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Thesis Final
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Effects of enclosure type and social environment on
stereotypical behaviours and interactions in
African Elephants (Loxodonta africana)
Sarah Grayson
MSc Animal Behaviour
7,508 words
630063815
“I certify that all material in this dissertation which is not my own work has been
identified with appropriate acknowledgement and referencing and I also certify that
no material is included for which a degree has previously been conferred upon me.”
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Abstract
Stereotypical behaviours are common amongst elephants kept in
captivity and consequently social and species typical behaviours tend
to be limited in comparison to their wild conspecifics. Various studies
have attributed these behaviours to enclosure types amongst other
factors; however the vast majority of elephant studies have been
conducted within a zoo environment. The present study investigated
how social enrichment and nocturnal enclosure types affected the
behaviour of a herd of nine African elephants living in a free range
captive environment at Knysna Elephant Park, South Africa.
Observations took place for approximately nine weeks from May to
August 2014 after an enclosure change from individually penned to
free roaming over two and a half hectares. General behaviours were
recorded using scan sampling every five minutes during certain periods
in the morning and evening and social and stereotypical behaviours
were recorded using continuous sampling.
Results indicated a significant effect following the change of nocturnal
enclosure type and social environment on stereotypical behaviours
and interactions. There was a significant reduction in both stereotypies
and interactions when the larger shared enclosure type was used
demonstrating that these factors are important in determining
elephant welfare.
This study is one of the first to find evidence of a link between
nocturnal husbandry management and stereotypical behaviours in a
herd of free range captive elephants, the implications of which are
discussed.
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Acknowledgements
It would not have been possible to conduct and write this research project without the help
and support of the many kind people around me, to only some of whom it is possible to give
particular mention here. In particular I would like to thank:
My internal supervisor, Dr Natalie Hempel de Ibarra (University of Exeter), for her
support and guidance from beginning to end.
My external supervisor, Lisa Olivier (African Elephant Research Unit, South Africa)
whose experience and guidance shaped my project.
Clare Padfield, (African Elephant Research Unit, South Africa) whose advice, kindness
and support made everything possible.
Dr Debbie Young, who made me feel welcome and at home and whose enthusiasm
and compassion for elephants was contagious.
My research assistants (in alphabetical order): Darina Brejtrova, Jenn Claeys, Marina
Loch, Clare Padfield, Andrew Schuster, Jessica Smith, Nicky Webber, Kirsty Wilson
and Cloe Wood without their generous donations of time, effort and enthusiasm this
project would not have been the same.
The staff and guides at Knysna Elephant Park who made me feel welcome and
inspired me with their continued compassion and care for their elephants.
The volunteers working with AERU, in particular Darina Brejtrova whose enthusiasm
and dedication to my project was incredible and Karin Baumgartner who was a loyal
friend and supported me throughout.
My boyfriend Ben Ainsworth, who has taught me to believe in myself, without him
this project would never have been possible.
My parents, for their continued advice and support.
And finally, Sally, Nandi, Keisha, Shungu, Mashudu, Thato, Shaka, Clyde and Thandi
who deepened my love for elephants and gave me an experience I will never forget.
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Introduction
The African elephant (Loxodonta africana) is one of the most iconic and majestic animals on
earth, they are well known for their strong social groups, emotion and intelligence. The
welfare of captive elephants is something that remains one of the greatest challenges for
zoos and wildlife parks worldwide. Captive elephants can be used to support the
conservation and behavioural research (Hosey, 1997) that is vitally important for their long
term survival as a species and can further our understanding of this magnificent and
complex species. As the worlds’ largest land mammal, African elephants in captivity due
unfortunately to their sheer size, high level of intelligence and sophisticated social networks
are especially vulnerable to various welfare problems.
Elephant welfare refers to their state of psychological wellbeing and has been notoriously
difficult to measure (Mason & Mendl, 1993). Zoos and parks rely upon indirect physiological
and behavioural indicators as well as external variables such as environment to measure an
elephant’s mental state. Behavioural observations and research tend to focus on
behaviours that are not recognised as typically normal for the species, also known as
stereotypic behaviours or stereotypies. Stereotypies often develop in captive animals and
are defined as repetitive, unvarying behaviours with no obvious goal or function (Clubb &
Mason, 2002), such as the weaving, swaying and bar biting often seen in captive elephants.
Stereotypies are often used as indicators of poor welfare associated with boredom,
frustration, injury or cramped environments to name a few. Other indicators of reduced
welfare could be external or environmental variables, restrictions in movement, confined
into small enclosures and a lack of social opportunities (Veasey, 2006).
Nowadays there is often a lot of debate around captive animals housed in zoos with many
people considering it wrong to do so and zoos and their advocates disagreeing with this.
This is also known as the ‘zoo debate’. One of the most common arguments used in this
debate relates to the amount of space available to animals housed in captivity. Zoos often
argue that they provide as much space as is possible in the confines of their establishment
or that the space offered is enough with opponents arguing that enclosures are far too
small. Previous research has found links between enclosure size and stereotypical
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behaviours in a vast array of species (Paulk et al., 1977; Harris et al., 2008; Draper and
Bernstein, 1963) and so in order to maintain high standards of welfare in zoos, research into
optimum enclosure types should be used a tool to provide effective management solutions.
