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TESTING THE CORRELATION BETWEEN BIRD SPECIES RICHNESS AND THE PRESENCE OF LIONS OR LARGE UNGULATES IN AN ECOSYSTEM. A CASE STUDY OF BIRDS SPECIES IN DAMBWA FOREST WITH LIONS AND MOSI-OA-TUNYA NATIONAL PARK WITH LARGE UNGULATES. 2013 i DECLARATION I, Mathan Chikambwe do hereby declare that this work is of my own and that all the work of other people used in this research have been dully acknowledged. To the best of my knowledge, this work has not been previously presented at this institution or any other for similar purpose. Author’s signature …………………………….. Date …………………….. MATHAN CHIKAMBWE Supervisor’s signature ……………………........ Date ………….....………... DR. LACKSON CHAMA

Mathan chikambwe, THE COPPERBELT UNIVERSITY OF ZAMBIA

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TESTING THE CORRELATION BETWEEN BIRD SPECIES RICHNESS AND THE PRESENCE OF LIONS OR LARGE UNGULATES IN AN ECOSYSTEM. A CASE STUDY OF BIRDS SPECIES IN DAMBWA FOREST WITH LIONS AND MOSI-OA-TUNYA NATIONAL PARK WITH LARGE UNGULATES.

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TESTING THE CORRELATION BETWEEN BIRD SPECIES RICHNESS AND THE PRESENCE OF LIONS OR LARGE UNGULATES IN AN ECOSYSTEM. A CASE STUDY OF BIRDS SPECIES IN DAMBWA FOREST

WITH LIONS AND MOSI-OA-TUNYA NATIONAL PARK WITH LARGE UNGULATES.

2013

i

DECLARATION

I, Mathan Chikambwe do hereby declare that this work is of my own and that all the

work of other people used in this research have been dully acknowledged. To the best

of my knowledge, this work has not been previously presented at this institution or any

other for similar purpose.

Author’s signature …………………………….. Date ……………………..

MATHAN CHIKAMBWE

Supervisor’s signature ……………………........ Date ………….....………...

DR. LACKSON CHAMA

TESTING THE CORRELATION BETWEEN BIRD SPECIES RICHNESS AND THE PRESENCE OF LIONS OR LARGE UNGULATES IN AN ECOSYSTEM. A CASE STUDY OF BIRDS SPECIES IN DAMBWA FOREST

WITH LIONS AND MOSI-OA-TUNYA NATIONAL PARK WITH LARGE UNGULATES.

2013

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ABSTRACT

Changes in ecological variables such as habitat structure, water chemistry, fruit abundance,

predator abundance, or disturbance levels are known to explain changes in bird abundance.

Basically, such factors that influence birds abundance and diversity, have important indirect

effects on community structure and function. Ecosystems with large mammals can affect

vegetation structure and composition directly, with potential indirect effects on birds. The

objective of this study was to test the correlation between bird species richness and the presence

of lions or large ungulates in an ecosystem in order to establish what is driving the evolution

of the bird communities in the Mosi-oa-tunya national park and Dambwa forests. This study

revealed that, both bird species richness and abundance were not significantly different across

habitats implying that the presence or absence of top predators (lions) or large ungulates in

both ecosystems did not directly influence birds’ community. It is expected that, available food

resources for birds in both habitats were comparable and were not fully affected by the types

of large mammals present in these habitats. High species diversity across habitats may have

also been achieved by different management regimes. Due to the fact that both habitats are

protected areas, the relationship in bird species diversity, abundance and richness may be

attributed to the protection measures by management in both habitats which may have

substantially improved the habitat quality for avian species to persist, and not directly on the

types of large mammals present. Consequently habitat quality or ability of these two

ecosystems to provide conditions appropriate for avian individuals and population persistence

were comparable.

TESTING THE CORRELATION BETWEEN BIRD SPECIES RICHNESS AND THE PRESENCE OF LIONS OR LARGE UNGULATES IN AN ECOSYSTEM. A CASE STUDY OF BIRDS SPECIES IN DAMBWA FOREST

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ACKNOWLEDGEMENT

This thesis reflects the research conducted during my stay as an intern at Lion Encounter for

three months from January to March 2013. During this period I worked as a Young Researcher

with Lion Encounter in conjunction with African Lion and Environmental Research Trust

(ALERT).

The writing of my thesis was supervised by Dr. L. Chama, whom I would like to express my

gratitude for his constant willingness to comment on my ideas and drafts and carefully

reviewing my written English at every stage.

Working with Lion Encounter provided me with a lively research environment. In particular,

it gave me the possibility of browsing through several conservation activities, and of getting in

touch with a number of people that constantly encouraged me during this work in various ways.

Among these people, I am especially grateful to Jacqui Kirk (Lion Encounter research

coordinator), for her constant encouragement and suggestions throughout the different stages

of my data collection.

Interning at Lion Encounter for three months was certainly enriched by the friendliness of the

Lion Encounter members of staff including Richard, Nicola, Mulenga, Carah, Jacqui, all the

workers and the volunteers I had the luck to meet including, Jos van der Maiden, William

Meech, Patrick Martz, Callie Price, Genevieve Zock, Alexandra Skenteri, Vanessa Buchweitz,

Ashley Blannin, Peter Dockens, Tarah and many others, to whom I am deeply grateful for

making my experience even nicer.

TESTING THE CORRELATION BETWEEN BIRD SPECIES RICHNESS AND THE PRESENCE OF LIONS OR LARGE UNGULATES IN AN ECOSYSTEM. A CASE STUDY OF BIRDS SPECIES IN DAMBWA FOREST

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DEDICATION

This research paper is dedicated to the persons that received less attention than deserved during

its writing in particular, my parents including My mother Chinyama Ndeleki, Mr. S

Chikambwe, Mr. and Mrs. E. M. Chikambwe, my brothers and sisters, my friends and above

all, the Lord almighty for giving me strength through every stage in my life.

TESTING THE CORRELATION BETWEEN BIRD SPECIES RICHNESS AND THE PRESENCE OF LIONS OR LARGE UNGULATES IN AN ECOSYSTEM. A CASE STUDY OF BIRDS SPECIES IN DAMBWA FOREST

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Contents

DECLARATION.......................................................................................................................................... i

ABSTRACT ................................................................................................................................................. ii

ACKNOWLEDGEMENT ......................................................................................................................... iii

DEDICATION............................................................................................................................................ iv

LIST OF FIGURES .................................................................................................................................. vii

LIST OF TABLES ................................................................................................................................... viii

ACRONYMS .............................................................................................................................................. ix

CHAPTER ONE ......................................................................................................................................... 1

1.0 INTRODUCTION ................................................................................................................................. 1

1.1 BACKGROUND ................................................................................................................................... 2

1.2 PROBLEM STATEMENT .................................................................................................................. 4

1.3 JUSTIFICATION ........................................................................................................................... 4

1.4 OBJECTIVES ....................................................................................................................................... 5

1.4.1 MAIN OBJECTIVE ...................................................................................................................... 5

1.4.2 Specific Objectives ......................................................................................................................... 5

1.5 ASSUMPTIONS .................................................................................................................................... 6

1.6 SCOPE OF THE STUDY ..................................................................................................................... 6

CHAPTER TWO ........................................................................................................................................ 7

2.0 LITERATURE REVIEW .................................................................................................................... 7

CHAPTER 3 .............................................................................................................................................. 13

3.0 RESEARCH METHODOLOGY ...................................................................................................... 13

