Undergraduate Dissertation

NOTTINGHAM TRENT UNIVERSITY
OBSERVATIONS OF TERRITORIAL BEHAVIOUR IN A CAPTIVE POPULATION
OF MADAGASCAR FODIES (Foudia madagascariensis)
by
ALEXANDER M. WILLEY
A dissertation submitted to Nottingham Trent University in partial
fulfilment of the requirements for the degree of Bachelor of Science with
Honours in Zoo Biology
School of Animal, Rural & Environmental Sciences
Nottingham Road
Southwell
Nottinghamshire
NG25 0QF
April 2013

i
ABSTRACT
The Durrell Wildlife Park is one of a number of institutions to maintain a collection of
Madagascar fodies (Foudia madagascariensis). Though not threatened with extinction,
this species serves to raise awareness of conservation efforts for the endangered and
closely related Mauritius fody (Foudia rubra). As such, Madagascar fodies were
maintained in the Kirindy Forest Aviary; an immersion exhibit, based on Madagascan
forest, that housed a range of indigenous bird species in a mixed, free-flying
environment. The Madagascar fody is a naturally territorial species during certain
times of the year, however the extent to which the species’ territorial behaviours
persisted in a shared, captive environment was poorly researched. As such, zoo
managers became concerned that an excessive density of mature males during this
period would result in aggressive intraspecific competition. This study was requested
in order to determine whether territorial behaviour would restrict the number of
males that could safely share the limited space.
All five mature males in the aviary were observed over a fifty-day period, during
which time subjects’ behaviours and locations were recorded. These were analysed to
determine whether subjects were operating within a territory, and whether
behaviours were affected by the onset of territoriality. The study showed that
subjects behaved territorially, as indicated by the loss of two subjects through
intraspecific conflict and by the localised movements of remaining subjects within the
aviary. Territories formed were of significantly different size between individuals
when compared using one-way ANOVA (F(2,9) = 64.645, p<0.0005). In contrast, the
size of observed territories was not found to significantly impact aggressive (p=0.679)
or reproductive (p=0.399) behaviours when compared using Spearman’s rank-order
correlation.

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These results imply that the conditions of the Kirindy Forest Aviary, though restricted,
do not prevent territorial behaviour. As such, the presence of more than three
individuals may result in harmful intraspecific aggression and should be avoided.
Further study is needed to support these findings, and to assess the applicability of
these findings to other captive populations.

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ACKNOWLEDGEMENTS
With thanks to Eluned Price and the Durrell Wildlife Park Bird Department, for
providing this research project and for their invaluable support and advice throughout.
Thanks also to Dr Samantha Bremner-Harrison for her help and support, and for
keeping me on the right track.
Thanks finally to family for their never-ending support, intellectual, financial and
emotional, during this project and beyond. It will never be forgotten.

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TABLE OF CONTENTS
ABSTRACT.....................................................................................................................................i
ACKNOWLEDGEMENTS................................................................................................................ iii
TABLE OF CONTENTS................................................................................................................... iv
1.0 INTRODUCTION ...................................................................................................................... 1
1.1 Aims................................................................................................................................... 2
1.2 Hypothesis.......................................................................................................................... 2
1.2.1 Null Hypothesis............................................................................................................. 3
2.0 LITERATURE REVIEW............................................................................................................... 4
2.1 The Issues of Behaviourin Ex Situ Housing............................................................................ 4
2.1.1 Physical Environment................................................................................................... 5
2.1.2 Social Structure ............................................................................................................ 7
2.2 The Effect of Housing on Conservation................................................................................. 9
2.3 Background on the Study Species........................................................................................11
2.3.1 Madagascar Fody.........................................................................................................11
2.3.2 Mauritius Fody ............................................................................................................12
2.4 Previous Studies into Fody Behaviour..................................................................................13
2.5 Territoriality.......................................................................................................................17
2.5.1 Determinants of Territory Size......................................................................................18
2.5.2 Methods of Calculating Territory Size............................................................................20
2.6 Research Rationale.............................................................................................................21
3.0 MATERIALS AND METHODS ....................................................................................................22
3.1 Subjects.............................................................................................................................22
3.2 Housing.............................................................................................................................22
3.3 Management Strategies .....................................................................................................24
3.4 Preliminary Investigation....................................................................................................26
3.5 Enclosure Mapping.............................................................................................................26
3.6 Ethogram...........................................................................................................................28
3.7 Investigation Procedure......................................................................................................31
3.8 Variables ...........................................................................................................................32
3.9 Ethical Considerations........................................................................................................32
3.10 Risk Assessment...............................................................................................................33
3.11 Data Analysis ...................................................................................................................33
3.11.1 Statistical Analysis......................................................................................................34

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4.0 RESULTS................................................................................................................................35
4.1 Territories..........................................................................................................................35
4.1.1 The Existence and Distribution of Territories .................................................................35
4.2 Enclosure Level Preference.................................................................................................43
4.3 Occurrence of Aggressive Behaviour ...................................................................................45
4.4 Occurrence of Courtship Behaviour.....................................................................................48
5.0 DISCUSSION...........................................................................................................................52
5.1 The Existence of Territory...................................................................................................52
5.2 Territory Size .....................................................................................................................53
5.3 Enclosure Level Preference.................................................................................................55
5.4 Aggressive Behaviour.........................................................................................................56
5.5 Courtship Behaviour...........................................................................................................57
5.6 Additional Limitations and Future Study..............................................................................58
6.0 CONCLUSION AND APPLICATION OF FINDINGS........................................................................60
7.0 REFERENCES..........................................................................................................................61
8.0 APPENDICES..........................................................................................................................74
8.1 Appendix 1 – Ethical Review Form.......................................................................................74
8.2 Appendix 2 – Risk Assessment.............................................................................................77

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1.0 INTRODUCTION
The preferred layout of zoo enclosures has seen drastic change in recent decades. As
public perception of the appropriate treatment of captive animals has shifted, zoo
housing has evolved from the barren, concrete housing of the ‘first -generation’ to the
naturalistic replications of natural environments of the ‘third generation’ that are
seen commonly today (Shettel-Neuber, 1988). This has been accompanied by a
greater focus on animal welfare (Hosey, Melfi and Pankhurst, 2009), including the
provision of an appropriate social structure within the constraints of the captive
environment.
One such enclosure is the Kirindy Forest Aviary, built and maintained by the Durrell
Wildlife Conservation Trust (DWCT) at the Durrell Wildlife Park, Jersey. The Kirindy
Forest Aviary is designed to simulate a Madagascan forest environment, to increase
awareness of the DWCT’s conservation efforts in Madagascar (DWCT, 2009). As such,
the aviary houses a number of native bird species, including the Madagascar Fody
(Foudia madagascariensis). The Madagascar Fody is a territorial species during the
breeding season (Garrett et al., 2007), therefore to ensure the safety of the captive
population, and to further knowledge of appropriate management techniques, the
organisation has requested that the territorial behaviours of this particular captive
population be examined.
The primary objective of this study is to investigate the impact of natural territorial
behaviours on the mature males in the captive population of Madagascar fodies at
the Durrell Wildlife Park, Jersey, UK, through behavioural observations. The sample
population was observed routinely over a fifty-day period, during which time the
movements and behaviours of each subject were recorded.

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1.1 Aims
The primary aim of this study is to determine which areas of a shared enclosure are
utilised by resident mature male Madagascar fodies as territory, and to determine
whether the sizes and locations of these territories vary between individuals. Also in
relation to spatial utilisation, this study aims to determine whether variations exists
in the enclosure level (height of the enclosure) used most commonly by subjects.
Additional aims relate to determining how the size of subjects’ territories impact on
the frequency of expression of certain behaviours. These include:
 Measuring frequency of aggressive interactions with conspecifics, to determine
if a relationship exists between aggression and territory size
 Measuring frequency of courtship behaviours to determine if a relationship
exists between reproductive behaviour and territory size
1.2 Hypothesis
The existence of territories will be determined through observations of the
movements of each individual around the enclosure. The current hypotheses are that,
while territories will be formed, not all individuals in the study will maintain territories
due to spatial constraints, and that subjects will maintain territories of different size
to one-another. Subjects are also expected to show a preference towards a particular
enclosure level.
Additionally, increased territory size is expected to relate to more frequent
aggressive behaviour, and increased territory size is expected to result in more
frequent courtship behaviour.

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1.2.1 Null Hypothesis
The null hypothesIs for this investigation is that no subjects will be observed to form
or defend a territory. Additional null hypotheses are that no subjects will demonstrate
a preference towards a single enclosure level, that no significant relationship will
exist between the amount of space utilised by subjects and the occurrence of
aggressive behaviour, and that no significant relationship will exist between the
amount of space utilised by subjects and the occurrence of courtship / reproductive
behaviour.

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2.0 LITERATURE REVIEW
2.1 The Issues of Behaviour in Ex Situ Housing
The differences between the conditions experienced by animals in the in situ
environment and the conditions experienced in an ex situ environment are well-
recognised. In the specific instance of zoos, Hosey (2005) proposed that the factors
most dramatically distinguishing the ex situ from the in situ environment were the
limitations to available space, the frequent presence of humans and the management
of the animal’s lifestyle.
Considering these differences, the subject of how the alterations to environment
affect the behaviour of captive animals has been frequently researched. As the World
Association of Zoos and Aquaria (WAZA) operates with a core principle of allowing
captive animals to exhibit natural behaviours (WAZA, 2003), this research is often
conducted with a view to determine how to best facilitate natural behaviours in spite
of the restrictions of captivity, generally through modifications to housing and
management strategies. This policy therefore gives the indication that the housing
for animals in WAZA institutions is intended to have no modifying effect on behaviour,
and that resident animals are intended to perform their natural behaviour repertoires
(NBRs) uninterrupted.
The impact of this policy on animal welfare has been a subject of debate in the zoo
community over recent years. Researchers such as Mason and Burn (2011) have
argued that, as life in the in situ environment is filled with challenges and threats,
motivation of the behaviours developed to cope with such challenges is unnecessary
ex situ to the point of being a detrimental to welfare. An example of this was
documented by Wielebnowski et al. (2002a), who found evidence of increased stress
in North American clouded leopards (Neofelis nebulosa) when an anti-predator
response was motivated by potential predator species housed nearby. In the United