There have been many studies investigating the effects of enrichment on stereotypical
behaviours in captive elephants (Swaisgood & Shepherdson, 2005; Shyne, 2006; Thorne et
al., 2005) which have proven extremely useful, however these to some extent patch over an
underlying cause of these behaviours, such as suboptimal enclosure size. Harris et al. (2008)
investigated the effects of various factors on the welfare, housing and husbandry of
elephants in UK zoos and found over half displayed stereotypies during the day and those
with a small amount of indoor space displayed more over a 24 hour period than those with
a medium amount of indoor space. They found that larger amounts of outdoor space were
associated with reduced stereotypical activity and this space was clearly linked to improved
welfare. With more space allowance, there is an increased opportunity for enrichment and
exercise, thus reducing the importance for enrichment devices. Draper and Bernstein (1963)
investigated the effects of enclosure size on rhesus monkeys and found that those housed in
small cages exhibited more stereotypical behaviours than when they were housed in larger
cages. Paulk et al. (1977) found similar evidence in both wild born and captive bred rhesus
monkeys, highlighting that prolonged housing in smaller enclosures leads to stereotypical
behaviours. Wiedenmayer (1995) found that when they un-tethered a stable group of
captive elephants in Zurich zoo and allowed them to move freely within their enclosure
stereotypical behaviours reduced dramatically and in some cases was no longer observed.
These studies highlight the importance of enclosure size and restriction on captive animal
welfare.
In addition to enclosure size, elephants are often kept in countries with suboptimal
temperatures resulting in indoor heated housing. Rees (2004) found the frequency of
stereotypies increased with colder temperatures in elephants prone to stereotypical
behaviours. According to the American Zoo and Aquarium association (AZA) (2011) indoor
temperatures should be heated to a minimum of 55o
F (13o
C) with one room holding a
temperature of at least 70o
F (21o
C) and outdoor temperatures not dropping below 40o
F
(5o
C) if the elephants are to be outdoors overnight.
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Social conditions are often ignored but should however be considered important in captive
elephant welfare. Elephants are social animals and often participate in and perform certain
activities such as feeding, resting, foraging, walking and bathing as a herd. As elephants are
active for approximately 20 hours a day (Moss, 1982) it can be assumed this is the time
when most social interactions occur within the herd. Elephants will also generally herd
together during periods of sleep and use a variety of sensory modalities to communicate
with one other including sound, olfaction, vision and touch (Berg, 1983; Payne, 2003). The
AZA Standards for Elephants Management and Care (2011) suggests that each zoo housing
elephants should have a minimum of three females, two males or three of mixed gender to
try and maintain some form of social group. They recognise that elephants are a highly
social species and state that elephants should have the opportunity for olfactory, tactile,
visual and auditory interaction with other elephants. Despite this there are zoos that
continue to keep single elephants; this will almost certainly contribute to impoverished
welfare. Many zoos house elephants individually overnight for approximately 12 hours,
restricting normal social behaviour and interactions during this time. A survey by the AZA
(2011) found 20% of the elephant’s studied did not have access to the rest of the herd
overnight and 28% were restrained using leg chains (Bashaw et al., 1999). Brockett et al.
(1999) found individuals within a group of three female African elephants spent half of the
night within one body length of each other, supporting the idea that group housing may be
more beneficial and that individual housing may be a welfare concern. Wilson et al. (2006)
conducted a follow up study from Brockett et al. (1999) and found the highest duration of
social behaviour occurred during the hours of 23:00 and 05:00; concluding that unrestricted
social access at night is an effective management strategy.
The manner in which elephants are managed in captivity has a substantial effect on their
welfare and their lives. In captive settings elephants should be allowed to exhibit natural
behaviours as often as possible together with as little restriction on social interactions and
movement as possible. It therefore seems prudent to promote the provision of large, high
quality enclosures that encourage locomotion and allows the elephant’s to have full use for
as much time as they wish. Taking into consideration the implications of enclosure type and
social access on the welfare of African elephants an effective management solution would
be to combine an enclosure of a substantial size that allows for both indoor and outdoor
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access with unrestrained social environment during the night. This should not only reduce
the amount of health and psychological related problems, such as stereotypies, arising from
restricted movement but would also allow for normal social interactions which are a proven
indicator of welfare (Veasey, 2006).
This study explored whether nocturnal enclosure type and social environment in a captive
herd of African elephants are related to levels of stereotypical and social behaviours. Until
December 2013 the herd were housed individually and unchained in an indoor enclosure
overnight. More recently a shared, outdoor enclosure was created for use during the night,
with constant access to the indoor enclosure and an area of approximately 2.5 hectares to
roam thus allowing the herd the option of displaying the full range of social interactions. A
previous study when the elephants were individually penned was conducted in 2013 by
Padfield (2013), this recorded stereotypical behaviours and general social interactions. The
present study builds upon that study. It was hypothesised that the change to the enlarged
enclosure and the less restricted social environment could lead to a reduction in
stereotypical behaviour and an increase in social interactions. Comparing the data recorded
in the new enclosure to previously collected data when the same elephants were kept in
smaller individual enclosures with restricted social access allows examination of whether
these factors have implications for elephant welfare.
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Method
Subjects
The study animals were a herd of nine African elephants (Loxodonta africana) aged between
six and twenty-six years old, housed at Knysna Elephant Park (KEP) between Knysna and
Plettenberg Bay in South Africa. The elephants are all habituated to humans and are part of
on-going studies by the African Elephant Research Unit (AERU), who are also based at KEP.
The elephants are from mixed backgrounds with two being biologically related and together
they form two herds; a matriarchal herd consisting of five females (one juvenile) and two
juvenile males, and the bachelor herd consisting of two bulls. During the day from 07:30
until 16:30, the matriarchal herd are walked out of the 2.5 hectare camp and allowed to
roam over 100 hectares with handlers present. Tourist’s visit the park throughout the day
and at a metal barrier that the elephants are trained to stand behind tourists can touch,
have photographs taken with and hand feed fruit to the elephants under the guidance of
the elephant handlers. The bachelor herd spend the day in the 2.5 hectare camp that the
matriarch herd use during the night and are not involved in any interactions with the public.