3.1 DAMBWA FOREST STUDY AREA ................................................................................................ 13

3.1.1 Dambwa forest Location ............................................................................................................. 14

3.1.2 Protection of Dambwa Forest ..................................................................................................... 14

3.1.3 Climate .......................................................................................................................................... 15

3.1.4 Vegetation ..................................................................................................................................... 15

3.2.0 MOSI-OA-TUNYA NATIONAL PARK STUDY AREA ............................................................ 16

3.2.1 LOCATION OF MOSI-OA-TUNYA NATIONAL PARK ...................................................... 17

3.2.2 Protection of Mosi-oa-tunya ........................................................................................................ 17

3.2.3 VEGETATION ............................................................................................................................ 17

TESTING THE CORRELATION BETWEEN BIRD SPECIES RICHNESS AND THE PRESENCE OF LIONS OR LARGE UNGULATES IN AN ECOSYSTEM. A CASE STUDY OF BIRDS SPECIES IN DAMBWA FOREST

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3.3.0 RESEARCH DESIGN ..................................................................................................................... 18

3.3.1 Distance sampling ........................................................................................................................ 18

3.3.2 Point counts .................................................................................................................................. 19

3.3.3 Line transects ............................................................................................................................... 19

3.4.0 DATA ANALYSIS ........................................................................................................................... 20

CHAPTER FOUR ..................................................................................................................................... 21

4.0 RESULTS AND DISCUSSION ......................................................................................................... 21

4.1 RESULTS ............................................................................................................................................ 21

4.1.2 SPECIES RICHNESS AND DIVERSITY ACROSS HABITATS .............................................. 22

4.1.1 The birds’ species abundance in Dambwa forest and Mosi-oa-tunya national park ................. 23

4.1.3 Bird species nutritional functional group categories in Mosi-oa-tunya national park .......... 24

4.1.4 Bird species nutritional functional group categories in Dambwa forest ................................. 25

4.2 DISCUSSION ...................................................................................................................................... 26

4.2.1 BIRD GUILDS WITHIN AND ACROSS HABITATS ............................................................ 27

4.2.2 ANIMAL INFLUENCE AND PROTECTION REGIMES ON HABITAT QUALITY ........... 31

CHAPTER FIVE ...................................................................................................................................... 32

5.0 CONCLUSION AND RECOMMENDATIONS .............................................................................. 32

5.1 RECOMMENDATIONS .................................................................................................................... 33

6.0 REFERENCES .................................................................................................................................... 34

Appendix 1 ................................................................................................................................................. 40

TESTING THE CORRELATION BETWEEN BIRD SPECIES RICHNESS AND THE PRESENCE OF LIONS OR LARGE UNGULATES IN AN ECOSYSTEM. A CASE STUDY OF BIRDS SPECIES IN DAMBWA FOREST

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LIST OF FIGURES

Figure 1: Dambwa forest lion enclosure……………………………………….……………..13

Figure 2: The map of Mosi-oa-tunya National Park in Livingstone………………………….16

Figure 3: The numbers of birds identified in both habitats one and two…………………….. 22

Figure 4: Birds’ species abundance in Mosi-oa-tunya national park (denoted as H1) and in

Dambwa forest (H2)………………………………………………………………………....23

Figure 5: The Birds species according to their nutritional functional groups in Mosi-oa-tunya

national park…………………………………………………………………………………24

Figure 6: Birds species according to their nutritional functional groups of birds of Dambwa

forest…………………………………………………………………………………………25

Figure 7: Birds Nutritional Functional Groups Proportions across Habitats………………..28

TESTING THE CORRELATION BETWEEN BIRD SPECIES RICHNESS AND THE PRESENCE OF LIONS OR LARGE UNGULATES IN AN ECOSYSTEM. A CASE STUDY OF BIRDS SPECIES IN DAMBWA FOREST

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LIST OF TABLES

Table 1: The birds’ species identified in Dambwa forest and in Mosi-oa-tunya national park..21

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ACRONYMS

ALERT African Lion and Environmental Research Trust

CBNRM Community Based Natural Resources Management

CRBs Community Resource Board

CITES Convention on International Trade in Endangered Species

GMA Game Management Area

HEC Human-Elephant Conflicts

KNP Kafue National Park

NGOs Non-Governmental Organizations

PA Protected Area

WWF World-wide Fund for Nature

ZAWA Zambia Wildlife Authority

TESTING THE CORRELATION BETWEEN BIRD SPECIES RICHNESS AND THE PRESENCE OF LIONS OR LARGE UNGULATES IN AN ECOSYSTEM. A CASE STUDY OF BIRDS SPECIES IN DAMBWA FOREST

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CHAPTER ONE

1.0 INTRODUCTION

The causes of variation in avian species richness at large spatial scales are intensively debated

such that in the recent past, a variety of models and hypotheses have been proposed to explain

patterns of species richness and variations across different ecosystems. Different species have

significantly different effects on the environment because they differ in the amounts and

combinations of resources that they utilize. For this reason, many ecosystem properties and

processes can only be understood on the basis of the organisms that are present in the

ecosystem. Based on high correlations with species richness, contemporary climate and energy

variables (e.g. precipitation, temperature and evapotranspiration) are often thought to explain

spatial variation in species richness better than any other non-climatic variable (Currie et al.

2004). There is a strong temptation to try to explain such changes in diversity on the basis of

the physical factors that are changing along same gradient. In fact, most of the explanations’

that have been proposed to explain species diversity are based on physical conditions that

change along gradients of species diversity. However, it is important to realize that natural

gradients are almost always complex, with many different factors changing along the gradient,

some increasing while others decreasing.

Although there is vigorous debate about the relative importance of different factors across

landscapes and beyond that regulate species diversity, it is widely accepted that habitat extent

and configuration at the landscape level can be important for many ecological processes

(Villard et al. 1999). Landscape influence on community diversity has particularly been a

strong area of recent interest (Debinski et al. 2001). The influences of the rate of disturbance,

TESTING THE CORRELATION BETWEEN BIRD SPECIES RICHNESS AND THE PRESENCE OF LIONS OR LARGE UNGULATES IN AN ECOSYSTEM. A CASE STUDY OF BIRDS SPECIES IN DAMBWA FOREST

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habitat extent, and habitat configuration at the landscape level on community diversity are also

central to several core ideas of ecological theory (Chesson and Case 1986). We may also agree

that in nature, food items are rarely mixed homogeneously across the landscape visited by

forager. Basically, this seems to suggest the reasons for wildlife species to choose habitats

carefully, to enhance the opportunities for feeding at the same time reducing the risk of being

eaten. Habitat associations relate bird distribution data (e.g. presence, abundance, or nest site

location) to habitat data. This research uses a comparative mensurative landscape-level

experiment to quantify the relative importance of Dambwa forest area (Lion enclosure) and

Mosi-oa-tunya national park in influencing avian community diversity.

1.1 BACKGROUND

The Mosi-oa-tunya national park is found in Livingstone, Southern province of Zambia. It is

the smallest park and has an area of 660ha and located approximately 450Kilometers South of

Lusaka. The park, is best known for its vast numbers of herbivores which include Elephants,

Buffaloes, Giraffe, Eland, Wildebeest, Lechwe, Impala, Warthog, and also the Monkeys,

implying great herbivore activity (browsing and grazing) in the park ecosystem. Associated

with such herbivore activities and the absence of the large carnivores are certain birds’ species

adapted to such an ecosystem.