5
Kingdom, the Secretary of State’s Standards of Modern Zoo Practice (SSSMZP)
attempt to avoid this by instructing that a captive animal’s housing be designed in
such a way as to avoid stress (Department for Environment, Food and Rural Affairs
(DEFRA), 2012). Based on the evidence discussed above, this would appear to be in
conflict with the goal of housing animals to allow natural behaviours.
Overall, the way animals are housed could be considered to be intended to encourage
only behaviours that are representative of good welfare. Some specific elements of
housing that are manipulated to achieve this will be discussed below.
2.1.1 Physical Environment
In the development of animal enclosures, modern zoos show a preference towards
the use of naturalistic materials and the simulation of the natural environment of the
resident species. Enclosures are designed this way in large pa rt to facilitate the
resident animal’s natural behaviours, as discussed earlier. As an example, enclosures
for orang-utans (Pongo) at Chester Zoo, UK, and Tiergarten Schönbrunn, Austria,
contain large climbing frames in a naturalistic setting to encourage natural arboreal
behaviour (ZooLex, 2009; 2012).
The physical environment can, however, have a pronounced effect on animal
behaviour in cases where the environment is insufficiently complex, or wherein
resident animals cannot perform motivated behaviours. The frustration caused by
these conditions is known to cause abnormal stereotypic behaviour; patterns of
behaviour that are repetitive, invariant and which serve no obvious function (Ödberg,
1978; Mason, 1991). These behaviours vary widely between species and causative
factors are not always clearly understood. Large carnivores such as lions ( Panthera
leo), for example, are known to pace repetitively within their enclosures (Bashaw et
al., 2003); a behaviour thought by some researchers to be stimulated by frustration

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with the limited space in the captive environment (Clubb and Mason, 2007; Rodden
et al., 2012) or with the inability to escape from external stressors (Wielebnowski et
al., 2002a; Collaham et al., 2012). The appearance of such behaviours, therefore, is
generally considered to be an indication of suffering to some degree (Mason and
Latham, 2004). A further study by Latham and Mason (2010) gave evidence that
these behavioural abnormalities manifest more severely in animals (in this case mice
(Mus musculus)) that experience a change from enriched housing to more barren
housing, suggesting the effects of enriched, complex housing conditions on behaviour
are both dramatic and long-term.
To make housing conditions less likely to influence abnormal behaviours, many zoos
fill enclosures with various enrichment devices. The effect of these devices on animal
behaviour varies; a study by Gottlieb et al. (2011) found a reduction in abnormal
behaviours in rhesus macaques (Macaca mulata) presented with puzzle balls
containing food, requiring object manipulation and encouraging natural foraging
behaviour. However, the same study by Gottlieb et al. (2011) found that ‘shakers’
(containers shaken to gradually release food) had the opposite effect on the
macaques, in that stereotypic behaviours increased. This indicated that the device
caused the animals stress, either through frustration at the device’s operation or
through apprehension of the novel object. As the devices were presented to different
subject groups in this study, subjects’ individual preferences could also have
potentially influenced their response to the presented device.
In instances where enrichment devices have achieved the desired effect, there is
evidence that the behavioural impact of the devic e on the animal decreases over time
(Lutz and Novak, 2005). This suggests that the novelty of enrichment devices is a
major determinant of the device’s effect on animal behaviour, and that the effect of
such devices on behaviour may only be short-term.

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2.1.2 Social Structure
Many animal species that are held in captivity are gregarious, and would naturally
form social groups of various size. In situ, the number of individuals in a social
grouping would be determined based on the value of potential benefits, such as co-
operative anti-predation strategies and co-operative foraging / hunting (Krauss and
Ruxton, 2002) versus the value of associated costs, such as intraspecific competition
(Price and Stoinski, 2007) in relation to the species’ ecological niche. The importance
of replicating these social conditions in captive environments is recognised, but is
restricted by environmental constraints and limited knowledge of a species’ full range
of in situ social behaviour (Swaisgood and Schulte, 2010). Additionally, the captive
environment imposes a different selection of pressures; while food competition and
risk of predation are removed, spatial restriction prevents members of captive
populations from regulating their social structure to alleviate social tension or breed
(Price and Stoinski, 2007).
When these restrictions lead to inappropriate social groupings, the result may be the
onset of abnormal behaviour, as with instances of inappropriate physical
environments. Early research into the effects of social isolation on a normally
gregarious species was conducted by Harlow and Harlow (1962), using rhesus
macaques. These studies showed that macaques reared without maternal influence
or the presence of conspecifics developed severe behavioural abnormalities, including
self-injurious behaviour (SIB) and aggression, which continued into later life. More
recent studies have found similar behavioural abnormalities in chimpanzees (Pan
troglodytes) reared in social deprivation (Bradshaw et al., 2008; Lopresti-Goodman,
Kameka and Dube, 2013). This relates to a theory postulated by Mason and Latham
(2004) that early deprivation of animals (social or otherwise) resu lts in a
psychological ‘scar’ that persists into later life; a theory supported by evidence that

8
early deprivation negatively impacts neurological development (Kraemer and Clarke,
1990; Lewis et al., 2006).
Alternatively, there is evidence that housing animals that would be solitary in situ in
unnatural social conditions may also have a negative impact on behaviour. In the
case of cheetahs (Acinonyx jubatus), for example, females are naturally solitary.
Consequently, a study by Wielebnowski et al. (2002b) identified an increase in
stereotypies and aggression in female cheetahs housed together in captivity.
Similarly, Morgan and Tromborg (2007) described the tense behaviour exhibited by a
captive group of Japanese macaques (Macaca fuscata) comprised entirely of males,
as opposed to the natural social structure of multiple individuals of both sexes
(though these behaviours were not observed as part of a controlled study). This
implies that captive populations should be structured as similarly as possible to in
situ populations to maximise welfare and natural behaviour. This structure may not
be uniform for all individuals of a species; male cheetahs, for example, differ from
females in that they benefit from being housed socially (Caro, 1993). Groups should
also be structured so as to avoid overcrowding in the limited space, as this has also
been linked to heightened aggression between conspecifics (Blanc and Thériez, 1998;
Boyce et al., 1998).
In contrast; orang-utans are frequently maintained in conspecific social groups ex
situ despite living solitarily in the wild (Price and Stoinski, 2007). Rather than
negatively influencing welfare, this style of housing has induced enriching social
behaviours and uncharacteristic paternal care of offspring (Bond and Watts, 1997;
Munn and Fernandez, 1997). As a result, Price and Stoinski (2007) theorise that the
captive environment may accommodate a more flexible of social groupings if
housing is planned effectively, and with an awareness of the species’ needs.

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2.2 The Effect of Housing on Conservation
The WAZA code of ethics and animal welfare dictates that all member institutions
contribute towards the goal of conserving species (WAZA, 2003). Many institutions
work towards this goal by implementing captive breeding programs for their species,
with the intention of increasing the number of individuals in the captive population.
The way in which an animal is housed affects the success of these programs in a
number of ways. Carlstead and Shepherdson (1994) noted that the increase in
reproductive success in zoo populations has corresponded with the increased
consideration of basic requirements for reproduction as zoos have developed; the
importance of conditions such as climate, available substrates, nesting facilities and
social groupings is evidenced by the fact that many species, such as gorillas ( Gorilla
gorilla) and flamingos (Phoenicopterus), yielded no successful captive births before
1951 (Carlstead and Shepherdson, 1994), after which an awareness of such
necessities became more common. This trend suggests that barren housing, or
housing which deprives animals of the requirements to trigger reproductive
behaviour, may inhibit conservation efforts by preventing an increase in the captive
population.
Reproduction may also be inhibited by the issues associated with irregular social
housing discussed earlier. A well-documented example of this is the hand-rearing of
the kakapo (Strigops habroptilus), particularly the hand-rearing of chicks as a means
of protection from predators and threatening environmental conditions (Elliott,
Merton and Jansen, 2001). Elliott, Merton and Jansen (2001) observed that strong
habituation to humans in hand-reared individuals was unavoidable and that, at the
time, none had successfully reproduced. A later study by Eason and Moorhouse
(2006) also confirmed that, at that point, some males considered humans to be
correct sexual partners, and that none had reproduced(though natural kakapo
breeding hierarchy may have contributed to this) . This implies that conservation

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may be inhibited if the way an animal is housed prevents correct reproductive
behaviour.
Released or reintroduced animals may also be affected by the physical environment
of pre-release housing. As the IUCN regulations for the reintroduction of species
dictate that reintroduction subjects from captive stock be given the opportunity to
develop the necessary skills to survive in the wild (IUCN, 1998), many individuals are
placed in environments that allow development in such areas as physical fitness,
locomotion and predator evasion (Reading, Miller and Shepherdson, 2012). For
example, Biggins et al. (1999) documented the housing of black-footed ferrets
(Mustela nigripes) in semi-natural pens for the development of natural behaviours.
These individuals consequently spent less time above-ground upon release,
improving predator avoidance. In contrast, early release programs for golden lion
tamarins (Leontropethicus rosalia) resulted in the loss of released individuals due to
poor locomotion between branches; the result of the provision of static climbing
structures in pre-release housing (Kleiman et al., 1986). Overall, the pre-release
housing of captive animals appears to contribute positively or negatively towards
conservation based on how completely the housing prepares the animal for in situ
challenges.

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2.3 Background on the Study Species
Fodies are small weaver birds and members of the Ploceidae family, genus Foudia.
This genus contains six recognised species, all of which are endemic to various
regions of the western Indian Ocean (Garrett et al., 2007). This study focuses
specifically on the Madagascar fody, which will be discussed in more detail below.
2.3.1 Madagascar Fody
The Madagascar fody, also known as the red fody, is endemic to Madagascar, though
populations have been introduced to multiple regions, including the Mascarene
Islands, St Helena and Bahrain (International Union for the Conservation of Nature
(IUCN), 2009). In situ this species is generally found in regions of second growth
vegetation but avoids evergreen forest areas (Brickell, 2006), and are often seen
occupying areas of human habitation in order to acquire food (Haydock, 1954; Penny,
1992). This implies a long association with humans.
Due to the species’ large geographical range and apparently stable population trend,
the Madagascar fody is listed as ‘Least Concern’ for conservation by the IUCN (IUCN,
2009). The population has not, however, been accurately quantified, therefore this
classification is based on assumption. In spite of this classification, captive
populations of Madagascar fodies are held in multiple zoological institutions across
Europe (including the Durrell Wildlife Park and Chester Zoo, UK), and in North
America (Louisville Zoological Garden and Bronx Zoo, USA) and Asia (Jurong Bid Park,
Singapore). Some of the reasons for this include the species’ usefulness as a
representative of native wildlife in Madagascar-themed exhibits, and the use of the
Madagascar fody as a model species for the more endangered Mauritius fody (Foudia
rubra), which will be discussed in greater detail below.

