Mathan chikambwe, THE COPPERBELT UNIVERSITY OF ZAMBIA by Mathan Chikambwe

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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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 ……………………........ DR. LACKSON CHAMA

Date ………….....………...

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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 birdsâ&#x20AC;&#x2122; 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.

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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.

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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.

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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 v

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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

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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

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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

Protected Area

WWF

World-wide Fund for Nature

ZAWA

Zambia Wildlife Authority

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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 explanationsâ&#x20AC;&#x2122; 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, 1

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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 birdsâ&#x20AC;&#x2122; 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

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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. 3

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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, 4

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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 birdsâ&#x20AC;&#x2122; 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.

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1.5 ASSUMPTIONS This research assumes that the physical (vegetation and climatic) conditions between the Mosioa-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-oatunya 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.

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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 7

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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 whitetailed 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 8

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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 MacArthurâ&#x20AC;&#x2122;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, 9

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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 10

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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).

CHAPTER 3 3.0 RESEARCH METHODOLOGY 3.1 DAMBWA FOREST STUDY AREA

Figure 1: Dambwa forest lion enclosure (source: Jucqui Kirk, lion Encounter) 13

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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-oatunya 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 Mosioa-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 19

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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 birdsâ&#x20AC;&#x2122; species identified in Dambwa forest and in Mosi-oa-tunya national park. Number of birds identified Species found only in H1 (Mosi-oa-tunya)

Species found only in H2 (Dambwa forest)

Species found in both H1 and H2

Total number of bird Species

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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IDENTIFIED BIRDS SPECIES

Species found Species found in only in H1 27, both H1 and H2, 31% 35, 39%

Species found only in H2, 27, 30%

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, Simpsonâ&#x20AC;&#x2122;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).

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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 birdsâ&#x20AC;&#x2122; 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 grounddwelling 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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NUTRITIONAL FUNCTIONAL GROUPS 23

15 9

No. Of Bird Species

5 0 GRANIVORES

INSECTIVORES

FRUGIVORES

No. in Habitat 1

RAPTORS

SCAVANGERS

No. in Habitat 2

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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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 Mรกrquez 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, Mรกrquez 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 29

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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 worldâ&#x20AC;&#x2122;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 scavengersâ&#x20AC;&#x2122; 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. 30

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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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6.0 REFERENCES Anderson, B.W., Ohmart, R.D., and Rice, J. (1983). Avian and vegetation community structure and their seasonal relationships in the lower Colorado River Valley. Condor, 85. Bennun, L. and Howell, K. (2002). African Forest biodiversity: a field survey manual for vertebrates, eds. G. Davies, and M. Hoffmann, Earthwatch Europe, Earthwatch, Oxford. Block, W. M., And L. A. Brennan. (1993). The habitat concept in ornithology: theory and applications. Current Ornithology 11:35–91. Brown, J.L. (1969). The buffer effect and productivity in tit populations. American Naturalist 103:347–354. Chesson, P. L., and T. J. Case. (1986). Overview: non-equilibrium community theories, chance, variability, history and coexistence. Pages 229-239 in J. Diamond and T. J. Case, editors. Community ecology. Harper and Row, New York, USA. Chomba C., Mwenya A.N., and Nyirenda V. (2011). Journal of Sustainable Development in Africa (Volume 13, No.6, 2011) Clarion University of Pennsylvania, Clarion. ISSN: 15205509 Currie, D. J. (2004). Predictions and tests of climate-based hypotheses of broad-scale variation in taxonomic richness. Ecol. Lett. 7, 1121–1134. DeAngelis, D.L., Post, W.M., and Travis, C. (1986). Positive feedback in Natural systems. Heidelberg: Springer-Verlag.

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Debinski, D.M., Ray C., and Saveraid E.H., (2001). Species diversity and the scale of the landscape mosaic: do scales of movement and patch size affect diversity? Biological Conservation 98:179-190. Donnelly R., Marzluff J.M., (2004). Importance of reserve size and landscape context to urban bird conservation. Conservation Biology 18:733-745 Duffy, J.E. (2002). Biodiversity and ecosystem function: the consumer connection. Oikos, 99, 201–219. Forestry Department, (2003). Final Draft Joint Forest Management Plan for Dambwa Local Forest, Forestry Department, Zambia. Groves, C., Valutis L., Vosick D., Neely B., Wheaton K., Touval J., and Runnels B. (2000). Designing a geography of hope: a practitioner’s handbook for eco-regional conservation planning. The Nature Conservancy. Hager S. (2009). Human-related threats to urban raptors. Journal of Raptor Research, 43, 210226. Herremans M. (1995). Effects of woodland modification by African elephant (Loxodonta Africana) on bird diversity in northern Botswana. Ecography 18:440-454 Houston, D.C. (1979). The adaptations of scavengers. In Sinclair, A.R.E. & Griffiths, M.N. (eds) Serengeti, Dynamics of an Ecosystem: 263–268. Chicago, IL: University of Chicago Press.