On the other hand, approximately 5km north of the Mosi-oa-tunya national park is Dambwa

forest. As we may be aware, the African lion population is on its way to extinction due to

increased threats from human activities. In an effort to restore the declining African lion

population, a lion rehabilitation program through breeding and release introduced one male

TESTING THE CORRELATION BETWEEN BIRD SPECIES RICHNESS AND THE PRESENCE OF LIONS OR LARGE UNGULATES IN AN ECOSYSTEM. A CASE STUDY OF BIRDS SPECIES IN DAMBWA FOREST

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and six female lions into the Dambwa Forest Reserve in an enclosure (fig 1). The presence of

lions in Dambwa forest enclosure has with time contributed to the modification of the

ecological processes that occur within Dambwa forest, such as grass vegetation structural

changes. The type and structure of vegetation is important to birds in providing nest sites, roost

locations, refuge from predators, acting as food for herbivorous birds or providing herbivores

for carnivorous birds, and by its structure, enabling or constraining foraging. Changes in

landscape structure and function through time have important effects on the distribution of

resources, with resulting influences on the survival of species. The main focus of this research

is to test the potential influence of introduction of Lions in an ecosystem on bird’s species

richness, by firstly, assessing bird’s species in an ecosystem which excluded the presence of

Lions in Mosi-oa-tunya national park, followed by an ecosystem to which Lions were

introduced in Dambwa forest. Comparison of the first ecosystem with the second ecosystem

will allow for the evaluation on whether introduction of Lions had a significant effect on bird’s

species richness in Dambwa, or whether it only acted upon bird’s species richness through

causal relationships with other environmental variables.

An intellectual understanding of the natural regulation of the biological diversity-especially in

this case birds-may seem like an irrelevant luxury in a world where 140 hectares of tropical

forest are being cut or destroyed every minute, resulting in the extinction of an estimated 50 to

150 species per day (Reid and Miller, 1989). The preservation and maintenance of multiple

levels of organization, including species, populations, communities, and ecosystems, require

an understanding of how these various levels interact with each other and their environment

across a range of spatial scales.

TESTING THE CORRELATION BETWEEN BIRD SPECIES RICHNESS AND THE PRESENCE OF LIONS OR LARGE UNGULATES IN AN ECOSYSTEM. A CASE STUDY OF BIRDS SPECIES IN DAMBWA FOREST

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1.2 PROBLEM STATEMENT

Landscape-level mensurative studies are faced with numerous challenges (McGarigal et al.,

2002). Changes in landscape structure and function through time have important effects on the

distribution of resources, with resulting influences on the survival of species. A number of

studies have shown that the size, number, and distribution of habitats has an important

influence on species diversity (Bennun and Howell, 2002). In the same vein, an insight into

the part played by the various predators and large ungulates in attracting other species will be

important for our ultimate understanding of the interactions that occur among many species.

This is imperative for the formulation of a sound management plans directed at conservation

of the vast number of animals.

1.3 JUSTIFICATION

Ecological interactions between animal species are known to exist, yet given little attention by

researchers. Most of the research carried out on factors influencing local bird diversity has

focused either on the effects of vegetation structure (e.g. Macarthur and Macarthur, 1961) or

the effects of human-induced habitat changes, such as habitat fragmentation and conversion

(Donnelly and Marzluff, 2004). In nature, more than one type of interaction usually occurs at

the same time (Huston, 1994). In many cases, the outcome of one type of interaction is

modified or even reversed when another type of interaction is also occurring. Sometimes

species are not able to occupy their entire niche because of the presence or absence of other

species. By quantifying the interactions between the bird species, their food and habitat

preferences, this study will provide important insights into what is driving the evolution of the

bird communities in the Mosi-oa-tunya national park and Dambwa forests. At landscape level,

TESTING THE CORRELATION BETWEEN BIRD SPECIES RICHNESS AND THE PRESENCE OF LIONS OR LARGE UNGULATES IN AN ECOSYSTEM. A CASE STUDY OF BIRDS SPECIES IN DAMBWA FOREST

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many factors co-vary in their influences on species diversity and abundance. The comparison

in bird species found in Dambwa forest and Mosi-oa-tunya national park is intended to test

whether the occurrence of top predators in Dambwa forest may be more efficient biodiversity

indicators than other species. Moreover, diversity correlates are widely used as justifications

for various conservation planning strategies (Groves et al. 2000).

1.4 OBJECTIVES

1.4.1 MAIN OBJECTIVE

To test the correlation between bird species richness and the presence of lions or large

ungulates in an ecosystem. A case study of birds’ species in Dambwa forest with lions and

Mosi-oa-tunya national park with large ungulates.

1.4.2 Specific Objectives

To determine the types of bird species found in Dambwa forests where there are Lions

and in Mosi-oa-tunya national park where there are no lions from January to March

(rain season).

To determine if the presence of lions directly or indirectly influenced the types of bird

species in Dambwa forests as compared to those found in Mosi-oa-tunya national park.

To establish the ecological interaction that exist between the birds species found in

these two ecosystems, Dambwa forest with lions and Mosi-oa-tunya with large

ungulates.

TESTING THE CORRELATION BETWEEN BIRD SPECIES RICHNESS AND THE PRESENCE OF LIONS OR LARGE UNGULATES IN AN ECOSYSTEM. A CASE STUDY OF BIRDS SPECIES IN DAMBWA FOREST

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1.5 ASSUMPTIONS

This research assumes that the physical (vegetation and climatic) conditions between the Mosi-

oa-tunya national park and Dambwa forest are the same, implying that the bird species must

be the same. Hence, any differences in bird species richness are a result of the types of

ecological interactions that exist between the birds and animals found in a given area.

There is a difference in some birds species found in Dambwa forest and in Mosi-oa-

tunya national park because of the presence of lions in Dambwa forest.

The presence of Lions has an influence on some of the birds species found in Dambwa

forest.

1.6 SCOPE OF THE STUDY

This study is centered strictly on establishing an understanding as to why some bird species

occur where they do (thus, in Dambwa forest where there are lions or in Mosi-oa-tunya national

park with large ungulates species) in order to explain whether the occurrence of top predators

in Dambwa forest could be used to detect biodiversity hotspots, and also to establish whether

top predators may be more efficient biodiversity indicators than other species.

TESTING THE CORRELATION BETWEEN BIRD SPECIES RICHNESS AND THE PRESENCE OF LIONS OR LARGE UNGULATES IN AN ECOSYSTEM. A CASE STUDY OF BIRDS SPECIES IN DAMBWA FOREST

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CHAPTER TWO

2.0 LITERATURE REVIEW

Regular patterns of zonation, latitudinal gradient of species diversity, and the temporal patterns

of diversity found in succession beg for an explanation. Often times, it is obvious that changes

in physical properties of the environment such as temperature, precipitation, or soil are

correlated with the changes in diversity across landscape through time. This however may fuel

a temptation to try to explain species diversity changes across landscape on the basis of the

physical factors neglecting the different and more essential components that regulate

populations of species in various ecosystems. Autecological and behavioral studies of bird

species attempt to identify the environmental factors that influence population processes and

behavior. Applied research directed at bird conservation is important to improve our

understanding of habitat preferences and the relationships between demographic rates or

population density and habitat area and quality.

HABITAT ASSOCIATIONS

Habitat associations relate bird distribution data (e.g. presence, abundance, or nest site

location) to habitat data. One of the main methods, area comparisons, involves the selection

of a range of areas and relate abundance or presence to habitat. An example would be to select

a number of landscapes and quantify both the habitat and the number of birds in each.