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During breeding periods, male individuals of this species are easily identifiable by
brightly-coloured plumage and a black patch around the eye (Morris and Hawkins,
1998). At this time, plumage is generally vermillion red in colour, or occasionally
orange or yellow, around the head and underparts. Outside of the breeding season
this plumage returns to an olive-brown colour, similar tio the year-round plumage
colouration of females and juveniles of the species (Morris and Hawkins, 1998).
2.3.2 Mauritius Fody
Unlike the Madagascar fody, the Mauritius fody is categorised as ‘Endangered’ by the
IUCN (IUCN, 2012). This is due to the species’ extremely small population, estimated
at 108 breeding pairs in 2012 (IUCN, 2012). A la rge historical decline in population
size and habitat range has been attributed excessive predation by introduced species,
in particular the crab-eating macaque (Macaca fascicularis) and the black rat (Rattus
rattus) (Safford, 1997), and to the clearing of forests for plantations (IUCN, 2012).
Though the species naturally occupies forested regions, the threat of these invasive
predators has caused an increased reliance on plantations as protective nesting space
(Cristinacce et al., 2009). As a result of these threats, the species is now restricted to
south-west Mauritius; the extent of habitat loss can be seen in Figure 2.1, with red
regions indicating areas where the Mauritius fody is regionally extinct and yellow
areas indicating present range.

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Figure 2.1 The current geographical range of the Mauritius fody, represented by
yellow regions, compared to areas where the Mauritius fody is regionally extinct,
represented by red regions (BirdLife International and Natureserve, 2012)
A major role in the conservation of the Mauritius Fody is taken by the Gerald Durrell
Endemic Wildlife Sanctuary, Mauritius, which undertakes captive breeding of the
species in a private, forested environment. This program has seen some success,
producing 47 individuals in 2005 (Anon., 2005). Additionally, although no populations
of the Mauritius fody are kept outside of Mauritius a number of institutions including
the Durrell Wildlife Park maintain collections of the closely-related Madagascar fody
as a model species. This is useful for the development of captive breeding techniques
and for the use in public education on species conservation activities in Madagascar
and the Mascarene Islands.
2.4 Previous Studies into Fody Behaviour
Studies have determined various differences between the behaviour of members of
the fody family, particularly in relation to territorial behaviour. The Mauritius fody, for
example, is known to maintain a territory throughout the year, whereas the

14
Madagascar fody has been observed to only maintain territories during the breeding
season (Garrett et al., 2007). Though this season is generally thought to be between
the months of December and June (Garret et al., 2007), Madagascar fody
populations have been recorded to breed from November - March / April on the
Seychelles and as early as September in Madagascar (Brickell, 2006). Brickell (2006)
suggests that the species breed primarily during rainy periods, therefore the variation
in breeding season may be attributed to climatic differences experienced by
geographically separate populations.
During breeding, as a weaver bird, nests constructed by the Madagascar fody are
woven loosely together into a spherical shape, using strips of palm leaves and
grasses, and are built c.1m-3m from the ground (Brickell, 2006). In situ, these nests
are additionally lined with woolly fibres taken from the kapok tree (Ceiba pentandra).
The entrance to these nests is generally constructed near the top of the structure,
forming a downward-curving tunnel (Brickell, 2006). An example of this structure can
be seen in Figure 2.2
Figure 2.2 The nest of the Madagascar fody (Krejčík, 2009)

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In a comparative study of territories of Madagascar and Mauritius fodies on Ile aux
Aigrettes, Garret et al. (2007) determined that the male Mauritius fodies each
maintained considerably larger territories (c. 1 hectare) than the male Madagascar
fodies (c. 0.07 hectares). This was attributed the larger population density of
Madagascar fodies in the study area, hence less space available per individual. Only a
portion of Madagascar fodies were recorded accurately, however, and only for the
first period of Garret et al.’s (2007) study, therefore the number and size of defended
Madagascar fody territories may not be accurately represented. As more Mauritius
fodies had established territories by the second stage of Garrett et al.’s (2007) study,
the combined territories had expanded to utilise almost all of the available space on
Ile aux Aigrettes (illustrated in Figure 2.3) and existed congruently to one-another.
This implies that territory size is flexible, adjusting to accommodate as much space
as can be acquired by each individual based on the overall space available.
Figure 2.3 (a) Observed territories of Mauritius fodies and Madagascar fodies
defended during April-June 2005 on Ile aux Aigrettes. (b) Observed Mauritius fody
territories on Ile aux Aigrettes defended in December 2005 (Garret et al., 2007).

16
In terms of territory defence, a study conducted by Kraaijeveld and Komdeur (2003)
identified responses of various intensity by territory-holding Seychelles fody (Foudia
sechellarum) pairs, one male and female, to the presence of a decoy male (in full
breeding colouration) under controlled conditions. These responses varied based on
the activity of the territory-holding individuals; responses were generally
unaggressive during nest-building activities, but escalated to physical altercation
during incubation periods (Kraaijeveld and Komdeur, 2003). This may imply that
territories are most aggressively defended in the protection of offspring. While these
finding may not be applicable to other species of fody, due to behavioural differences,
Kraaijeveld and Komdeur’s (2003) study also recorded a reaction of equal intensity to
the presence of Madagascar fody decoy subjects as to conspecifics, suggesting that
interspecific violence may occur as a result of fody breeding behaviour.
In contrast to behaviours observed during breeding seasons, fody social structure
may differ during other times of the year. Madagascar fodies, for example, have been
observed forming flocks composed of several hundred individuals in situ outside of
the breeding season (Brickell, 2006). In spite of this, Brickell (2006) suggests
optimal ex situ housing for Madagascar fodies to be a mixed sex pair or a single male
with two females, indicating that the non-breeding season social structure of the
Madagascar fody is not thought to be of importance for ex situ welfare. To this end,
Brickell (2006) recommends an allocation of an aviary of around 4m long x 3m wide
x 3m high as suitable housing space for the social grouping of Madagascar fodies
identified above.

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2.5 Territoriality
Territory has been defined as a more-or-less exclusive area that is defended by an
individual or group (Davies and Houston, 1984). Though a territory may or may not
incorporate an animal’s full home range, territory differs from home range in that,
through territorial defence, the territory-holder has exclusive or priority access to
that area’s resources (Powell, 2000), benefitting fitness. As such, territorial behaviour
is only thought to occur when a population is subjected to a limiting resource (Brown,
1964). Based on the relationship between territoriality and limiting resources, Brown
(1964) proposed the idea of ‘economic defendability’. This theory suggests that a
territory should only be defended if the resources within the area are sufficiently
valuable to justify the costs of defence. Carpenter and MacMillen (1976) further
suggested that a territory should only be formed when the limiting resource is
sufficiently abundant to compensate for the effort of defence, but is sufficiently
scarce that access to an exclusive supply is necessary. Consequently, territorial
behaviour is known in a number of species to vary with season (Hunt et al., 1995;
Wikelski, Hau and Wingfield, 2000) or resource abundance (Carpenter and MacMillen,
1976; Powell, Zimmerman and Seaman, 1997).
In some species, territories are shared and defended by more than one individual. A
well-known example of such a species is the pied wagtail, observed by Davies and
Houston (1981). As the territory owner cyclically feeds on insects washed onto river
banks then patrols his territory, a secondary or ‘satellite’ male follows the same
procedure half a cycle behind (Davies and Houston, 1981). In this way the territory is
more closely defended, hence the satellite male is tolerated. This relationship
dissolves, however, if resources become sufficient for only the territory holder
(Davies and Houston, 1981).
As with the pied wagtail, food is among the most common limiting resource for
territory formation (Powers and McKee, 1994; Toobaie and Grant, 2012), however

18
reproductive resources such as nest sites (Aebischer et al., 1996; Lindstrōm and
Pampoulie, 2001) may also contribute. In these instances, territory may directly
relate to reproduction; mate selection by females may be influenced on the
availability of such resources in the male’s territory (Andersson, 1994).
Alternatively, territory may influence mate selection by affecting the physical quality
of the territory-holder. The Madagascar fody, as an example, displays breeding
plumage through the use of carotenoid-based pigments (Estep, Shawkey and Hill,
2006). These pigments naturally occur in various types of vegetation, and
consequently must be consumed by the animal to be utilised in colouration (Goodwin,
1984). Based on the fact that the acquisition of this resource is energetically costly,
the quality of the plumage colouration (or scale colouration in fish species) is
considered to be an honest indicator of breeding quality (Andersson, 1994; Hill,
2002). This relates to territory in that a territory-holding individual will have
preferential access to resources and will consequently display superior physical
condition / breeding quality. Males may alternatively demonstrate fitness through
other means, such as the construction of nests to attract females (Collias and Collias,
1984; Szentirmai, Komdeur and Székely, 2008).
2.5.1 Determinants of Territory Size
As discussed previously, the formation of a territory is dependent on the abundance
of the desired resource(s) within the area (Brown, 1969; Carpenter and MacMillen,
1976). A study by Simon (1975) gave evidence that the territory sizes of wild iguanid
lizards (Sceloporus jarrovi) fluctuated based on the concentration of food in the area;
when the resource is more concentrated, less space must be defended to meet the
territory-holder’s requirements. Resource abundance has also been found to have
similar impacts on territory size in more recent studies (Marshall and Cooper, 2004;
Zharikov and Skilleter, 2004), supporting Simon’s (1975) findings.

