BINDURA UNIVERSITY OF SCIENCE EDUCATION DEPARTMENT OF

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BINDURA UNIVERSITY OF SCIENCE EDUCATION DEPARTMENT OF ENVIRONMENTAL SCIENCE RESPONSE OF BLUE DUIKERS TO UNFAMILIAR CONSPECIFICS’ FAECAL PELLET DEPOSITS OF VARIOUS AGES . REJOICE NYABASA (B0621730) A DISSERTATION SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS OF THE BACHELOR OF ENVIRONMENTAL SCIENCE HONOURS DEGREE IN WILDLIFE AND RANGELAND MANAGEMENT. (JANUARY 2010)

Transcript of BINDURA UNIVERSITY OF SCIENCE EDUCATION DEPARTMENT OF

BINDURA UNIVERSITY OF SCIENCE EDUCATION

DEPARTMENT OF ENVIRONMENTAL SCIENCE

RESPONSE OF BLUE DUIKERS TO UNFAMILIAR CONSPECIFICS’

FAECAL PELLET DEPOSITS OF VARIOUS AGES .

REJOICE NYABASA

(B0621730)

A DISSERTATION SUBMITTED IN PARTIAL FULFILLMENT OF THE

REQUIREMENTS OF THE BACHELOR OF ENVIRONMENTAL SCIENCE

HONOURS DEGREE IN WILDLIFE AND RANGELAND MANAGEMENT.

(JANUARY 2010)

i

DEDICATION

To my dad who loves animals.

ii

ACKNOWLEDGEMENTS

I would like to thank my industrial supervisor Miss N. Lunt, for all her guidance,

advice and support. Surely you are a superb tutor and I believe I am now equipped to

pursue any kind of research specifically in the field of wildlife conservation. To my

academic supervisors Mr J. Muvengwi and Mr V. Muposhi, I am ever grateful for

constant advice given throughout the attachment period. I would also like to express

gratitude to S. Williams and A. Mudungwe for knowledge and encouragement given

while I was at Marwell Zimbabwe Trust. The mention of Ms V. Bowman (MZT

director), Mrs. A Edwards (logistics manager) and Mr C. Davison (station manager) is

inevitable for their boundless support during my stay at MZT.

A warm thanks to my friend, Mr I. Mavhurere, for inspiring me and making my ten

months stay at DAMBARI Field Station joyous and worthwhile. I will always be

grateful for valuable academic discussions we did together. To my colleague, Miss N.

Sithole, it was my greatest pleasure to work and stay with you during the attachment

period. I also wish to sincerely thank the general staff at Marwell Zimbabwe Trust, I

am ever grateful that I had an opportunity to work with you.

My special and sincere thanks proceed to my family for standing as a pillar of

strength and motivation in all aspects of my life. With your support, surely the sky is

my limit!

Forever, I thank God the Almighty for everything.

Nyabasa Rejoice.

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ABSTRACT

Blue duikers (Cephalophus monticola) are territorial animals and both males and

females mark out their territory continually using urine, faecal pellets and preorbital

gland secretions. The aim of the research was to assess the behavioural response of

blue duiker to unfamiliar conspecific’s faecal pellet deposits of varying ages in its

territory. To determine the intensity and type of blue duikers’ responses to foreign

conspecifics’ faecal pellets deposits, the following parameters were recorded for each

recipient: dung age, recipient sex and donor sex. Captive adult blue duikers kept in a

semi-natural environment at DAMBARI Field Station were used for the research

experimental observations. Experimental treatments consisted of dung piles from

donor animals that had aged outdoors to 1 hr, 10 hrs, 34 hrs, and 58 hrs. Two types of

controls were used: one comprised stones (“dummy” dung) similar in appearance to

faecal pellets while the second involved faecal piles from the recipient enclosure that

were aged in the same manner as donor faeces and reintroduced to the same

enclosure. Faecal pellets from donor animal enclosures were aged in the same natural

environment and later deposited in recipient pens for observations. To determine how

strongly (i. e. intensity) and in what ways (i. e. type) blue duikers responded to

foreign conspecifics' faecal pellets the following measurements and observations were

recorded for each recipient: response lag time (minutes), number of times the deposit

was approached, time spent investigating introduced pellets (seconds), type of

response exhibited and frequency of scent marking. The main type of response to

foreign faecal pellets deposits was sniffing and preorbital scent marking. Consistent

trends were detected in response intensity whereby fresh to intermediate dung were

investigated more than old dung. The results indicate that dung scent retains a high

degree of efficacy for some time (up to 34 hours) and afterward the scent signal starts

to fade out. A slight increase was noted when recipient animals investigated dung

from the same sex donor animals, most probably due to perceived threat to mates. The

results from this study suggest that the interaction of donor sex, dung age and

recipient sex had a significant effect on the intensity of response by blue duiker

recipient animal.

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TABLE OF CONTENTS

DEDICATION ......................................................................................................................................... I

ACKNOWLEDGEMENTS ................................................................................................................... II

ABSTRACT .......................................................................................................................................... III

TABLE OF CONTENTS ..................................................................................................................... IV

LIST OF FIGURES .............................................................................................................................. VI

LIST OF TABLES .............................................................................................................................. VII

LIST OF ACRONYMS AND ABBREVIATIONS ........................................................................... VIII

APPENDICES ...................................................................................................................................... IX CHAPTER 1: INTRODUCTION........................................................................................................... 1

1.0 BACKGROUND TO THE STUDY .......................................................................................... 1 1.1 PROBLEM STATEMENT .......................................................................................................... 1 1.2 JUSTIFICATION ...................................................................................................................... 2 1.3 AIM ....................................................................................................................................... 2 1.4 OBJECTIVES .......................................................................................................................... 2 1.5 RESEARCH HYPOTHESES ....................................................................................................... 3

CHAPTER 2: LITERATURE REVIEW ............................................................................................... 4

2.0 INTRODUCTION..................................................................................................................... 4 2.1 INTERNATIONAL AND LOCAL DISTRIBUTION OF DUIKERS ..................................................... 4 2.2 IMPORTANCE OF DUIKERS ..................................................................................................... 5 2.3 THE CONSERVATION STATUS OF DUIKERS ............................................................................ 6 2.4 ECOLOGY AND BIOLOGY ASPECTS OF BLUE DUIKERS ......................................................... 10

CHAPTER 3: RESEARCH METHODOLOGY.................................................................................. 17

3.0 STUDY AREA ........................................................................................................................ 17 3.1 RESEARCH METHOD ............................................................................................................ 17 3.2 DATA ANALYSIS................................................................................................................... 19 3.3 LIMITATIONS OF THE STUDY ............................................................................................... 19