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Huston A.M. (1994). Biological Diversity: The coexistence of species. Cambridge University Press. IUCN. (2006a). Conservation strategy for the lion in West and Central Africa. Yaounde, Cameroon. IUCN. (2006b). Regional conservation strategy for the lion Panthera leo in Eastern and Southern Africa. Katrin Bohning-GaeseSource, (1997). Determinants of Avian Species Richness at Different Spatial Scales. Journal of Biogeography, Vol. 24, No. 1 (Jan., 1997), pp. 49-60. Kissling W.D., Rahbek C., and Katrin Bรถhning-GaeseSource. (2007). Food plant diversity as broad-scale determinant of avian frugivore richness. Proc. R. Soc. B 2007 274, 799-808. Krebs, J.R. (1971). Territory and breeding density in the great tit, Parus major L. Ecology 52:222. MacArthur R.H., and MacArthur J, (1961). The bird species diversity. Ecology. 42:594598. Magurran A.E. (1988). Ecological diversity and its measurement. Cam-bridge University Press, Cambridge Mรกrquez A.L., Real R., and Vargas, M.J. (2004). Dependence of broad-scale geographical variation in fleshy-fruited plant species richness on disperser bird species richness. Global Ecology and Biogeography, Blackwell Publishing, Ltd. 13, 295-304.

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McGarigal, K., Cushman, S.A., Neel, M.C. and Ene, E. (2002). FRAGSTATS: Spatial Pattern Analysis Program for Categorical Maps. Computer software program produced by the authors at the University of Massachusetts, Amherst. McShea W.J., Rappole J.H. (2000). Managing the abundance and diversity of breeding bird populations through manipulation of deer populations. Conserv Biol 14:1 161-1 170. Ogada D.L., Gadd M.E., Ostfeld R.S., Young T.P., and Keesing F. (2008). Impacts of large herbivorous mammals on bird diversity and abundance in an African savanna. Ogada, D.L., Torchin, M.E., Kinnaird, M.F. & Ezenwa, V.O. (2012). Effects of vulture declines on facultative scavengers and potential implications for mammalian disease transmission. Conserv. Biol. 26: 453–460. Olson, Z.H., Beasley, J.C., DeVault, T.L., and Rhodes, O.E. Jr. (2012). Scavenger community response to the removal of a dominant scavenger. Oikos 121: 77–84. Pedro G.B., and Ricardo A.O., (2001). Seed supply as a limiting factor for granivorous bird assemblages in the Monte Desert, Argentina Biodiversity Research Group, CC 507, 5500. Pulliam H.R., and Parker III T.A., (1979). Population regulation of sparrows. Fortschr. Zool. 25, 137–47. Ralph, C.J. (1985). Habitat association patterns of forest and steppe birds of northern Patagonia, Argentina. The condor, 87,471-83. Reid, W.V., and Miller, K.R. (1989). Keeping options Alive: the scientific basis for conserving biodiversity. Washington, D.C.: world resources institute. 37

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Ribeiro, N., Shugart, H., and Washington-Allen, R., (2008). The effects of fire and elephants on species composition and structure of the Niassa Reserve, northern Mozambique. Forest Ecology and Management 255: 1626–1636 Robinson, R.A. (1997). Ecology and Conservation of seed-eating birds on farmland. Unpublished PhD thesis, University of East Anglia. Ruxton, G.D., and Houston, D.C. (2004). Obligate vertebrate scavengers must be large soaring fliers. J. Theor. Biol. 228: 431–436. Schluter D., and Repasky R. R. (1991). Worldwide limitation of finch densities by food and other factors. Ecology 72, 1763–74. Sekercioglu, C. H., G. C. Daily, and P. R. Ehrlich. (2004). Ecosystem consequences of bird declines. Proceedings of the National Academy of Sciences USA 101:18042–18047. Sergio, F., Newton, I., Marchesi, L., and Pedrini, P. (2006). Ecologically justified charisma: preservation of top predators delivers biodiversity conservation. British Ecological Society, Journal of Applied Ecology, 43, 1049-1055 Sinclair R.E., Fryxell M.J., and Coughley G. (2006). Wildlife ecology, conservation, and management. Second edition, Blackwell Publishing Ltd. Sinclair, I., Hockey, P., Tarboton, W. (2002). Birds of southern Africa. The region’s most comprehensively illustrated guide by Peter Hayman &Norman Arlott. Third edition, struik publishers.