Housing
From 17:30 to 07:30 the matriarchal herd (with the exception of one, see below) are housed
in the 2.5 hectare camp overnight with access to their old indoor enclosure for shelter (13m
x 13m) (Appendix 3). The elephants are not chained and have full tactile, olfactory, auditory
and visual access to each other. The bachelor herd are penned individually, as they are
solitary animals in the wild, from 17:30 to 07:30 as well as Thandi, a mature female elephant
from the matriarchal herd as she was having difficulties mixing with the juvenile males and
was re-penned in April 2014. From 17:30 both herds are fed a mixture of branches and
straw, which are placed in small piles in the outdoor camp and in the individual pens. The
lights in the indoor enclosure are switched off at 21:20 and back on at 06:20. During the
winter months two infra-red heat lamps are switched on inside the indoor enclosure from
17:30 and are switched off at 07:00. The floor of the indoor enclosure is covered with a thick
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layering of sawdust. The outdoor camp is mainly grass and dirt with a small dam for the
elephants to drink and bathe in. Until late 2013 the herd were housed separately and
unchained in individual 4m x 7m pens in the indoor enclosure from 17:15 to 07:15
(Appendix 4). These enclosures allowed each elephant full visual access to at least two other
members of the herd. In late 2013 the outdoor enclosure was created for use during the
night and on 31st
December 2013 all individual pens except three had been knocked down
and the matriarchal herd were allowed access to both the indoor enclosure and outdoor
camp.
Figure 1: Current indoor enclosure with three individual pens at the back
Figure 2: Old indoor enclosure with nine individual pens
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Figure 3: Outdoor camp (approx. 2.5 hectares)
Study design and procedure
Before live data collection was started preliminary observations were carried out in order to
refine the ethogram (Appendix 1), familiarise with general activity patterns and identify
individuals. A pilot study was conducted one week prior to live data collection to ensure the
methodology worked and to identify any potential problems. The preliminary period and
pilot study found no social interactions or stereotypical behaviours were displayed in the
outdoor camp and so observations were only carried out on the indoor enclosure from the
indoor observational lounge.
Figure 4: Indoor Observational Lounge
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The research was purely observational and was conducted at specified times during the
morning and evening. It consisted of recording general behaviours, stereotypical behaviours
and social interactions (affiliative and agonistic) using an ethogram and a behaviour
recording sheet (Appendix 2). The ethogram allowed examination and comparisons to
behaviours observed in the previous study. Following a preliminary period and pilot study,
data was collected over an eight week period between 2nd
June and 27th
July 2014, during
the southern winter, approximately the same dates as were recorded in 2013. Data
collection was carried out approximately three to four times per week (Including weekends),
starting at 17:30 and concluding at 21:00 and 05:30 until 07:00 the following morning to
ensure any behaviours that occurred as a result of the evening was captured the following
morning. The hours differed slightly from the previous study, due to a change in elephant
management times and the elephants sleeping patterns. The pilot study and a previous
study (Johnson, 2013) using this specific elephant population found after feeding and
during the later times at night stereotypical behaviours were absent therefore this made it
unnecessary to collect data for the whole night.
A repeated measures design was used with both instantaneous and continuous sampling.
Instantaneous sampling every five minutes allowed for general and social behaviour
observations of each elephant within the herd during the night (Martin & Bateson, 1995).
Continuous sampling allowed for more detailed collection of stereotypical and social
behaviours that occurred during the five minute scans. Stereotypical behaviours were
defined using an ethogram adapted from Harris et al. (2008) and Padfield (2013) (Appendix
1). Social behaviours were defined using an ethogram adapted from Wilson et al. (2006) and
included both affiliative and agonistic behaviours. Initiators and recipients of all social
behaviours and stereotypical behaviours were recorded using all-occurrence sampling
(Altmann, 1974). Distance was recorded as a measure of how much time the elephants
spent together proximate to a focal elephant that changed every five minutes, in contact
with the focal elephant or distant from the focal elephant. Contact was defined as when one
elephant was in physical contact with another. Proximate was defined as when an elephant
was within the indoor enclosure within proximity to the focal elephant. Distant was defined
as when an elephant was outside when the focal elephant was inside or vice versa.
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‘Unknown’ was recorded when both the focal elephant and another elephant was outside at
the same time as their distance from one another could not be seen.
Once observations started, a stop watch was used to prompt a time period of five minutes
and the behaviour of each elephant was recorded at the end of the five minute period on
the data sheet as well as their distance from a particular focal elephant. Continuous
observations took place during the 5 minute scans and any stereotypical or social behaviour
was recorded on the data sheet noting the initiators and recipients. There was a separate
comments column on the data sheet, which was used for stating any significant behaviour
not already listed on the ethogram to ensure all key behaviours were recorded. All nine
elephants were observed in the same order on each data collection signal to ensure data
consistency. Initiators and recipients of all social behaviour types were recorded as well as
the distance from one focal elephant each five minutes. The researcher observed from the
same view point each night and temperatures and weather data was collected on each night
as extraneous variables (see below).
Observers
Following a three day pilot study it was found that observing continuously for a 3.5 hour
period could result in unreliability due to tiredness and so for this reason volunteers were
recruited to help with 1.5 hours of the total observation period. The evening observations
took place from 17:30 to 21:00, where the primary researcher conducted 17:30 to 19:30
and an assistant observed and collected data from 19:30 to 21:00. This was varied randomly
throughout the eight weeks in order to ensure accuracy. No assistants were needed for
morning observations. A total of nine assistants were recruited and used from a pool of
willing volunteers and AERU staff members. Assistants were trained and carried out practice
sessions before completing inter-observer reliability tests, where the researcher and
assistant would observe and record consecutively with no conferring and the data was
compared for accuracy. Tests were varied over different hours of data collection to ensure
reliability and consistency. Before collecting live data, assistants were asked if they felt
confident or needed more time and inter-observer reliability had to be reached at 80% or
more. Index of concordance scoring was used to calculate the reliability percentage by
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looking at the total frequencies of behaviours in one session. Concordance across all
assistants and all tests, varied between 83% and 98%, with a mean of 93% overall. There
were no failed tests.