CHAPTER 4: RESULTS ...................................................................................................................... 21

4.0 TYPE OF RESPONSE............................................................................................................ 21 4.1 RESPONSE OF RECIPIENT ANIMALS TO DONOR ANIMAL DUNG ........................................... 23 4.2 THE EFFECT OF DUNG AGE, RECIPIENT SEX AND DONOR SEX TO RECIPIENT ANIMALS’ RESPONSE ......................................................................................................................................... 23 4.3 TRENDS IN BLUE DUIKER RESPONSE (LAG) TIME TO INTRODUCED TREATMENTS............... 25 4.4 TIME SPENT BY BLUE DUIKERS INVESTIGATING INTRODUCED TREATMENTS ..................... 26 4.5 PREORBITAL SCENT MARKING BY BLUE DUIKERS TO INTRODUCED TREATMENTS IN THEIR TERRITORIES .......................................................................................................................... 26 4.6 INVESTIGATION FREQUENCY BY BLUE DUIKERS TO TREATMENTS INTRODUCED INTO THEIR TERRITORIES .......................................................................................................................... 28

5.0 DISCUSSION .......................................................................................................................... 29

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CHAPTER 6: CONCLUSION AND RECOMMENDATIONS .......................................................... 31

6.0 CONCLUSION ....................................................................................................................... 31 6.1 RECOMMENDATIONS ........................................................................................................... 32

REFERENCES ..................................................................................................................................... 33

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LIST OF FIGURES FIGURE 1: BLUE DUIKER URINATION SITE WITHIN ENCLOSURE ................................................................... 21 FIGURE 2: BLUE DUIKER DEFECATION SITE ADJACENT TO NEIGHBORING ENCLOSURE ............................ 22 FIGURE 3: MARKS OF PREORBITAL GLAND SECRETION LEFT BY A TERRITORIAL BLUE DUIKER ON THE

POLE. ............................................................................................................................................. 22 FIGURE 4: TRENDS IN MALE RECIPIENTS RESPONSE TIMES TO INTRODUCED TREATMENTS ........................... 25 FIGURE 5: TRENDS IN FEMALE RECIPIENTS RESPONSE TIMES TO INTRODUCED TREATMENTS...................... 25 FIGURE 6: MEAN TIME TAKEN BY MALE RECIPIENTS SNIFFING INTRODUCED TREATMENTS .......................... 26 FIGURE 7: MEAN TIME TAKEN BY FEMALE RECIPIENTS SNIFFING INTRODUCED TREATMENTS ...................... 26 FIGURE 8: TRENDS IN PREORBITAL SCENT MARKING BY MALE RECIPIENTS ................................................. 27 FIGURE 9: TRENDS IN PREORBITAL SCENT MARKING BY FEMALE RECIPIENTS .............................................. 27 FIGURE 10: FREQUENCY OF INVESTIGATION BY MALE RECIPIENTS TO INTRODUCED TREATMENTS ............... 28 FIGURE 11: FREQUENCY OF INVESTIGATION BY FEMALE RECIPIENTS TO INTRODUCED TREATMENTS............ 28

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LIST OF TABLES TABLE 1: SUMMARY OF THE CONSERVATION STATUS OF SUB-SAHARAN AFRICAN ANTELOPE ...................... 6 TABLE 2: SOURCES OF VARIATION FOR TOTAL SNIFF DURATION SHOWING THAT MALE RECIPIENTS

SNIFFED FOR LONGER DURATION 10HR FEMALE DUNG COMPARED TO 1HR, 34HR, AND 58HR AGED DUNG FROM CONTROL, MALE AND FEMALE DONORS......................................................................... 24

TABLE 3: SOURCES OF VARIATION FOR INVESTIGATION FREQUENCY. MALE RECIPIENTS INVESTIGATED MORE 10HR FEMALE DUNG COMPARED TO 1HR AND 58HR AGED DUNG FROM CONTROL AND FEMALE DONOR. ............................................................................................................................. 24

TABLE 4: SOURCES OF VARIATION FOR LAG TIME RESPONSE VARIABLE SHOWING THAT FEMALE RECIPIENTS TOOK LONGER TIME TO START INVESTIGATING INTRODUCED 58HR FEMALE DUNG COMPARED TO 1HR AGED DUNG. ..................................................................................................... 25

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LIST OF ACRONYMS AND ABBREVIATIONS

CAMPFIRE Communal Areas Management Programme For Indigenous Resources

DAMBARI Duikers and Mini-antelope Breeding and Research Institute

MZT Marwell Zimbabwe Trust

ODA Overseas Development Administration

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APPENDICES APPENDIX 1: SAMPLE DATA RECORD SHEET ............................................................................................ 39

1

CHAPTER 1: INTRODUCTION

1.0 Background to the Study

Scent marking is a common form of signaling in mammals and it usually occurs in

complex spatial patterns, often involving several marks that are regularly replenished

in marking and remarking sessions by the signaler. According to Ralls (1971), most

mammals have a highly developed olfactory sense, which they employ in social

communication using signals originating in urine, faeces or scent glands. Several

territorial mammals apply scented secretions and excretions to areas or objects in their

environment as a way of demarcating their territories or home ranges (ibid).

Scent marks are a form of status signal, advertising territory ownership, and recent

evidence suggests that they may also be condition-dependent signals of quality and

competitive ability (Gosling et al., in press). A unique characteristic of scent marks as

a social signal is that receivers often encounter the signal in the absence of the

signaler. Studies which have examined the distribution of scent marks within a

territory have usually found that they are either clustered at the territory boundary or

near its centre (Roberts, 1992).

Blue duikers are the smallest antelopes found in the southern African subregion

(Smithers, 1983). The average height of adult males is generally 35.5 cm with a mass

of about 4, 5 kg, while females are slightly larger with an average height of 36.2 cm

with a mass of 5.4 kg (Best and Best, 1977). Blue duikers are territorial (Bowland and

Perrin, 1995) and not easily driven from their territories. Direct interactions between

neighbours are rare, but territorial occupancy is continually advertised by visual,

olfactory and sometimes by auditory means (Estes, 1997). They scent mark with a

pair of preorbital glands, urine or faecal pellets to demarcate their territory boundary

and pedal glands mark their movements in their forest habitats (Smithers, 1983).

1.1 Problem Statement

Scent marking is a widespread but poorly understood form of communication in

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duikers (Johnson, 1973; Stoddart, 1980; Brown and Macdonald, 1985). In their

natural environment, blue duikers’ territorial behaviour is very difficult to observe and

study in detail due to the reason that they are shy and secretive (Wilson, 2005). In

comparison to other aspects of scent marking and territoriality in mammals, the

strength of behavioural response of blue duikers to the introduction of unfamiliar

conspecifics’ faecal pellet deposits of varying ages has received relatively little

attention.