Abundance or presence could then be related to habitat. Area comparisons are more likely to

reveal the habitat associations if a wide diversity of sites are used. This approach can be carried

out on a range of scales. At a patch scale the frequency with which different foraging patches

TESTING THE CORRELATION BETWEEN BIRD SPECIES RICHNESS AND THE PRESENCE OF LIONS OR LARGE UNGULATES IN AN ECOSYSTEM. A CASE STUDY OF BIRDS SPECIES IN DAMBWA FOREST

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are used could be quantified and related to habitat, while at a site scale the density of birds in

different forest blocks or on different landscapes could be related to habitat quality.

In some ecosystems, large mammals can affect vegetation structure and composition directly,

with potential indirect effects on birds. Cattle generally have negative effects on bird diversity

and abundance (Taylor 1986). In one study, McShea and Rappole (2000) found that white-

tailed deer (Odocoileus virginianus) did not affect bird diversity at sites in eastern North

America, although the deer did depress bird abundance through their effects on understory

vegetation. In Africa, elephants (Loxodonta africana) have been found to reduce bird

abundance and diversity in sites with high levels of elephant impact (Herremans, 1995).

TOP PREDATOR INFLUENCE ON BIODIVERSITY

Basically, according to Sergio, (2006) sites occupied by top predators have higher diversity of

vulnerable and non-vulnerable bird species than sites that are randomly chosen or are based on

lower trophic level species. He further established that, on average, there was a 1.5 to 2-fold

difference in mean biodiversity values between predator sites and control sites with no

predators. In addition to the overall biodiversity benefits, sites selected on the basis of predators

held greater densities of individual birds and butterflies (all species combined) than other sites

(Sergio et al. 2006).

TEMPORAL VARIATION IN SPECIES DIVERSITY

One of the complexities associated with studying diversity is that, diversity is constantly

changing at many different scales. In some groups of organisms, diversity measured at one

time of year is very different from diversity measured at another time of year. This emphasizes

TESTING THE CORRELATION BETWEEN BIRD SPECIES RICHNESS AND THE PRESENCE OF LIONS OR LARGE UNGULATES IN AN ECOSYSTEM. A CASE STUDY OF BIRDS SPECIES IN DAMBWA FOREST

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the extreme importance of carefully defining how diversity is evaluated in a specific situation.

The diversity of bird communities undergoes great seasonal fluctuations as a result of

migrations between summer breeding areas and winter feeding areas. The increase in temperate

bird species diversity that results from the influx of birds from the tropics to breed during the

temperate summer is proportionately much greater than the changes in tropical bird diversity

that result from the comings and goings of these same species. Evaluation and interpretation

of bird species diversity obviously require careful definition of which component of the bird

community is to be measured and when (Weins, 1989a).

DIVERSITY AND THE SPATIAL HETEROGENEITY OF THE SAMPLE AREA

As suggested by the species-area relationship, the spatial heterogeneity of an area is strongly

correlated with the number of species that are found there. The number of factors that

contribute to spatial heterogeneity is virtually innumerable, and can even include organisms

themselves, which offers the theoretical possibility of a positive feedback cycle of indefinite

increasing diversity (DeAngelis et al., 1986). One of the classic generalizations of ecology is

based on MacArthur and MacArthur’s (1961) observation that bird species diversity is

positively correlated with the structural complexity of the vegetation, which can be quantified

with the statistic called foliage height diversity. Since this observation, similar patterns have

been reported for the diversity of many different kinds of animals in relation to vegetation

structure or other aspects of environmental heterogeneity (Anderson, et al., 1983). There has

also been reports of bird species diversity decreasing with increasing foliage height diversity

(Ralph, 1985). Animals themselves can further increase heterogeneity created by plants by

eating or killing plants, by disturbing the soil with burrows, trails and wallows, by defecating,

TESTING THE CORRELATION BETWEEN BIRD SPECIES RICHNESS AND THE PRESENCE OF LIONS OR LARGE UNGULATES IN AN ECOSYSTEM. A CASE STUDY OF BIRDS SPECIES IN DAMBWA FOREST

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and by dying. In general, there are many causes of environmental heterogeneity, and

differences in heterogeneity are almost always correlated with differences in species diversity.

PREY AVAILABILITY

A major problem in relating foraging behavior to prey density is that not all prey are available.

Thus, prey may seem abundant to the observer, but is largely inaccessible to the bird.

Availability is always difficult to quantify and sometimes impossible. The usual first step is to

analyze the prey species, size classes, and locations in which birds feed and restrict the study

to these prey. For long-billed shorebirds, the depths at which prey are taken can be assessed by

comparing the probing depth to bill length. The depth from which immobile prey can be

extracted can be determined within artificial feeding sites by experiments on captive or free

living birds. Thus Robinson (1997) marked seeds with felt tip pens and placed the seeds in

different depths within trays of soil placed in the wild. He watched to see which species fed

there. After birds had fed he sieved the soil and recorded the seeds left and could thus determine

the proportion taken from different depths.

RESOURCE SELECTION AS THE CAUSE FOR HABITAT PREFERENCE

Most species have a suite of other needs to meet, including obtaining shelter from inclement

weather, gaining access to water, or locating suitable breeding sites such as cavities in dead

trees or burrows. The logical basis of virtually all measures of selective use is comparison

between the frequency of use of a particular resource (habitat) and its availability in the

environment. According to Sinclair et al (2006), a resource (habitat) is preferred when its use

by animals exceeds its availability and conversely that a resource (habitat) is avoided when its

TESTING THE CORRELATION BETWEEN BIRD SPECIES RICHNESS AND THE PRESENCE OF LIONS OR LARGE UNGULATES IN AN ECOSYSTEM. A CASE STUDY OF BIRDS SPECIES IN DAMBWA FOREST

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use is less than that expected from its availability in the environment. The purposeful

intermingling of “resource” and “habitat” is because similar analyses can be used for

determining whether animals (in this case birds) preferentially choose diets as for determining

whether animals show preferential habitat selection.

SOCIAL BEHAVIOR AND FORAGING

Given that there are differences in the intrinsic suitability of habitats, due to variation in

resources, cover, and risk from predators, one might expect animals to concentrate in the most

favorable habitats. The attractiveness of particular habitats is likely to depend, however, on the

density of foragers already present. Birth rates tend to fall and mortality rates to climb as

forager density increases (Sinclair et al. 2006). As a consequence, habitat suitability is often

negatively associated with density. Density dependent declines in habitat suitability can be

extended to multiple habitats. Individuals should concentrate in the best habitat until the

density in that habitat reduces its suitability to that of the next best alternative.

The ideal free hypothesis predicts that most individuals should be found in preferred habitats

when foragers population density is low, spilling over into less preferred habitats when forager

density is high. This pattern has been demonstrated several times in different bird species. One

of the earliest examples was Brown’s (1969) pioneering studies of great tits (Parus major) in

the woodlands near oxford, England. Brown showed that adult birds nested predominantly in

woodlands near habitats in years with low bird abundance, expanding outward into less

attractive hedgerows only when densities were high. Krebs (1971) tested the assumption that

this distribution stemmed from differences in fitness, by experimentally removing birds from

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woodlands habitats, resulting in vacancies that were filled rapidly by former hedgerow

“tenants.” Group versus solitary foraging behavior is a highly debated topic in behavioral

ecology in general and birds in particular (Sekercioglu et al. 2004). It has been claimed that

birds departing without prior knowledge of profitable foraging grounds may reduce their

search time and gain advantage by following other birds to foraging grounds (Ward and Salz,

1994). On the other hand, it has been debated that given the unpredictability of patchily

distributed prey, group foraging increases efficiency because individual birds can follow active

flight lines or search commonly exploited areas and locate currently profitable patches of

foraging habitat by cueing on the presence of other birds (Magurran,1988).

