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Natural Sciences Master Dissertations

2020 Spatial-temporal variation of pteropus voeltzkowi population and its potential role in restoring forest connectivity in Pemba Island

Mwinyi, Ali Mbarouk

The University of Dodoma

Mwinyi, A. M. (2020). Spatial-temporal variation of pteropus voeltzkowi population and its potential role in restoring forest connectivity in Pemba Island (Master's dissertation). The University of Dodoma, Dodoma. http://hdl.handle.net/20.500.12661/2727 Downloaded from UDOM Institutional Repository at The University of Dodoma, an open access institutional repository. SPATIAL-TEMPORAL VARIATION OF PTEROPUS VOELTZKOWI POPULATION AND ITS POTENTIAL ROLE IN RESTORING FOREST CONNECTIVITY IN PEMBA ISLAND

ALI MBAROUK MWINYI

MASTER OF SCIENCE IN NATURAL RESOURCES MANAGEMENT THE UNIVERSITY OF DODOMA DECEMBER, 2020 SPATIAL-TEMPORAL VARIATION OF PTEROPUS VOELTZKOWI POPULATION AND ITS POTENTIAL ROLE IN RESTORING FOREST CONNECTIVITY IN PEMBA ISLAND

BY ALI MBAROUK MWINYI

A DISSERTATION SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE IN NATURAL RESOURCES MANAGEMENT OF THE UNIVERSITY OF DODOMA

THE UNIVERSITY OF DODOMA DECEMBER, 2020 DECLARATION AND COPYRIGHT I, Ali Mbarouk Mwinyi, hereby declare that this dissertation is my own original work and that it has not been presented and will not be presented to any other university for a similar or any other degree award.

Signature

No part of this dissertation may be reproduced, stored in any retrieval system, or transmitted in any form or by any means without prior written permission of the author or the University of Dodoma. If transformed for publication in any other format shall be acknowledged that, this work has been submitted for degree award at the University of Dodoma.

i CERTIFICATION The Undersigned certifies that she has read and hereby recommends for acceptance by the University of Dodoma a dissertation entitled ‘’Spatial-Temporal Variation of Pteropus voeltzkowi Population and Its Potential Role in Restoring Forest Connectivity in Pemba Island, Zanzibar’’ in partial fulfilment of the requirements for the Degree of Master of Science in Natural Resources Management of the University of Dodoma.

Dr. Happiness J Nnko

Signature Date

(Supervisor)

ii DEDICATION This dissertation is first dedicated to God under whose care I did my studies safely and successfully. Secondly, the work is dedicated to my mother Msim Ali Ahmed, and my son Nadir, Nasihuddin, Nashal and Irshad who bore the consequences but remained my unfaltering source of inspiration and encouragement.

iii ACKNOWLEDGEMENTS Glory, honour, adoration, excellency are ascribed into the Almighty God, who in His infinite love guided me to the success of this work. Enthusiastically, words are inadequate to express my appreciation to Him. I avail myself of this opportunity, with pleasure, to follow the good tradition of recognizing all those who, in one way or the other contributed to make my studies a success.

I take this opportunity to express my gratitude to my supervisor, Dr Happiness J Nnko for her patience, guidance, encouragement, constructive criticisms, and readiness to assist. Her support and understanding kept me going up to completion of this work. I also express my sincere gratitude to my brother Mwinyi Othman for supporting me over the whole course time. I would have not achieved this stage, if it had not been for moral and material support that had been extending to me from time to time. Also, I would like to thank all staff and management in the Department of forestry in Pemba, special thanks to Mr. Suleiman Mohd Salim, Mr. Salum, Mr. Said Juma Ali, Mr Ali Hamad, Mr Hamad Juma, and Mrs Sada. To the very special people in my life, my mother Msim Ali, my son, my brother Ibrahim Mbarouk, my lovely sisters Asha Mbarouk.

I appreciate the contribution of my family towards the success of this work. They missed me throughout the period of my study and yet were patient to see that, this academic endeavour is realized. I thank them all.

Finally, I have to recognize my friends, Mr. Madalisto Henry Mwale, Mr Juma Ali Juma, Mr Rajab Hamad and Mr Amir Mohamed Amir your kindness is highly appreciated and God bless you.

iv ABSTRACT This study examined spatial and temporal variation of Pteropus voeltzkowi population and its potential roles in restoring forest connectivity in Pemba Island where four villages including; Wete, Ngezi, Kidike, and Msitu mkuu were involved. The study adopts longitudinal research design whereby the data were collected once in a dry season and once in a wet season. The collected data were then processed and analysed using R computer software. Specifically, the study aimed at assessing variation of Pteropus voeltzkowi population during dry and wet season in these four villages, habitat preference and potential roles of Pteropus voeltzkowi in restoring forest connectivity in Pemba Island. The findings revealed that population of Pteropus voeltzkowi significantly varied between the seasons with high relative abundance in dry season compared to wet season and between villages. Further there was significant decrease of Pteropus voeltzkowi relative abundance during wet season (β= -63.5, SE = 25.4) compared to dry season. On the other hand, there was a significant increase of Pteropus voeltzkowi relative abundance in Wete village (β= 142.1, SE=46.6, df= 138, p - value = 0.002) and significant decrease of Pteropus voeltzkowi relative abundance in Ngezi (β = -105.3, SE=46, df=138, p-value=0.0237) compared to Kidike. In addition, there was an increase in relative abundance of Pteropus voeltzkowi on every increase of roosting trees canopy diameter (β= 10.4, SE = 2.9, df = 138, p- value= 0.0005) and height (β= 4.5, SE = 1.95, df = 138, p- value= 0.0219). Furthermore, observations revealed that, out of twelve plant species only ten plant families were identified as sources of food for Pteropus voeltzkowi during the study and the preferred food type by Pteropus voeltzkowi were fruits followed by nectar and leaves. Lastly, experimental results revealed that, all the seeds that were processed by Pteropus voeltzkowi significantly showed better germination rate than those extracted from unprocessed when they were planted in a plastic beaker under natural condition (X2 = 16.266, df= 3, P- value =0.001) These findings can be used to inform spatial temporal species distribution models intended to cover extensive area and longer time. In addition, it can help to develop a village specific conservation strategy to alleviate potential human-bats conflicts. It highlight the potential role of Pteropus voeltzkowi in enhancing forest connectivity and thus additional reason for conserving this species.

v TABLE OF CONTENTS DECLARATION AND COPYRIGHT ...... i CERTIFICATION ...... ii DEDICATION ...... iii ACKNOWLEDGEMENTS ...... iv ABSTRACT ...... v TABLE OF CONTENTS ...... vi LIST OF TABLES ...... ix LIST OF FIGURES ...... x LIST OF APPENDICES ...... xi LIST OF ACRONYMS AND ABBREVIATIONS ...... xii

CHAPTER ONE ...... 1 INTRODUCTION ...... 1 1.1 Chapter Overview ...... 1 1.2 Background ...... 1 1.3 Statement of the problem and justification ...... 2 1.4 Objectives ...... 4 1.4.1 Overall objectives...... 4 1.4.2 Specific Objectives...... 4 1.4.3 Hypothesis ...... 4 1.5 Research Questions: ...... 4 1.6 Significance of the Study ...... 4 1.7 Scope of the study ...... 5

CHAPTER TWO ...... 6 LITERATURE REVIEW ...... 6 2.1 Chapter Overview ...... 6 2.2 General description of the Pteropus voeltzkowi ...... 6 2.2.1 Physical Description...... 6 2.2.2 Taxonomy ...... 7 2.2.3 Population of Pteropus voeltzkowi in Pemba Island ...... 7 2.2.4 Lifespan of Pteropus voeltzkowi ...... 8

vi 2.2.5 Habitat of Pteropus voeltzkowi ...... 8 2.2.6 Ecosystem Roles ...... 8 2.2.7 Conservation Action of Pteropus voeltzkowi ...... 8 2.2.8 IUCN Justification ...... 10 2.3 Definition of the key terms ...... 11 2.3.1 Seasonality ...... 11 2.3.2 Roosting ...... 12 2.3.3 Forest restoration ...... 12 2.3.4 Habitat Destruction and Fragmentation ...... 13 2.4 Geographical distribution of the genus Pteropus ...... 13 2.5 Ecological functions of Pteropus species ...... 13 2.5.1 Seeds Dispersal ...... 13 2.5.2 Pest Regulator ...... 14 2.5.3 Soil Fertility and Nutrient Distribution ...... 14 2.6 Empirical Literature Review ...... 16 2.6.2 Spatial- temporal population dynamic of Pteropus species ...... 16 2.6.3 Flying fox – Potential forest restoration...... 17 2.7 Research Gap ...... 18 2.8 Conceptual Frame work ...... 18

CHAPTER THREE ...... 20 RESEARCH METHODOLOGY ...... 20 3.1 Chapter Overview ...... 20 3.2 Selection of the Study Area and Justification ...... 20 3.3 Climatic Conditions of the Study Areas...... 20 3.4 Research design and data collection procedures ...... 21 3.5 Objective one ...... 22 3.6 Objective two ...... 23 3.7 Objective three ...... 24 3.8 Data Processing and Analysis ...... 25 3.9 Data quality and Control ...... 26 3.9.1 Validity and Reliability ...... 26 Validity ...... 26

vii Reliability ...... 26 3.10 Ethical Consideration ...... 27

CHAPTER FOUR ...... 28 RESULTS AND DISCUSSION ...... 28 4.1 Chapter Overview ...... 28 4.2 Spatial-temporal variation of the Pteropus voeltzkowi population in study area . 28 4.3 Habitat and diet preference of Pteropus voeltzkowi ...... 38 4.4 Potential roles of the Pteropus voeltzkowi in restoring forest connectivity through germination success ...... 40