1.2 Justification

Little has been done to investigate on the response of blue duikers to the introduction

of unfamiliar scent of varying ages, therefore this study will enable a better

understanding of this species’ territorial behaviour. The study will also facilitate the

provision of detailed information on blue duiker territorial behaviour for present and

future academic purposes especially in wildlife related research work. Territoriality is

one of the most important behavioural traits affecting the spatial organization of

animal populations (Mizutani, 1993). As a result, successful re-introductions of blue

duiker species require the general knowledge of the animal’s territorial behaviour so

as to be able to facilitate the maintenance of social organization, space use patterns,

resource utilization and also reduce energy demanding direct encounters among

animals of the same species. The availability of captive blue duikers in semi-natural

enclosures at DAMBARI Field Station helped in providing a conducive environment

for clear observations of individual animal territorial behaviour thus accurate results

and conclusions can be deduced.

1.3 Aim

To determine the responses of blue duikers to unfamiliar conspecifics’ faecal pellets

in their territories.

1.4 Objectives

i) To determine how intensity and type of response by recipient animals varies with

sex of donor, dung age and sex of recipient.

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1.5 Research Hypotheses

H0: The intensity and type of response by recipients do not vary with sex of donor.

H0: The intensity and type of response by blue duikers do not vary with sex of

recipient.

H0: The intensity and type of response by recipients do not vary with dung age.

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CHAPTER 2: LITERATURE REVIEW

2.0 Introduction

The life history of duikers is relatively unstudied in natural conditions due to the fact

that they are secretive and shy thus difficult to observe for long periods in forest

environments (Newing, 1994). However, they follow the general patterns of forest

animals including small body size, short horns, frugivorous diet and small group size

(Dubost, 1983). According to Estes (1997), observations of captive blue duikers

maintained under semi-natural conditions confirm that pair bonds are probably life

long.

All duikers defend their territories against intruders thus they are classified under

territorial animals (Kingdon, 1997). They scent mark with a pair of preorbital glands,

urine or faecal pellets to demarcate their territory boundary and pedal glands mark

their movements in their forest habitats (Smithers, 1983). Usually there is no joint or

family defense of the territory, but each resident adult and subudult selectively

excludes interlopers of its own sex (Estes, 1997).

This chapter outlines the ecological aspects of duikers including habitat preference,

social organizations and also feeding, breeding and territorial behaviors. It also gives

an overview of the general distribution of duikers internationally and locally and also

the ecological, economic and social importance of duikers. The conservation status of

duikers in Africa is also summarized incorporating the present survival threats and the

conservation strategies in use to reduce continuing decline of duiker populations.

2.1 International and Local Distribution of Duikers

There are 16 species of duikers found in Africa (Wilson, 2005). Nearly every African

forest and woodland is occupied by at least one species of duiker (Kingdon, 1997) and

the majority of duiker species live in the Equatorial Rain Forest (Estes, 1997). The

blue duiker has a very wide distribution in Africa south of the Sahara desert with its

overall range extending from the east bank of the Cross River in Nigeria stretching

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through Central Africa to the coastal areas of the Cape Province of South Africa

(Wilson, 2005). In Zimbabwe, blue duikers are confined to the evergreen forests in

the eastern highlands specifically areas like Chirinda forest, Chipete forest, Vumba

Botanical Garden, Bunga forest, Bridal Veil Falls forest and also Nyanga and Honde

valley (Wilson, 2005). However, the largest population of blue duiker in Zimbabwe

occurs in the Chirinda evergreen forest on Mt. Selinda (ibid).

2.2 Importance of Duikers

Conservationists have long recognized the necessity of examining not only the

ecological but also the economic and social importance of wildlife to communities (Fa

et al., 1995; Fitzgibbon et al., 1995). Duikers are generally shy and secretive animals

thus difficult to study in their natural environment. Due to this reason, they have been

assigned minor roles in many ecological studies or even ignored. However, they play

important roles in many ecosystems including structuring understorey vegetation,

recycling nutrients, seed dispersal and forming a major part of the prey base for a

wide range of predators (N. Lunt, pers. comm.).

Ecologically, duikers play an important role in the diet of certain tropical forest

predator species including leopard (Panthera pardus) and golden cat (Felis aurata)

(Hart et al., 1996). The threat to Africa’s predator populations caused by dramatic

declines in their prey base is likely to be significant without duikers (ibid).

Duikers are primarily frugivorous although they also browse selectively and

occasionally take animal matter (Feer, 1995). About 150 different species of fruit

were found among the rumen contents of forest duikers, which suggest that they may

play an important role as seed dispersal agents (ibid).

Rural communities are highly dependent on forest resources for both nutritional and

economic subsistence (Lahm, 1993; Noss, 1995). Traditionally, duikers are hunted by

humans for consumption as a primary source of protein. The majority of bushmeat

studies conducted to date involve market surveys, many of which indicate that duikers

comprise the majority (40-80%) of meat sold in bushmeat markets throughout Central

Africa (Chardonnet and Fritz, 1995; Bowen–Jones, 1998).

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2.3 The Conservation Status of Duikers

Despite their widespread range, nearly all species of duiker are currently projected to

decline considerably in the near future while several species are currently considered

on the point of extinction if immediate protection measures are not put into effect

(East and IUCN/SSC/ASG, 1999).

The antelope data presented in Table 2.1 is completely taken from the African

Antelope Database 1998 (AAD 1998) compiled by Rod East of the IUCN/SSC

Antelope Specialist Group. However, the information given in AAD 1998 is not

complete due to lack of detailed information available from many parts of Africa. The

term threatened in this case is used specifically to refer to those species classified as

vulnerable, endangered or critically endangered (ibid).

According to East and IUCN/SSC/ASG (1999), the percentage of small antelope that

are currently threatened is approximately the same as that for large antelope. However

within the next 15 years the percentage of small antelope forms that will be threatened

will more than double, whereas other threatened antelope will only increase by 38%.

Small antelope will become more threatened than the average African antelope.

Perhaps the most worrying of all is that an astounding 92% of small antelope are

currently undergoing population declines or show a tendency to decline compared to

66% of other antelope.