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CHAPTER 3

3.0 RESEARCH METHODOLOGY

3.1 DAMBWA FOREST STUDY AREA

Figure 1: Dambwa forest lion enclosure (source: Jucqui Kirk, lion Encounter)

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3.1.1 Dambwa forest Location

The study site for this research is Dambwa Forest Reserve which is located adjacent to the

Mosi-oa-Tunya National Park in Livingstone, Southern province of Zambia. Dambwa Forest

Reserve is located approximately 5 km north of Livingstone town center along the Great North

Road and approximately 470 km south of Lusaka, the capital city of Zambia. The forest reserve

has an area of 10,690 hectares and it is located between latitudes 17o and 18o South and between

longitudes 25o and 26o East, with an altitude of 1,000 m above sea level (Forest Department,

2003).

3.1.2 Protection of Dambwa Forest

The area was set aside and gazetted as a protected forest area (Forest Reserve No. 22) in 1976

for the purpose of supplying timber, fire wood and other forest products to the local

communities in Livingstone. The ownership of the forest reserve is vested in the Republican

President, as provided for under section 3 of the Forests Act Cap 199 of the laws of Zambia

(GRZ, 1973; GRZ, 1998). The area was further declared a joint forest management area

through the Statutory Instrument No. 47 of 2006 (GRZ, 2006). The central government through

the Forestry Department administered the management and protection of the forest reserve

before the introduction of joint forest management.

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3.1.3 Climate

Dambwa forest reserve falls within a semi-arid zone with a mean annual rainfall around 700

mm that usually falls between November and March. The area lies at an altitude of between

900 m and 1,000 m above sea level. The average maximum temperature range is 26°C to 37°C,

and the average minimum temperature range is 6°C to 19°C. The mean annual temperature is

20°C. On average, relative humidity is 56.8% throughout the year.

3.1.4 Vegetation

The predominant vegetation types of the forest reserve are Mopani (Colophospermum mopane)

in the northern boundary; Miombo woodland (Brachystegia) also found in the northern portion

of the forest; Baikiaea remnants are present throughout the forest, but more concentrated in the

north; and grasslands occur on the southern part of the forest with Baikiaea remnants. In the

northern part of the forest, there is a concentration of Baikiaea plurijuga, Pterocarpus

angolensis and Brachystegia (miombo) species. In the south, Schinziophyton rautanenii

(Mungongo), Afzelia quanzensis, Strychnos cocculloides, Diplorhyncus condylocarpon,

Combretum spp, Ochna spp, and Albizia species are common. Lannea species, Ochna species,

and Pseudolachnostylis maprouneifolia are common understorey trees in Dambwa Forest

Reserve. Diplorhyncus shrubs dominate much of the forest area demonstrating high human

activity, which had affected the forest in the past (Forest Department, 2003). Wildlife species

such as elephants, monkeys, kudu, impala, warthogs and even buffalo, are also found in the

forest reserve, as it is adjacent to Mosi-oa-Tunya National Park (Forest Department, 2003;

IUCN, 2006).

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3.2.0 MOSI-OA-TUNYA NATIONAL PARK STUDY AREA

Figure 2;The map of Mosi-oa-tunya National Park in Livingstone:(www.zambiatourism.com)

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3.2.1 LOCATION OF MOSI-OA-TUNYA NATIONAL PARK

Mosi-Oa-Tunya National Park is located in Livingstone on the upper part of the Zambezi

River. The Park covers an area of 66km2 with GPS latitude of 170 52’’ 00’ south and 250 49’’

59’ east. Climate is generally hot with low Precipitation figures. Daytime temperatures are

just above 30 degrees Celsius on average.

3.2.2 Protection of Mosi-oa-tunya

In relation to the protection and conservation of the Victoria Falls, an area about 66 square

kilometers in extent encompassing the Victoria Falls was declared the Mosi oa Tunya National

Park in 1972. Under the provisions of the National Parks and Wildlife Act of 1968, the removal

from or damage within any National Park, without the consent of the Director of any object of

geological, prehistoric, historical, archeological, scientific or aesthetic interest were prohibited.

In general terms, the 1968 Act contained more restrictions and regulations intended to conserve

wildlife and protect National parks (chomba et al. 2011).

3.2.3 VEGETATION

It includes the Miombo woodlands on the upper part, riverine vegetation on the Zambezi river

banks, grasslands, Mopane and palm trees. The wildlife park includes tall riverine forest with

palm trees, miombo woodland and grassland with plenty of birds, and animals including

giraffe, zebra, warthog, sable, eland, buffalo, impala and other antelope.

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3.3.0 RESEARCH DESIGN

Field work consisted of structured inventory methods in particular, point counts conducted at

two sites selected for bird diversity in Dambwa forest and Mosi-oa-tunya national park and

less structured methods designed to detect the types of species available during the inventory

seasons (rain season) including day surveys and walking/line transects. Conducting

experiments on top vertebrate predators is usually impossible for logistical, ethical and legal

reasons (Duffy 2002). According to Sutherland, (2006), line and point transect methodology

may be applied to such situations during birds inventory because it is particularly suitable for

animals that are difficult to catch, especially the birds, that can be observed directly in the field

but detected with any certainty only if they are close.

3.3.1 Distance sampling

The theory of distance sampling assumes that all animals at the observation point or on the line

are detected: it is the comparison of the number of animals detected at greater distances with

those detected at the point or line that allows detectability to be modelled (Sutherland, 2006).

The line and point transect requires that distances from the observation point or line to each

animal are recorded, as distance sampling. A pair of 10 x 42 binoculars was the instrument

which was used during the bird survey and a Birds identification guide book, for ease of

identification of bird species on both study sites. The route of each survey was consistent within

each site. Bird presence and abundance were recorded using both visual and aural methods.

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3.3.2 Point counts

Point counts were the most regimented method of collecting bird data at both the Mosi-oa-

tunya national park and Dambwa forest lion enclosure. Point-counts are usually conducted by

a team of observers or an individual who visit(s) each point in the field several times. However,

it should be noted that point-counts were conducted by one individual because the presence of

other observers could have caused a significant amount of disturbance, and thus influence the

frequency of contacts and the types of species detected by the counts. Six points were selected

in both Dambwa forest and Mosi-oa-tunya national park from which birds surveys were

conducted during late January to mid-march in 2013. Three points were selected in the Mosi-

oa-tunya national park, while the other three were likewise selected in Dambwa forest. The

protocol for these counts entailed standing at the center of a 100meter diameter plot and

identifying all birds heard and seen for 10 minutes; birds were recorded as occurring at one of

four distance intervals (< 25 m; 25-40 m; 40-100 m; and >100 m) or as flyovers (Hamel et al.

1996). The plots used for point counts were selected in a nonrandom manner due to the nature

of the sites from which the research was done.