CHAPTER FIVE ...... 42 SUMMARY, CONCLUSION AND RECOMMENDATIONS ...... 42 5.1 Chapter Overview ...... 42 5.2 Summary of the Finding ...... 42 5.3 Conclusion ...... 43 5.4 Recommendations ...... 44 5.5 Recommendation for Further Studies ...... 44 REFERENCES ...... 45 APPENDICES ...... 49

viii LIST OF TABLES Table 4. 1: Linear mixed-effects model between Pteropus voeltzkowi, village, and season DBH, Canopy diameter and Tree Height...... 38 Table 4. 2: Food providing plant for Pteropus voeltzkowi...... 39 Table 4. 3: Average germination time and rate selected plant species that are frequently consumed by Pteropus voeltzkowi ...... 41

ix LIST OF FIGURES Figure 2. 1: Signboard at Msitu Mkuu roost site indicating that, bats are protected species and hunting is not allowed...... 9 Figure 2. 2: Conceptual frame work...... 19 Figure 3. 1: Map of the Pemba Island showing the study areas...... 21 Figure 3. 2: Research Assistants take measurements on dbh of the plant at Msitu mkuu roost site...... 23 Figure 4. 1: The variation of Pteropus voeltzkowi relative abundance between dry and wet season...... 28 Figure 4. 2: Variation of Pteropus voeltzkowi relative abundance across study villages...... 29 Figure 4. 3: Variation of Pteropus voeltzkowi relative abundance between habitat types studied...... 31 Figure 4. 4: Seasonal variation of Pteropus voeltzkowi relative abundance between (A) habitat types and (B) villages...... 32 Figure 4. 5: Relative abundance of Pteropus voeltzkowi across various tree/plant species ...... 34 Figure 4. 6: Relative abundance of Pteropus voeltzkowi across various tree species in different villages...... 35 Figure 4. 7: Relative abundance of Pteropus voeltzkowi across various tree species in both residential and forested habitats...... 36 Figure 4. 8: Roosting site canopy diameter (M) and height (M) variation between habitat types and villages...... 37 Figure 4. 9: Preferred plant parts as source of foods for Pteropus voeltzkowi...... 40

x LIST OF APPENDICES Appendix 1: Assessment on spatial-temporal variation of Pteropus voeltzkowi population...... 49 Appendix 2: Habitat preference of Pteropus voeltzkowi ...... 53 Appendix 3: Estimate germination success of seeds taken from bats droppings (Processed) with those from fresh fruits (Unprocessed) under progressively more natural conditions...... 54 Appendix 4: Number of species counted during dry season at Ngezi roost site...... 55 Appendix 5: Number of species counted during wet season at Ngezi roost site ...... 57 Appendix 6: Number of species counted during dry season at Msitu Mkuu roost site ...... 59 Appendix 7: Number of species counted during wet season at Msitu Mkuu roost site ...... 61 Appendix 8: Number of species counted during dry season at Kidike roost site ...... 63 Appendix 9: Number of species counted during wet season at Kidike roost site ...... 65 Appendix 10: Number of species counted during dry season at Wete roost site...... 67 Appendix 11: Number of species counted during wet season at Wete roost site...... 69 Appendix 12: Habitat preference of P.voeltzkowi...... 71 Appendix 13: Mean number of days of seed germination between processed and unprocessed seeds from different plant species and number of seeds germinating when they planted in a plastic beaker...... 72 Appendix 14: Result from Linear mixed-effects model...... 73 Appendix 15: Research Clearance from the University of Dodoma...... 76

xi LIST OF ACRONYMS AND ABBREVIATIONS DBH Diameter at breast height

FFI Fauna and flora international

IUCN International union for conservation of nature.

LDD Long dispersal distance.

PFF Pemba flying fox

SDD Small dispersal distance.

xii CHAPTER ONE INTRODUCTION 1.1 Chapter Overview The focus of the study was to assess the spatial-temporal variation of Pteropus voeltzkowi population and its potential role in restoring forest connectivity in Pemba Island. This chapter gives a glimpse on what the study was about. It comprise of the background of the study, statement of the problem, objectives of the study, research question, research hypothesis, significance of the study and scope of the study.

1.2 Background The Old World fruit bats are among the world old fruit eating bats and the most important ecosystem services providers in the animal kingdom (Abedi-lartey & Dechmann, 2016). Pteropus take vital roles in ecosystem including pollination, seed spreading and insect regulator and different ecological and commercial importance of plant depend on flying fox (Entwistle & Corp, 1997).

The flying fox prefers to roost in tropical and sub-tropical zones, normally close to coastal zones (Carter, 2005) and live in huge and well-exposed plants, voluminous and mechanically unchanging (Granek, 2000). The size of the camp vary seasonally but at the time of reproduction, they group themselves as in a single roost (Mathur, Priya & Kumar, 2012).

Pteropus species are significant for the existence of more than 114 plant species of the biosphere (Mickleburgh, Hutson & Racey, 1992) and are widely acknowledged for positive roles such as plant pollinators, seed dispersers, and mediators for keeping plant communal diversity (Mickleburgh et al.,1992). For instance, Pteropus plays a great role of enhancing regeneration, dispersal of plant seeds and enhance community diversity in Island (Giannini & Acosta, 2012).

Generally, the Pteropus species is considered an ecosystem builders of the Islands due to the fact that, they are main pollinators and large seed-fruit disperser. According to Kunz (2011) flying fox contributes to sustainable ecological amenities where they occur, they are fundamental for tropical forest maintenance because they disperse at least 139 plant genera from 58 families as they travel up to 40 km from

1 roosting to foraging sites and they transport up to 200 g of the fruit/seed over a longer distance and thus ensuring vegetation connectivity between fragmented forest area.

Likewise, it has been reported that, fruit bats help to increase the rate of the plant germination by depositing the seeds with manure (Djossa et al., 2008), hence are important in structuring vegetation communities. Despite the ecological importance of these species, continue habitat modification through forest clearing for agriculture is leading to loose of roosting and foraging sites for Pteropus voeltzkowi (Daszak Tabor, Kilpatrick, Epstein & Plowwright, 2004).

Flying fox of the species Pteropus voeltzkowi are endemic to the Island of Pemba, Zanzibar, and was first documented by the scientist in 1912 (Entwistle & Corp, 1997). According to Seehausen (1991) Pteropus voeltzkowi are more abundant in western part of the Island which is dominated with rainforest compared to eastern drier part (Entwistle & Corp, 1997). Mangoes are the main diet sources, but they also prefer to eat other cultivated and native fruits such as breadfruit, figs, flower and leaves (Entwistle & Corp, 1997).

Pteropus voeltzkowi are social animals and their functional characteristic is evident in high population density. Like other plant-visiting bats, Pteropus voeltzkowi is believed to play a key role in structuring the forestry community (Boyles, Cryan & Kunz, 2011).

1.3 Statement of the problem and justification The Pteropus voeltzkowi is considered a key species that play a significant ecological functional role of structuring the ecological community in Pemba Island. The species is important pollinator and major disperser of seed and fruits, however, these ecological functional behaviour is an integral part of population density (Boyles et al., 2011).

For the past few decades, Pteropus voeltzkowi population estimates have varied considerably in space and time in the Pemba Island (Robinson, Bell, Saleh, Suleiman & Barr, 2010). In early 1990s population was drastically low and the species was considered to be on the edge of extinction, thus International union for conservation 2 of nature (IUCN) listed it as critically endangered. Due to the decline the ecological importance of Pteropus voeltzkowi, conservation efforts began on the Island, concentrating on ecological importance of the Pteropus voeltzkowi and also general environmental education to primary school kids, grassroots communities living adjacent to the roosting sites and establishment of community based and environmental projects that targeted reduction of Pteropus voeltzkowi protein dependency (Robinson et al., 2010).

According to Entwistle (1997) the Phoenix Zoo in Arizona also established in the same decade a captive breeding platform to safeguard the survival of the Pteropus voeltzkowi. All these efforts yielded an increasing population and thus in 2005 IUCN and CITIES Appendix II re-listed Pteropus voeltzkowi as a vulnerable species (Robinson et al., 2010). Despite the efforts by various conservation stakeholders, Pteropus voeltzkowi is still threatened by habitat loss, disturbance and to a small extent unsustainable hunting for food. Loss of habitat is mainly caused by clearance of forest for expansion of agriculture and human settlement (Mickleburgh et al., 2008).

Most of Island primary forests have declined and only remnants remain. While IUCN red list categorized Pteropus voeltzkowi as vulnerable species, the pace of development in the Pemba Island may worsen its population status. Their greater ecological functional roles of pollination, dispersal and restoring forest connectivity in the Island necessitate the need to balance development and the conservation objectives in the Island. It is expected that, without Pteropus voeltzkowi, the forest ecosystem of Pemba Island may start to crumble and hence communities wellbeing due to lose of food plant harvest, fire wood, medicinal plants and habitat for the numerous other forest dependent species.

The reconciliation of development and conservation objectives can be enriched by clear understanding of the species spatial and temporal population dynamics, which is the interest of this study. Potential this can inform the status of Pteropus voeltzkowi and can serve as the bio indicator for health ecosystem in the Island of Pemba.

3 1.4 Objectives 1.4.1 Overall objectives The overall objective of this study was to assess the spatial-temporal variation of Pteropus voeltzkowi population and its potential role in restoring forest connectivity in the Pemba Island.

1.4.2 Specific Objectives The specific objectives of the study were to: i) Assess spatial-temporal variation of Pteropus voeltzkowi population ii) Assess habitat preference of Pteropus voeltzkowi iii) Assess the potential roles of the Pteropus voeltzkowi in restoring forest connectivity in the Pemba Island

1.4.3 Hypothesis

H0 There is no variation in spatial-temporal distribution of Pteropus voeltzkowi in the study

H1 There is variation in spatial-temporal distribution of Pteropus voeltzkowi in the study

1.5 Research Questions: This study asked the following questions: i) How population of Pteropus voeltzkowi varies in space with wet and dry season? ii) What are the preferred habitat for Pteropus voeltzkowi in Pemba Island? iii) Does the Pteropus voeltzkowi have a potential for restoring forest connectivity?

1.6 Significance of the Study The findings in this study provide information that can inform stakeholders about the space and time of Pteropus voeltzkowi relative abundance and thus devising a fit for the purpose of sensitization program.

