain-link fence next to public walkway | 4.85 |
3 | Borders raised walkway along back wall; next to entrance to indoor area | 3.88 |
4 | Surrounds wooden platform | 4.85 |
5 | Borders raised walkway along back wall; includes metal gate to male’s section of the enclosure | 3.88 |
6 | Front left corner of original area of enclosure; contains small covered area under rock wall | 3.88 |
7–10 | Located indoors | N/A |
11 | Lower level of wooden platform | 1.46 |
12 | Mid-level of wooden platform; often used to climb up to Zone 13 | 0.97 |
13 | Top level of the wooden platform; offers high viewpoint | 1.46 |
14 | Top of a concrete slab in front of the entrance to indoor area | 1.46 |
15 | Area underneath Zone 14 | 1.46 |
16 | Grass-covered platform in front of Zone 6; overlooks moat | 0.97 |
17 | Located under Zone 16 | 0.97 |
18 | Thin zone bordering edge of moat | 8.74 |
19 | Start of new area of enclosure; contains rocky ledge along back wall | 6.80 |
20 | Covers right side of the 360 area; right side looks over the moat | 4.85 |
21 | Contains section of trees and bushes | 17.48 |
22 | Covers left side of 360 area | 8.74 |
23 | Back left corner of new area of enclosure | 4.85 |
24 | Allows access to Zone 25 | 8.74 |
25 | Covered area containing heated platforms (“Hot rocks”); where training occurs | 4.85 |
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26 | Faces access area where staff often walk; where outdoor training occurs | 15.09 |
27 | Contains access door for indoor area | 9.43 |
28 | Also faces access area where staff walk; contains part of small hill in middle of enclosure | 20.75 |
29 | Contains old train car/boxes; borders raised walkway | 26.42 |
30 | Borders mongoose enclosure | 13.21 |
31 | Contains train car where feeding sometimes occurs; allows access to train station platform with large public viewing windows | 15.09 |
Zone descriptions of Whipsnade zoo.
Zone | Features | % of Total area |
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1 | Back right corner; away from walkways | 24.24 |
2 | Front right corner; contains sleeping platform; walkway along front edge | 22.73 |
3 | Back left corner; walkway bordering side edge | 24.24 |
4 | Front left corner; walkway along front and side edges; contains training platform | 22.73 |
5 | Sleeping platform | 1.52 |
6 | Training platform | 1.52 |
7 | Area underneath sleeping platform | 1.52 |
8 | Area underneath training platform | 1.52 |
Personality questionnaires for each female were highly reliable. For Rubi, the average measure intraclass correlation (ICC) was .761 with a 95% confidence interval from .577 to .886. For Heidi, the average measure ICC was .805 with a 95% confidence interval from .652 to .907. For Indi, the average measure ICC was .857 with a 95% confidence interval from .744 to .932. According to a published interpretation scale [
Personality profiles for each female as determined by the questionnaires and behavioral observations are portrayed separately and combined in Figures
Personality profiles for each female compiled from questionnaires completed by keepers. Aggr = Aggressive, Fam = Familiar, Unfam = Unfamiliar, and CS = Conspecific.
Personality profiles for each female compiled from observation data. Aggr = Aggressive, CS = Conspecific.
Personality profiles compiled from both keeper questionnaires and observation data for each female. Aggr = Aggressive, CS = Conspecific.
Enclosure map of London Zoo.
Enclosure map of Whipsnade Zoo.
Overall, the lionesses were inactive for the majority of the time (56.7%), with Rubi displaying the most inactivity (74.0%). The high percentage of inactivity is appropriate as wild lions can sleep up to 21 hours per day [
Observed stereotypic behavior decreased from Whipsnade Zoo to London Zoo. Combining the females’ daytime data, stereotypies comprised 24% of the Whipsnade time budget, while only 11% of the London time budget. In part, this may be related to the difference in enclosure size. Lyons et al. [
Most of the lions’ pacing at London Zoo occurred along a chain link fence on the edge of Zone 2. Lyons et al. [
Much of the observed pacing behavior occurred before or after their morning feed. The postfeed stereotypic behavior could be due to a short feeding period, which implies that appetite behaviors were not fully expressed [
Although lions are typically most active during morning and evening hours [
Time budgets varied little between control nights and Sunset Safaris. The lions were inactive for the majority of their time during Sunset Safaris, even with higher overall decibel levels during these events. Rubi and Heidi displayed more activity during Sunset Safari than control nights, but that may have been affected by multiple scatter feeds during one Sunset Safari and the addition of multiple new forms of enrichment to the enclosure before a separate Sunset Safari. This small difference in behavior displayed between observation periods is likely a positive indicator of adjustment to the evening social events. One instance of aggression toward the public occurred during the first Sunset Safari, in which Heidi banged on the glass in front of large group of visitors, at least one of whom was a young child. However, this was the only occurrence of aggression toward the public observed during Sunset Safaris.
During daytime observations, the lions spent most of their time in Zones 1-2, where they were mainly inactive. They likely designated these zones as the core area of their territory, in which they felt most secure, with the rest of the enclosure being used for other purposes (e.g., playing, exploration, and occasionally resting) [
The lions also spent a large portion of their time in Zones 12-13, which make up a raised platform in the original part of the enclosure. These zones gave them a higher viewpoint of the visitor areas and surrounding animal enclosures. Lyons et al. [
When considering the change in enclosure use over time, the lions used more of the enclosure as the study progressed. During Week 1, the lions were not observed in the new part of the enclosure, but were observed in that area with increasing frequency over time. This could partially be due to changed training and enrichment practices used in order to influence the lions’ use of that area more often. Nonetheless, these husbandry practices may increase the lions’ comfortability with that area and may lead to them using the new part of the enclosure more often on their own accord.