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Strong A.M., Sherry T.W., and Holmes R.T., (2000). Bird predation on herbivorous insects: indirect effects on sugar maple saplings. Oecologia 125:370-379 Sutherland, J. William (2006). Ecological Census Techniques, 2nd edition. Cambridge University Press. Taylor D.M. (1986). Effects of cattle grazing on passerine birds nesting in riparian habitat. J Range Manage 39:254-258. Villard, M.A., Trzcinski M.K., and Merriam G., (1999). Fragmentation effects on forest birds: relative influence of woodland cover and configuration on landscape occupancy. Conservation Biology 13:774–783. Ward, D. & Salz, D. (1994). Foraging at different spatial scales: dorcas gazelles foraging for lilies in the Negev Desert. Ecology 75:48–58. Weins, J.A. (1989). The Ecology of Bird Communities. Vol. 1. Foundations and patterns. Cambridge: Cambridge University Press. Wilson, E.E., and Wolkovich, E.M. (2011). Scavenging: How carnivores and carrion structure communities. Trends Ecol. Evol. 26: 129–135. Wilson, W.J., Berndt, J., van Rensburg, B.J., Ferguson, J.W.H., and Keith, M. (2008). The relative importance of environment, human activity and space in explaining species richness of South African bird orders. Journal of Biogeography, 35: 342–352.

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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

(African) White backed vulture

Gyps africanus

0.69%

African broadbill

Smithornis capensis

0.07%

African golden oriole

Oriolus auratus

0.03%

0.69%

African green pigeon

Treron calvus

0.13%

African grey hornbill

Tockus nasutus

2.13%

2.30%

African harrier hawk (gymnogene)