The understanding of spatial and temporal variations of Pteropus voeltzkowi can deliver important information needed when developing predictive models of the spatial-temporal dynamics of Pteropus voeltzkowi and their ecological and human 4 wildlife conflicts potentials roles. The information can as well be useful in adding on available data of Pteropus voeltzkowi population trends that can inform the conservation strategies for this species.

1.7 Scope of the study Flying fox comprises huge number of species in Animal kingdom. This study discusses only one species of Pteropus who is Pteropus voeltzkowi found in Pemba Island, the study was conducted in four villages including Wete, Ngezi, Msitu Mkuu and Kidike. The study villages were selected based on; (i) Presence of suitable habitats for Pteropus voeltzkowi which was a target population, (ii) Easy accessibility of the villages, (iii) Unknown distribution of Pteropus voeltzkowi in space and time and (iv) Presence of potential threats to Pteropus voeltzkowi.

5 CHAPTER TWO LITERATURE REVIEW 2.1 Chapter Overview This chapter present literature review about spatial-temporal variation of Pteropus voeltzkowi population based on the various studies conducted within and outside of Zanzibar, Tanzania. The main aspects that are discussed in this part include, general description of Pteropus voeltzkowi, definition of the key terms, geographical distribution, ecological function, review of empirical studies, identification of the research gaps and conceptual framework.

2.2 General description of the Pteropus voeltzkowi The purpose of this sub-section is to address general description of Pteropus voeltzkowi, taxonomy, conservation status, habitat type, ecology and population.

2.2.1 Physical Description Pteropus voeltzkowi belong to the order Chiroptera (Mickleburgh et al., 2008). Pteropus voeltzkowi is pervasive to the Island of Pemba in Zanzibar having wing span of 1.6 meters (5 ft 3 in) and are unique among the largest species of fruit bat, they have fox-like appearance, black ear, nose and wings (Mickleburgh et al., 2008). Pteropus voeltzkowi are phytophagous species, feeding primary on floral resource and fruits in an extensive variety of vegetation communities, comprising sealed forest, gallery forest, eucalypt open forest, and woodland, mangroves forest and profitable fruits crops.

This highly mobile species forages at night and can disperse seeds and pollen over large distance. In Pemba Island, Pteropus voeltzkowi are found at all region from North to South region.

6

Figure 2. 1: Image of Pteropus voeltzkowi

2.2.2 Taxonomy Pteropus voeltzkowi is scientifically classified as follows: Kingdom Phylum Class Order Family Genus Species Animalia Chordate Mammalia Chiroptera Pteropodidae Pteropus Pteropus voeltzkowi

Taxon Name: Pteropus voeltzkowi Common Name: Pemba flying fox

2.2.3 Population of Pteropus voeltzkowi in Pemba Island In the early 1990s there was evidence, that the population of this species had been reduced to a few hundred animals (Seehausen, 1991) with an approximation of 2,800 to 3,600 individuals. In 1992, the mainstream being establish at just two roost sites (Carter, 2005). In 1995 Entwistle (1997) predicted a population of 4,600 to 5,500 individuals, with 94% of bats originate at 10 roosts of 41 in total. Above the consequent year’s, surveys showed by local teams on the Island conveyed growing population counts (Trewhella, Rodriguez, Corp, Entwistle, Garrett & Granek, 2005). A complete review in 2008 delivered a population estimation between 18,200 and 22,100 individuals (Robinson et al., 2010) and they were dispersed through 44 active

7 roost sites, with 87% of the population originate at four roost sites. Roosts reached from unsociable individuals to clusters of up to 5,040 bats

2.2.4 Lifespan of Pteropus voeltzkowi No evidence is available on the life span of Pteropus voeltzkowi (Mickleburgh et al., 1992). Nevertheless, other members of the same genus are described to have lived as long as 30 years (Carter, 2005).

2.2.5 Habitat of Pteropus voeltzkowi Pteropus voeltzkowi occur throughout coastal regions as well as arid landscapes of Pemba Island. Limited knowledge from recent studies suggests that these species often congregate at camps, such as saltwater mangroves, bamboo, and closed forests. Selection of such congregation sites may be determined by seasonal variation, as well as by other factors; such as human hunting, and climatic fluctuations (Robinson et al., 2010).

2.2.6 Ecosystem Roles Pteropus voeltzkowi take vital roles in ecosystem facilities including pollination, seed spreading and insect regulator and different ecological and commercial importance of plant depend on flying fox (Boyles et al., 2011).

2.2.7 Conservation Action of Pteropus voeltzkowi Current alertness raising on the significance and exclusivity of the endemic fruit bat, and the essential for hunting controls and avoidance of roost site disruption, has been taken on through environmental education programmes (Seehausen,1991) Communities are straight leading active conservation platforms for their local roosts, and three of the four largest roosts were associated with community protection schemes (Robinson et al., 2010). The native community has been reinforced to develop bat related ecotourism activities.

8

Figure 2. 2: Signboard at Msitu Mkuu roost site indicating that, bats are protected species and hunting is not allowed

Figure 2. 3: Signboard at Kidike roost site which explain community conservation effort on Pteropus voeltzkowi

9 2.2.8 IUCN Justification The status of all Pteropus species were assessed using the IUCN Red List Categories (IUCN, 1994), twelve Pteropus species are confirmed as Extinct. A further 238 species, almost a quarter of the total, are threatened, Critically Endangered, Endangered or Vulnerable. In 2005 IUCN and CITIES Appendix II re-listed Pteropus voeltzkowi as a vulnerable species (Robinson et al., 2010).

2008 – Vulnerable (VU)

2004 – Vulnerable (VU)

1996 – Critically Endangered (CR)

1994 – Endangered (E)

1990 – Endangered (E)

1988 – Vulnerable (V)

10

Figure 2. 4: Show community effort in conservation of Pemba flying fox at Mjini Ole Source: Kidike community conservation

2.3 Definition of the key terms The purpose of this sub-section is to define and describe various key concepts related to the study, including seasonality, roosting, forest restoration and habitat fragmentation.

2.3.1 Seasonality Seasonality refer to the variations that happen at exact regular intervals less than a year, such as weekly, monthly, or trimestral. In this study dry season was defined as a period of short rainfall between January and February.

11 Also wet season in this study was defined as the period in a year where average rainfall occur particularly from March and May.

2.3.2 Roosting Roosting refers to the animal behaviour where a group of individual typical of the same species congregates on the same area (Carter, 2005), Roost are vital to flying fox since is the place where they actively get involved in mating, rearing offspring and facilitate the social living. Bats select roost that provides suitable thermal condition which is necessary for metabolic and reproductive demand (Boyles et al., 2011).

Figure 2. 5: Pteropus voeltzkowi roosting in Antiaris toxicaria roost tree

2.3.3 Forest restoration Forest restoration is well-defined as “actions to re-instate ecological processes, which speed up reclamation of forest structure, ecological operational and biodiversity levels, Restoration of tropical forest depend in large part on seed dispersal by fruit- eating animals that transport seeds into planted forest patches (Schupp & Jadano, 2010).

12 2.3.4 Habitat Destruction and Fragmentation Habitat destruction is a process which happens when a natural habitation such as forest or wetland is changed so dramatically that no longer support the species in it originally persistent (Pimm & Raven, 2000). Habitat damage is considered as the most key drivers of the species disappearance worldwide, agriculture is the one among the biggest causes of habitat destruction.

2.4 Geographical distribution of the genus Pteropus The genus Pteropus constitutes the largest order of Megachiroptera, with 57 species which are distributed from Australia and south-east Asia to the coast of Africa. Pteropus is largely on Island, genus with 35 species endemic to Islands or Island groups (Mickleburgh et al., 1992).

Pteropus voeltzkowi are distributed widely across all areas in Pemba Island including Wete port, Kidike, Msitu Mkuu, Mbiji, Mchanga wa Kwale, Ngezi forest reserve, Kiuyu Minungwini, Kiwani Mchanga Mdogo and Makombeni (Mickleburgh et al., 1992).

2.5 Ecological functions of Pteropus species Pteropus species are the animal of strange ecological and economic importance and they are succeeding most species rich order of mammal with great ecological variety in the tropics (Kunz, Elizabeth, Dana, Lobova & Fleming, 2011). Pteropus have long been postulated to play important ecological roles in prey and predator, arthropod suppression, seed dispersal, pollination, material and nutrient distribution, and recycle (Kunz et al., 2011)

2.5.1 Seeds Dispersal According to Duncan (1999) seeds dispersal is the biggest approach in which animals subsidize for ecosystem succession by dropping seeds from one area to another, the role played by Pteropus species in dispersing the seed is wonderful, a tropical trees are modified for seed dispersal by animal particular by flying fox and birds. Night-time foraging fruit flying fox are more obedient than birds since them covering long distance each night defecating in flight and scattering for more seeds across cleared area (Horner, Fleming & Sahley, 1998).

13 Unlike most of seeds dispersal by vertebrate that disperse close to parent plant with only 100-1000 m away, the seeds dispersed by flying fox relative far away 1-2 km (Horner et al., 1998), this condition assist to maintain species diversity by hosting seeds from outside disturbed areas (Muscarella & Fleming, 2007).

2.5.2 Pest Regulator There are over 1200 species of Pteropus and those above two third are insectivorous to some degree (Kunz et al., 2011). Pteropus species particularly pregnant and lactating female which have great energy demand, can eat large number of Arthropods and many of these species consume agricultural pest and disease vectors (Kunz et al., 2011). The influence of Pteropus species to pest regulator has only recently started to be evaluated (Boyles et al., 2011).

2.5.3 Soil Fertility and Nutrient Distribution Pteropus voeltzkowi play an important ecological role in soil fertility and nutrient spreading due to their relatively high mobility and the use of different habitats for roosting and foraging, which facilitates nutrient transfer within ecosystems (Kunz et al., 2011). However, the suspected importance of nutrient transfer by Pteropus in overall ecosystem function is probably relatively low when compared with microhabitat conditions (Boyles et al., 2011). For soil fertility and nutrient distribution, Pteropus voeltzkowi has a great ecological potential as it contributes a lot in nutrient redistribution, from nutrient rich sources to nutrient-poor regions (Muscarella & Fleming, 2007).