Extraneous Variables
Previous research by Rees (2004) found the frequency of stereotypies increased with colder
temperatures in those elephants prone to stereotypical behaviours and so weather was
recorded as a possible extraneous variable. A reliable weather site, World Weather Online
(2014), was used to obtain temperatures at 07:00 and 20:00 during data collection times.
During the pilot study rain was found to drive the elephants in to the indoor enclosure and
so for this reason the presence or absence of rain was recorded during data collection as
another possible extraneous variable.
Ethical Consideration
Ethical clearance was granted by the University of Exeter’s ethical review group on 4th
December 2013 (Appendix 5). As the research was purely observational there was no
physical or psychological harm suffered by the elephants.
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Results
Stereotypical Behaviours
Last year, in 2013 when the elephants were housed in individual pens, stereotypical
behaviours were displayed by Sally, Keisha, Shungu, Clyde and Thandi. Sally had the most
instances of stereotypical behaviours and Shungu the least. This year stereotypical
behaviours were displayed by only Sally, Keisha and Clyde. Stereotypical behaviours were no
longer present in Shungu and Thandi. Stereotypical behaviours in both Sally and Clyde have
dropped substantially from 2013, with Sally having now 20 instances compared to 140 in
2013 and Clyde only 2 compared to 56 in 2013. Stereotypical behaviours in Keisha however
have risen from 5 instances in 2013 to 36 instances in 2014. Table 1 shows the breakdown
of the instances of stereotypical behaviours in 2013 and 2014.
Table 1: Total frequency of stereotypical behaviours in 2013 and 2014
Sally Nandi Keisha Shungu Mashudu Thato Shaka Clyde Thandi
Stereotypical
2013
140 0 5 4 0 0 0 56 24
Stereotypical
2014
20 0 36 0 0 0 0 2 0
Stereotypical behaviour was compared using Generalised Estimating Equations (GEE) (IBM
SPSS Statistics v.20). As the dependent variable (total frequency of stereotypical behaviour)
displayed a heavy positive skew towards zero, a Wald distribution was used to allow for a
valid interpretation of these results. Furthermore, the data were offset by 1.1 and then
transformed using a base-10 logarithm in order to change the zero values into positive
values which would then be accepted by the GEE model.
In a model where the enclosure type was the independent variable with covariates rain and
time spent inside as an interaction there was a significant effect of enclosure type on
stereotypical behaviours (Wald χ2
(1)=4.05, p=0.044), with stereotypical behaviours reducing
significantly from 2013 to 2014 (β= -0.17). This supports the hypothesis that the larger,
shared enclosure would significantly reduce stereotypical behaviours compared to when the
elephants were individually penned. The interaction between rain and the time the
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elephants spent inside was important due to the fact that the rain caused the elephants to
move to the inside enclosure from outside; this approached significance (Wald χ2
(2)=5.16,
p=0.076). The main effects of rain (Wald χ2
(1)=1.35, p=0.245), time spent inside (Wald χ2
(1)=0.07, p=0.782) and high (Wald χ2
(1)=0.01, p=0.918) and low (Wald χ2
(1)=1.16, p=0.282)
temperatures were removed from the model as they were none significant.
Social Interactions
Scan samples every 5 minutes were recorded both in 2013 and 2014 for social interactions
(Table 2). Interactions were recorded when two or more elephants were engaged in some
form of social contact on the 5 minute mark. Although some elephants displayed more
interactions in 2014 than 2013, the total number of interactions was higher in 2013 than
2014. Interactions were higher for Sally and Nandi in 2014, remained the same for Keisha,
however were higher for the rest of the herd in 2013. Shungu and Thandi displayed the least
amount of interactions in 2014 whereas Nandi displayed the most. In 2013, Shungu
displayed the most and Nandi displayed the least. Table 2 shows a breakdown of the
average number of interactions on a typical night by each elephant over the two
observation periods.
Table 2: Average number of interactions per night. The group of elephants was scanned every 5
minutes to record all behaviours. Data were collected in 2013 and 2014.
Sally Nandi Keisha Shungu Mashudu Thato Shaka Clyde Thandi
Interactions
2013 1.71 0.06 0.24 4.29 1.71 3.12 1.06 2.82 3.00
Interactions
2014 1.80 2.04 0.24 0.16 0.28 0.28 0.52 0.36 0.16
Standard
Deviation 0.07 1.40 0.00 2.92 1.01 2.01 0.38 1.74 2.01
The normalised frequencies were analysed in IBM SPSS Statistics v.20 using a Generalised
Estimating Equations (GEE) model. As the interactions (dependant variable) displayed a
heavy positive skew towards zero, a Wald distribution again was used and data transformed
as described above.
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In a model where the enclosure type was the independent variable with the covariate rain
there was a highly significant effect of enclosure type on interactions (Wald χ2
(1)=6.41,
p=0.011), with social interactions reducing significantly from 2013 to 2014 (β= -0.24). This
does not support the hypothesis that the larger, shared enclosure would significantly
increase the number of interactions compared to when the elephants were individually
penned. Rain had a significant effect on interactions (Wald χ2
(1)=21.04, p=>.001) with more
interactions occurring when there was no rain present. The main effects of time spent inside
(Wald χ2
(1)=0.05, p=0.822) and high (Wald χ2
(1)=0.68, p=0.409) and low (Wald χ2
(1)=0.95,
p=0.331) temperatures were removed from the model as they were none significant as was
any interaction between rain and time spent inside (Wald χ2
(1)=0.193, p=0.660). The
significant effect of rain was unexpected and so post-hoc analysis was carried out in order to
investigate this further. A GEE was conducted on the continuous interaction data from 2014
only, in order to eliminate any effects from the elephants being 100% indoors in 2013 and
due to the fact this data was more concise. A significant effect of rain on interactions was
found (Wald χ2
(1)=3.99, p=0.046) with interactions decreasing when rain was present. More
interestingly there was a significant reduction in agonistic behaviours when there was rain
(Wald χ2
(1)=5.81, p=0.016), there was no significant effect of rain on affiliative behaviours
(Wald χ2
(1)=3.52, p=0.061).