19
Territory size is additionally determined by population density. This is well-illustrated
in Figure 2.3 (Garrett et al., 2007); with the increase in territory-holding males,
Mauritius fody territories changed in size and position during Garrett et al.’s (2007)
study. This relationship has also been observed in other studies of territory (Turpie,
1995; Kwiatkowski and Sullivan, 2002). This reflects the idea by Huxley (1934), that
territories are subject to a degree of elasticity, able to expand and contract, but that
contraction is resisted by territorial defence at boundaries. Such resistance may
occur through non-aggressive methods, such as scent marking (Wronski et al., 2006;
Jordan, Cherry and Manser, 2007) and vocalisations (Mennil, 2006; Darden an
Dabelsteen, 2008), or through physical confrontation (Reaney et al., 2008; Arnott
and Elwood, 2009). In the case of leks, individual territories may be very small in
relation to a large population density, such density being necessary to draw widely-
dispersed females for breeding (Balmford, 1991).
Finally, Verbeek et al. (1999) relate the acquisition of territory to social dominance,
this being affected by such physiological strengths as size and weight. The
relationship between body size and territory size has been suggested in other stud ies
(Adams, 2001), however this is not always the case (Strohm and Lechner, 2000) and
some argue that access to a larger territory with greater resources may itself
influence increased body size in the holder (Both and Visser, 2000; Candolin and
Voigt, 2001). In this way, holders of bountiful territories may also be more likely to
win conflicts than an invader. Verbeek et al. (1999) also imply a relationship between
social dominance and individual behavioural qualities, particularly aggression. Though
the impact of temperament on success in natural populations has not historically
been extensively researched (Wilson et al., 1994), Réale et al. (2007) suggest that
certain ecological conditions may indeed be favourable to individuals with particular
behavioural dispositions, and that differences in temperament may result in varying
territory use between individuals.

20
2.5.2 Methods of Calculating Territory Size
A number of methods have been developed to calculate a subject’s overall territory
size, or home range, based on observations of the subject’s movements. In a study
of wild chimpanzee territories, Herbinger, Boesche and Rothe (2001) employ and
compare a number of these methods. The most simplistic of these is the grid cell
method; calculating the sum of all grid cells visited by subjects on a map of subjects’
movements (Horner and Powell, 1992; Zoellick and Smith, 1992; Herbinger, Boesche
and Rothe, 2001). Alternatively, the minimum-convex polygon (MCP) method
determines distribution by calculating the area of a convex polygon formed by joining
the outer limits of subjects’ observed positions (Hayne, 1942; Wiktander, Olsson and
Nilsson, 2001; Dillon and Kelly, 2008).
While favoured for their simplicity, these methods are vulnerable to inaccuracies
through overestimation; the MCP method assumes that distributions form a convex
polygon shape, which is unlikely in heterogeneous habitats, and consequently may
include unused areas in the calculated distribution (Anderson, 1982; Kool and Croft,
1992; Burgman and Fox, 2003). Similarly, the grid cell method is heavily influenced
by the size of cells used (Herbinger, Boesche and Rothe, 2001; Fieberg and
Kochanny, 2003). Herbinger, Boesche and Rothe (2001) suggest methods of
improving these techniques’ reliability, such as creating smaller grid cells in relation
to the sample area or assuming that unused areas included by the MCP method
compensate for missed observations.
A potentially more accurate alternative is the statistical Fourier method, developed by
Anderson (1982) using the Fourier series. Using this method, observed locations are
recorded as spikes over the third dimension of an x-y plane (Powell, 2000). The
spikes are then smoothed, using the Fourier transform estimator, into a surface to
determine distribution. The advantage of the Fourier method is that the form of the
calculated territory is not assumed, as with the MCP method. In Herbinger, Boesche

21
and Rothe’s (2001) study, the Fourier method calculated subject territory sizes to be
far smaller than when calculated using the grid cell or MCP methods. This implies the
large extent to which the non-statistical methods overestimate areas of distribution.
2.6 Research Rationale
Madagascar fodies demonstrate seasonal territoriality in situ, and are known to
demonstrate aggression towards conspecifics during such times (Brickell, 2006).
While some research has been conducted determining the average sizes of the in situ
territories of this species (Crook, 1961; Garrett et al., 2007), the spatial restrictions
of the captive environment prevent the allocation of equal space ex situ (Hosey, 2005;
Kagan and Veasey, 2010). Consequently, the sample population of this study
experience a higher population density than may occur naturally during breeding
periods. In spite of this, the occurrence of territorial behaviour by this species in a
shared, captive environment has not been researched.
This study investigates which areas of a shared aviary are utilised by each mature
male in a captive population of Madagascar fodies. This is in order to determine
whether the captive population exhibits territorial behaviour, and consequently
whether a finite number of mature males can be safely housed in the available space.
This study additionally used behavioural observation techniques to assess the
relationships between territory size and aggression, and between territory size and
courtship behaviour. In terms of management, this information is essential to ensure
the welfare of the captive population, based on the potential risk of territorial
aggression, and to determine whether certain individuals are more likely to reproduce.
This additionally benefits conservation, in that a successful, healthy population of
Madagascar fodies will continue to raise awareness of conservation efforts for the
Mauritius fody, and other endangered species native to the Madagascar fody’s natural
range.

22
3.0 MATERIALS AND METHODS
3.1 Subjects
Four adult male Madagascar fodies and one juvenile that was expected to mature
during the course of the investigation were observed for this study. These subjects
were housed at the Durrell Wildlife Park, Jersey, UK. All subjects were born and
reared at the Durrell Wildlife Park, but were of varying ages. One individual lacked
any identification rings, and was suspected to be an unrecorded, unmanaged birth
(an egg, or clutch of eggs, being unnoticed and the offspring being reared
successfully). As a result, this individual was of unknown age, parentage or breeding
quality but was permitted to operate within the aviary.
Prior to the study all subjects appeared to be healthy, with no indication of physical
complaints or abnormal behaviour. The four adult males displayed full breeding
season plumage colouration, while the younger male displayed a mixed plumage of
non-breeding season colouration with sparse patches of red.
3.2 Housing
Subjects were housed in the Kirindy Forest walkthrough aviary (c. 30m x 20m
including entranceways). This aviary was first opened to the public in November 2009,
and is designed as an ‘immersion experience’ simulation of the dry forest regions of
western Madagascar (DWCT, 2009). As such the main area of the aviary is heavily
vegetated, but contains a wide central path with seating areas to accommodate
visitors. Rope cordons run along the length of this path, on either side, to deter
visitors from entering restricted parts of the enclosure. While subjects can be
observed from any point along the path, allowing for very close observation, areas of

23
the enclosure off the path are difficult to access due to heavy vegetation and three
ponds situated throughout the enclosure. This also makes these difficult areas to
conduct observations around due to the concealment they provide to smaller species.
Birds also have access to multiple smaller indoor shed areas which may be sealed if
isolation becomes necessary. Subjects are effectively concealed in these areas.
Additionally, high wooden awnings are situated at various points throughout the
aviary to provide shade. Subjects are also concealed when perched on these
structures.
As a mixed exhibit, Kirindy Forest aviary contains a number of bird species found in
Madagascar in a shared environment. These species included Madagascar ibis
(Lophotibis cristata), Hottentot teals (Abas hottentota), Hamerkop (Scopus umbretta)
and others at the time of observation. The enclosure is also occupied by a number of
female and juvenile Madagascar fodies which were not subjects of this study. Though
all resident birds are free-flying, escape is prevented using a large tent of netting
over the entire enclosure, supported by large central posts. The aviary is otherwise
open to the environment, having no roof over the main area. A single entrance and a
single exit, at either end of the path, are normally used by both visitors and keepers.
Each of these doors adjoin to smaller enclosed rooms before entering or leaving the
aviary, with the function of preventing escape; instructions to open only one door at
once are clearly posted nearby. During the study the exit door was sealed to avoid
visitors causing stress to a neighbouring narrow-striped mongoose (Mungotictis
decemliniata) that had recently given birth, therefore visitors were required to leave
via the way they came in.
A diagram of the Kirindy Forest Aviary (not to scale) is shown in Figure 3.1.

24
3.3 Management Strategies
The aviary is open to the public between the hours of 9am and 4.30-5pm (varying
based on season). During this time visitors have unrestricted access to the aviary,
leading to a random variation of no human presence to the presence of large groups
of various ages. This had resulted in a degree of habituation, and resident birds are
generally unaffected by human presence provided visitors do not leave the path.
Regardless, visitors are requested to behave sensibly and quietly, and to make no
attempt to touch or feed the animals.
Food is routinely distributed into numerous bowls spread at elevated locations
throughout the aviary (generally attached to trees). Several varying feeds are
provided around the enclosure due to the varying diets of different resident species.
In addition, millet sprays are suspended in vegetation throughout the enclosure as an
added feeding enrichment. No water bowls are supplied, however the three ponds
around the aviary provide a constant source of fresh water.

25
Key
- Path
- Vegetation
- Sheds
- Ent / Exit
- Ponds
- Utility area
Figure 3.1 Layout of the Kirindy Forest Aviary (not to scale)
IN
OUT

26
3.4 Preliminary Investigation
Preliminary observations of each subject were carried out over the course of two
weekends prior to the study. Each individual was observed for thirty minutes per day
between the hours of 12pm to 2.30pm. The purpose of these observations was to
familiarise the observer with the behaviours displayed by each individual, both for
easier future recognition and for the construction of an ethogram to record
behaviours during the study. Additionally, the preliminary study allowed proper
observation techniques to be developed based on the layout of the aviary, the usual
movements of the subjects and the number of observers.
3.5 Enclosure Mapping
Due to the absence of an accurate map of the aviary prior to the study, one was
constructed specifically for the task. This was achieved by taking measurements of
the distance of each corner of the aviary, as well as prominent structures and
features, from three other points in the enclosure. Following this, a mathematical
compass was used to plot each point on a 1:100 scale to construct an accurate
diagram.
A grid was then laid over the constructed diagram, with each cell measuring 2c m2
and representing 2m2 of the aviary, for use in recording the location of a subject at
each interval during observations. This completed grid can be seen in Figure 3.2.
Areas for the recording of a subject’s location (grid reference) were also added to the
recording sheet for the study (Appendix 1), which will be elaborated on below.

27
Figure 3.2 An accurately plotted diagram of the Kirindy Forest aviary (1:100 scale), including prominent structures, overlaid with a
grid for the recording of subject location

28
3.6 Ethogram
During the preliminary investigation, observed behaviours were recorded and sorted
into relevant categories. These categories included territorial, courtship, feeding,
vocalisation and neutral behaviours. The criteria for a behaviour to apply to these
categories can be seen in Table 3.1. Once catalogued, these behaviours were used to
construct an ethogram to record their frequency during the main study (Appendix 1).
The ethogram included the option to record previously unobserved behaviours and to
record instances where a subject was not visible. The full description of behaviours
recorded during the study can be seen in Table 3.2.