3.3.3 Line transects

Line transects of 1 to 2.5 km were walked along the Dambwa lion release enclosure and also

at the Mosi-oa-tunya national park, noting all bird species both previously and not previously

recorded. Bird surveys were conducted starting from around 07:00hrs to 09:00hrs (local time)

and the late afternoon surveys were also conducted from 15:00 h to 16:30hrs at a slow pace of

1 km/h. Much of morning surveys were conducted in Mosi-oa-tunya national park and the late

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afternoon surveys were conducted in Dambwa forest due to transport problems. Surveys were

not conducted throughout the middle of the day due to the fact that temperatures were very

high and bird activity becomes very low.

3.4.0 DATA ANALYSIS

Data collected was analyzed both qualitatively and quantitatively using two computer software

packages; thus Microsoft excel 2013 and ‘R’-statistical package version 2.15.1 (2012-06-22)

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CHAPTER FOUR

4.0 RESULTS AND DISCUSSION

4.1 RESULTS

From January through to March (2013), 2375 birds of 89 different species were identified

across the two study sites, each habitat having 62 species of which, 35 were found to occur in

both habitats.

Table 1: The birds’ species identified in Dambwa forest and in Mosi-oa-tunya national park.

Number of birds identified

Species found only in H1 (Mosi-oa-tunya) 27

Species found only in H2 (Dambwa forest) 27

Species found in both H1 and H2 35

Total number of bird Species 89

Of all the bird species identified in habitat one and habitat two, 39% were recorded in both

habitats including species like the White-browed sparrow weaver (Procepasser mahali), Cape

turtle dove (Streptopelia capicola) and Flappet larks (Mirafra rufocinnamomea). On the other

hand, 31% of the birds were only recorded in habitat one including species like the Red billed

oxpeckers (Buphagus erythrorhynchus), Bearded woodpecker (Thripias namaquus) and Cattle

egrets (Bubulcus ibis) and 30% were recorded only in habitat two including the African white

backed vulture (Gyps africanus), African harrier hawk/Gymnogene (Polyboroides typus) and

Magpie (African long tailed) Shrike (Urolestes melanoleucus).

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Figure 3: The numbers of birds identified in both habitats one and two.

4.1.2 SPECIES RICHNESS AND DIVERSITY ACROSS HABITATS

Overall species richness was 29 for Mosi-oa-tunya national park and 28 for Dambwa forest.

Because of unequal numbers of individuals counted among habitats, Simpson’s diversity

values (H) were 0.886 for Mosi-oa-tunya national park and 0.965 for Dambwa forest lion

enclosure respectively. Based on CHI SQUARE TEST, bird diversity between Mosi-oa-tunya

national park and Dambwa forest was not significantly different (df = 1, P = 0.954).

Species found only in H1 27,

31%

Species found only in H2, 27, 30%

Species found in both H1 and H2,

35, 39%

IDENTIFIED BIRDS SPECIES

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4.1.1 The birds’ species abundance in Dambwa forest and Mosi-oa-tunya national park

Overall, the mean bird species richness in habitat one (16.90 ± 46.35 (i.e. Mean ± SD if not

otherwise noted)) and habitat two (9.79 ± 19.02), did not differ across habitats (P > 0.05). The

birds species abundance between habitat one and habitat two was not significantly different (F

=1.9128 and P =0.1692; fig 4).

Figure 4; Birds’ species abundance in Mosi-oa-tunya national park (denoted as H1) and in

Dambwa forest (H2).

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4.1.3 Bird species nutritional functional group categories in Mosi-oa-tunya national park

The birds’ species in Mosi-oa-tunya national park were grouped according to their nutritional

functional groups constituting of five categories, thus granivores (7.82 ± 34.38), insectivores

(8.48 ± 33.18), frugivores (1.07 ± 5.41), raptors (0.21 ± 1.04) and Scavengers (0.13 ± 0.86).

The overall distribution of bird nutritional functional groups, based on dietary guilds was

significantly different (df = 4, X2 = 34.77, P < 0.0001). Mosi-oa-tunya national park had 44%

Granivores species, 31% Insectivores, 14% frugivores, 8% raptors and 3% scavengers

respectively.

Figure 5: The Birds species according to their nutritional functional groups in Mosi-oa-tunya

national park.

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4.1.4 Bird species nutritional functional group categories in Dambwa forest

The birds’ species in Dambwa forest were grouped according to their nutritional functional

groups constituting of five categories, thus Granivores (5.29 ± 16.82), insectivores (3.34 ±

10.28), frugivores (0.58 ± 3.35), raptors (0.15 ± 0.79) and Scavengers (0.72 ± 4.45) (figure 4).

The overall distribution of bird nutritional functional groups, based on dietary guilds was

significantly different (df = 4, X2 = 36.71, P < 0.0001). Dambwa forest had 40% Insectivores

species, 37% Granivores, 10% raptors, 8% frugivores and 5% scavengers respectively.

Figure 6: Birds species according to their nutritional functional groups of birds of Dambwa

forest.

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4.2 DISCUSSION

Changes in variables such as habitat structure, water chemistry, fruit abundance, predator

abundance, or disturbance levels, explain changes in bird abundance. Bird species richness and

abundance are influenced by local resource availability and vegetation composition, in addition

to the size of habitat patches (MacArthur and MacArthur 1961). In some ecosystems,

protection measures by management as well as the presence of large mammals can affect

vegetation structure and composition directly with potential indirect effects on bird species

richness and abundance. The results obtained from this study show that birds species

abundance between habitat one and habitat two was not significantly different (F = 1.9128, P

= 0.1692). These results seem to suggest that there is a correlation in birds’ species richness

and abundance across habitats. This may mean that the presence of lions in Dambwa forest

where there are no large ungulates did not affect bird species richness and abundance.

Similarly, Mosi-oa-tunya national park containing large ungulates and no top predators (lions)

did not affect bird species richness and abundance. Both bird species richness and abundance

were unaffected by the presence or absence of either the large ungulates or top predators

(African Lion) in either of the two ecosystems or habitats. A similar study was conducted by

Ogada et al. (2008) who used a large scale, replicated exclusion experiment in Kenya to

investigate the impacts of different guilds of native and domestic large herbivores on the

diversity and abundance of birds over a 2-year period. Ogada et al. (2008) found no significant

effects of native large herbivores other than mega herbivores on bird diversity or abundance.

The research findings also indicated that, exclusion of large herbivorous native mammals,

including giraffes (Girqffa camelopardalis), elephants (Loxodonta africana), and buffalos

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(Syncerus coffer), increased the diversity of birds by 30%. Much of this effect was attributable

to the absence of elephants and giraffes; these mega herbivores reduced both the canopy area

of subdominant woody vegetation and the biomass of ground-dwelling arthropods, and both

of these factors were good predictors of the diversity of birds. The canopy area of subdominant

trees was positively correlated with the diversity of granivorous birds. The biomass of ground-

dwelling arthropods was positively correlated with the diversity of insectivorous birds. These

results suggest that most native large herbivores are compatible with an abundant and diverse

bird fauna, if they are at a relatively low stocking rate.

4.2.1 BIRD GUILDS WITHIN AND ACROSS HABITATS

Overall, granivores and insectivore species were marginally higher than the frugivores, raptor

and scavenger species in both habitat one (X2 = 34.77, P < 0.0001) and habitat two (X2 =

36.71, P < 0.0001) respectively. These results demonstrate that granivorous and insectivorous

functional groups activity was higher in both sites than the frugivores, raptors and scavengers.

The structure and dynamics of biological communities are usually influenced by the combined

effect of several ecological variables each operating at different spatial and temporal scales

(Katrin 1997).