Interactions over 25 days
Over the 25 days, the elephants were observed in continuous periods for a total of 125
hours in which continuous interactions were recorded (Table 3). Thato, the youngest of the
group was the recipient of most interactions and Thandi received the least. Sally, the 24-
year old matriarch initiated most interactions and again Thandi initiated the least, most
likely due to the fact Thandi is penned and so is limited to the amount of contact she can
access with the others.
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Table 3: Total number of social interactions during the whole observation period
Sally Nandi Keisha Shungu Mashudu Thato Shaka Clyde Thandi
Interactions
Received
64 103 82 74 87 120 45 45 32
Interactions
Initiated
181 140 101 30 49 65 59 47 19
When broken down into affiliative and agonistic behaviours (Table 4), it was found that Sally
received no agonistic behaviours from any of the other elephants over the 25 days studied;
she did however initiate the most. Thato was the recipient of most agonistic behaviours,
receiving 27 instances, followed closely by Keisha who received 26. Shaka, Clyde and Thandi
did not receive any agonistic behaviour, most likely as a result of being penned.
Table 4: Total amount of agonistic behaviours observed over 25 days
Sally Nandi Keisha Shungu Mashudu Thato Shaka Clyde Thandi
Agonistic
Received
0 16 26 16 21 27 0 0 0
Agonistic
Initiated
76 23 7 1 2 0 1 0 0
In terms of affiliative behaviours (Table 5), Nandi, initiated the most and Thato again was
the recipient of the most. Thandi received and initiated the least amount of affiliative
behaviours. Out of those elephants that were unpenned Shungu initiated the least and
Keisha received the least.
Table 5: Total Affiliative Behaviours observed continuously
Sally Nandi Keisha Shungu Mashudu Thato Shaka Clyde Thandi
Affiliative
Received
64 87 56 58 66 93 45 45 32
Affiliative
Initiated
105 117 94 29 47 65 58 47 19
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Affiliative
84%
Agonistic
16%
Overall, there was a much higher percentage of affiliative behaviours displayed (84%)
compared to agonistic behaviours (16%) which demonstrates a positive effect on behaviour
of the enclosure change (Figure 5).
Figure 5: Distribution of agonistic and affiliative behaviours in all elephants.
Location
Over the twenty-five days of observation the location of each elephant was determined
against a focal elephant every five minutes in order to examine how the new enclosure
space was being utilised and how often the elephants spent together as a herd, regardless
of the amount of space.
Figure 6 shows the percentage of time the elephants spent in their inside enclosure during
the hours observed. Shungu and Mashudu, the two juvenile males, spent the least amount
of time inside (average 55%) whereas the four females spent an average of 73% of their
time inside. All elephants slept in the indoor enclosure during the night of their own choice.
The females were always the first to enter the indoor enclosure from the outside camp
whereas Shungu and Mashudu were the last ones to come in. Figure 7 shows the average
amount of time the herd spent either inside or outside in the camp as a whole.
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0%
10%
20%
30%
40%
50%
60%
70%
80%
Sally Nandi Keisha Shungu Mashudu Thato
70%
75%
71%
51%
59%
74%
30%
25%
29%
49%
41%
26%
Inside Outside
Inside
67%
Outside
33%
Figure 6: Percentage of time each elephant spent inside the boma and outside in the camp
Figure 7: Average amount of time the herd spent inside and outside in the camp
In terms of proximity to one another the results show that as a herd the elephants spent
most of their time within close proximity to one another (45%), 30% of their time was spent
distant and 25% of the time their distance was unknown due to the focal elephant and the
elephant being observed being outside and out of sight. Very few direct contacts, where the
elephants were physically touching another were observed on the five minute mark.
Figure 8 shows the percentage of time the elephants as a herd spent either in contact,
proximate or distant from the focal elephant on each 5 minute scan.
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Discussion
This study illustrated the ways in which the nocturnal husbandry management of captive
free range elephants can affect welfare standards, in particular the effects that enclosure
type and social environment can have on stereotypical behaviours and social interactions.
Stereotypical behaviours were reduced in the larger enclosure allowing more social contacts
A significant effect of enclosure type on stereotypical behaviours was found, which supports
the experimental hypothesis that a larger, shared enclosure would reduce the amount of
stereotypical behaviours displayed by the elephants compared to when they were penned
individually. Five out of the nine elephants displayed stereotypies when individually penned
in 2013 and it was surprising to find that two of those five had no stereotypical behaviour at
all in the new enclosure. What is more surprising is that Clyde and Thandi who displayed
stereotypies in 2013, remained penned this year, however Thandi displayed no stereotypies
this year and Clyde had reduced from 56 instances in 2013 to only 2 this year. This shows
that the enclosure change not only affected those elephants that were given more space
but also those that remained penned. This may be due to the fact that they are still able to
interact more with the rest of the herd and have more social enrichment than they did last
year. Sally, the matriarch, who displayed the most stereotypical behaviours last year with
140 instances, reduced significantly this year to only 20 instances and stereotypies in
Shungu had gone completely. Keisha on the other hand was the only elephant to increase in
her instances of stereotypies from last year, increasing from 5 to 36. Although Keisha only
displayed stereotypical behaviours (mainly weaving) over 3 nights out of the 25, her
behaviours tended to last for longer periods with short breaks between each behaviour.