Table 1: Summary of the conservation status of sub-Saharan African antelope

Small Antelope Other Antelope Total Antelope

Threatened 10(27%) 23(32%) 33(30%)

Threatened by 2025 22(59%) 37(51%) 59(54%)

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Decreasing or showing a

tendency to decrease

34(92%) 47(66%) 81(74%)

Source: African Antelope Database, 1998

Eves (1996) states that one possible reason for the difference in population trend is

that the majority of other antelope are larger savannah species. These species have

already experienced dramatic declines from historical population sizes and are now

stabilizing at reasonably low levels in protected areas and areas that still have low

human densities. In contrast a large number of small antelope are forest dwellers. By

virtue of their habitat, these species have been less exposed to widespread human

influences until more recent times when human population continued to increase

(Bowen-Jones, 1998).

In Africa, the blue duiker is still widespread and common in spite of heavy hunting

pressure all over its range and is still regarded as non-endangered species (Kingdon,

1997). However, in Zimbabwe blue duikers are in danger of extinction and unlikely to

survive if the present population decline contributing factors continue operating

(Wilson, 2005).

2.3.1 Survival Threats of Duikers

Many populations of duikers are at risk of local or ecological extinction due to

excessively high off take rates and unnecessary wastage (Hart, 2000), whereby some

of the animals caught in snares are lost to scavengers and decomposition and some

escape with dreadful injuries. The major threats facing most of these species are bush

meat hunting and habitat loss or degradation (ibid). Both problems are complex and

cannot be solved easily and will require considerable funding, research and education

initiatives.

2.3.2 Commercial and Traditional Hunting for Bush Meat

Hunting of duikers for traditional markets has been undertaken for considerable time

and is economically, culturally and socially important to local communities (Eves,

1996). Due to demographic changes and better road network between the source and

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potential markets, hunting for commercial markets in towns and cities has also

increased dramatically in recent years. It is possible to hunt for these species at a

sustainable level but much more data are needed on most species to be able to

determine what the sustainable yields may be (Hart, 2000). There will be need for

education and capacity building to convince hunters that it is for their own advantage

to abide by quotas, low wastage hunting methods and proper policing and protection

to monitor hunting effort (ibid).

2.3.3 Habitat Loss and Degradation

In some areas particularly in East Africa, habitat loss or degradation may be an equal

or even greater threat to duikers than hunting (Masoud, 2000). This is largely due to

clearance for agriculture and commercial wood harvesting. Many duiker populations

are still outside of protected areas and are therefore at high risk of increased habitat

loss as human populations continue to expand in the future (Bowen-Jones, 1998).

2. 3. 2 Conservation Strategies for Small Antelopes

The survival of most species of duiker in the future significantly depends on increased

conservation efforts presently and in the coming years. Masoud (2000) states that

participation by local and international stakeholders is greatly helpful in allowing

successful implementation of conservation measures.

2.3.4 Law Enforcement

In the past, law enforcement was the major conservation tool whereby governments

would allocate enough resources to employ as many guards as required to deter

people from hunting (Masoud, 2000). However, changes in socio-economic

conditions of people, population pressure and lack of adequate financial resources

have created a more difficult environment for the governments to rely on law

enforcement alone (ibid.). Therefore, this strategy is often implemented through

community participation programmes like CAMPFIRE and ODA.

2.3.5 Community Management Programmes

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The community wildlife programmes started as a result of human-environment

interaction work done through research (Masoud, 2000). During this period, local

people expressed their desire to be involved in the management of natural resources

especially antelope since they believe that they are an important alternative protein

source. Community participation allow locals to plan and set aside their forestlands

for various purposes to include conservation and sustainable exploitation areas and in

return they gain revenue from hunting fees to support conservation and development

work in respective villages.

A case study of Zanzibar reveals a successful community management programme

whereby a new Forestry legislation was passed. It empowers the community to enter

into agreement with the Forest Administrator to manage their wildlife resources on

presentation of a management plan and by-laws. This has been done very successfully

with 12 village communities on southern Unguja, whereby through technical backup

from national and international staff, the by-laws where prepared approved and are

currently in force (ibid).

2.3.6 Translocation

Western and Pearl (1989) states that translocation is believed to be one of the

effective ways of saving a population from complete extinction within its bio-

geographical region. Translocations have been done in Zanzibar as rescue

programmes. Aders’ duikers were first translocated into Funzi Island in Pemba but the

population became locally extinct due to uncontrolled hunting. The second

translocation was a community managed initiative where suni antelope were first

introduced into Juja islet at Ukongoroni, Zanzibar. The community, except for socio-

cultural functions involving the entire community, strictly forbids hunting. This

translocation has been successful and the population of suni still exists at Juja. This

technique is common in New Zealand and has been in use with greater success (ibid).

2.3.7 Captive Breeding Programme

Captive breeding can be an important strategy to prevent species from extinction and

has grown enormously in recent years (Snyder et al., 1996). However, there are

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numerous drawbacks such as high costs involved in establishing conducive captive

breeding facilities and loss of heterozygosis in poorly managed captive populations

(Masoud, 2000).

2.4 Ecology and Biology Aspects of Blue Duikers

2.4.1 Habitat Preferences

Duikers generally flourish in a wide range of habitats in which requirements prescribe

an obligate balance between food resources and shelter (Bowland and Perrin, 1995),

of which a deficiency of either reduces habitat quality. Blue duiker habitats include

lowland and montane rainforests and also riverine and moist thickets (Kingdon,

1997). They mostly occupy areas with comparatively little undergrowth to minimise

obstruction of its movements or vision (Estes, 1997).

2.4.2 Activity Periods and Social Organization

All duikers are thought to be sedentary and territorial (Estes, 1997). Some species are

solitary, some live in pairs and others are occasionally found in groups with one adult

male and a number of adult females (Feer, 1989). At any rate, pairs of blue and

Maxwell’s duikers (Cephalophus maxwellii) apparently associate more closely than

most other duikers because they are more vulnerable to predators than larger species

that live in denser cover in more spacious home ranges. Blue and Maxwell’s duikers

are known to live in monogamous pairs within small territories that are jointly marked

and defended against foreign members of the same sex (Estes, 1997). In blue duikers

pair bonding is reinforced mainly by pressing the preorbital glands together and social

coat licking.

Activity periods of duikers vary with species. Some species such as the blue duiker

and the Maxwell’s duiker are active during the day (Newing, 1994). Others such as

the bay duiker (Cephalophus dorsalis) are nocturnal and the larger species like

yellow-backed duiker (Cephalophus sylvicultor), Abbott’s duiker (Cephalophus

spadix) and Jentink’s duiker (Cephalophus jentinki) are active both by day and by

night (ibid).