29
Table 3.1 Description of behaviour category criteria
Behaviour Category Identifying Criteria
Territorial Subject is actively defending territory, or engaging in a dispute with conspecifics
Aggressive Subject is engaged in an altercation with another individual
Courtship Subject is engaged in mating or mating practices
Feeding Subject is acquiring food from any source in the aviary
Vocalisation Subject is vocalising (may coincide with other behaviours)
Neutral Subject is inactive, travelling or engaged in behaviour unrelated to the above categories

30
Table 3.2 Ethogram of observed behav7iours
Behaviour Category Description
Physical displaying Territorial Subject stands tall and / or puffs out feathers to appear larger, often very exposed
Pursuing mature male Territorial / Aggressive Subject actively chases another mature male
Pursuing female / juvenile Territorial / Aggressive Subject actively chases female or juvenile (indistinguishable at a distance)
Fleeing mature male Territorial Subject is actively chased by another mature male
Fighting Territorial / Aggressive Subject engages in physical confrontation with another mature male
Nest building Courtship Subject engaged in construction of nest
Collecting nest material Courtship Subject acquiring nest-building material from any source
Mating Courtship Subject copulating with female
Food bowl feeding Feeding Subject feeding from any supplied food bowl around the aviary
Millet spray feeding Feeding Subject feeding on millet sprays supplied around the aviary
Enclosure vegetation feeding Feeding Subject feeding from any other natural food source in the aviary
Active vocalisation Vocalisation Rapid chirping, often while exposed or engaging in territorial behaviour
Passive vocalisation Vocalisation Slower, relaxed but continuous chirps, often while engaged in non-territorial behaviours
Preening Neutral Self-grooming of the feathers
Travelling (flight) Neutral In flight without pursuing another individual
Sheltering Neutral A period of inactivity, often when concealed
Other - Any behaviour not recorded during preliminary observation (will be specifically recorded)
Not visible - Subject concealed or lost, generally in shed areas, awnings or thick vegetation

31
3.7 Investigation Procedure
Observations took place over a period of 50 days, between the hours of 1pm and
2.15pm every day. Prior to every observation, a note was made of present weather
conditions.
Subjects were observed one at a time by a single observer, and all subjects were
observed each day. Observations were conducted from whichever area of the aviary
path was convenient to clearly view the current subject without disrupting the
subject’s behaviour. Consequently, this location varied between areas of the path
frequently during and between observations based on the subjects’ movements. Each
subject was observed for fifteen minutes per day, during which time the behaviours
exhibited by the observed subject were recorded at thirty second intervals. Therefore,
31 intervals were recorded per subject per day, totalling 1550 recordings for each
subject over the course of the study. Multiple behaviours may have been recorded at
each of these intervals, for example subjects may have vocalised whilst
simultaneously engaging in other behaviours.
Due to the limited number of observers, all recordings were made vocally, using a
Dictaphone, and were later transcribed into the constructed ethogram (a fresh
ethogram was used for each subject). This was to prevent inaccuracy in the event
that the observer lost track of a subject while making a written recording.
The location of the subject being observed was simultaneously recorded at each
thirty second interval using the appropriate grid reference from the constructed map.
As no devices could be employed to make physical markers of the grid in the aviary,
grid reference was determined by the observer’s interpretation. Finally, the elevation
of the subject being observed was recorded in conjunction with the subject’s location.
Due to the inability to install accurate measuring devices for this observation, the
elevation of the subject was recorded as being at ground level, lower level, middle

32
level or upper level based on the interpretation of the observer (generally influenced
by the height of vegetation or man- made structures in close proximity to the subject).
The occurrence of noteworthy events (such as irregular environmental conditions or
changes in weather mid-observation) were also recorded when appropriate.
3.8 Variables
A number of uncontrollable factors may have impacted upon the results by
influencing subject behaviour. In particular these included weather, human presence,
the behaviour of other residents of the aviary (including conspecifics), and potentially
influential activity outside of the aviary (such as construction noises).
In contrast, variables such as subjects, observation times, observation techniques
and husbandry procedures were kept constant throughout the study.
3.9 Ethical Considerations
This study was carried out with consideration of the ethical acceptability of
investigative procedures used. To this end, an ethical review was completed prior to
the commencement of the study. This review may be seen in Appendix I.
At no point during the study were any animals harmed or physically contacted, nor
were any alterations made to the sample population’s husbandry regime.

33
3.10 Risk Assessment
The potential risks to individuals involved in this study have been considered, and
appropriate control measures to reduce these risks have been determined prior to
the start of observations. The full risk assessment can be seen in Appendix II.
3.11 Data Analysis
Territories were calculated over four periods during the study. The time frames
represented by these periods are described in Table 3.3. The areas utilised by each
individual during each of these periods were marked on the enclosure map (Figure
3.2). The existence and location of territories were determined by examining the
distribution of each subject’s movements during each period, and through statistical
tests of cell use (which will be elaborated on below). Territory size was then
calculated as the sum area of all cells that formed the territory.
Table 3.3 Description of timeframe covered in each study period
Period Days Covered Dates Covered
1 1-13 07/05/2012 – 19/05/2012
2 14-25 20/05/2012 – 31/05/2012
3 26-37 01/06/2012 – 12/06/2012
4 37-50 13/06/2012 – 25/06/2012

34
3.11.1 Statistical Analysis
All statistical calculations were performed in SPSS® (Version 19). Results were
considered significant if p<0.05, indicating a confidence level of 95%, very significant
if p<0.01, indicating a confidence level of 99%, and highly significant if p<0.001,
indicating a confidence level of 99.9%.
To confirm the existence of territories within the enclosure, rather than shared use of
the entire area, an independent-samples t-test was used to determine whether a
significant difference existed between the number of cells (Figure 3.2) used by a
single individual and the number used by multiple individuals.
Two separate one-way analysis of variance (ANOVA) tests were then conducted. The
first was used to determine if territory size was significantly different between
individuals during the study. The second was used to determine if the amount of
space utilised as territory was significantly different between study periods. An
additional one-way ANOVA was conducted to determine if a significant difference
existed between the use of enclosure levels over the course of the study.
The relationship between territory size and the occurrence of aggressive behaviours
was determined using a scatterplot and Spearman’s rank-order correlation. A one-
way ANOVA was then used to determine if the occurrence of aggressive behaviour
was significantly different between subjects. These two tests were then repeated for
the occurrence of courtship behaviours.

35
4.0 RESULTS
Unexpectedly, within the first week of observations two subjects (B6482 and B6952)
were no longer viable for study. Despite no indication of abnormal behaviour during
the previous day’s observations, neither individual could be located on Day 6 of the
study (12/05/2012). Subject B6482 was located the following day, showing signs of
poor physical condition, preferring to remain concealed in the enclosure shed areas,
and was observed to be repeatedly, aggressively pursued by other subjects. Subject
B6482 was consequently isolated as a protective measure, in the enclosure sheds,
and was not observed for the remainder of the study. Subject B6952 was never
located following the initial disappearance.
As a result of the early departure of these subjects from the study, data collected
relating to changes in their territory size will not be included in statistical tests. This
is to prevent skewing of results. Data collected prior to these subjects’ departure will
still be displayed in results tables, and the impact of their departure from the study
will be considered in the discussion portion of this paper.
4.1 Territories
4.1.1 The Existence and Distribution of Territories
The distribution of subjects at the conclusion of each period may be seen in Figure
4.1 (Period 1), Figure 4.2 (Period 2), Figure 4.3 (Period 3) and Figure 4.4 (Period 4).
The total number of cells utilised by a single subject and the total number of cells
utilised by more than one subject, at the conclusion of each period, are listed in Table
4.1.
An independent-samples t-test showed that a greater number of cells were utilised
by a single subject (61.5 ± 6.25) than were utilised by more than one subject (22.5

36
± 8.51) over the course of the study, indicating a highly significant difference of 39
(95% Confidence Interval (CI), 26 to 52), t(2) = 7.392, p<0.0005. This is displayed
in Figure 4.5.
Table 4.1 The number of cells utilised by a single subject versus the number of cells
utilised by more than one subject over the course of each period.
Period Cells Used by Single Subject Cells Used by Multiple Subjects
1 54 35
2 59 19
3 68 20
4 65 16
Total 246 90

37
Figure 4.1 Distribution of subjects during Period 1 of the study (B6482 and B6952 prior to departure). Symbols indicate instances
where associated subject (see key) was observed outside of normal distribution.
Key
- B6481
- B6482
- B6804
- B6952
- No ID

38
Figure 4.2 Distribution of subjects during Period 2 of the study. Symbols indicate instances where associated subject (see key) was
observed outside of normal distribution.
Key
- B6481
- B6804
- No ID

39
Key
- B6481
- B6804
- No ID

40
Key
- B6481
- B6804
- No ID

41
Figure 4.5 Mean (±S.E.M.) frequency of cell use by single subjects and by multiple
subjects across study.
4.1.2 Variation of Territory Size
The calculated territory sizes for each subject (including areas of overlap) during
each study period are displayed in Table 4.2.

42
Table 4.2 The approximate size of each subject’s territory during each study period.
Subject
Approximate Territory Size (m2)
Period 1 Period 2 Period 3 Period 4
B6481 164 160 160 152
B6482* 48 - - -
B6804 100 80 100 88
B6952* 72 - - -
No ID 100 108 124 112
*Prior to departure from study
The results of the one-way ANOVA showed that the difference in territory size
between individuals was highly significant, F(2,9) = 64.645, p<0.0005. The largest
territory was held by subject B6481 (159 ± 5.03), followed by subject No ID (111 ±
10). Subject B6804 was found to have the smallest territory (92 ± 4.9). This is
illustrated in Figure 4.6.
The mean difference in territory size between subjects No ID and B6804 (19, 95% CI
(2.04 to 35.96)) was found to be statistically significant by Tukey post-hoc tests
(p=0.03). The differences between territory size of subjects B6481 and No ID (48, 95%
CI (31.04 to 64.96)) and between subjects B6481 and B6804 (67, 95% CI (50.04 to
83.96)) were both found to be highly significant (p<0.0005).

43
Figure 4.6 Mean (±S.E.M.) approximate territory size (m2) of each subject
(excluding subjects B6482 and B6952).
A separate one-way ANOVA determined that the difference in territory sizes between
study periods was not significant (F(3,8) = 0.07, p = 0.974) (data were not normally
distributed for the “Period 1” group).
4.2 Enclosure Level Preference
The total utilisation of each enclosure level by each subject, over the course of the
study, is displayed in Table3.3.