Polyboroides typus

0.80%

African mourning dove

Streptopelia decipiens

0.47%

1.26%

African pied wagtail

Motacilla aguimp

0.33%

Arrow marked babbler

Turdoides jardineii

1.15%

F10

Banded martin

Riparia cincta

16.56%

4.02%

F11

Barn (european) swallow

Hirundo rustica

0.53%

F12

Bearded woodpecker

Thripias namaquus

0.20%

F13

Black collared barbet

Lybius torquatus

0.13%

F14

Black coucal

Centropus grillii

0.11%

F15

Black tit

Parus niger

0.27%

0.57%

F16

Blue waxbill

Uraeginthus angolensis

4.45%

7.81%

F17

Broadbilled roller

Eurystomus glaucurus

0.07%

0.11%

F18

Broad-tailed Paradise-whydah

Vidua obtusa

0.69%

F19

Brown hooded kingfisher

Halcyon albiventris

0.23%

F20

Brown snake eagle

Circaetus cinereus

0.13%

F21

Cape turtle dove

Streptopelia capicola

4.45%

13.55%

F22

Cattle egret

Bubulcus ibis

12.03%

F23

Collared sunbird

Anthreptes collaris

0.23%

F24

Common qual

Coturnix coturnix

0.27%

0.23%

F25

Common scimitarbill

Rhinopomastus cyanomelas

0.34%

F26

Coppery tailed coucal

Centropus cupreicaudus

0.11%

F27

Crested barbet

Trachyphonus vaillantii

0.13%

0.11%

F28

Dark capped Bulbul

Pycnonotus tricolor

0.73%

F29

Dickinson's kestrel

Falco dickinsoni

0.53%

0.11%

F30

Emmerald spotted wood dove

Turtur chalcospilos

0.47%

1.15%

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F31

Eurasian golden oriole

Oriolus oriolus

0.07%

0.34%

F32

Flappet lark

Mirafra rufocinnamomea

2.86%

3.56%

F33

Fork tailled drongo

Dicrurus adsimilis

1.13%

F34

Gabar goshawk

Melierax gabar

0.13%

F35

Greater blue eared starling

Lamprotornis chalybaeus

0.23%

F36

Green wood hoopoe

Phoeniculus purpureus

0.80%

F37

Grey lourie/ go away bird

Corythaixoides concolor

1.13%

3.44%

F38

Hamerkop

Scopus umbretta

0.07%

F39

Helmeted guineafowl

Numida meleagris

0.33%

F40

Hooded vulture

Necrosyrtes monachus

0.27%

4.48%

F41

Jameson's firefinch

Lagonosticta rhodopareia

0.34%

F42

Lappet faced vulture

Torgos tracheliotos

0.47%

1.72%

F43

Lesser masked-weaver

Ploceus intermedius

0.80%

F44

Levaillant(stripped) cuckoo

Clamator levaillantii

0.69%

F45

Lilac breasted roller

Coracias caudatus

0.07%

F46

Little bee-eater

Merops pusillus

1.33%

2.07%

F47

Lizzard buzzard

Kaupifalco monogrammicus

0.33%

0.11%

F48

Longtailed paradise whydah

Vidua paradisaea

0.11%

F49

Loughing dove

Stigmatopelia senegalensis

0.86%

1.72%

F50

Madagascar bee-eater

Merops superciliosus

0.20%

F51

Magpie (African long tailed) shrike

Urolestes melanoleucus

5.63%

F52

Meve's starling

Lamprotornis mevesii

2.26%

F53

Meyer's parrot

Poicephalus meyeri

0.13%

0.23%

F54

Namaqua dove

Oena capensis

1.13%

F55

Orange-breasted bush shrike

Telophorus sulfureopectus

0.13%

F56

Pied (jacobin) cuckoo

Clamator jacobinus

0.13%

0.92%

F57

Pied kingfisher

Ceryle rudis

0.13%

F58

Purple indigobird (dusky indigobird)

Vidua purpurascens

0.47%

1.38%

F59

Queen (shaft tailed) whydah

Vidua regia

0.34%

F60

Racket tailed roller

Coracias spatulatus

0.20%

F61

Red billed firefinch

Lagonosticta senegala

0.80%

F62

Red billed francolin

Pternistes adspersus

0.23%

F63

Red billed hornbill

Tockus erythrorhynchus

2.93%

0.92%

F64

Red billed Quelea

Quelea quelea

7.45%

1.72%

2013

F65

Red eyed dove

Streptopelia semitorquata

0.92%

F66

Red faced mousebird

Urocolius indicus

3.13%

0.92%

F67

Red headed weaver

Anaplectes rubriceps

0.07%

F68

Red throated twinspot

Hypargos niveoguttatus

0.60%

1.15%

F69

Red-billed oxpecker

Buphagus erythrorhynchus

0.73%

F70

Red-billed wood hooper

Phoeniculus purpureus

0.13%

F71

Scarlet-chested sunbird

Nectarinia senegalensis

0.23%

F72

Southern red bishop

Euplectes orix

0.27%

F73

Spectacled weaver

Ploceus ocularis

0.33%

F74

Steppe eagle

Aquila nipalensis

0.11%

F75

Swainson's spurfowl (francolin)

Francolinus swainsonii

0.27%

0.46%

F76

Swallow tailed bee-eater

Merops hirundineus

0.11%

F77

Tawny-flanked prinia

Prinia subflava

1.86%

7.50%

F78

Trumpeter hornbill

Bycanistes bucinator

0.27%

F79

Village indigobird

Vidua chalybeata

1.60%

3.00%

F80

Violet backed starling

Cinnyricinclus leucogaster

0.60%

0.34%

F81

White bellied sunbird

Nectarinia talatala

0.34%

F82

White fronted bee-eater

Merops bullockoides

0.33%

0.46%

F83

White throated swallow

Hirundo albigularis

0.33%

F84

White winged widowbird

Euplectes albonotatus

0.33%

0.69%

F85

White-browed Robin-chat

Cossypha heuglini

0.11%

F86

White-browed sparrow-weaver

Procepasser mahali

19.48%

7.58%

F87

Yellow billed oxpecker

Buphagus africanus

1.06%

F88

Yellow throated longclaw

Macronyx croceus

0.12%

F89

Yellow-fronted Canary

Serinus mozambicus

1.15%