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2.4.3 Diet Selection

The common accepted view of duikers is that they are high concentrate feeders

(Hofmann, 1973 and Wilson, 2005). The requirement for easily assimilated and

highly nutritious foods is linked with the high metabolic demands of a small body

(Bell, 1971; Milton, 1982; Van Soest, 1983). The Jarman-Bell principle illustrates that

small mammalian ruminants feed on small, energy-rich, dispersed food sources at

relatively low density and low in plant fibre (Jarman, 1974; Spinage, 1986;

McNaughton, 1987).

Blue duikers are primarily frugivorous but also browse selectively (i.e. leaves, shoots,

and buds) and occasionally take animal matter in the form of insects (Bodmer, 1990;

Hanekom and Wilson, 1991). In areas where fruiting trees and shrubs are abundant, it

was found that upto 80% of blue duiker diet may be fruits with little of foliage,

flowers, gum and animal (Kingdon, 1997). Preferred fruits are the abundant small

berries or seeds that are often typical of dominant trees in secondary or colonizing

forest (ibid). Blue duikers have been seen to follow below monkeys and birds to pick

up fallen fruits. Probably duikers gain enough water from their food and rarely have

to drink, but are occasionally seen licking rainwater off foliage (Estes, 1997).

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2.4.4 Breeding

According to Estes (1997), all African bovids bear one young and breed at least once

a year (small species often twice), giving birth to well developed young after gestation

periods of about six months (small antelopes and gazelles) to eight or nine months in

medium and large species. Wilson (2005), states that female blue duiker age of sexual

maturity ranges from eight to twelve months whilst for males ranges from eleven to

fourteen months. A female’s maximum lifetime offspring production has been found

to be about 14 young ones for both small and large species. Most species have

extended but definite breeding seasons during the rainy period of the year and in

equatorial regions with two rainy seasons, many have a secondary calving peak

(Estes, 1997). However, records at DAMBARI Field Station indicate year round

breeding in blue duikers with a slight peak in the wet season (N. Lunt, unpub. data).

2.4.5 Scent Marking and Territoriality

Scent marking is a form of animal communication which is common in territorial

species (Roberts, 1997). It refers to the application of scented secretions and

excretions by an animal to areas or objects in its environment (Wells and Bekoff,

1981). A territory is a specific area within a home range in which an animal is

aggressive and supremely dominant with respect to certain categories of intruder

(Anderson, 1983).

2.4.6 Types of Scent Marks

Scent marks occur in complex patterns often involving several marks that are

regularly replenished in active marking and remarking sessions (Gosling and McKay,

1990). Many species demarcate territories using chemical signals originating from

urine, faeces and cutaneous scent glands (Ralls, 1971).

Several mammal species including antelopes use faeces for territory demarcation (Asa

et al, 1985). Faeces serve as substrate for glandular secretions and as visual

‘signposts’ to intruders. The use of faeces may be ideal for scent marking because

they have a minimal energetic cost to the signaler (Gosling, 1981) since they are

waste from metabolism. However, limited food sources may affect faecal pellets

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production thus limiting marking rates in species that use them to advertise territory

borders (Komers, 1996).

The use of gland secretions is also common with many animal species whereby

different scent glands secretions are used through varying marking methods. Many

duiker species scent mark using secretion from preorbital gland located just below

each eye (Ralls, 1971). Both males and females frequently rub the gland on objects in

their territory leaving a small, visible deposit of secretion (Roberts, 1994). Some

mammals like rabbits have two main scent gland secretions which are associated with

social dominance and territoriality (Ralls, 1971). These are submandibular or chin

gland secretion and anal gland secretion which is usually produced together with

faeces (ibid).

Urine serves a variety of functions besides territory demarcation, depending on the

species and the environment in which it is expressed (Brown, 1985). Bowyer and

Kitchen (1987) mentioned that scent urination is believed to contribute in information

transmission whereby it could act as a visual display and also as an olfactory signal.

Scent urination is commonly performed by males whereby they urinate onto

themselves and others urinate onto the ground and sometimes then wallow in the

urine.

2.4.7 Significance of Scent Marks

Scent marking may serve as a mechanism for advertising territory ownership and also

serve as a challenge ritual to potential intruders thus reducing the frequency of

fighting over territorial rights (Roberts, 1997). Scent marks provide an opportunity for

intruders to identify resource holders and then to modify their behaviour in a way that

takes account of the benefits of utilizing the defended resource and the costs of

meeting the resource holder (Gosling, 1990). A receiver could examine deposited

scent marks and it would be able to determine if the opponent was of high rank and,

depending on its own relative competitive ability and the value of the marked

resource, it could withdraw from the encounter at an appropriate stage (Gosling,

1982). As a result, resource holders can avoid or reduce the costs of contests with a

large proportion of opponents (ibid).

14

Scent marks also contribute in sex recognition and also serve as an indicator of sexual

condition and maturity thus helping in mate selection (Wells and Bekoff, 1981).

During the breeding season, many male ungulates use urine as a medium to transmit

information to conspecifics (ibid). Female white- tailed deer investigate and scent

mark at male marking sites when they are in estrus suggesting that males use marking

sites to assess potential mates (Sawyer et al., 1989). Female mice also use male mice

odor originating from urine to distinguish parasitized and unparasitized males thus

avoiding mating with infected males (Gosling, 1985).

Scent marks also help in providing internal information to orient members of the same

resident group (Wells and Bekoff, 1981). Estes (1997) states that scent trails left by

the pedal glands and the preorbital gland secretion left on trees probably help family

members keep track of one another. Blue duikers also perform mutual gland pressing

whereby mate pairs scent mark each other or their offspring on projections such as the

forehead and tail base as a social behaviour (ibid).

2.4.8 Temporal Variation in Scent Mark Efficacy

Scent marks are placed in the environment often in the absence of a potential receiver

and only detected some time later often when the signaler is no longer present

(Gosling and Roberts, 2000). The two crucial parameters of temporal variation for

signalers are the rise time, which is the interval between deposition and achievement

of maximal detection and the mark’s persistence time which is interval between

deposition and the time when the mark can no longer be detected (Ferkin et al., 1995).

The efficacy of animal scent marks is not constant but varies with time after

deposition (Alberts 1992). Recipient animals use the relative proportions of volatile

sub-components of the scent to estimate the age of the scent mark (Johnston and

Schmidt 1979; Apps et al. 1989; Ferkin et al. 1995). Information about the age of

scent marks may allow receivers to modulate their response to scent encountered in

proportion to the perceived likelihood of encountering the signaler (Roberts, 1998).

Owners encountering foreign scent marks in their territory may take evasive action if

the scent is very fresh compared to old scent perhaps to avoid physical conflict since

the probability that the signaler will be in the vicinity is high (Maynard Smith, 1974).

15

It can be expected to over mark its area with own scent to maximise scent matching

when the intruder assess the area again (Gosling, 1985).