2.5.4 Pollination Some Pteropus species primarily the two families (Pteropodidae in the Old World and Phyllostomidae in the New World) play important roles in plant pollination (Kunz et al., 2011). Although bat pollination is relatively uncommon when compared with bird or insect pollination, it involves an impressive number of economically and ecologically important plants (Kunz et al., 2011). Particularly, beyond the economic value of plant pollination and seed dispersal services, plant-visiting bats provide important ecological services by facilitating the reproductive success and the recruitment of new seedlings (Fleming, Geiselman & Kress, 2009). Bat pollination occurs in more than 528 species of 67 families and 28 orders of angiosperms

14 worldwide (Fleming et al., 2009). Pteropodidae bats are known to pollinate flowers of about 168 species of 100 genera and 41 families and phyllostomid bats pollinate flowers of about 360 species of 159 genera and 44 families (Fleming et al., 2009).

Figure 2. 6: flower pollination by Pteropus species Source: The World Technology Network.

2.5.5 Disease Transition and Contamination Pteropus species are hosts to a range of zoonotic and potentially zoonotic pathogens. They differ from other disease reservoirs because of their unique and diverse lifestyles, including their ability to fly, often highly gregarious social structures, long life spans, and low fecundity rates (Calisher, Childs, Field, Holmes & Schountz, 2006). They represent a potential epidemiologic of several diseases that can be fatal to humans, including rabies, Ebola, leptospirosis, histoplasmosis, and pseudotuberculosis (Calisher et al., 2006). Pteropus are reservoirs of several pathogens, whose spread may be related to physiological stress associated with habitat loss or alteration (Calisher et al., 2006). Human activities that increase exposure to bats will likely increase the opportunity for infections (Calisher et al., 2006).

15 Like bird droppings, Pteropus can contain a potentially infectious fungus Histoplasma capsulatum that causes lung infection known as histoplasmosis (Calisher et al., 2006).

2.6 Empirical Literature Review This section examines various literature on spatial-temporal population dynamic of Pteropus voeltzkowi. The section also discuss empirical study that address potential role of Pteropus voeltzkowi in restoring forest connectivity in order to provide an empirical supports of the parameters used in this study.

2.6.1 Status of Pteropus species World Wide Currently Pteropus species population are decline worldwide as result of numeral factors such as habitat damage and destruction, disturbance to roost, exposure to toxin, human hunting pressure, introduce predators and disease outbreak (Jone, Purvi & Gittleman, 2009). This make more difficult to makes it a general conclusion about their population dynamics (Mickleburgh et al., 2008). Flying fox of genus “Pteropus” are the more threatened bats than any other bats type with the largest quantity of critically rare species (Schipper, Chanson & Chiozza, 2008). Pteropus are vulnerable to environmental change affecting longevity and reproductive success and they result species to change in location based on seasonality (Schipper et al., 2008).

Many southwest Asia countries consists of high level of bats abundance at the same time lack of legal regulation resulting in huge bats decline issue (Schipper et al., 2008). Pteropus species much affected by hunting process (Mickleburgh et al., 2008) which stemmed in the extinction of many Pteropus species. Flying fox are targeted in the bush meat trade since of their large body mass and their propensity to aggregate in large groups which could increase easy of capture (Mickleburgh et al., 2008).

2.6.2 Spatial- temporal population dynamic of Pteropus species Pteropus species provide idyllic models for studying the effects of spatial and temporal variation. Spatial-temporal movement of Pteropus species play an essential role in modelling biodiversity patterns across spatiotemporal scales. It affects biodiversity directly and indirectly (Schweiger, Heikkinen, Harpke, Hickler & Klotz,

16 2012). Seasonal movements frequently follow seasonal fluctuations in resource availability, and can cover thousands of kilometres. However, spatial and temporal variability in environmental circumstances may affect all types of movement across all scales, from local to global, generating new occasions for the evolution of movement parameters. The ability to move rapidly away from unsuitable conditions, or from threats can potentially have considerable adaptive value.

2.6.3 Flying fox – Potential forest restoration About 300 plants species really depend on bat for seed dispersed and these plants offer almost 500 products such as foods, medicines and timber. More over bats play important part in forest reinforcement because of their capability to return feasible seeds in their gut for numerous hours (Carter, 2005).

Flying fox are the important in seeds dispersal and pollinators in several economically important trees (Carter, 2005). The ability of bats to carry pollen over longer distance facilitates crossing pollination (Carter, 2005), since the bats are highly mobile their movement is expected to affect the quantity and quality of pollen flow. The role of bats in the ecosystem functioning is undervalued because of their nocturnal cryptic life cycles (Fujita & Tuttle, 1991).

Bats have long been involved in imperative ecosystem service such as cross- pollination, seed dispersal and insect suppression. It is estimated that about 99% of potential crop pest are restricted by natural ecosystem of which some fraction can be endorsed to predation by bats (Fujita & Tuttle, 1991).

Bats plant interaction such as crosspollination and seed dispersal are mutualistic population interaction in which plant deliver a nutritional reward (nectar, pollen and fruit pulp) for useful services through the highly specialized bats flower mutualism In addition the flower visited by bat reportedly produce about seven time more pollen grain than the flower catering to humming birds but the reasons behind the increasing not well understood (Schweiger et al., 2012).

Determinations should be made to safeguard the bats are safe in order to keep our forest ecosystem healthy for the future. Therefore to sympathetic local perception on bats is essentially for effective long term effort in protection (Schweiger et al., 2012). 17 2.7 Research Gap In the review of the available literature it is revealed that many studies available at Pemba Island have concentrated on the recovery of the vulnerable Pteropus voeltzkowi (Robinson et al., 2010). Some of literature focuses on assessing diet of Pteropus voeltzkowi, current status and population fluctuation (Aziz & Olival, 2014).

In spite of the knowledge gained from the previous studies on a Pteropus voeltzkowi, there are still number of research gaps that characterize Pteropus voeltzkowi and its potential role in restoring forest connectivity. Firstly, there is still a debate on the reliability of the data used to explain the current status and population fluctuation of Pteropus voeltzkowi (Entwistle et al., 1997).

Second, the spatial-temporal variation of Pteropus voeltzkowi population and potential role in restoring forest connectivity is still unclear. The second research gap is the driving force that made the necessity of this study.

2.8 Conceptual Frame work The conceptual framework (Figure 2.1) explains and guides the key variables, concepts and the variable relationships of the study. The framework show that due to the change in seasonality (wet and dry), diet sources, plant species, plant parts (fruits, seeds, flower, pollen, leaves and nectar) the spatial-temporal variation of Pteropus voeltzkowi is intensified and these are independent variable which cause spatial- temporal variation of Pteropus voeltzkowi.

Furthermore, the frame work shows the factors like plant relative abundance, distribution in space and time, seed dispersal and germination rate as a dependent variables. Lastly, the relationship between variables is illustrated by one directional arrow showing a two-ways relationship.

18

Figure 2. 7: Conceptual Frame Work

19 CHAPTER THREE RESEARCH METHODOLOGY 3.1 Chapter Overview This chapter describes the research design, study area and methodology that were adopted in data collection and method of data analysis. The chapter also discusses data quality and control applied the study including the ethical consideration.

3.2 Selection of the Study Area and Justification This study was conducted in four villages within Pemba Island, Zanzibar. The Island is located approximately 50 km from the mainland and it lies between 40˚52' and 60˚31' South of equator. The four villages recruited in this study were Wete, Ngezi, Micheweni (Msitu mkuu) and Kidike (Figure 3). The study villages were selected based on (i) Presence of suitable habitats for Pteropus voeltzkowi which was a target population (ii) Easy accessibility of the villages (iii) Unknown distribution of Pteropus voeltzkowi in space and time and (iv) Presence of potential threats to Pteropus voeltzkowi.

3.3 Climatic Conditions of the Study Areas The Pemba Island is characterized by tropical climate with long rain seasons occurring between March and June and the short rain seasons starting in October through December. The average rainfall for Island is about 1500 mm per annual. The highest temperature occurs during the short dry season with maximum of 290C and minimum of 21.10C.

20

Figure 3. 1: Map of the Pemba Island showing the study areas

3.4 Research design and data collection procedures To understand the spatial-temporal variation of Pteropus voeltzkowi, the study employed longitudinal research design whereby data were collected once in dry season (January and February 2020) and once in rainy season (March and May 2020). The design was adopted to give an indication on whether seasonality and habitat types affects spatial and temporal distribution. Data were collected from February to April to mark for dry and wet season respectively.

21 Depending on the objectives, a variety of methods and instrument were used in data collection and they included total count, seed germination experimentation, direct/physical observation, and indirect field observation as detailed in the subsequent sub-section.

First of all, reconnaissance survey was done in all four villages known roosting sites and surrounding Pteropus voeltzkowi potential habitats in order to generally assess the occurrence of Pteropus voeltzkowi and their habitats conditions. The abundance and distribution of Pteropus voeltzkowi population were determined by using total count method due to small population size and the study took advantage of roosting behaviour since they could be found in the same place most of the time. The general ecology of the area including habitat type’s observation of Pteropus voeltzkowi was done on each counting day. The restoration potential role was implemented through collection and germination rate assessment of both Pteropus voeltzkowi processes (taken from bat droppings) and unprocessed (ripe fresh fruits) seed while habitat preference was assessed through direct field observation.

3.5 Objective one Assessment of spatial and temporal variation of Pteropus voeltzkowi population To assess the spatial temporal variations of Pteropus voeltzkowi, the data was collected through counting of Pteropus voeltzkowi in each visited roosting site. Specifically, the direct/ Ground-counts technique was used. Ground counts involves observer walking through roost sites during the day when Pteropus voeltzkowi are roosting and estimate the total number. Ground counts only focused on roosting sites where the animal were abundant and efficiently monitored. The advantage of this technique is the fact that, it can be rapidly implemented at a large geographical scale by relatively fewer human resources of about two-person team.