One plausible explanation of Keisha’s stereotypical behaviours, currently under
investigation with the African Elephant Research Unit (AERU), is that they are likely to be
related to her oestrus cycle and the possible discomfort caused by this (C Padfield 2014,
pers. comm., 10 September). Data from last year has highlighted a similar pattern in
Keisha’s stereotypies and previous research into stereotypical behaviours has revealed that
animals are likely to display more stereotypical behaviours as they get older (Philbin, 1998),
which would provide an explanation for the increase in stereotypies.
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The fact that there was a significant effect of enclosure type on stereotypical behaviours
raises important questions for elephant welfare and management practices not only in free
range captive elephants as in this instance but in captive environments also. The findings
from this research could have powerful and positive implications for elephants in captivity
all over the world. The reduction in stereotypical behaviours between the two years
provides compelling evidence that a larger, unrestricted enclosure type is an effective
nocturnal management strategy for these elephants and supports previous research that
larger enclosures are associated with lower stereotypical behaviours (Harris et al, 2008). It
would not be possible for zoos to provide space for captive elephants equivalent to natural
home ranges; however as more space is clearly linked to improved welfare then as much
space should be provided as possible. Caution should be exercised however when
concluding that the relationship between enclosure size and stereotypic behaviour is a
causal one. It could be that when there was less space there were other aspects of that
environment that were suboptimal that influenced the display of stereotypic behaviours.
For example the increased opportunities for social enrichment this year may have led to the
reduction of stereotypies. Additionally, although stereotypical behaviours have reduced
significantly and for some elephants gone completely, for some elephants stereotypies are
still present indicating that some areas of management may not be ideal. Stereotypical
behaviours have been found to occur around the time of scheduled events (Bloomsmith et
al., 2001; Rushen, 1985) and so changing the timing of these events such as when they are
fed and when the handlers arrive may help to further eradicate stereotypical behaviours in
these elephants. Further research on this particular population of elephants over a longer
period of time could highlight possible influencing factors such as the time of year or the
time spent in the new enclosure. Although these may be unlikely influencing factors they
should still warrant further investigation.
Social Interactions decreased in the larger, shared enclosure
Social interactions observed every five minutes significantly decreased from 2013 to 2014,
which does not support the initial hypothesis that the larger, shared enclosure would
significantly increase social interactions. The reasoning for this may be mainly due to the
fact that the elephants now have much more to do and much more space to explore than
24. 630063815
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they did last year. In the 2.5 hectare enclosure the elephants are able to roam, forage, eat
and display more species typical behaviours over a much larger area. Location data (Figure
8) shows that although the elephants spent 45% of their time proximate to one another,
30% of their time was spent distant and in the outdoor enclosure, thus a decrease in social
interactions was displayed. When the elephants were individually penned indoors, there
was a limited amount of behaviours they could display. Once their evening food had been
eaten there was only an area of approximately 28ft2
to explore and may have been more
likely to interact with their neighbours. Their new outdoor enclosure provides
approximately 2.5 hectares of grass and mud with a damn and approximately 169ft2
of
indoor space, thus encouraging species typical behaviours such as foraging and feeding.
The amount of interactions displayed by each elephant has interestingly varied between the
two years. Sally, the matriarch and Nandi, the second most dominant female were the only
ones to have increased in their number of interactions from 2013. This may be to be
expected from the most dominant of the herd and the fact they spent a lot of time together
interacting. Keisha’s amount of scan interactions seemed to have stayed roughly the same
as last year, whereas the rest of the herd, including those still individually penned, have
decreased in their number of interactions. Last year Shungu had the highest number of
interactions, however this year had the least along with Thandi, perhaps due to the fact he
spent the most time outside in the outdoor enclosure (Figure 6) and Thandi lacked the
opportunity to interact with other members of the herd at will as she remains penned.
Interactions are a large part of natural behaviour amongst elephants in the wild and
although this study showed that a larger, shared enclosure decreased the number of
interactions in comparison to when the elephants were penned, it does however
demonstrate the need for elephants to display more species typical behaviours such as
foraging and feeding over a larger enclosure space and that the new enclosure allowed for
elephants to choose who they interacted with. A further study examining the elephant’s
behaviours both in the indoor enclosure and in the outdoor camp would prove highly useful.
Looking specifically at species typical behaviours in both parts of the enclosure would shed
more light on how the elephants utilise their time whilst housed nocturnally.
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Social interactions observed continuously over 25 days showed high rates of affiliative
behaviours
In order to assess the effectiveness of the new enclosure type descriptive statistics were
used to analyse the number and type of continuous social interactions occurring within the
herd. Results showed that Sally, the matriarch, initiated the majority of the interactions. This
is typical of a matriarch in the wild who would use physical interactions to maintain a herd.
Touch has been found to be an extremely important interaction in many species (Franz,
1999; Schino, 2001; Nakamura, 2003) and is considered salient in elephant communication
(Makecha et al, 2012) in particular when engaging in playful interactions. Mothers usually
engage in a high rate of contact behaviours with their calves (Berg, 1983; Lee, 1986) and
higher ranking members use touch to discipline lower ranking members (Freeman et al,
2004; Langbauer, 2000) which may be a possible explanation as to why Thato, the youngest
female, was the recipient of most interactions and Sally, the matriarch, initiated most.
Social behaviours were collapsed into two categories for discussion: agonistic and affiliative
(see appendix 1 for details). The continuously observed social interactions consisted largely
of affiliative behaviours (84%), with agonistic behaviours accounting for only 16% (See
Figure 5). In addition to each elephant initiating significantly different amounts of each
behaviour, differences were seen in the amount of agonistic and affiliative behaviours each
elephant initiated and received.