A receiver can examine deposited scent marks and it will be able to determine if the

opponent was of high rank (i.e. physical and sexual condition) and, depending on its

own relative competitive ability and the value of the marked resource, it can withdraw

from the encounter at an appropriate stage (Gosling, 1982). Several studies conducted

before revealed that when receivers approach scent marks, they often sniff and lick

the scent deposit (Brown and MacDonald, 1985; Idris, 1994; Roberts, 1998) or they

over mark on or around the deposited marks (Melemis and Falls, 1982).

2.4.9 Territoriality in Blue Duikers

Territoriality is one of the most important behavioural traits affecting the spatial

organization of animal populations (Mizutani, 1993). Particularly in blue duikers,

direct interactions between neighbours are rare thus territorial occupancy is

continually advertised by visual, olfactory and probably also by auditory means

(Estes, 1997). Odendaal and Bigalke (1979) indicated that the blue duiker is strongly

territorial and both males and females mark out their home range continuously using

urine, faecal pellets and preorbital gland secretion.

Rubbing the preorbital glands on tree trunks and other objects, wherever the duiker

goes, is the main form of demarcation (Estes, 1997). The scent oozes from glands just

below their eyes. The preorbital gland is equally productive in both sexes and the

secretion is an inhomogeneous mixture of a clear, colourless liquid and white mucous

material which in females, tends to have a bluish-grey colour (Kingdon, 1997).

According to Estes (1997), the other form of territorial demarcation is tree horning, in

which both sexes engage from the time their horns emerge. Permanent, visible marks

are made on target trees, chiefly young saplings with stems small enough to fit into

the fork between the horns. The duikers always horn at shoulder level after first

sniffing and licking and sometimes scraping the spot with the incisors. Audible alarm

signals may also serve as an auditory reminder that a territory is occupied (ibid).

17

CHAPTER 3: RESEARCH METHODOLOGY

3.0 Study Area

DAMBARI Field Station is a research institution situated 25 km along Bulawayo-

Gwanda road southeast of Bulawayo, Zimbabwe. The station property covers a total

area of 25 ha, situated at an altitude of 1050 m. The area receives an annual average

rainfall of 600 mm, which normally falls between November and March. The

vegetation type is mainly Acacia woodland interspersed with shrubs. The land slopes

down to a small seasonal river called Umguza (Bowman & Plowman, 2002), and the

vlei area near the river is dominated by Hyparrhenia species.

3.1 Research Method

Captive blue duikers (Cephalophus monticola) were observed at DAMBARI Field

Station between November 2008 and April 2009. Eight adult blue duikers (six

females and two males) in their prime were maintained either singly or in pairs in six

150 m2 fenced, semi-natural pens that are arranged in a hexagonal pattern around a

double-storey observation platform. Minimization of interactions between animals in

adjacent enclosures was facilitated by the presence of black opaque plastic sheeting

attached to fences to a height of 1 m.

Donor animals were maintained singly in pens located at least 40 m from the pens of

recipient animals. They were composed of four adult blue duikers (two females and

two males) in their prime, which where also kept in a similar semi-natural

environment to recipient animals. Faecal pellets were collected when fresh, and aged

in the same natural environment or immediately deposited in recipient pens,

depending on the treatment requirement. Clean polythene bags were used to collect

faeces, to prevent scent contamination by the researcher.

Experimental treatments consisted of dung piles from donor animals that had aged

outdoors to 0-1 hr, 9-10 hr, 33-34 hr or 57-58 hr. Two types of controls were used:

one comprised stones (“dummy” dung) similar in appearance to faecal pellets while

the second involved faecal piles from the recipient enclosure that were aged in the

18

same manner as donor faeces, and reintroduced to the same enclosure. The dummy

dung control, which was scent-neutral, was used to determine if animals used visual

cue to identify unfamiliar objects in their pens.

Treatments were given in a randomized fashion in a repeated measures design such

that each enclosure received each treatment once. An interval of at least 24 hours was

allowed before introducing a different treatment to the same enclosure to ensure that

there was no temporal autocorrelation in recipient responses.

Treatments were introduced to experimental enclosures at 1630 hrs, deposited at a site

away from the animals’ usual latrine site/s but readily visible to the observer.

Observation periods were restricted to times when duikers were most active (Bowman

and Plowman 2002; pers. obs.), i.e. from 1630 hrs to 2000 hrs. During observation

periods, territory-defence related activities occurring in one pen were observed and

recorded.

3.1.1 Territorial Behaviour Assessment

The number of times a recipient individual approached an introduced unfamiliar

conspecifics’ faecal deposit was recorded, with the type of response stated.

Responses consisted of investigation (sniffing) and pre-orbital scent marking within

the range of 1 m from the introduced deposit. To determine how strongly (i. e.

intensity) and in what ways (i. e. type) blue duikers responded to unfamiliar

conspecifics’ faecal pellets, the following measurements and observations were

recorded for each individual:

1) Response lag time (number of minutes from faecal introduction to first

investigation).

2) Type of response exhibited (for example approach and sniff; approach and

scent-mark).

3) Time (in seconds) spent investigating faecal pellets on each occasion the pile

was approached.

4) Frequency of scent marking on each occasion the pile was approached.

Other important factors like name of donor and recipient animal, weather conditions,

19

starting and ending time of observations were also recorded.

3.2 Data Analysis

Data collected were entered into Microsoft Excel (Microsoft Corporation, 2003)

spreadsheets and pivot tables were used to summarize data. Data from the pivot table

were then analyzed using Minitab 15 (Minitab Inc., 2006). Response variables were

(i) lag time, (ii) total sniff duration (sum of seconds spent sniffing an introduced

deposit), (iii) mean sniff duration (mean number of seconds spent sniffing a deposit)

and (iv) scent-mark frequency (total number of occasions that pre-orbital scent-marks

were applied to structures within 1 m of the introduced deposit). Predictor variables

were dung age, recipient sex and donor sex.

To assess the validity of using ANOVA for analysis, Levene’s test and normal

probability distributions were run for each response variable against predictors. Data

were non-normal (Anderson-Darling tests, p < 0.05), but variances were not

significantly different (Levene’s test, p > 0.05 for all tests). To be conservative,

permutational multivariate analysis (which is not affected by non-normal data) was

used to test the effects of donor sex, recipient sex and dung age on each response

variable with DISTLM (Anderson, 1999). To ensure that responses to donors were

random, Sign tests were run for each response. Line plots of responses against dung

age for each donor/recipient group were run using SigmaPlot 8.0 (Brannan et al.,

2002).