With the help of pairs of binocular of 7×50, Pteropus voeltzkowi were observed when at the roosting site and the counting was done while standing under a roosting site and all the activities performed by the Pteropus voeltzkowi were recorded. All surveys were conducted from 6:00 am to 6:00 pm and the same procedures were repeated in each counting day. During each visit, the sequence of the survey was randomly chosen to avoid bias. Counting was conducted every day of the first week

22 of February and April 2020 for dry and wet season respectively. On each counting day a team of five people participated in counting and their average counts, time and GPS location was recorded.

3.6 Objective two Assessment of Pteropus voeltzkowi habitat and diet preferences Assessment of habitat and food was carried out during the ground counting time and the night respectively. With the help of pairs of binocular of 7×50 field angles of 6.60 116 m/1000 m Pteropus voeltzkowi were observed when they carry food away from feeding roost. Habitat types were categorized based on types of vegetation, size of the roosting sites (both canopy diameter and diameter at breast height) and plant species. On the other hand diet source preferences, were categorized based on plant species being eaten and plant parts (seed, flower, fruits, leaves and nectar) being consumed. Food consumption was recorded as frequent, or infrequent. This method is also supported by (Mukherjee, 2010) in his study of Dietary energy from fruit preferences of Cynopterus sphinx.

Figure 3. 2: Research Assistants take measurements on dbh of the plant at Msitu Mkuu roost site

23 3.7 Objective three Assessment of the potential roles of the Pteropus voeltzkowi in restoring forest connectivity in the Pemba Island To understand the potential role of Pteropus voeltzkowi in restoring forest connectivity, a viable seed from four plant species including Ficus sur, Psidium guajava, Syzygium comini and Vitex doniana were collected, the seeds from the chosen plant were selected because they take relatively shorter time to germinate under natural condition. Furthermore, seeds were planted in to plastic beaker of 250 mil with height of 10 cm and diameter of 6 cm and each beaker contained a total of

25 seeds. The beaker containing processed seeds were labelled A1, B1, C1 and D1, while a beaker containing ripen unprocessed seeds were labelled A2, B2, C2, and D2.

The loaded beakers were placed on to the table that has height of 100 cm width 100 cm and length of 300 cm and were exposed to natural environment to get natural air and light. The seeds were watered twice a day in the morning and evening. This technique was also used by Izhaki and Arad (1995) who studied germination rate of the ingested seeds by species in the Old World fruit bats. The germination rate of seeds taken from Pteropus voeltzkowi droppings and ripe fresh fruits under natural conditions were recorded and compared so as to assess the potential role of Pteropus voeltzkowi in regenerating forest for enhanced connectivity.

Figure 3. 3: Unprocessed seeds collected from different roosting site

24 3.8 Data Processing and Analysis Various methods were used in this study to process and analyse data collected. The data need to be systematic well organized and well processed before they are subjected to any useful analysis (Creswell, 2007). The initially stage data was well organized and transformed into understandable and useful information. Furthermore, data were coded and finally data interpretation was done. Data analysis was thematically categorized and focuses to address the main specific objectives and their corresponding research question. The following are the methods of data analysis for each specific objective:

To understand the spatial and temporal variation of Pteropus voeltzkowi population, data was presented as the mean ± standard error (SE) and R software was used to estimate the relative abundance of Pteropus voeltzkowi. To understand how seasonality and vegetation type affect the dynamics of Pteropus voeltzkowi, a generalized linear mixed effect model (equation 1) was used, collection/counting sites were treated as random factor to take into account site variability and the re- sampling on the same site over the two seasons.

풀 = 푴휷 + 풁휸 + Ԑ where Y is a N x 1 column Pteropus voeltzkowi of observation; M is a N x p design matrix for fixed effects relating observation Y to β; β is a p x 1 column Pteropus voeltzkowi of the fixed-effects; Z is the N x q design matrix for the q random effects relating observation Y to γ; γ is a q x 1 Pteropus voeltzkowi of the random effects; and ε is a N x 1 column Pteropus voeltzkowi of the residuals, that part of Y that is not explained by the model, Mβ+Zγ.

To understand diet and habitats preferences, types of habits and food preferences were descriptively analyzed using percentages and frequencies in R software platform. Furthermore, potential roles of Pteropus voeltzkowi in restoring forest connectivity in the Pemba Island was descriptively assessed by percentages of germination rates of processed and unprocessed seeds.

25 3.9 Data quality and Control Data quality control is the important feature in any research in order to make a research perceived. To ensure this action two important aspect were observed those are validity and reliability.

3.9.1 Validity and Reliability Validity and Reliability are the two important quality control objects in research design (Kumar, 2011). Validity and reliability are two component used to address the issue concerning the quality of the data and appropriate of the methods used in carrying out a research project therefor should be consider when designing of the study, analyzing and during presentation of the result.

Validity Validity refers to the ability of the scale to measure what is supported to measure, and it concerns whether the measurements provide the information needed to answer the research question. The significance of formulating validity is to prove that the variable being study, methodology and the reached conclusion can be generalized beyond the specific study. In this study the validity was obtain through carefully counting of Pteropus voeltzkowi in a field, carefully observation of Pteropus voeltzkowi when they feed and through carefully germination experiment of the seed. Secondly the collected data was well checked and all the mistake error was eliminated for the purpose of attain the accuracy and uniformity.

Reliability Reliability of the study is the extent in which other researcher comes to similar result if they undertake the study with the same case using exactly the same procedure as the first researcher (Creswell, 2007). Reliability is concerned with the repeatability of the study, whereby the same results can be obtained if the same data collection and analysis are used in a new study. In this study the researcher was spend more time during data collection to ensure the accuracy of the data. Moreover, the reconnaissance was conducted for the purpose of testing the instrument before they are engaging in the actual study.

26 3.10 Ethical Consideration The study methodology was reviewed and approved by Pemba Wildlife Management Committee (PWMC), the body responsible for making recommendations on application for the uses of wildlife for scientific research. It was also reviewed by Second Vice President Office for given permission of data collection in Pemba Island.

27 CHAPTER FOUR RESULTS AND DISCUSSION 4.1 Chapter Overview This chapter presents results and discussion of data collected on the field sites. The results are presented and discussed objective wise and cover spatial-temporal variation of Pteropus voeltzkowi population, habitat preference of Pteropus voeltzkowi and potential roles of the Pteropus voeltzkowi in restoring forest connectivity in the Pemba Island.

4.2 Spatial-temporal variation of the Pteropus voeltzkowi population in study area Field observation in this study showed that, there was variation in relative abundance of Pteropus voeltzkowi between the villages as well as between wet and dry season, Figure 4.1 present the variation in relative abundance between dry and wet season.

Figure 4. 1: The variation of Pteropus voeltzkowi relative abundance between dry and wet season

Dry season recorded significantly higher relative abundance of Pteropus voeltzkowi compared to wet season, due to the fact that, wet seasons lower security of Pteropus voeltzkowi by not only fallings off roosting trees but also strong winds may create

28 uncomfortable living conditions for this species (Aryal, Brunton & Raubenheime, 2014). Moreover, seasonal food resource accessibility influence population variation of flying fox in different seasons (Aryal et al., 2015). In addition, studies by Tidemann and Nelson (2004) have also reported a declining trend of various species in the genus Pteropus during wet season.

It was further revealed that, there was significant variation of Pteropus voeltzkowi between villages (χ2 =307.29, df = 264, p-value = 0.03449). Compared to other villages, highest number of Pteropus voeltzkowi was recorded in Wete village. Figure 4.2 present the relative abundance of Pteropus voeltzkowi across the study areas.

Figure 4. 2: Variation of Pteropus voeltzkowi relative abundance across study villages

Apparently, the high relative abundance of Pteropus voeltzkowi was reported in Wete village (roost site) which had abundant preferred feeding resources such as Mangifera indica, Carica papaya, Terminalia catappa, Vitex doniana, Ficus sur, Musa paradisiaca, Psidium guajava, and Syzygies cumini. Availability of feeding 29 resources reduce Pteropus voeltzkowi flight distance and thus possibility of this species aggregating in one location. High relative abundance of Pteropus voeltzkowi in Wete roosting site and the fact that there were relatively more food resources in Wete village could also mean that Pteropus voeltzkowi move from other villages to Wete roosting site, consequentially high abundance of Pteropus voeltzkowi in Wete village than in Ngezi, Msitu Mkuu and Kidike roosting sites which had relatively fewer preferred feeding resources.

Generally, many studies have simply provided evidence that, the area having abundance of preferred fruits are more likely to have large numbers of old world fruit bats, particularly Pteropus species (Mathur & Kumar, 2012). Furthermore, findings from observation indicated that relative abundance of Pteropus voeltzkowi was higher in residential habitat types compared to forest type. Figure 4.3 shows the result of relative abundance of Pteropus voeltzkowi between forest and residential habitant types.

30

Figure 4.3: Variation of Pteropus voeltzkowi relative abundance between habitat types studied

It was revealed that, relative abundance of Pteropus voeltzkowi was high in residential area and Wete roost site (Figure 4.4A & B) irrespective of the season.

31

Figure 4. 4: Seasonal variation of Pteropus voeltzkowi relative abundance between (A) habitat types and (B) villages

Pteropus voeltzkowi commonly roost in residential habitat near human settlement compared to forest. This can be explained by the fact that preferred fruit trees were abundant near human settlement compared to the forest and where feeding resource competition could be high while almost absent in the human settlement. This observation has also been reported in other Pteropus species example P. poliocephalus and P. scapulatus. Furthermore, field observation shows that, most of the roosting trees having large canopy diameter caries large number of Pteropus voeltzkowi since it provide enough space for resting during day time, compared to those having small canopy diameter which are commonly found in forest habitat. Based on these finding and the results in Table 4.1, null hypothesis is rejected and 32 thus we affirm that there is spatial temporal variation in Pteropus voeltzkowi population as it has been reported by other scholars elsewhere and some parts of Pemba Island that the entire genus Pteropus varies in space and time with possible explanations being habitat variation (food resources and roosting sites) and seasons.