Agonistic behaviours are a normal part of an elephant’s behavioural repertoire and
dominance status and hierarchy plays a salient role in the display of these behaviours.
Higher-ranking animals initiate more agonistic behaviours than do lower-ranking animals,
with these interactions serving the purpose of disciplining other members, especially those
younger in the group (Freeman et al, 2004). It is therefore not surprising that Sally, the
matriarch, did not receive any agonistic behaviours from any other elephant, she was
however the initiator of most and Thato and Keisha the recipients of most. In addition lower
ranking members are less likely to initiate agonistic behaviours towards the matriarch
(Makecha et al, 2012) which is consistent with the findings from this study as the three
juvenile elephants initiated the least amount of agonistic behaviours. Thato did not initiate
any, with Shungu and Mashudu, the two juvenile males only initiating one and two. Clyde
26. 630063815
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and Thandi also did not initiate any agonistic behaviours and Shaka only one, with none of
them receiving any, mainly due the fact these three elephants remained individually penned
and so these types of interactions would have been difficult to display from behind the
enclosure bars.
The amount of affiliative behaviours displayed within the herd was substantially higher than
the amount of agonistic behaviours. It was surprising to find that Nandi, the second oldest
female, was the initiator of most affiliative interactions and Thandi, her daughter, initiated
and received the least. This is inconsistent with previous research that suggests young
elephants initiate the majority of affiliative interactions (Gutham, 2014). Thato again was
the recipient of most affiliative interactions within the herd.
Post hoc tests revealed an interesting finding; that there was a significant interaction
between rain and the amount of social interactions that occurred. To my knowledge this is
the first finding of a relationship between elephant’s social interactions and rain. There was
a significant decrease in interactions when it was raining compared to when it was dry,
more interestingly there was a significant decrease in agonistic behaviours when it was
raining. The reasons for this remain unknown; however we know from the pilot study that
the elephants went in to the indoor enclosure when it was raining. If we were to speculate
then the decrease in interactions may be due to the noise on the metal roof of the indoor
enclosure as a distraction or possibly, similar to some humans, when it was raining the
herd’s mood may have fallen and made them less likely to interact. This however may not
fully explain as to why there was a significant decrease in agonistic behaviours but not
affiliative behaviours. Why would elephants be less agonistic when it was raining? In order
to answer this interesting question further investigations are needed into the relationships
between social interactions and weather, in particular the effects of rain.
Perhaps the most important finding from the continuous interaction data is that the amount
of affiliative behaviours greatly outweighs the amount of agonistic behaviours. Should the
results have been reversed then this may have suggested that the new enclosure type was
not beneficial in terms of the welfare of the elephants and that individual pens may have
been more appropriate. The amount of affiliative behaviours displayed provides persuasive
evidence that unrestricted social access is an appropriate nocturnal management strategy
27. 630063815
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for this particular herd of elephants. Agonistic behaviours were infrequent and none of
those observed resulted in injury as a consequence. These findings are consistent with
previous research that suggests serious fighting amongst wild matriarchal herds is rare
(Moss, 1982) and that the majority of social behaviours within a herd are primarily affiliative
(Moss & Poole, 1983).
The elephants spent most of their time together as a herd in the indoor enclosure
Location data was recorded in order to assess the utilisation of the new enclosure type and
to examine how much of the space was being used overnight. On average the elephants as a
herd (not including the three individually penned) spent more time (67%) in the indoor
enclosure than the outdoor camp (33%) (See Figure 7). Although more time was spent inside
this may have been due to various factors. One possible reason may be that Nandi’s
daughter, Thandi, was penned. Thandi was often heard vocalising to which Nandi responded
by coming inside and as such the other female elephants would often follow. Additionally,
the time of year may have been an influencing factor. As the study was conducted during
the winter months it would be interesting to see if these values changed over the summer
when the weather is warmer and dryer, as an interaction was found between rain and the
amount of time the elephants spent indoors. The indoor enclosure also had infra-red heat
lamps which may have encouraged them to come in to get warm on the colder evenings. It
was observed that all elephants came in to the indoor enclosure to sleep at night, which is
consistent with previous research that suggests elephants always sleep together as a herd
(Makecha et al, 2012). In general the elephants spent most of their time proximate to each
other (45%) which demonstrates that given the freedom of choice to move around the 2.5
hectares they chose the majority of time to spend together which supports the new
enclosure type in terms of sociality. Should the elephants have spent most of their time
distant from one another then this may have been suggestive of the fact the elephants
found the unrestricted access to each other compromising to their needs. The four females
spent an average of 73% of their time inside during the hours observed compared to the
males who only spent an average of 55% of their time inside. This is suggestive of normal
male elephant behaviour in the wild where as juvenile males get older they tend to spent
less time within the herd and more time on the outskirts, becoming more independent in
preparation for their separation from the herd as bulls (Keet, 2013). Overall both the indoor
28. 630063815
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enclosure and outdoor camp were utilised by the elephants during the observation periods
and gave the elephants substantial opportunities to either spend time together as a herd or
time on their own foraging and feeding. This space reduces the need for manmade
enrichment devices as the elephants are able to source their own enrichment, be that social
or the ability to express species typical behaviours.
Conclusions
As African elephants are the earth’s largest land mammal it remains a difficult task trying to
maintain an optimum environment for these highly social creatures in captivity. The current
study aimed to demonstrate the impact and importance that enclosure types can have on
the behaviour and welfare of captive elephants and to my knowledge is the first of its kind
to show a clear link between stereotypical behaviours and enclosure type in the same
population of African elephants over a period of time. Although the population used in this
study may have benefited from the abundance of land they had access to during the day,
the way they are housed nocturnally can be compared to those housed in zoos worldwide.