3.3 Limitations of the Study

Variation in weather conditions could have affected the results when experiments

were conducted during bad weather conditions (i. e. cold and wet). During cold and

wet weather, recipients tend to become inactive, so the intensity of response to

introduced faecal deposits is expected to be underestimated. Inaccurate conclusions

20

can also be deduced if donor females in estrus were involved, because recipient males

may have responded more to faecal pellets of females in estrus resulting in an

overestimate of response intensity.

21

CHAPTER 4: RESULTS

4.0 Type of Response

Sniffing and scent marking with preorbital gland secretion were the only responses

noticed for both male and female recipients. Recipient animals sniffed at or around

the deposited foreign faecal pellets then sometimes followed by deposition of

preorbital gland secretions on nearby tree and grass stems, poles and rocks. Urine and

faeces were also deposited during observation periods, but this was never directly

associated with the introduced deposits; rather, faeces and urine were deposited at

fixed locations along borders shared with other animals and sometimes randomly

within the pen (Fig. 1 and 2). Animals also did not scent mark exclusively at

introduced pellet sites, but continued to mark their usual marking sites such as

perimeter poles (Fig. 3).

Figure 1: Blue duiker urination site within enclosure

22

Figure 2: Blue duiker defecation site adjacent to neighboring enclosure

Figure 3: Marks of preorbital gland secretion left by a territorial blue duiker on the pole.

23

4.1 Response of Recipient Animals to Donor Animal Dung

Recipient animal responses were not significantly linked with specific individuals

within the same sex category (Sign test; p > 0. 05 for all tests), indicating that there

was no preferential response to one donor over another for example male Nyathi over

male Secret.

4.2 The Effect of Dung Age, Recipient Sex and Donor Sex to Recipient

Animals’ Response

The interaction of donor sex, dung age and recipient sex had a significant effect on the

intensity of response by recipient animals (permutation ANOVA, d.f = 3, 164;

p<0.05) for all response variables tested.

To precisely determine the source of variation, Tukey’s HSD tests were run, and the

significant results were recorded (tables 4.1 – 4.3). Male recipients sniffed for longer

duration (table 4.1) and investigated more (table 4.2) 10 hr female dung compared to

1 hr, 34 hr and 58 hr aged dung. Female recipients reacted faster and sniffed longer to

fresh female dung than to old dung from either male or female donor (table 4.1 and

4.3).

The tables below represent sources of variation derived from Tukey’s HSD tests for

specific response variables showing experimental combinations with significant

different results. Combinations which were not significantly different were excluded

from the results listed in tables below.

24

Table 2: Source of variation for total sniff duration showing that male recipients

sniffed for longer duration 10hr female dung compared to 1hr, 34hr, and 58hr aged

dung from control, male and female donors.

Donor Recipient Sign Donor Recipient P-Value

Female Female (1 hr) > Female Female (58 hr) 0.004

Female Female (1 hr) > Male Female (58 hr) 0.045

Female Male (10 hr) > Male Female (1 hr) 0.013

Female Male (10 hr) > Control Female (1 hr) 0.016

Female Male (10 hr) > Control Female (10 hr) 0.038

Female Male (10 hr) > Female Female (34 hr) 0.046

Female Male (10 hr) > Male Female (34 hr) 0.042

Female Male (10 hr) > Female Female (58 hr) 0.001

Female Male (10 hr) > Male Female (58 hr) 0.009

Female Male (10 hr) > Control Female (58 hr) 0.038

Table 3: Source of variation for investigation frequency. Male recipients investigated more 10hr female dung compared to 1hr and 58hr aged dung from control and female donors.

Donor Recipient Sign Donor Recipient P-Value

Female Male (10 hr) > Control Female (1 hr) 0.036

Female Male (10 hr) > Female Female (58 hr) 0.005

25

Table 4: Source of variation for lag time response variable showing that female recipients took longer time to start investigating introduced 58hr female dung compared to 1hr aged dung. Donor Recipient Sign Donor Recipient P-Value

Female Female (58 hr) > Female Female (1 hr) 0.015

4.3 Trends in Blue Duiker Response (lag) Time to Introduced Treatments Consistent trends were detected in response whereby both male and females took

longer to start investigating fresh (1 hr) to intermediate (10 hr and 34 hr) control dung

compared with foreign dung. Both recipient sexes investigated fresh foreign dung

faster than old foreign dung (58 hr) from same sex donor animals.

Recipient male

Dung age (hrs)

0 20 40 60 80 100

Lag

time

(min

s)

0

20

40

60

80

100

120

140

160

180

200

Donor male, Recipient male Recipient male, Donor female Control, Recipient male Stones, Recipient male

Recipient female

Dung age (hrs)

0 20 40 60 80 100

Lag

time

(min

s)

0

20

40

60

80

100

120

140

160

180

200

Recipient female, Donor female Donor male, Recipient female Control, Recipient female Stones, Recipient female

Figure 4: Trends in male recipients response times to introduced treatments

Figure 5: Trends in female recipients response times to introduced treatments

26

4.4 Time Spent by Blue Duikers Investigating Introduced Treatments

Male and female recipients sniffed control dung less than foreign dung. Very fresh

dung from same sex donor animals was sniffed more compared to that from opposite

sex donor animals. However, the longest sniff duration recorded for both sexes was

for introduced stones. Recipient Male

Dung age (hrs)

0 20 40 60 80 100

Mea

n sn

iff d

urat

ion

(sec

)

0

2

4

6

8

Donor male, Recipient male Control, Recipient male Recipient male, Donor female Stones, Recipient male

Recipient female

Dung age (hrs)

0 20 40 60 80 100

Mea

n sn

iff d

urat

ion

(sec

)

0

2

4

6

8

Recipient female, Donor female Donor male, Recipient female Control, Recipient female Stones, Recipient female

Figure 6: Mean time taken by male recipients sniffing introduced treatments

Figure 7: Mean time taken by female recipients

sniffing introduced treatments

4.5 Preorbital Scent Marking by Blue Duikers to Introduced Treatments in

their Territories

27

Both sexes scent marked less in response to control dung than to introduced foreign

dung. Male recipients never scent marked against 1 hr and 10 hr control dung whilst

female recipients never scent marked to 1 hr, 34 hr or 58 hr control dung. Both

recipient sexes never scent marked to the stone treatments. The recipient animals

showed more interest in scent marking intermediate (10 hr) foreign dung compared to

the rest of dung age categories.