In general, observation in this study revealed that, there were more Pteropus voeltzkowi on Terminalia catappa roost trees, compare to other roosting tree species. Figure 4.5 present relative abundance of Pteropus voeltzkowi in different roosting plant.

33

Figure 4. 5: Relative abundance of Pteropus voeltzkowi across various tree/plant species

34 Moreover, Wete village and Residential habitat recorded more Terminalia catappa (Mikungu) with relatively higher abundance of Pteropus voeltzkowi. Figure 4.6 and 4.7 shows relative abundance of Pteropus voeltzkowi across various tree species in different villages and in both residential and forested habitats.

Figure 4. 6: Relative abundance of Pteropus voeltzkowi across various tree species in different villages

35

Figure 4. 7: Relative abundance of Pteropus voeltzkowi across various tree species in both residential and forested habitat

In addition, in residential habitat and Wete village there were more Pteropus voeltzkowi roosting trees with higher canopy diameter compared to forest habitat and other villages while roosting trees height were relatively similar in both habitat but they varies between villages. Figure 4.8 present the result of the canopy diameter, roosting height and variation between habitat types and village.

36

Figure 4. 8: Roosting site canopy diameter (M) and height (M) variation between habitat types and villages

From linear mixed effect model, it was revealed that there was significant decrease of Pteropus voeltzkowi relative abundance during wet season (β= -63.5, SE = 25.4 p- value = 0.0134) when compared to dry season. There was also significant increase of Pteropus voeltzkowi relative abundance in Wete village (β= 142.1, SE=46.6, df= 138, p - value = 0.002) and significant decrease of Pteropus voeltzkowi relative abundance in Ngezi (β = -105.3, SE=46, df=138, p-value=0.0237) compared to Kidike. In 37 addition, the finding shows that, there was significant positive relationship between relative abundance of Pteropus voeltzkowi and roosting trees canopy diameter (β= 10.4, SE = 2.9, df = 138, p- value= 0.0005) and roosting trees height (β= 4.5, SE = 1.95, df = 138, p- value= 0.0219) (Table 4.1).

Table 4. 1: Linear mixed-effects model between Pteropus voeltzkowi, village, and season DBH, Canopy diameter and Tree Height Coef ±Std Error Df p-value Intercept -61.26340 ±61.57427 138 0.3215 Msitu Mkuu village 29.90845±41.28668 138 0.4700 Ngezi village -105.32894±46.06087 138 0.0237 Wete village 142.12659±46.80272 138 0.0029 Wet season -63.54058±25.47593 138 0.0138 Roosting trees Dbh (cm) 0.03481±1.15095 138 0.9759 Roosting trees Canopy 10.46915±2.95584 138 0.0005 diameter (m) Roosting trees Height (m) 4.52979±1.95344 138 0.0219 Random effect: Site 19.62 AIC value 2318.258

Note: Kidike village and dry season have been taken as reference parameters

4.3 Habitat and diet preference of Pteropus voeltzkowi From field observation, it was revealed that, out of 19 plant species recorded, only 12 plant species from 10 families were identified as sources of food for Pteropus voeltzkowi during the study time. Table 4.2 show food providing plants and parts of the plant being consumed. Furthermore, it was revealed that, preferred food type for Pteropus voeltzkowi were fruits followed by nectar and leaves. Figure 4.9 show percentage of food preference for Pteropus voeltzkowi.

38 Table 4. 2: Food providing plant for Pteropus voeltzkowi Sn Family Plant Species Consumption preference Preferred Part Consumption frequency 1 Anacardiac Mangifera indica Fr Frequently 2 Bombacac Ceiba pentandra N/FL Infrequent 3 Caricaceaeeae Carica papaya Fr Frequently 4 Combretaceae Terminalia catappa Fr Frequently 5 Lamiaceae Vitex doniana Fr Infrequent 6 Moraceaeeae Ficus sur Fr Frequently Artocarpus altilis Fr/N Frequently 7 Musaceae Musa paradisiaca N/Fr Infrequent 8 Myrtaceae Psidium guajava Fr/L Infrequent Syzygies cumini Fr Frequently 9 Phyllantha Uapaca guineensis Fr Infrequent 10 Sapindacea Litchi chinensis Fr Frequently FLcceae = flower, Fr= fruits, N= nectar, L= leaves 0 e The recorded plant species were preferred by Pteropus voeltzkowi as a source of food because they carry fruits, nectar and flower petals which contain high water and carbohydrate content. Among all consumed plants parts that were recorded, observation showed that, fruits were the most preferred food resource possibly because of its ability to meet nutritional demand compared to flower petals, nectar and leaves (Izhaki & Arad, 1995).

It was further revealed that Pteropus voeltzkowi feeds on plant leaves which can be explained by calcium supplementation as reported by (Pierson, Elmqvist, Rainey & Cox, 1996). Fruit preference as observed in this study concur with (Mukherjee, 2010) in his study on dietary energy from fruit preferences of Cynopterus sphinx which is a species among the Old World fruit bats. Preference of fruits might imply the potential human-bats conflicts associated with fruits raiding and potential zoonotic diseases (Panthi, Coogan, Aryal & Raubenheimer, 2015).

39

Figure 4. 9: Preferred plant parts as source of foods for Pteropus voeltzkowi

4.4 Potential roles of the Pteropus voeltzkowi in restoring forest connectivity through germination success An experiment on seeds germination under natural condition showed that processed seeds had highest germination rate (Table 4.3). Chi-square tests showed that, germination success differ significantly between processed and unprocessed seed (X2 = 16.266, df= 3, P- value =0.001).

40 Table 4.3: Average germination time and rate selected plant species that are

frequently consumed by Pteropus voeltzkowi Sno. Plant Germination Time Number of seeds Germinated species (Average) (N=50) Processe Unprocesse Processed seeds Unprocessed d seeds d seeds seeds N (%) N (%) 1 Ficus sur 9 14 33 66 10 20 2 Psidium 21 33 21 42 8 16 guajava 3 Syzygium 10 Nil 13 26 0 0 comini 4 Vitex 13 Nil 9 18 0 0 doniana

All the seeds that were processed by Pteropus voeltzkowi had better germination rate than unprocessed one. This is attributed to the fact that, Pteropus voeltzkowi remove the fruit pulp “typically contains germination inhibitors which cause seed dormancy” (Entwistle & Speakman, 1997) hence lead to better germination rate. In addition to removal of germination inhibitors, processed seeds from fecal matter germinated relatively faster compared to others because, some amount of Pteropus voeltzkowi fecal material that remained on the seeds act as fertilizer which can potentially influence germination. Similar findings have been reported by Izhaki and Arad (1995), who studied germination rate of the ingested seeds by species in the Old World fruit bats. The enriched germination rate of processed seed implies that Pteropus voeltzkowi can potentially facilitate secondary vegetation succession and consequently improve vegetation/forest patchy connectivity provided that other factors are kept constant.

41 CHAPTER FIVE SUMMARY, CONCLUSION AND RECOMMENDATIONS 5.1 Chapter Overview This chapter provides summary, conclusion and recommendations based on the findings of the study.

5.2 Summary of the Finding The main focus of this study was to assess the spatial- temporal variation of Pteropus voeltzkowi population and its potential role in restoring forest connectivity. The study was conducted at the Pemba Island.

The first study question that was answered was “how population of Pteropus voeltzkowi varies in space in wet and dry season”? The study findings show that, population of Pteropus voeltzkowi were varying widely between the seasons and was significantly higher in dry season compared to wet season. The results of this study also reject the null hypothesis, since the entire genus Pteropus, varies in space and time with possible explanations being habitat variation (food resources and roosting sites) and seasons.

The second study question which was answered was “what are the preferred habitat of Pteropus voeltzkowi in Pemba Island”? Although the diet of Pteropus voeltzkowi may vary throughout its range. This study reveals that plant species that contribute a significant part to its diet include Mangifera indica, Ceiba pentandra, Carica papaya, Terminalia catappa, Vitex doniana, Ficus sur, Artocarpus altilis, Musa paradisiaca, Psidium guajava, Syzygies cumini, Uapaca guineensis and Litchi chinensis. Whereby fruits were more preferred diet for Pteropus voeltzkowi because contained highly water content and carbohydrate, frequently eating fruits by Pteropus voeltzkowi help to increase water and carbohydrate intake in their body system resulting gaining of energy help them to fly in a long distance.

Furthermore, the third objective of the study investigated the potential roles of the Pteropus voeltzkowi in restoring forest connectivity. This objective answers the following study question ‘’does Pteropus voeltzkowi have potential role for restoring forest connectivity”? The study findings show that all the seeds that were processed

42 by Pteropus voeltzkowi significantly showed better germination rate than those from unprocessed.

Unique reason for the positive effect of Pteropus voeltzkowi on germination rates could be due to fruit selection by the Pteropus voeltzkowi since fruits attractive to Pteropus voeltzkowi may contain higher numbers of mature and viable seeds.

5.3 Conclusion Based on the results, it can be concluded that spatial-temporal variation of Pteropus voeltzkowi population is not only the matter of season, but there is lot of factors including food availability and human factors such as habitat destruction and hunting process. Findings in this study provide information that can inform stakeholders about the space and time of Pteropus voeltzkowi relative abundance and thus devising a fit-for-purpose of sensitization program. The information obtained in this study can as well be useful in adding on available data of Pteropus voeltzkowi population trends that can inform the conservation strategies for this species.

Furthermore, the understanding of spatial and temporal variations of Pteropus voeltzkowi population can deliver important information needed when developing predictive models of the spatial-temporal dynamics of Pteropus voeltzkowi and their ecological and human wildlife conflicts potential roles.

It is important to note that Pteropus voeltzkowi was observed feeding on flowers, fruit, leaves and nectars. Thus Pteropus voeltzkowi, potentially have an important role as pollinators for a wide range of plant species, but perhaps perform a dual role as both a pollinator and seed disperser. In addition, the results from germination success performed in this study imply that Pteropus voeltzkowi could be important for dispersing seeds between forest patches and involved in the successful regeneration and establishment of forestry in Pemba Island.