44
Table 4.3 Total number of times each subject was recorded at each level of the
enclosure during the study
Subject
Enclosure Level
Ground Lower Middle Upper
B6481 20 112 867 438
B6482* 0 27 154 1
B6804 12 181 1011 238
B6952* 1 17 114 19
No ID 8 95 874 338
Total 41 432 3020 1034
* Prior to departure from study
One-way ANOVA indicated that the difference between the mean frequency of
utilisation of enclosure levels was highly significant, F(3,8) = 103.5, p<0.0005.
Middle level was utilised most frequently (917.3 ± 81.2), followed by Upper level
(338 ± 100) then Lower level (129.3 ± 45.5). Ground level was utilised the least
frequently (13.3 ± 16.1). This is shown in Figure 4.7.
Tukey post-hoc tests determined that the difference in utilisation between Middle
level and Upper level (579.3, 95% CI (400.54 to 758.13)), Middle level and Lower
level (788, 95% CI (609.2 to 966.8)) and Middle level and Ground level (904, 95%
CI (725.2 to 1082.8)) were all highly significant (p<0.0005). The difference between
Upper level and Lower level (208.7, 95% CI (29.87 to 387.46)) was also found to be
significant (p=0.024), and the difference between Upper level and Ground level
(324.7, 95% CI (145.9 to 503.5)) was found to be very significant (p=0.002). The
difference between Lower level and Ground level (116, 95% CI (-62.8 to 294.8)) was
not found to be significant (p=0.239).

45
Figure 4.7 Mean (±S.E.M.) number of time each enclosure level was utilised during
the study
4.3 Occurrence of Aggressive Behaviour
The occurrence of each observed behaviour that was considered to be ‘aggressive’
from each individual, during each period and overall, is displayed in Table 4.4.

46
Table 4.5 The occurrence of each aggressive behaviour, and overall aggressive behaviours, for each subject during each study period
(P) and in total throughout the study (T)
*Prior to departure from study
Occurrence of Aggressive Behaviours
Subject Pursuing Mature Male Pursuing female / juvenile Fighting Overall
P1 P2 P3 P4 T P1 P2 P3 P4 T P1 P2 P3 P4 T P1 P2 P3 P4 T
B6481
7 1 2 4 14 46 29 41 37 153 1 1 1 3 6 54 31 44 44 173
B6482*
0 0 0 0 0 2 0 0 0 2 0 0 0 0 0 2 0 0 0 2
B6804
1 3 4 4 12 12 6 9 4 31 0 1 2 2 5 13 10 15 10 48
B6952*
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
No ID
2 3 1 3 9 19 20 19 21 79 0 1 0 0 1 21 24 20 24 89

47
The results of a Spearman’s rank-order correlation showed that a very weak positive
correlation existed between territory size and the occurrence of aggressive behaviour,
however this was not statistically significant, rs(10) = 0.134, p = 0.679.
A one-way ANOVA determined that the number of occurrences of aggressive
behaviours were highly significantly different between individuals over the course of
the study, F(2,9) = 30.708, p<0.0005. Subject B6481 exhibited aggressive
behaviour the most frequently during each period (43.25 ± 9.4), followed by subject
No ID (22.25 ± 2.1). Subject B6804 exhibited aggressive behaviour the least
frequently (12 ± 2.4). This is shown in Figure 4.8.
Tukey post-hoc tests determined that the mean difference between the occurrence of
aggressive behaviour in subjects B6804 and No ID (12.25, 95% CI (-1.1 to 21.6))
was not significant (p = 0.076). The mean occurrence of aggressive behaviour by
subject B6481 was found to be very significantly greater than by subject No ID (21,
95% CI (9.7 to 32.4), p = 0.002) and highly significantly greater than by subject
B6804 (31.25, 95% CI (19.9 to 42.6), p<0.0005).

48
Figure 4.8 Mean (±S.E.M.) number of times each subject exhibited ‘aggressive’
behaviour
4.4 Occurrence of Courtship Behaviour
The occurrence of each ‘courtship’ behaviour by each individual, over each period and
overall, is displayed in Table 4.5.
Comparing territory size and courtship behaviour on a scatter graph identified only
an extremely weak, negative correlation between the two (R2 = 0.145). This is
displayed in Figure 4.9. This was confirmed, using Spearman’s rank-order correlation,
to not be statistically significant, rs(10) = -0.296, p = 0.399.

49
Table 4.5 The occurrence of each courtship behaviour, and overall courtship behaviours, for each subject during each study period (P)
and in total throughout the study (T)
* Prior to departure from study
Subject
Occurrence of Courtship Behaviours
Collecting Nest Material Nest Building Mating Overall
P1 P2 P3 P4 T P1 P2 P3 P4 T P1 P2 P3 P4 T P1 P2 P3 P4 T
B6481 23 2 0 1 26 15 3 2 3 23 2 0 0 0 2 40 5 2 4 51
B6482*
0 0 0 0 0 3 0 0 0 3 0 0 0 0 0 3 0 0 0 3
B6804
42 3 27 18 90 71 7 44 16 138 0 0 1 0 1 113 10 72 34 229
B6952*
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
No ID
13 12 19 1 45 22 23 23 0 68 0 0 0 0 0 35 35 42 1 113

50
Figure 4.9 Relationship between territory size and the overall occurrence of
courtship behaviours
Additionally, one-way ANOVA determined that there was no significant difference in
the mean exhibition of courtship behaviours between subjects, F(2,9) = 2.263, p =
0.160. This is shown in Figure 4.10.

51
Figure 4.10 Mean (±S.E.M.) occurrence of times each subject exhibited ‘courtship’
behaviours

52
5.0 DISCUSSION
In this section, the findings of this study will be discussed and evaluated to assess
the territorial behaviour of the study species in captivity. Other findings relating to
enclosure use, including enclosure level preference and interactions with conspecifics,
will also be evaluated. Additionally, this section will discuss the limitations of the
study, and suggest areas of further research.
5.1 The Existence of Territory
Significantly more cells (p<0.0005) were utilised by a single individual than by
multiple individuals during the study. Additionally, the distributions of subjects
(Figures 4.1 – 4.4) indicate that subjects utilised separate areas of the enclosure to
one-another. Combined, these findings strongly support the hypothesis, which stated
that territories would be formed in the captive environment. Furthermore, the loss of
subjects B6482 and B6952 from the study supports the hypothesis that not all
subjects would successfully form a territory.
The formation of territories can be attributed to the onset of the breeding season,
coinciding with Garrett et al.’s (2007) observations of a December – June breeding
period. This notion is supported by the author’s observations of multiple reproductive
behaviours during the study period, including mating and the hatching of offspring.
The precise time that territories were formed, prior to the study, is unknown; all
subjects already displayed breeding plumage as early as the preliminary study (with
the exception of the juvenile B6952), and operated within established territories
throughout. As the species’ breeding season has been observed to vary based on
geographical location (Brickell, 2006; Garrett et al., 2007), or based on the

53
occurrence of rain (Brickell, 2006), the breeding season of captive UK populations
may commence at a different time to that of in situ populations.
Also in relation to breeding, the (assumed) intraspecific aggression leading to the
loss of subjects B6482 and B6952 may be attributed to competition for breeding sites
(Andersson, 1994). There would seem to be little basis to suggest competition for
other resources due to the even distribution of food and water throughout the aviary
(author’s observations, however subject B6952 was observed to utilise a food bowl
within the territory of subject B6481
The loss of subjects B6482 and B6952 can be assumed, though not absolutely proven,
to be the result of intraspecific conflict over territory. In this case, the conflicted
limited resource is most likely to be breeding / nesting space. There is less basis to
suggest competition for other resources, based on the even distribution of food and
ad libitum water throughout the enclosure, however the close proximity of subjects
B6482, B6952 and No ID may still have resulted in some conflict over access to food.
5.2 Territory Size
The findings of this study show that the subjects formed territories of significantly
different sizes (p<0.0005), coinciding with the hypothesis that subjects would
maintain differently-sized territories to one-another. Once established, however,
these territories were not found to change significantly in size during the study (p =
0.974).
No existing research could be found relating to the size of territories he ld by birds in
a zoo environment. The size of territories during in situ studies of the Madagascar
fody have been recorded as having a radius of c. 10-30m (Crook, 1961; Garrett et al.,
2007), translating to an area of c. 314 – 2827m2. This range suggests that the
between-subject variation in territory size found in this study reflects the territorial

54
habits of in situ populations. The sizes of territories were considerably lower in this
study than those observed in situ (Crook, 1961; Garrett et al., 2007), with the
largest recorded territory of this study measuring 164m2 and the smallest measuring
only 80m2 (excluding subjects B6482 and B6952)(Table 4.2). The fact that subjects
functioned normally in such a comparatively small area may indicate that the species
is flexible, and is able to cope with a certain degree of spatial restriction.
Alternatively, the ability to cope with spatial restriction may be an effect of being
reared in captivity, and only ever having experienced less spac e than would be
available in situ.
Also similar to Garrett et al.’s (2007) observations is the distribution of territories to
utilise most of the available space. Though Garrett et al. (2007) observed this trend
in Mauritius fodies (Figure 2.3), both observations appear to conform with Huxley’s
(1934) idea of territories expanding until met with resistance from another territory-
holder. In that instance, the occurrence of unused areas during a period may
potentially be attributed to coincidence, based on the relatively short observation
time per subject per day. Alternatively, Figures 4.1-4.4 indicate that the three
territories maintained throughout the study are distributed with relative similarity to
the aviary’s vegetated zones (see Figure 3.1). This may imply that the size and
distribution of territories seen in this study are regulated by the availability of this
vegetation based on the desire for cover and perching locations, with the barren path
acting as a barrier to territory expansion. Future studies may gain better insight into
this area by increasing observation time for increased location recordings, and using
more accurate analytical techniques to determine precise distribution.
Regardless of flexibility or tolerance, the loss of subjects B6482 and B6952 indicates
that there is still a limit to the spatial restriction that can be tolerated. While this limit
cannot be accurately determined from this study, it could be assumed that the space
shared by subjects B6482, B6952 and No ID at the beginning of observations (Figure
4.1) was of insufficient size to accommodate three individuals. This may be

55
interpreted to support the previous idea of the path acting as the ‘territory divider’,
as the three subjects occupied the same undivided body of vegetation. In any case,
the minimum space that can be tolerated may be an important area of future study,
which will be discussed further below.
5.3 Enclosure Level Preference
The collected results indicate that there was a highly significant difference between
the usage of different enclosure levels (p<0.0005). The Middle Level of the enclosure
was used by far the most frequently, confirming the hypothesis that subjects would
show a preference to a particular level. Ground level was used the least frequently,
and while the results do not indicate a significant difference between Ground and
Lower level usage (p = 0.239), Lower level was used far more frequently during the
study than Ground level (Table 4.3). Post-hoc test results may, therefore, have been
skewed by the large difference between the use of the Middle level and other levels.
The frequency with which subjects utilised the Middle level of the enclosure was
almost certainly affected by the presence of food bowls on that level; the Middle level
was consequently utilised for most observed feeding behaviours. Furthermore, the
Middle level was favoured for nest-building (author’s observation) coinciding with the
in situ preference of Madagascar fodies to construct nests 1m-3m from the ground
(Brickell, 2006). This implies that enclosure level preference is influenced by natural
behavioural requirements. Consequently, future studies may benefit from
investigating the relationship between enclosure level and the occurrence of specific
behaviours.