Similar to the time budget comparison, there was little difference between the lions’ enclosure use during control nights and Sunset Safaris. The greater variety of zone use seen during Sunset Safaris may be related to multiple scatter feeds and addition of new enrichment during separate evenings, which caused increased movement through the enclosure. Zone 25, an area containing heated platforms for the lions and offering great viewing experience for visitors, was used more during control nights than during Sunset Safaris. Visitor sound levels and behavior (e.g., banging on the windows next to the heated rocks) may have influenced the lions to not spend much time there during Sunset Safaris.
SPI values demonstrate that the lions used the enclosure unevenly, which is supported by the charts separating enclosure use into original and new parts of the enclosure. However, the change in enclosure use over time suggests the lions may continue to spend more time in the new part of the enclosure. Overall SPI values were the same for Whipsnade Zoo and London Zoo. This uneven enclosure use reinforces the previously described idea of felids having core areas of their territory and has been similarly described in prides of wild lions [
Both sociograms indicate that Heidi and Indi have a slightly stronger bond with Rubi than with each other, but differences in AI values are minimal. Schaller found that there was no consistent lioness leadership of an African lion pride [
The profiles created from keeper questionnaires do not differ much between the lions, which was not the expected result. However, the interrater reliability results show that the method is reliable, as has been found in other studies [
The personality profiles created from behavioral observations were affected by having three subjects and a small data set, which makes the difference in trait ratings appear as large deviations in personality between the three females. Rather, these profiles may best be viewed as the lion that exhibited the most, least, or mid-amount of a trait. However, personality profiles from observations are a reliable and objective method that would be even more useful with a larger, long-term data set [
Previous events in the lions’ lives may have greatly influenced the results of the personality questionnaires. In 2014, the females lost both of their parents within a few months. Shortly after this, they were transferred to Whipsnade Zoo. These experiences may have been traumatic for the lions and possibly have affected their behavior for an extended period of time. For example, the large amount of pacing behavior exhibited by the lions at Whipsnade may have been a response to these stressful events. As previously discussed, this pacing behavior may have carried over to the current study as a “scar” from these traumatic experiences [
Rubi had the highest average rating on keeper questionnaires for “Solitary,” but comparatively Heidi was the most solitary according to observation data. This may be connected to Heidi’s high ratings in “Curious” and “Playful” in that she often investigated or played with objects. For instance, after the addition of new enrichment to the enclosure, Heidi spent more time interacting with the items compared to her siblings. Considering Heidi’s time budget, she also exhibited more “Exploratory” behavior than her sisters. This increased time exploring and interacting with objects may indicate that she spent less time near her sisters, therefore increasing her rating for “Solitary.”
As expected considering the lions’ time budgets, Indi had the highest rating for “Eccentric” on her profile created from observation data, which is due to her exhibiting the most stereotypic behavior. Before moving to Whipsnade Zoo, Rubi was the first lionesses to begin pacing after the loss of their parents. Heidi and Indi soon joined Rubi in this behavior, which then continued at Whipsnade Zoo. The fact that they followed Rubi in her display of stereotypic behavior may be an example of social facilitation and would support the aforementioned idea of Rubi as the leading female of the pride. Conversely, during observations at London Zoo, Indi often initiated the pacing behavior, and sometimes Rubi and/or Heidi would join her. Evidently, pride sociality plays a role in their behavioral patterns and preferences, but that role may be dynamic depending on their circumstances.
The personality profiles create opportunities for more individualized management of the lions, as demonstrated by Marieke Cassia and David’s study on snow leopard personality [
These conclusions stem from observations and from the valuable perspectives of the keepers. In the past year, some of the keepers have spent more time with the lions and may be better able to distinguish personality differences between the females. Although personality is consistent overtime, the questionnaires may have been influenced by previous traumatic events and by keeper knowledge of the animals. Now that the lions have settled into Land of the Lions and the questionnaires have been expanded, it would be ideal to repeat the questionnaires.
This research provided valuable behavioral and personality profiles for the lionesses at London Zoo. There was little difference in behavioral data between Sunset Safaris and control nights, which may be an indicator of little negative impact on the lions because of increased human social interaction. The personality questionnaires were found to be a reliable method of assessing personality. The personality profiles created by keeper questionnaires showed little difference between the females, therefore making individual comparisons difficult. However, the profiles created from behavioral observations showed more of a distinction between the lions. Combined together, these profiles offer some opportunities for individualized management of the lions, including varied enrichment methods. This research provides useful information for these specific lions to support current and future management decisions, and an interesting case study on individual animal adjustment to new environments. A personality study of all captive Asiatic lions would enable a comparison of lion personality across a variety of captive management systems and further development of methodologies for felid personality research.
The authors declare that they have no conflicts of interest.
The Royal Veterinary College generously provided the funding for this study. The authors would like to thank the Asiatic lion keepers at London and Whipsnade Zoos and for their support with project logistics and for completing the personality questionnaires.