This study provides important information on spatial temporal variation and habitat preferences required by Pteropus voeltzkowi. Future research may put more focus on assessing the critical habitat to the survival of the Pteropus voeltzkowi.

43 5.4 Recommendations Based on the results documented in this study, the researcher wants to recommend the following to the Department of Forest at Pemba Island

i) Improve knowledge of population dynamics of Pteropus voeltzkowi including their movement, distribution and behavior

ii) Improve the management and conservation status of Pteropus voeltzkowi

iii) Establish community-based conservation to all areas whereby Pteropus voeltzkowi are found

iv) Establish and implement plan for the management of Pteropus voeltzkowi roosting site

v) Establish and publish information to the community to build their capacity to co-exist with Pteropus voeltzkowi

vi) Promote education to the community for the purpose of increasing public awareness on Pteropus voeltzkowi value

5.5 Recommendation for Further Studies The study suggests the following recommendations where further research might be particularly relevant

i) Undertake a study to assess the effects of increasing heat on juvenile Pteropus voeltzkowi during dry season

ii) Initiate a study on assessing the critical habitat to the survival of the Pteropus voeltzkowi

iii) Undertake a study to assess the perception of the local people on Pteropus voeltzkowi

iv) Undertake the study to assess the way Pteropus voeltzkowi recognize their permanent and temporal roosting site and the method used for foraging

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48 APPENDICES Appendix 1: Assessment on spatial-temporal variation of Pteropus voeltzkowi population. Wete Roosting Site GPS Location……………………………

Date Time Roost Number of GPS location Elevation Plant Height of Canopy Seasonality Types of habitant from site ID species Coordinate species the pant diameter counted South East Latitude Longitude

49 Assessment on spatial-temporal variation of Pteropus voeltzkowi population.

Kidike Roosting Site GPS Location……………………………

Date Time Roost Number of GPS location Elevation Plant Height of Canopy Seasonality Types of habitant from site ID species Coordinate species the pant diameter counted South East Latitude Longitude

50 Assessment on spatial-temporal variation of Pteropus voeltzkowi population.

Ngezi Roosting Site GPS Location……………………………

Date Time Roost Number of GPS location Elevation Plant Height of Canopy Seasonality Types of from site ID species Coordinate species the pant diameter habitant counted South East Latitude Longitude

51 Assessment on spatial-temporal variation of Pteropus voeltzkowi population.

Msitu Mkuu Roosting Site GPS Location……………………………

Date Time Roost Number of GPS location Elevation Plant Height of Canopy Seasonality Types of from site ID species Coordinate species the pant diameter habitant counted South East Latitude Longitude

52 Appendix 2: Habitat preference of Pteropus voeltzkowi SN Plant Part of the Preference Location Elevation Season species plant being being eaten consumed

53 Appendix 3: Estimate germination success of seeds taken from bats droppings (Processed) with those from fresh fruits (Unprocessed) under progressively more natural conditions. Number of seeds that is germinate S/N Plant species Processed Percentage % Unprocessed Percentage %

54 Appendix 4: Number of species counted during dry season at Ngezi roost site. Assessment on spatial-temporal variation of Pteropus voeltzkowi population. Site, Ngezi roost site GPS Location, S05.06182, E039.71945 Elevation 1m Date Time Roost Number GPS location Elevati Plant species Height dbh Canopy Seaso Types from site ID of species (Coordinate) on of the (cm) diameter nality of counted plant habitat Latitude Longitude

12/2/2020 10:25am 01 55 04.93855 039.70825 31 Antiaris toxicaria 41 57 10 Dry Forest ” 10:40am 02 95 04.93845 039.70821 34 Milicia excels 35 34 13 ” ” ” 10:53am 03 79 04.93813 039.70824 34 Erythrophloem suaveolens 20 23.5 11 ” ” ” 11:09am 04 27 04.93801 039.70830 35 Dypsis pembanus 15 11.5 13 ” ” ” 11:23am 05 25 04.93797 039.70821 36 Dypsis pembanus 20 16.8 17 ” ” ” 11:40am 06 35 04.93790 039.70825 36 Bombax rhodognaphalon 16 18.5 12 ” ” ” 12:00pm 07 66 04.93789 039.70821 36 Antiaris toxicaria 35 64 15 ” ” ” 12:17pm 08 195 04.93751 039.70840 34 Quassia indica 35 65 10 ” ” ” 12:32pm 09 96 04.93756 039.70841 36 Milicia excels 30 28.5 09 ” ” ” 12:50pm 10 78 04.93767 039.70856 37 Erythrophloem suaveolens 28 25 16 ” ” ” 1:09pm 11 105 04.93756 039.70858 35 Antiaris toxicaria 40 38 09 ” ” 13/2/2020 8:30am 12 110 04.93761 039.70858 35 Antiaris toxicaria 40 29 07 ” ” ” 8:45am 13 55 04.93769 039.70869 34 Antiaris toxicaria 36 27.5 13 ” ” ” 9:01am 14 86 04.93773 039.70872 34 Maesopsis eminii 44 35.8 08 ” ” ” 9:18am 15 43 04.93779 039.70893 35 Quassia indica 25 17.8 13 ” ” ” 9:33am 16 135 04.93771 039.70886 34 Antiaris toxicaria 38 27 16 ” ” ” 9:52am 17 63 04.93750 039.70897 35 Antiaris toxicaria 43 29 09 ” ” ” 10:05am 18 125 04.93758 039.70889 36 Antiaris toxicaria 45 40.8 07 ” ” 55 ” 10:24am 19 100 04.93754 039.70889 36 Antiaris toxicaria 44 61 12 ” ” ” 10:41am 20 70 04.93746 039.70895 37 Antiaris toxicaria 42 44 15 ” ” Date Time Roost Number GPS location Elevati Plant species Height dbh Canopy Seaso Types from site ID of species (Coordinate) on of the diameter nality of counted plant habitat Latitude Longitude

10:58am 21 80 04.93748 039.70900 38 Antiaris toxicaria 47 49.8 09 ” ” 14/2/2020 8:15am 22 195 04.93744 039.70894 38 Antiaris toxicaria 43 55.5 08 ” ” ” 8:35am 23 55 04.93752 039.70961 39 Maesopsis eminii 25 26 13 ” ” ” 8:47am 24 63 04.93740 039.70958 38 Milicia excelsa 28 23.7 17 ” ” ” 9:05am 25 85 04.93767 039.70917 37 Antiaris toxicaria 45 60 09 ” ” ” 9:23am 26 65 04.93772 039.70907 37 Maesopsis eminii 35 39.8 17 ” ” ” 9:42am 27 52 04.93799 039.70909 37 Maesopsis eminii 28 41.2 10 ” ” ” 9:57am 28 35 04.93796 039.70908 37 Antiaris toxicaria 15 16 07 ” ” ” 10:10am 29 40 04.93804 039.70910 38 Antiaris toxicaria 16 23.3 08 ” ” ” 10:35am 30 45 04.93805 039.70917 38 Antiaris toxicaria 30 57.6 16 ” ”

56 Appendix 5: Number of species counted during wet season at Ngezi roost site Assessment on spatial-temporal variation of Pteropus voeltzkowi population. Site, Ngezi roost site GPS Location S04.93996, E039.71086 Elevation 1m Date Time Roos Number GPS location Elevati Plant species Height dbh Canopy Season Types from t site of species (Coordinate) on of the diameter ality of ID counted plant habitat Latitude Longitude

15/3/2020 8:15am 01 40 04.93848 039.70829 32 Antiaris toxicaria 40 57 12 Wet Forest ” 8:32am 02 80 04.93849 039.70819 34 Milicia excelsa 35 34 8 ” ” ” 8:47am 03 70 04.93815 039.70829 34 Erythrophloem suaveolens 20 23.5 11 ” ” ” 9:00am 04 15 04.93799 039.70827 34 Dypsis pembanus 15 11.5 8 ” ” ” 9:18am 05 8 04.93795 039.70821 36 Dypsis pembanus 15 16.8 7 ” ” ” 9:33am 06 60 04.93788 039.70821 36 Bombax rhodognaphalon 35 64 13 ” ” ” 9:47am 07 20 04.93785 039.70825 36 Antiaris toxicaria 16 18.5 12 ” ” ” 10:00am 08 120 04.93747 039.70837 34 Quassia indica 35 65 20 ” ” ” 10:15am 09 95 04.93753 039.70842 35 Milicia excels 30 28.5 10 ” ” ” 10:32am 10 70 04.93766 039.70859 38 Erythrophloem suaveolens 28 25 14 ” ” ” 10:49am 11 85 04.93757 039.70858 35 Antiaris toxicaria 40 38 11 ” ” ” 11:07am 12 73 04.93760 039.70863 33 Antiaris toxicaria 40 29 9 ” ” 16/3/2020 9:25am 13 38 04.93769 039.70874 35 Antiaris toxicaria 36 27.5 13 ” ” ” 9:39am 14 88 04.93770 039.70869 34 Maesopsis eminii 44 35.8 8 ” ” ” 9:49am 15 33 04.93778 039.70892 35 Quassia indica 25 17.8 13 ” ” ” 10:05am 16 130 04.93773 039.70890 34 Antiaris toxicaria 38 27 14 ” ” ” 10:19am 17 55 04.93754 039.70889 34 Antiaris toxicaria 43 29 17 ” ” ” 10:34am 18 100 04.93756 039.70895 35 Antiaris toxicaria 45 40.8 13 ” ” 57 ” 10:48am 19 95 04.93753 039.70884 36 Antiaris toxicaria 44 61 15 ” ” ” 11:05am 20 60 04.93746 039.70894 37 Antiaris toxicaria 42 44 18 ” ” Date Time Roost Number GPS location Elevati Plant species Height dbh Canopy Season Types from site of Coordinate on of the diameter ality of ID species plant habitat counted Latitude Longitude