56
5.4 Aggressive Behaviour
No significant correlation was found between territory size and the occurrence of
aggressive behaviour (p = 0.679), conflicting with the hypothesis that increased
territory size would relate to increased aggressive behaviour. However, the difference
in the exhibition of aggression between subjects was still found to be highly
significant (p<0.0005), with the most ‘aggressive’ individual being subject B6481.
This result may relate to Verbeek et al.’s (1999) ideas of social dominance. Subject
B6481 meets a number of indicators of social dominance listed by Verbeek et al.
(1999), being an older, successful male who has potentially held a territory in the
past. If Verbeek et al.’s (1999) notion of aggressiveness also increasing social
dominance is accurate, then this may account for why subject B6481 maintained the
largest territory, based on the theory of certain ecological conditions favouring
particular dispositions (Réale et al., 2007). By contrast, the least aggressive male,
subject B6804, maintained the smallest territory, further supporting this argument.
Alternatively, the male with the largest territory was also the oldest and most
experienced of the surviving subjects; this finding does not determine whether
aggression or any other contributor to social dominance (Verbeek et al., 1999) is
more influential. As such, further studies may wish to examine each subject closely,
in terms of size, weight, age and condition, to better understand this outcome.
Furthermore, the failure of subject B6482, the second oldest male, to maintain a
territory suggests that seniority / experience alone is not sufficient to ensure
territorial success, particularly when multiple males come into conflict over a single
territory, as appears to have been the case with subjects B6482, B6952 and No ID.

57
5.5 Courtship Behaviour
No significant relationship was found between territory size and the occurrence of
courtship behaviours (p = 0.399), conflicting with the hypothesis that increased
territory size would relate to more frequent courtship behaviour. While the difference
in the exhibition of courtship behaviours between subjects was not found to be
significant (p = 0.160), subject B6804 was observed to perform the most courtship
behaviours during the study (Table 4.5) and subject B6481 to perform courtship
behaviours the least frequently (Table 4.5).
These results directly oppose findings relating to aggressive behaviour, seen above.
If subject B6481 is considered to have an ‘aggressive temperament’, then this may
relate to Réale et al.’s (2007) idea of differential use of territory between conspecifics
based on temperament. In this case, therefore, an aggressive temperament could be
assumed to negatively influence the occurrence of courtship behaviours. This
assumption is heavily limited by this study’s small sample size, therefore future
studies may wish to examine the relationship between aggression and courtship
behaviours in a larger population. Additionally, as noted in section 3.2 of this paper,
heavily vegetated areas resulted in numerous missed observations during this study,
therefore courtship behaviour conducted by subject B6481 may have been obscured
and missed.
Alternatively, a possible contributor towards subject B6804’s high frequency of
courtship behaviour was the inclusion of the keeper utility area in B6804’s territory
(Figure 3.1). This area was noted to contain a supply of rope, for enclosure
maintenance, which subject B6804 was frequently observed to exploit as a source of
nesting material. By comparison, other subjects most commonly acquired nesting
material from vegetation or pre-existing nests. Based on this observation, the impact
of material accessibility on nesting behaviour may be an area of future study.

58
Regardless of the theories postulated above, both subjects B6481 and B6804 were
observed to copulate with similar frequency during the study (Table 4.5). This
behaviour was extremely brief (author’s observation), therefore a great number of
copulations may have gone unobserved, even during observation periods. If this
limitation is ignored, then this finding questions whether mate selection by females in
the sample population was more heavily influenced by nest construction (B6804) or
social dominance (B6481). Previous studies into weaver bird reproductive behaviour
would seem to support the former (Crook, 1964), however the relationship between
social dominance and reproductive success is also recognised (Verbeek et al., 1999).
This finding may be better understood if future studies observed the behaviour of
females in the population, though continuously identifying females may be difficult,
over a longer constant period to more accurately catalogue mate and nest selection.
5.6 Additional Limitations and Future Study
A number of factors limited this study, which may have reduced the accuracy or
reliability of the collected results. Further studies into this area should be accordingly
developed to minimise or remove the impact of these limitations.
In regards to the subjects used in the study, results were particularly limited by the
small sample size. The loss of two subjects over the course of this study strongly
suggests that housing an increased number of individuals together would be neither
safe for subjects, due to potential territorial disputes, nor accurate if increasing this
number influenced an inclination towards aggressive territorial behaviour. Sample
size could instead be increased by observing a number of different populations across
multiple institutions. If that became the case, however, the difference on
environmental variables would need to be considered. For example, other populations
may not be housed in an enclosure accessible to the public.

59
In this case the close contact between the subjects and humans was an another
limiting factor to the investigation. As discussed in section 3.3 of this paper, the
continuous presence of visitors had resulted in a level of habituation in residents of
the aviary. The true impact of this on the behaviour of the subjects is unknown, and
the behaviour of subjects in this study may not be reflective of individuals housed in
isolation from close human contact. Based on this, the behavioural impact of human
presence may be an area for future study, as an indication of significantly altered
behaviour may influence the future enclosure design for fodies and similar species.
Finally, in terms of enclosure design, this study was severely limited by the inability
to perceive subjects in certain regions of the aviary. This caused the obvious
detriment of not being able to observe the subject’s behaviour at intervals,
potentially leading to an inaccurate representation of the subject’s activities during
the observation period. In cases where this was the result of thick vegetation, the
issue of visibility may be unavoidable. This is particularly the case if enclosures are
designed to simulate a natural environment, which is known to be sometimes
necessary to avoid stressing the resident (Wielebnowski et al., 2002a; Collaham et
al., 2012). One potential measure for future study would be the use of video
recording equipment in areas where visibility is poor, potentially establishing the
equipment at a better angle to observe the area, and analysing the recording later.
This would be minimally intrusive once set up, and has been successful in other
studies (Vanak and Gommper, 2007; Rowcliffe et al., 2008) , however inaccurate
observations may be caused in the angle or image quality is poor.

60
6.0 CONCLUSION AND APPLICATION OF FINDINGS
To conclude, this study has found strong evidence that the population of Madagascar
fodies in the Durrell Wildlife Park’s Kirindy Forest aviary engage in normal territorial
behaviour, regardless of the spatial restrictions and provision of resources in the captive
environment, including the aggressive defence of territories against conspecifics. This is
based on the loss of subjects during the study, and on the observed movements of
remaining subjects, and supports the main hypothesis.
These findings imply that, to prevent potential detriments to the welfare of resident
animals, the number of mature males of this species housed in the Kirindy Forest aviary
should not exceed three individuals during the breeding period. If this number is
exceeded, animals may be injured or lost through the consequent competition for
territory. Otherwise, individuals can be expected to operate preferentially within the
boundaries of their territory.
This study has additionally determined that, in the sample population, territory size was
not significantly related to the frequency of expression of aggressive or reproductive
behaviours, though these findings may be limited by the small sample size and limited
observation time. As such, the perceived size of an individual’s territory, in this population,
cannot be taken as a direct indicator of aggressive disposition or reproductive success.
Overall, the territorial behaviour of the sample population, and the subjects’ preference
for similar elevation to that utilised in situ, suggest that the captive environment does not
supress natural behaviours. Further studies should examine a larger sample size across
multiple institutions to determine whether this conclusion is applicable to all captive
populations.

61
7.0 REFERENCES
ADAMS, E.S., 2001. Approaches to the Study of Territory Size and Shape. Annual
Review of Ecology and Systematics. 32: pp.277-303.
AEBISCHER, A. et al.,1996. The role of territory choice, mate choice and arrival date
on breeding success in the Savi’s Warbler Locustella luscinioides. Journal of Avian
Biology. 27: pp.143-152.
ANDERSON, D.J., 1982. The home range: A new non-parametric investigation
technique. Ecology. 63: pp.103-112.
ANDERSSON, M. 1994. Sexual Selection. Princeton: Princeton University Press.
ANONYMOUS, 2005. Mauritius Fody being released. MWF Newsletter: 2.
ARNOTT, G. and ELWOOD, R.W., 2009. Assessment of fighting ability in animal
contests. Animal Behaviour. 77(5): pp.991-1004.
BALMFORD, A., 1991. Mate choice on leks. Trends in Ecology and Evolution. 6(3):
pp.87-92.
BASHAW et al., 2003. To Hunt or Not to Hunt? A Feeding Enrichment Experiment
With Captive Large Felids. Zoo Biology. 22: pp.189-198.
BIGGINS, D.E. et al., 1999. influence of prerelease experience on reintroduced
black-footed ferrets (Mustela nigripes). Biological Conservation. 89: pp.121-129.

62
BIRDLIFE INTERNATIONAL AND NATURESERVE (2012). Bird species distribution
maps of the world 2012: Foudia rubra [online]. Available at:
<http://maps.iucnredlist.org/map.html?id=106008576> [Accessed 17 March 2013].
BLANC, F. and THÉRIEZ, M., 1998. Effects of stocking density on the behaviour and
growth of farmed red deer hinds. Applied Animal Behaviour Science. 56: pp.297-307.
BOND, M. and WATTS, E., 1997. Recommendations for infant social environment. In:
SODARO, C., ed. Orangutan Species Survival Plan Husbandry Manual. Chicago:
Chicago Zoological Park, 1997, pp.77-78.
BOTH, C. and VISSER, M.E., 2000. Breeding territory size affects fitness: An
experimental study on competition at the individual level. Journal of Animal Ecology.
69(6): pp.1021-1030.
BOYCE, W.T. et al., 1998. Crowding stress and violent injuries among behaviorally
inhibited rhesus macaques. Health Psychology. 17: pp.285–289.
BRADSHAW, G.A. et al., 2008. Building an Inner Sanctuary: Complex PTSD in
Chimpanzees. Journal of Trauma and Dissociation. 9(1): pp.9-34.
BRICKELL, N., 2006. Collated data on the Madagascar Fody Foudia madagascariensis.
Avicultural magazine. 112(4): pp.169-173.
BROWN, J.L., 1964. The Evolution of Diversity in Avian Territorial Systems. Wilson
Bulletin. 76(2): pp.160-169.
BROWN, J.L., 1969. Territorial behaviour and population regulation in birds: A review
and re-evaluation. Wilson Bulletin. 83: pp.293-329.