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Figure 7: Birds Nutritional Functional Groups Proportions across Habitats

a) Granivores

Seed supply may also be an important regulator of granivore density in other ecosystems of

the world, either directly, or indirectly by promoting competitive interactions among coexisting

species (Pulliam and Parker 1979). Local abundance of granivore species is strongly regulated

by food availability (Schluter and Repasky 1991). The granivore species marginal increase in

both ecosystems are expected to have been due to the abundance of seed supply. Seed supply

is a proximate factor that limits the abundance of granivorous birds but does not limit species

richness (Pedro and Ricardo, 2001).

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9

5

2

23

25

5

6

3

0

5

10

15

20

25

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G R A N I V O R E S I N S E C T I V O R E S F R U G I V O R E S R A P T O R S S C A V A N G E R S

No

. Of

Bir

d S

pec

ies

NUTRITIONAL FUNCTIONAL GROUPS

No. in Habitat 1 No. in Habitat 2

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b) Insectivores

Insectivorous birds have significant behavioral, ecological, and evolutionary effects on their

invertebrate prey species by affecting their behavior and limiting their movements, as well as

the damage they cause to plants (Strong et al., 2000). The marginal increase in the number of

insectivore species in both habitats seems to suggest that more prey species occur in both

habitats. This may be attributed to the habitat quality as a result of the large mammals and the

protection measures put in place by management in both ecosystems.

c) Frugivores

Kissling et al. (2007) found a positive relationship between food plant and frugivores species

richness. The number of observed frugivore species in both ecosystems may have been

influenced by various factors such as the types of fruit plants available and the season for fruit

ripening in both ecosystems. According to Márquez et al (2004) biotic dispersal of fruits

strongly affects broad-scale geographical trends of fleshy-fruited plant species richness,

whereas richness of fruited plants moderately affects frugivore richness. Further, Márquez et

al (2004) suggested that frugivores are important because they offer various ecosystem

services and economic benefits which include the removal of seeds from the parent tree, escape

seeds from herbivores and seed predators, improved germination, increased economical yield,

increased gene flow, re-colonization and restoration of disturbed ecosystems.

d) Raptors

Although raptors as a group have a lower percentage of extinction-prone species than most

other functional groups, large raptor species are more sensitive to disturbance and are more

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threatened than average (Hager, 2009). The raptors observed could have been influenced by

the availability of their prey species given the habitat quality as a result of the existing

protection regimes in both habitats. The raptors play a vital role in terrestrial ecosystem through

controlling unwanted species, nutrient deposition, soil formation in polar environments,

indicators of fish stocks in aquatic ecosystems, and are considered as environmental monitors

(Hager, 2009).

e) Scavengers

Compared to other avian functional groups, the obligate scavenger guild is tiny (fig 5) and

comprised of only a few species whose food consumption is predominantly based on

scavenging. According to Wilson and Wolkovich (2011), scavenging patterns depend on many

interacting factors related to the carcass, the consumer or extrinsic variables. With respect to

consumers, the presence of specialist scavengers and the number of scavenger species might

be particularly important. Vultures are the only scavenger species which were observed in both

habitats. Vultures are the world’s only obligate terrestrial vertebrate scavengers, which display

many adaptations for efficient carrion consumption (Ruxton and Houston 2004). These

adaptations make vultures strong competitors in the terrestrial scavenger guild (Houston,

1979), and the presence of obligate scavengers can increase interspecific trophic competition,

and in turn influence carrion consumption patterns (Ogada et al. 2012). Due to the types of

animals that occur in both habitats, it is likely that the numbers of the scavengers’ where

influenced by the carcass availability. However, Olson et al. (2012) showed experimentally

that reduction of facultative scavenger abundance resulted in reduced carcass consumption

rates in an area where the vultures were absent.

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4.2.2 ANIMAL INFLUENCE AND PROTECTION REGIMES ON HABITAT

QUALITY

Comparison of bird species richness between Dambwa and Mosi-oa-tunya national park was

confounded by variation in habitat area and species composition which to some extent, was

expected to have impacted on structure of vegetation and habitat quality. According to the

results obtained, bird richness, diversity and abundance were not directly affected by the

presence of large ungulates, despite the potential for large ungulates to significantly reduce

grass cover through browsing and grazing in habitat one. Similarly, bird diversity and

abundance in habitat two was not directly affected by presence of top predator despite the

abundant grass cover due to the absence of grazers and browsers. It is however expected that

large ungulates and/or top predators to some extent, impacted on habitat quality which

ultimately contributed to the increase in avian species richness and abundance across the two

habitats. On the other hand, it is important to note that, Mosi-oa-tunya national park has been

under protection from as far as 1972 under the provisions of the National Parks and Wildlife

Act of 1968. Dambwa forest was also set aside and gazetted as a protected forest area (Forest

Reserve No. 22) in 1976. Proper management of protected areas is important in fostering

species richness and diversity (Sinclair et al, 2006). The fact that these are protected areas, the

relationship in bird species diversity, abundance and richness may also be attributed to the

protection measures by management in both ecosystems which may have directly or indirectly

improved the habitat quality for avian species to persist, and not directly on the types of large

mammals present. The vegetation structure in both protected areas, provides some interesting

insights into the relationships between bird species diversity and vegetation structure.

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CHAPTER FIVE

5.0 CONCLUSION AND RECOMMENDATIONS

To conclude, results from this study broadly suggest that the correlation in bird species

richness, abundance and diversity in both habitats is not directly due to the presence or absence

of ungulates or top predator in an ecosystem. The effects of fragmentation and habitat

modifications are better understood by comparing species richness across a range of sites.

Basically, both study areas were assessed to determine the effect of habitat associations (e.g.

ecosystem changes by large ungulates in Mosi-oa-tunya) and species introduction (Lions

introduction in Dambwa Forest) in order to document changes over time. It is expected that,

available food resources for birds in both Dambwa forest and Mosi-oa-tunya national park

were actually not fully affected by the types of large mammals present in both habitats under

consideration. Based on the findings in this study, it is likely that the habitat quality or ability

of these two habitats to provide conditions appropriate for avian individuals and population

persistence were comparable. High species diversity at different spatial scales may have also

been achieved by different management regimes. Due to the fact that these are protected areas,

the relationship in bird species diversity, abundance and richness may be attributed to the

protection measures by management in both ecosystems which may have improved the habitat

quality for avian species to persist, and not directly on the types of large mammals present.

Applied research directed at bird conservation will improve the understanding of habitat

preferences and the relationships between demographic rates or population density and habitat

area and quality. Monitoring changes in species diversity, across ecosystems helps in

understanding the roles that the animals play in conserving other wildlife species.

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5.1 RECOMMENDATIONS

Changes or introductions of carnivore species can affect profoundly the density of other

species in an ecosystem. Further research should be conducted in Dambwa forest to

assess the impact of introduction of predators (African lion) on the density of other

species.

Parallel studies on Birds foraging behavior should be conducted in both habitats in

order to answer a wide range of questions like, what the species eat, why they prefer

certain areas for feeding, competition for food among particular species or among age

classes or sexes within a species, how many individuals a site can sustain, and the

consequences of habitat change.

The co-existence of the different members of a guild is a prerequisite for biodiversity

to exist. Given the potential importance of birds in ecological systems, future research

should address the ecological consequences of changes in bird diversity brought about

by large mammals. Future research is to go beyond documenting that species interact,

but also to determine exactly how different species co-exist in the presence of these

interactions.