The overlap between the results from this study and those from wild populations suggests
that the study of elephant social interactions in captivity can provide insights into elephant
behaviour in the wild and vice versa. Perhaps more importantly, comparing social
interactions in captive environments with those from the wild may provide an important
analytical tool when assessing and evaluating the welfare of elephants in captive settings. If
the social behaviour of elephants is substantially different from those of their wild
conspecifics this is suggestive of a suboptimal environment. In addition to this, examining
the ways in which stereotypical behaviours vary between enclosure types can have a huge
impact on the ways in which elephants are housed in captivity worldwide. If we are to
accept that stereotypical behaviours are a proven indicator of welfare (Clubb & Mason,
2002) then the current study has shown that a larger enclosure with unlimited social access
is an effective nocturnal management solution and can almost certainly improve elephant
welfare by decreasing stereotypical behaviours. Caution, however must be exercised when
interpreting the results from this particular study, not only due to the small sample size but
the fact this is a study of only one population of elephants, which are free ranging during
the day, something most captive elephants do not experience.
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This study has demonstrated that the way in which elephants are housed nocturnally can
have a substantial effect on the type and number of interactions that occur within a herd as
well as significantly reducing stereotypical behaviours. Additionally it highlights the effect of
enclosure type on the opportunity to perform species typical behaviours. It is important for
elephants to be given the opportunity to interact with other elephants and the new
enclosure type studied gives these elephants that opportunity. The importance of continued
research in this field is imperative as elephants are particularly vulnerable to impoverished
welfare as a result of inadequate environmental conditions. Additional studies into the
effects of enclosure types on social interactions and stereotypies of elephants in captivity
would contribute greatly to elephant welfare around the globe. Additionally examining the
effects of enclosure type not only on psychological welfare but also on the physical welfare
of elephants would contribute greatly to overall elephant welfare in captivity. An increase in
nocturnal enclosure size may also lead to decreased obesity, arthritis and foot problems as a
result of increased activity and exercise. Creating more opportunities for elephants to
exhibit species typical behaviours in larger enclosures and the introduction of greater
opportunities for social interaction, may reduce the frequency of stereotypical behaviours
and increase the overall welfare of captive elephants.
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Appendices
Appendix 1 – Ethogram
Behaviour Code Description
Feeding Behaviours
FB Feeding Branches
FOR Foraging: Exploring the environment with trunk,
searching for food
FS Feeding Straw
FO Feeding Oats
FP Feeding Pellets
FF Feeding Fruit
FG Feeding Grazing
SF Stealing Food: Elephant takes food either from
another individual or from an adjacent enclosure
Locomotion Behaviours
S Standing: Individual is stationary in an upright
position whilst awake
W Walking: Using two or more steps to change location
in environment. Not in a stereotypic pattern, playing,
feeding, or exhibiting any other behaviour
simultaneously.
SL Sleeping lying down: Asleep
SS Sleep standing: Standing but sleeping with eyes
closed, body parts usually resting against individual
pen
Stereotypical Behaviour
SW Sway: Elephant sways from side to side without
moving feet for five seconds or more
WE Weave: More pronounced version of sway using feet
for five seconds or more
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Hbob Head bob: Moving the head up and down for five
seconds or more
PA Pace: Elephant walks repetitively in the same route
around the enclosure for five seconds or more
RO Rocking: Elephant moves backwards and forth in the
same position five seconds or more
BB Bar Biting: Biting the bars of the enclosure for five
seconds or more
TR Tusk Rub: Moving tusks against a part of the
enclosure for five seconds or more.
Vocalisation
R Rumble
T Trumpet
VO Vocalise: High pitched
Social
Affiliative
TC Trunk Contact: Trunk to other elephant anywhere on
body
BC Body Contact: Body to body contact (not trunk) inc
leaning etc
Ap Approach: Approach other with social intent
Agonistic
P Push: Push other with head or shoulders resulting in
1 step by recipient
Ch Charge: Charge or chase another
Dis Displace: One approaches with intent and recipient
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leaves
Other
I Interaction: Interacts with other members of the herd
using any body part
OD Other Dusting: Rolling in dirt/sawdust or using the
trunk to pick up and throw substrate on any part of
the body.
OOS Out of Sight: No recorded can be given due to an
obstructed view of the elephant
NIS Not In Sight: Elephants that are part of the study are
not in view of the observer but behaviours are being
recorded elsewhere
O Other: A Behaviour not in the aforementioned,
possibly due to a rare event. Write description of
behaviour in comments box
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= all other free elephants proximity to this elephant (Contact, Proximate, Distant)
Common
Behaviours:
Any other behaviours not listed
Time Sally Nandi Keisha Shungu Mashudu Thato Shaka Clyde Thandi Time Name(s) Behaviours Other
Weather / Temp:
SW = Sway / BB = Bar bite / R = Rumble / T = Trumpet /
TC = Trunk Contact / BC = Body Contact / A = Approach / P = Push / C = Charge / Dis = Diplacement
Social & Stereotypical Behaviours
(contact, trunk touch, body touch, sway, weave, bar bite etc.)
Behaviour Recording Sheet - Loxodonta africana
End Time:Start Time:
Behaviours and Proximity
(C = contact, P = 0 - 13 meters, D = > 13 meters (13m = length of boma))
OBSERVATIONS
Date: Location (In /Out):
Observer Name:
Appendix 2 – Behaviour recording sheet
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Appendix 5 - Ethical Approval
Ethics Approval System <D.M.Salway@exeter.ac.uk>
Thu 09/01/2014 11:48
Ethical Approval system
Your application (2014/488) entitled An investigation into the effects of enclosure size and herd
access on species typical behaviour in African Elephants (Loxodonta Africana) has been accepted
Please visit http://www.exeter.ac.uk/staff/ethicalapproval/
Please click on the link above and select the relevant application from the list