63
BURGMAN, M.A. and FOX, J.C., 2003. Bias in species range estimates from minimum
convex polygons; implications for conservation and options for improved planning.
Animal Conservation. 6: pp.19-28.
CANDOLIN, U. and VOIGT, H.R., 2001. Correlation between Mal Size and Territory
Quality: Consequence of Male Competition or Predation Susceptibility?. Oikos. 95(2):
pp.225-230.
CARLSTEAD, K. and SHEPHERDSON, D., 1994. Effects of Environmental Enrichment
on Reproduction. Zoo Biology. 13: pp.447-458.
CARO, T.M., 1993. Behavioral solutions to breeding cheetahs in captivity: Insights
from the wild. Zoo Biology. 12: pp.19-30.
CARPENTER, F.L. and MACMILLEN, R.E., 1976. Threshold model of feeding
territoriality and test with a Hawaiian honeycreeper. Science. 194: pp.634-642.
CLUBB, R. and MASON, G.J., 2007. Natural behavioural biology as a risk factor in
carnivore welfare: How analysing species differences could help zoos improve
enclosures. Applied Animal Behavioural Science. 102: pp.303-328.
COLLAHAM, H. et al., 2012. Lion (Panthera leo) Care Manual. (s.l.): Association of
Zoos and Aquariums.
COLLIAS, N.E. and COLLIAS, E.C., 1984. Nest Building and Bird Behavior. Cambridge:
Harvard University Press.
CRISTINACCE, A. et al., 2009. The release and establishment of Mauritius Fodies
Foudia rubra on Ile aux Aigrettes, Mauritius. Conservation Evidence. 6: pp.1-5.

64
CROOK, J.H., 1961. The fodies of the Seychelles Islands. Ibis. 103(a): pp.517-548.
CROOK, J.H., 1964. The evolution of social organisation and visual communication in
the weaver birds (Ploceinae). Behaviour. Supplement (10).
DARDEN, S.K. and DABELSTEEN, T., 2008. Acoustic territorial signalling in a small
socially monogamous canid. Animal Behaviour. 75: pp.905-912.
DAVIES, N.B. and HOUSTON, A.I., 1981. Owners and satellites: the economics of
territory defence in the pied wagtail, Motacilla alba. Journal of Animal Ecology. 50:
157-180.
DAVIES, N.B. and HOUSTON, A.I., (1984). Territory economics. In: KREBS, J.R. and
DAVIES, N.B. eds. Behavioural Ecology: an Evolutionary Approach. 2nd ed. Oxford:
Blackwell Scientific Publications, 1984, p.p. 148-169.
DEPARTMENT FOR ENVIRONMENT, FOOD AND RURAL AFFAIRS, 2012. Secretary of
State’s Standards of Modern Zoo Practice [online]. Available at:
<http://www.defra.gov.uk/publications/files/standards-of-zoo-practice.pdf>
[Accessed 13 March 2013].
DILLON, A. and KELLY, M.J., 2008. Ocelot home range, overlap and density:
Comparing radio telemetry with camera trapping. Journal of Zoology. 275(4):
pp.391-398.
DURRELL WILDLIFE CONSERVATION TRUST, 2009. Kirindy Forest [online]. Available
at: <http://www.durrell.org/Visit/Our-wildlife-park/Exhibits/Kirindy-forest/>
[Accessed 05 March 2013].

65
EASON, D.K. and MOORHOUSE, R.J., 2006. Hand-rearing kakapo (Strigops
habroptilus), 1997-2005. Notornis. 53(1): pp.116-125.
ELLIOTT, G.P., MERTON, D.V. and JANSEN, P.W., 2001. Intensive Management of a
Critically Endangered Species: The Kakapo. Biological Conservation. 99(1): pp.121-
133.
ESTEP, L.K., SHAWKEY, M.D. and HILL, G.E., 2006. Carotenoid-based breast
plumage colour, body condition and clutch size in red fodies (Foudia
madagascariensis). Ostrich. 77(3-4): pp.164-169.
FIEBERG, J. and KOCHANNY, C.O., 2003. Quantifying home range overlap: The
importance of the utilisation distribution. The Journal of Wildlife Management. 69(4):
pp.1346-1359.
GARRETT, L.J.H. et al., 2007. Competition or co-existence of reintroduced, critically
endangered Mauritius fodies and invasive Madagascar fodies in lowland Mauritius?.
Conservation Biology. 140(1-2): pp.19-28.
GOODWIN, T.W., 1984. The Biochemistry of Carotenoids. New York: Chapman and
Hall.
GOTTLIEB, D.H. et al., 2011. Efficacy of 3 Types of Foraging Enrichment for Rhesus
Macaques (Macaca mulatta). Journal of the American Association of Laboratory
Animal Science. 50(6): pp.888-894.
HARLOW, H.F. and HARLOW, M. K., 1962. Social deprivation in monkeys. Scientific
American. 207: pp.136–146.
HAYDOCK, E.L., 1954. A survey of the Birds of St. Helena. Ostrich. 25(2): pp.62-75.

66
HAYNE, D.W., 1949. Calculation of size of home range. Journal of Mammalogy. 30:
pp.1-18.
HERBINGER, I., BOESCHE, C. and ROTHE, H., 2000. Territory Characteristics among
Three Neighboring Chimpanzee Communities in the Taї National Park, Côte d’Ivoire.
International Journal of Primatology. 22(2): pp.143-167.
HILL G.E., 2002. A Red Bird in a Brown Bag: the Function and Evolution of
Ornamental Plumage Coloration in the House Finch. New York: Oxford University
Press.
HORNER, M.A. and POWELL, R.A., 1990. Internal structure of home ranges of black
bears and analyses of home-range overlap. Journal of Mammalogy. 30: pp.402-410.
HOSEY, G., 2005. How does the zoo environment affect the behaviour of captive
primates?. Applied Animal behaviour Science. 90: pp.107-129.
HOSEY, G., MELFI, V. and PANKHURST, S., 2009. Zoo Animals: Behaviour,
Management and Welfare. New York: Oxford University Press.
HUNT, K. et al., 1995. Temporal Patterns of Territorial Behavior and Circulating
Testosterone in the Lapland Longspur and Other Arctic Passerines. American
Zoologist. 35: pp.274-284.
HUXLEY, J.S., 1934. A natural experiment on the territorial instinct. British Birds. 27:
pp.270-277.
INTERNATIONAL UNION FOR THE CONSERVATION OF NATURE, 1998. IUCN
Guidelines for Re-introduction. Cambridge: IUCN.

67
INTERNATIONAL UNION FOR THE CONSERVATION OF NATURE, 2009. Foudia
madagascariensis [online]. Available at:
<http://www.iucnredlist.org/details/summary/106008573/0> [Accessed 05 March
2013].
INTERNATIONAL UNION FOR THE CONSERVATION OF NATURE, 2012. Foudia rubra
[online]. Available at: <http://www.iucnredlist.org/details/106008576/0> [Accessed
17 March 2013].
JORDAN, N.R., CHERRY, M.I. and MANSER, M.B., 2007. Latrine distribution and
patterns of use by wild meerkats: implications for territory and mate defence. Animal
Behviour. 73: pp.613-622.
KAGAN, R. and VEASEY, J., 2010. Challenges of Zoo Animal Welfare. In:
KLEIMAN,D.G., THOMPSON, K.V. and BAER, C.K., eds. Wild Mammals in Captivity:
Principles and Techniques for Zoo Management. 2nd ed. Chicago: University of
Chicago Press, 2010, pp.329-343.
KLEIMAN, D.G. et al. 1986. Conservation program for the Golden Lion Tamarin:
studies, educational strategies, and reintroduction. In: BENIRSCHKE, K. ed. Primates:
the Road to Self-Sustaining Populations. New York: Springer-Verlag, 1986, pp.959-
979.
KOOL, K.M. and CROFT, D.B., 1992. Estimators for home range areas of arboreal
colobine monkeys. Folia Primatologica. 58: pp.210-214.
KRAAIJEVELD, K. and KOMDEUR, 2003. Observations on the breeding biology of the
Seychelles Fody on Cousine Island. Ostrich. 74(1-2): pp.117-124.

68
KRAEMER G.W. and CLARKE, A.S., 1990. The behavioral neurobiology of self-
injurious behavior in rhesus monkeys. Progress in Neuro-Psychopharmacology and
Biological Psychiatry. 14(1): pp.141-168.
KRAUSE, J. and RUXTON, G.D., 2002. Living in Groups. Oxford: Oxford University
Press.
KREJČÍK, S., 2009. Foudia madagascariensis [online]. Available at:
<http://www.biolib.cz/en/image/id77106/> [Accessed 19 March 2013].
KWIAT KOWSKI, M.A. and SULLIVAN, B.K., 2002. Mating system structure and
population density in a polygynous lizard, Sauromalus obesus ( = ater). Behavioural
Ecology. 13(2): pp.201-208.
LATHAM, N. and MASON, G., 2010. Frustration and perseveration in stereotypic
captive animals: Is a taste of enrichment worse than none at all?. Behavioural Brain
Research. 211(1): pp.96-104.
LEWIS, M.H. et al., 2006. The Neurobiology of Stereotypy I: Environmental
Complexity. In: MASON, G. and RUSHEN, J., eds. Stereotypic Animal Behaviour:
Fundamentals and Applications to Welfare. UK: CABI, 2006, pp.190-214.
LINDST RŌM, K. and PAMPOULIE, C., 2005. Effects of resource holding potential and
resource value on tenure at nest sites in sand gobies. Behavioural Ecology. 16(1):
pp.70-74.
LOPRESTI-GOODMAN, S.M., KAMEKA, M. and DUBE, A., 2013. Stereotypical
Behaviours in Chimpanzees Rescued from the African Bushmeat and Pet Trade.
Behavioural Science. 3: pp.1-20.

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