Recipient male

Dung age (hrs)

0 20 40 60 80 100

Scen

t mar

king

freq

uenc

y

0

1

2

3

4

5

Donor male, Recipient male Recipient male, Donor female Stones, Recipient male Control, Recipient male

Recipient female

Dung age (hrs)

0 20 40 60 80 100

Scen

t mar

king

freq

uenc

y

0

1

2

3

4

5

Recipient female, Donor female Donor male, Recipient female Control, Recipient female Stones, Recipient female

4

Figure 8: Trends in preorbital scent marking by male

recipients

Figure 9: Trends in preorbital scent marking by female recipients

28

4.6 Investigation Frequency by Blue Duikers to Treatments Introduced into

their Territories

Male recipients investigated more fresh foreign dung compared to control dung but

the reverse was recorded for old dung where control dung was investigated more

frequently than foreign dung. On the other hand, female recipients showed more

interest in foreign dung compared to control dung. For both recipient sexes, the

investigation frequency decreased with the increase in dung age.

Recipient male

Dund age (hrs)

0 20 40 60 80 100

Inve

stig

atio

n fr

eque

ncy

0

2

4

6

8

10

12

Donor male, Recipient male Recipient male, Donor female Control, Recipient male Stones, Recipient male

Recipient female

Dung age (hrs)

0 20 40 60 80 100

Inve

stig

atio

n fr

eque

ncy

0

2

4

6

8

10

12

Recipient female, Donor female Donor male, Recipient female Control, Recipient female Stones, Recipient female

Figure 10: Frequency of investigation by male recipients to introduced treatments

Figure 11: Frequency of investigation by female recipients to introduced treatments

29

CHAPTER 5: DISCUSSION

5.0 Discussion

Unlike other antelope species such as klipspringer Oreotragus oreotragus (Roberts

and Lowen, 1997) and oribi Ourebia ourebi (Brashares and Arcese, 1999), blue

duikers were never observed licking or over marking with their own faecal pellets on

deposited foreign dung. Sniffing and scent marking with preorbital gland secretion

were the only forms of response noticed from both male and female blue duiker

recipient animals. This observation concurred with Estes (1997) who specified that

rubbing the preorbital glands on tree trunks and other objects within its environment,

is the main form of territory demarcation by blue duikers. However, scent marking

was not carried out in response to all introduced treatments especially control dung

suggesting that territory owners are able to distinguish between familiar and foreign

scent through scent matching (Gosling and McKay, 1990).

The study revealed that both sexes play a role in territory defense related activities

which is in agreement with Odendaal and Bigalke (1979) who mentioned that the blue

duiker is strongly territorial and both males and females mark out their territory

continually using urine, faecal pellets and preorbital gland secretion. Recipient

animals were reacting faster and investigating more to introduced foreign dung than to

control dung which suggest that they are territorial, and they scent mark more in

response to immediate perceived threats.

Neither urine nor faecal pellets were deposited in immediate response to foreign

faecal pellets deposited in their territory. However, faecal pellets were occasionally

seen deposited in enclosures, but mostly at fixed locations along borders shared with

other animals, which may be border marking to intimidate potential intruders and

cause them to withdraw from the marked territory (Roberts and Lowen, 1997).

Considering that defecation was mostly along territory boundaries, the possible reason

might be that the placement of marks is prioritized in areas where they can easily be

detected by the intended receiver, thus maximizing their value for territory

advertisement (ibid).

30

The fact that fresh and intermediate dung were investigated more than old dung

indicates that dung age had a significant effect on the intensity of response by

recipients. Consistent trends of decreasing response with increasing dung age which

were detected from this study concur with Ferkin et al., (1995). The other possible

reason for recipients showing more interest in fresh and intermediate dung compared

to old dung could be for defense purpose. If an owner notices foreign scent which is

fresh in its territory, it is possible that the intruder is still in the vicinity.

Consequently, the territory resident is expected to increase its marking rate in order to

cover a greater area with its own scent marks to maximize the probability of scent

matching by the intruder (Gosling, 1982).

Little sniffing was observed in response to control dung compared to foreign dung as

suggested by Gosling and MacKay (1990) that territory residents have got the ability

to differentiate between foreign and familiar scent. Consequently, they modulate their

response by spending more time investigating foreign scent so that they extract any

useful information needed about the signaler identity.

Blue duikers sniffed introduced stones regardless of them being scent neutral,

indicating that they also make use of visual cue in addition to olfactory and auditory

stimuli in territory maintenance and defense (Estes, 1997). Animals did not scent

mark in response to stones, signifying that they did not regard them as a territorial

threat but rather as something novel in their environment. This suggests that blue

duikers regulate the usage of scent marks and prioritize the placement of marks to

circumstances and areas where their value is worthy.

The findings of this research highlighted a slight increase in response (reduced lag

time, increased sniffing duration, increased scent mark frequency) when recipient

animals investigated dung from the same sex donors, most probably due to perceived

threat for mates. Basing on the findings by Wells and Bekoff (1981), scent marks

were reported to contribute in sex recognition which may suggests that same sex

animals might not tolerate each other’s company to reduce competition over potential

mates.

31

CHAPTER 6: CONCLUSION AND RECOMMENDATIONS

6.0 Conclusion

Blue duikers are primarily monogamous animals in which pair bonding is probably

life long. Direct interactions between animals of different social groups are rare

indicating that they are territorial. They defend their territory against potential

intruders using urine, faecal pellets and preorbital gland secretion.

Sniffing on or around the deposited dung pellets occasionally followed by preorbital

scent marking is the main form of response by blue duiker territory residents. The

marking densities generally increase in areas close to the deposited foreign faecal

pellets since the territory owner will be trying to mark with own scent to maximize

scent-matching probability. Urine and faeces are also deposited either at fixed

locations or randomly within the recipient animal pens but is not done in immediate

counter response to foreign substances found within the territory.

The efficacy of animal scent marks is not constant but varies with time after

deposition thus recipient animals react more to fresher dung compared to old one.

Dung scent retains a high degree of efficacy for some time and afterward the scent

signal starts to fade out.

Sex of either recipient or donor affects response intensity to a lesser extent because

both males and females are almost equally involved in territory defense related

activities. A slight increase in response intensity was noted when fresh strange scent

was introduced to recipient animals of the same sex to donor animals compared to

scent from donors of the opposite sex.

32

6.1 Recommendations

It is recommended that more experimental studies specifically on blue duiker

territorial behaviour must be conducted in the natural wild in order to have more

detailed and reliable information. There is also need to conduct a similar study using

dung aged more than 58 hours to see how long it takes for dung scent to be

undetectable.

33

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Appendix 1: Sample Data Record Sheet

DATE: ______________ TIME IN: __________ TIME OUT:

______________

TREATMENT: ________ PEN: ______________ WEATHER:

______________

Time Recipient

Animal

Sniff

duration(s)

Scent

marking

Notes

Type Frequency