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 partialfulfilment of the requirements for the degree of Bachelor of Science with

Honours in Zoo Biology

School of Animal, Rural & Environmental SciencesNottingham Road

SouthwellNottinghamshire

NG25 0QF

April 2013

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.

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

i

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 Behaviour in 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

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3.11.1 Statistical Analysis.............................................................................................................34

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.

1.1 Aims

1

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.

1.2.1 Null Hypothesis

2

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

4

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 part 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

with the limited space in the captive environment (Clubb and Mason, 2007; Rodden et

5

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 device 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.

2.1.2 Social Structure

6

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) results in a

psychological ‘scar’ that persists into later life; a theory supported by evidence that

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

7

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 may

be inhibited if the way an animal is housed prevents correct reproductive behaviour.

9

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.

11

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 large 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

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

13

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).

15

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.

2.5 Territoriality

16

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

reproductive resources such as nest sites (Aebischer et al., 1996; Lindstrōm and

Pampoulie, 2001) may also contribute. In these instances, territory may directly

17

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.

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,

18

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 studies

(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.

2.5.2 Methods of Calculating Territory Size

19

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 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.

20

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.

21

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

the enclosure off the path are difficult to access due to heavy vegetation and three

22

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.

3.3 Management Strategies

23

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.

24

Figure 3.1 Layout of the Kirindy Forest Aviary (not to scale)

25

Key

- Path

- Vegetation

- Sheds

- Ent / Exit

- Ponds

- Utility area

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 2cm2

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.

26

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

27

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.

28

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

29

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

30

recorded)

Not visible - Subject concealed or lost, generally in shed areas, awnings or thick vegetation

Table 3.2 Ethogram of observed behav7iours

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

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).

32

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.

3.10 Risk Assessment

33

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

3.11.1 Statistical Analysis

34

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.

38

Key

- B6481

- B6482

- B6804

- B6952

- No ID

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.

39

Key

- B6481

- B6804

- No ID

Figure 4.3 Distribution of subjects during Period 3 of the study. Symbols indicate instances where associated subject (see key) was

observed outside of normal distribution.

40

Key

- B6481

- B6804

- No ID

Figure 4.4 Distribution of subjects during Period 4 of the study. Symbols indicate instances where associated subject (see key) was

observed outside of normal distribution.

41

Key

- B6481

- B6804

- No ID

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.

Table 4.3 Total number of times each subject was recorded at each level of the

enclosure during the study

44

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

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

B64817 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

B68041 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 ID2 3 1 3 9 19 20 19 21 79 0 1 0 0 1 21 24 20 24 89

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

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

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

B680442 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 ID13 12 19 1 45 22 23 23 0 68 0 0 0 0 0 35 35 42 1 113

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

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

occurrence of rain (Brickell, 2006), the breeding season of captive UK populations

may commence at a different time to that of in situ populations.

53

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 held 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

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

54

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 space 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

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,

55

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.

5.4 Aggressive Behaviour

56

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.

5.5 Courtship Behaviour

57

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.

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

58

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.

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

59

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

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8.0 APPENDICES

8.1 Appendix 1 – Ethical Review Form

Ethical Review Form for Scientific Procedures

(Animals)

School of Animal, Rural and Environmental SciencesNottingham Trent University

1. Name of Applicant

Alexander Willey

2. Position (eg. BSc year 3 Zoo Biology)

BSc year 3 Zoo Biology (Placement)

3. Contact details (email/telephone)

Email – n0251186@ntu.ac.ukPhone – 07******37

4. Purpose of project application (eg. BSc year 3 dissertation, PhD thesis, or paper for publication)

BSc year 3 dissertation

5. Name of supervisor/member of staff responsible

Samantha Bremner-Harisson

6. Details of planned method/procedure, including explanation of how the 5 freedoms will be maintained. Use a separate sheet if necessary.

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The procedure will involve the daily observation of three adult males and two males coming into season of the Madagascar Fody species (Foudia madagascariensis) in a captive environment. The environment in question is a walk-through aviary owned by the Durrell Wildlife Conservation Trust, therefore the observation will be carried out within the aviary rather than from outside.

The fodies will observed for a set period of fifteen minutes each, with their behaviours and locations within the aviary recorded at intervals of thirty seconds. These details will be recorded on an ethogram.

In regards to the five freedoms:

1 – The fodies are provided with regular, species-appropriate food by the staff of the Durrell Wildlife Conservation Trust (DWCT). This diet has been developed to maintain the health of these individuals. The enclosure also contains a number of flowing ponds, therefore water is available whenever the individuals require

2 – The enclosure provided by the DWCT is filled with natural vegetation, including trees and shrubbery which provide a suitable shelter for these individuals, as well as a wide variety of comfortable perches throughout the area. The individuals are unrestricted in their movements around this enclosure

3 – The staff of the DWCT dedicate regular time to the maintenance of the enclosure and of the health of the individuals. The enclosure is kept to a suitable standard of sanitation and safety, and individuals are observed frequently for signs of illness or distress

4 – The individuals are unrestricted within the enclosure, and may move around as they please. The enclosure has been developed to replicate a natural habitat, and as such contains such naturalistic features as varied vegatation, ponds and rocks, allowing the demonstration of natural behaviours. Finally, the enclosure houses a large number of individuals, fodies and birds of other species, all of whom are free to interact as they desire

5 – The enclosure in question is a walk-through aviary for the public. The resident birds, therefore, are used to human presence to a certain extent. As long as humans remain on the path provided whilst in the aviary, the birds are not distressed. To ensure the birds are not distressed, the observations will be carried out from this path, using binoculars when required. Minimum noise will be made, and the birds will not be interfered with in any way. In the event of any disruption, the layout of the aviary allows any bird room to distance themselves from the disruption and provides numerous location suitable for concealment

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8.2 Appendix 2 – Risk Assessment

Task or Activity Description Location:

Kirindy Forest Aviary, Durrell Wildlife Conservation Trust

Observation of Madagascar Fodies within a naturalistic public walk-through aviary, containing numerous birds of numerous species, including passerines and waterfowl

Persons at Risk - Affected Groups:

A – Myself (the observer) B – Animals within the aviary

C – Visitors to the aviary D – Staff operating within aviary

Potential Hazard Existing Controls Risk level with controls

Additional Controls or Required Action &Date

Falling or tripping over enclosure features

Enclosure contains a well-maintained path for human use, which is kept clear of vegetation and debris

Low None

Drowning in ponds located within enclosure

All ponds are separated from the path by rope barriers at a distance of several feet

Low None

Tripping over / colliding with animals in enclosure

Most animals within the enclosure instinctively avoid close proximity to humans

Medium Care will be taken when moving throughout the enclosure

Colliding with other individuals in the enclosure

The path for human use is sufficiently wide for passing other people in all areas

Low None

Causing dangerous obstruction of work in the enclosure

All staff, including myself, are trained to operate safely and sensibly when carrying out maintenance

Low None

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Potential Hazard Existing Controls Risk level with controls

Additional Controls or Required Action &Date

Zoonotic disease from animal faeces

Numerous hand-washing facilities are available throughout the zooA good standard of hygiene is maintained in enclosures

Low Work clothing and footwear will be worn during observations, and frequently cleaned between observations

Eye strain from prolonged observation at a distance

Binoculars will be provided to minimise straining

Medium UV protective sunglasses will be worn in bright conditions

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