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Appendix 1

The following symbols are interpreted as: A= 0%, S ≤ 1%, 1% < H > 5%, L ≥ 5%

Species Scientific name Habitat one

% occurrence Habitat two

% occurrence

F1 (African) White backed vulture Gyps africanus A 0% S 0.69%

F2 African broadbill Smithornis capensis S 0.07% A 0%

F3 African golden oriole Oriolus auratus S 0.03% S 0.69%

F4 African green pigeon Treron calvus S 0.13% A 0%

F5 African grey hornbill Tockus nasutus H 2.13% H 2.30%

F6 African harrier hawk (gymnogene) Polyboroides typus A 0% S 0.80%

F7 African mourning dove Streptopelia decipiens S 0.47% H 1.26%

F8 African pied wagtail Motacilla aguimp S 0.33% A 0%

F9 Arrow marked babbler Turdoides jardineii A 0% H 1.15%

F10 Banded martin Riparia cincta L 16.56% H 4.02%

F11 Barn (european) swallow Hirundo rustica S 0.53% A 0%

F12 Bearded woodpecker Thripias namaquus S 0.20% A 0%

F13 Black collared barbet Lybius torquatus S 0.13% A 0%

F14 Black coucal Centropus grillii A 0% S 0.11%

F15 Black tit Parus niger S 0.27% S 0.57%

F16 Blue waxbill Uraeginthus angolensis H 4.45% L 7.81%

F17 Broadbilled roller Eurystomus glaucurus S 0.07% S 0.11%

F18 Broad-tailed Paradise-whydah Vidua obtusa A 0% S 0.69%

F19 Brown hooded kingfisher Halcyon albiventris A 0% S 0.23%

F20 Brown snake eagle Circaetus cinereus S 0.13% A 0%

F21 Cape turtle dove Streptopelia capicola H 4.45% L 13.55%

F22 Cattle egret Bubulcus ibis L 12.03% A 0%

F23 Collared sunbird Anthreptes collaris A 0% S 0.23%

F24 Common qual Coturnix coturnix S 0.27% S 0.23%

F25 Common scimitarbill Rhinopomastus cyanomelas A 0% S 0.34%

F26 Coppery tailed coucal Centropus cupreicaudus A 0% S 0.11%

F27 Crested barbet Trachyphonus vaillantii S 0.13% S 0.11%

F28 Dark capped Bulbul Pycnonotus tricolor S 0.73% A 0%

F29 Dickinson's kestrel Falco dickinsoni S 0.53% S 0.11%

F30 Emmerald spotted wood dove Turtur chalcospilos S 0.47% H 1.15%

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F31 Eurasian golden oriole Oriolus oriolus S 0.07% S 0.34%

F32 Flappet lark Mirafra rufocinnamomea H 2.86% H 3.56%

F33 Fork tailled drongo Dicrurus adsimilis H 1.13% L 6%

F34 Gabar goshawk Melierax gabar S 0.13% A 0%

F35 Greater blue eared starling Lamprotornis chalybaeus A 0% S 0.23%

F36 Green wood hoopoe Phoeniculus purpureus A 0% S 0.80%

F37 Grey lourie/ go away bird Corythaixoides concolor H 1.13% L 3.44%

F38 Hamerkop Scopus umbretta S 0.07% A 0%

F39 Helmeted guineafowl Numida meleagris S 0.33% A 0%

F40 Hooded vulture Necrosyrtes monachus S 0.27% L 4.48%

F41 Jameson's firefinch Lagonosticta rhodopareia A 0% S 0.34%

F42 Lappet faced vulture Torgos tracheliotos S 0.47% L 1.72%

F43 Lesser masked-weaver Ploceus intermedius A 0% S 0.80%

F44 Levaillant(stripped) cuckoo Clamator levaillantii A 0% S 0.69%

F45 Lilac breasted roller Coracias caudatus S 0.07% A 0%

F46 Little bee-eater Merops pusillus H 1.33% H 2.07%

F47 Lizzard buzzard Kaupifalco monogrammicus S 0.33% S 0.11%

F48 Longtailed paradise whydah Vidua paradisaea A 0% S 0.11%

F49 Loughing dove Stigmatopelia senegalensis S 0.86% H 1.72%

F50 Madagascar bee-eater Merops superciliosus S 0.20% A 0%

F51 Magpie (African long tailed) shrike Urolestes melanoleucus A 0% L 5.63%

F52 Meve's starling Lamprotornis mevesii H 2.26% A 0%

F53 Meyer's parrot Poicephalus meyeri S 0.13% S 0.23%

F54 Namaqua dove Oena capensis H 1.13% A 0%

F55 Orange-breasted bush shrike Telophorus sulfureopectus S 0.13% A 0%

F56 Pied (jacobin) cuckoo Clamator jacobinus S 0.13% S 0.92%

F57 Pied kingfisher Ceryle rudis S 0.13% A 0%

F58 Purple indigobird (dusky indigobird) Vidua purpurascens S 0.47% H 1.38%

F59 Queen (shaft tailed) whydah Vidua regia A 0% S 0.34%

F60 Racket tailed roller Coracias spatulatus S 0.20% A 0%

F61 Red billed firefinch Lagonosticta senegala A 0% S 0.80%

F62 Red billed francolin Pternistes adspersus A 0% S 0.23%

F63 Red billed hornbill Tockus erythrorhynchus H 2.93% S 0.92%

F64 Red billed Quelea Quelea quelea L 7.45% H 1.72%

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F65 Red eyed dove Streptopelia semitorquata A 0% S 0.92%

F66 Red faced mousebird Urocolius indicus H 3.13% S 0.92%

F67 Red headed weaver Anaplectes rubriceps S 0.07% A 0%

F68 Red throated twinspot Hypargos niveoguttatus S 0.60% H 1.15%

F69 Red-billed oxpecker Buphagus erythrorhynchus S 0.73% A 0%

F70 Red-billed wood hooper Phoeniculus purpureus S 0.13% A 0%

F71 Scarlet-chested sunbird Nectarinia senegalensis A 0% S 0.23%

F72 Southern red bishop Euplectes orix S 0.27% A 0%

F73 Spectacled weaver Ploceus ocularis S 0.33% A 0%

F74 Steppe eagle Aquila nipalensis A 0% S 0.11%

F75 Swainson's spurfowl (francolin) Francolinus swainsonii S 0.27% S 0.46%

F76 Swallow tailed bee-eater Merops hirundineus A 0% S 0.11%

F77 Tawny-flanked prinia Prinia subflava H 1.86% L 7.50%

F78 Trumpeter hornbill Bycanistes bucinator S 0.27% A 0%

F79 Village indigobird Vidua chalybeata H 1.60% H 3.00%

F80 Violet backed starling Cinnyricinclus leucogaster S 0.60% S 0.34%

F81 White bellied sunbird Nectarinia talatala A 0% S 0.34%

F82 White fronted bee-eater Merops bullockoides S 0.33% S 0.46%

F83 White throated swallow Hirundo albigularis S 0.33% A 0%

F84 White winged widowbird Euplectes albonotatus S 0.33% S 0.69%

F85 White-browed Robin-chat Cossypha heuglini A 0% S 0.11%

F86 White-browed sparrow-weaver Procepasser mahali L 19.48% L 7.58%

F87 Yellow billed oxpecker Buphagus africanus H 1.06% A 0%

F88 Yellow throated longclaw Macronyx croceus A 0% S 0.12%

F89 Yellow-fronted Canary Serinus mozambicus A 0% H 1.15%