” 11:23am 21 70 04.93750 039.70903 38 Antiaris toxicaria 47 49.8 17 ” ” ” 11:45am 22 250 04.93746 039.70903 38 Antiaris toxicaria 43 55.5 10 ” ” 17/3/2020 2:15pm 23 45 04.93751 039.70960 39 Maesopsis eminii 25 26 13 ” ” ” 2:32pm 24 55 04.93735 039.70955 39 Milicia excelsa 28 23.7 15 ” ” ” 2:45pm 25 65 04.93768 039.70918 37 Antiaris toxicaria 45 60 9 ” ” ” 2:59pm 26 78 04.93768 039.70910 38 Maesopsis eminii 35 39.8 12 ” ” ” 3:18pm 27 43 04.93801 039.70910 38 Maesopsis eminii 28 41.2 16 ” ” ” 3:35pm 28 25 04.93807 039.70918 37 Antiaris toxicaria 15 16 13 ” ” ” 3:54pm 29 30 04.93804 039.70914 37 Antiaris toxicaria 16 23.3 10 ” ” ” 4:14pm 30 35 04.93805 039.70921 38 Antiaris toxicaria 30 57.6 17 ” ”

58 Appendix 6: Number of species counted during dry season at Msitu Mkuu roost site Assessment on spatial-temporal variation of Pteropus voeltzkowi population. Site, Msitu Mkuu roost site GPS Location, S05.00233, E039.82601 Elevation 8m Date Time Roost Number GPS location Eleva Plant species Height dbh Canopy Season Types from site of species (Coordinate) tion of the diameter ality of ID counted plant habitat Latitude Longitude

15/2/2020 12:30pm 01 150 05.00596 039.83389 18 Celosia trigyna 10 32 12 Dry Forest ” 12:45pm 02 200 05.00588 039.83398 18 Manilkara sulcata 22 39.5 10 ” ” ” 12:57pm 03 50 05.00578 039.83394 18 Manilkara sulcata 18 34.3 14 ” ” ” 1:03pm 04 180 05.00574 039.83397 18 Celosia trigyna 17 42 08 ” ” ” 1:20pm 05 300 05.00569 039.83397 18 Celosia trigyna 17 41.2 12 ” ” ” 1:33pm 06 350 05.00579 039.83415 19 Celosia trigyna 26 36.5 09 ” ” ” 1:50pm 07 200 05.00579 039.83411 19 Celosia trigyna 16 32 10 ” ” ” 2:06pm 08 500 05.00583 039.83417 19 Celosia trigyna 28 61 17 ” ” ” 2:24pm 09 20 05.00555 039.83449 19 Manilkara sulcata 13 27.2 08 ” ” ” 2:39pm 10 60 05.00549 039.83452 20 Ficus carica 25 61 15 ” ”

59 16/2/2020 10:15am 11 100 05.00552 039.83449 20 Celosia trigyna 26 41.9 10 ” ” ” 10:32am 12 70 05.00559 039.83461 21 Lennea acutifoliolate 18 42.8 07 ” ” ” 10:45am 13 30 05.00563 039.83464 20 Manilkara sulcata 18 46.5 11 ” ” ” 11:07am 14 20 05.00572 039.83462 20 Celosia trigyna 13 38.6 06 ” ” ” 11:35am 15 150 05.00572 039.83460 20 Celosia trigyna 15 44 12 ” ” ” 11:49am 16 150 05.00557 039.83479 18 Celosia trigyna 15 49 16 ” ” ” 12:04pm 17 18 05.00558 039.83478 19 Lennea acutifoliolate 20 43.5 12 ” ” ” 12:22pm 18 150 05.00560 039.83492 18 Afzelia quanzensis 14 39.5 12 ” ” ” 12:40pm 19 19 05.00588 039.83493 19 Lennea acutifoliolate 22 48.1 14 ” ” ” 12:52pm 20 20 05.00630 039.83501 17 Celosia trigyna 27 48.4 17 ” ”

60 Appendix 7: Number of species counted during wet season at Msitu Mkuu roost site Assessment on spatial-temporal variation of Pteropus voeltzkowi population. Site, Msitu mkuu roost site GPS Location, S05.00233, E039.82601 Elevation 8m Date Time Roost Number GPS location Elevati Plant species Height dbh Canopy Season Types from site ID of species Coordinate on of the diameter ality of counted plant habitat Latitude Longitude

18/3/20 2:15pm 01 130 05.00597 039.83388 18 Celosia trigyna 20 32 12 Wet Forest ” 2:32pm 02 180 05.00588 039.83400 18 Manilkara sulcata 22 39.5 10 ” ” ” 2:45pm 03 55 05.00577 039.83394 18 Celosia trigyna 18 34.3 14 ” ” ” 2:58pm 04 188 05.00573 039.83400 18 Celosia trigyna 17 42 08 ” ” ” 3:13pm 05 350 05.00570 039.83395 18 Celosia trigyna 17 41.2 12 ” ” ” 3:30pm 06 440 05.00578 039.83411 18 Celosia trigyna 26 36.5 09 ” ” ” 3:45pm 07 150 05.00580 039.83410 19 Celosia trigyna 16 32 10 ” ” ” 4:03pm 08 510 05.00581 039.83418 19 Celosia trigyna 28 61 17 ” ” ” 4:22pm 09 18 05.00556 039.83447 19 Manilkara sulcata 13 27.2 08 ” ” ” 4:41pm 10 55 05.00555 039.83448 20 Ficus carica 25 61 15 ” ”

61 ” 4:57pm 11 110 05.00556 039.83448 21 Lennea acutifoliolate 26 41.9 10 ” ” ” 5:13pm 12 80 05.00560 039.83461 20 Lennea acutifoliolate 18 42.8 07 ” ” 19/3/20 9:16am 13 25 05.00566 039.83461 20 Manilkara sulcata 18 46.5 11 ” ” ” 9:35am 14 15 05.00566 039.83464 20 Celosia trigyna 13 38.6 06 ” ” ” 9:50am 15 145 05.00571 039.83462 20 Celosia trigyna 15 44 12 ” ” ” 10:09am 16 140 05.00561 039.83475 18 Celosia trigyna 15 49 16 ” ” ” 10:27am 17 20 05.00558 039.83478 19 Lennea acutifoliolate 20 43.5 12 ” ” ” 10:44am 18 155 05.00558 039.83493 18 Afzelia quanzensis 14 39.5 12 ” ” ” 10:59am 19 130 05.00589 039.83491 18 Lennea acutifoliolate 22 48.1 14 ” ” ” 11:20am 20 245 05.00626 039.83499 17 Celosia trigyna 27 48.4 17 ” ”

62 Appendix 8: Number of species counted during dry season at Kidike roost site Assessment on spatial-temporal variation of Pteropus voeltzkowi population. Site, Kidike roost site GPS Location, S05.17259, E039.83035 Elevation 14m Date Time Roost Number GPS location Elevati Plant species Height Dbh Canopy Season Types from site of species Coordinate on of the diameter ality of ID counted plant habitat Latitude Longitude

17/2/20 9:30am 01 167 05.17419 039.83042 12 Antiaris toxicaria 13 32 6 Dry Forest ” 9:42am 02 262 05.17444 039.83049 11 Antiaris toxicaria 27 42.8 10 ” ” ” 9:57am 03 174 05.17414 039.82971 13 Antiaris toxicaria 16 35.7 12 ” ” ” 10:10am 04 155 05.17439 039.83010 13 Antiaris toxicaria 35 64.5 07 ” ” ” 10:28am 05 159 05.17442 039.83010 13 Antiaris toxicaria 32 61 05 ” ” ” 10:45am 06 144 05.17445 039.83012 12 Antiaris toxicaria 36 55 06 ” ” ” 10:57am 07 142 05.17445 039.83053 14 Antiaris toxicaria 20 26.5 08 ” ” ” 11:16am 08 127 05.17434 039.83049 15 Antiaris toxicaria 19 16 07 ” ” ” 11:33am 09 122 05.17396 039.83055 14 Borassus aethiopum 15 40 05 ” ” ” 11:52am 10 141 05.17343 039.83062 14 Antiaris toxicaria 14 53 06 ” ”

63 ” 12:13pm 11 139 05.17322 039.83055 14 Antiaris toxicaria 30 65 07 ” ” ” 12:30pm 12 142 05.17320 039.83059 13 Antiaris toxicaria 35 63.7 10 ” ” ” 12:48pm 13 129 05.17325 039.83058 13 Antiaris toxicaria 36 59 05 ” ” ” 1:07pm 14 322 05.17312 039.83079 13 Antiaris toxicaria 30 64.6 12 ” ” ” 1:26pm 15 147 05.17315 039.83075 13 Antiaris toxicaria 42 53.7 08 ” ” ” 1:44pm 16 362 05.17330 039.83005 14 Antiaris toxicaria 32 60 13 ” ” ” 1:59pm 17 137 05.17330 039.82991 15 Mangifera indica 15 65 22 ” ” ” 2:15pm 18 143 05.17395 039.82970 16 Antiaris toxicaria 25 55 10 ” ”

64 Appendix 9: Number of species counted during wet season at Kidike roost site Assessment on spatial-temporal variation of Pteropus voeltzkowi population. Site, Kidike roost site GPS Location, S 05.17233, E 039.83066 Elevation 15m Date Time Roost Number GPS location Elevati Plant species Height dbh Canopy Season Types from site of Coordinate on of the diameter ality of ID species plant habitat counted Latitude Longitude

20/3/20 8:15am 01 65 05.17413 039.83044 12 Antiaris toxicaria 13 32 06 Wet Forest ” 8:28am 02 160 05.17444 039.83048 11 Antiaris toxicaria 27 42.8 10 ” ” ” 8:45am 03 75 05.17416 039.82967 13 Antiaris toxicaria 16 35.7 12 ” ” ” 8:57am 04 50 05.17439 039.83009 13 Antiaris toxicaria 35 64.5 07 ” ” ” 9:18am 05 57 05.17441 039.83009 13 Antiaris toxicaria 32 61 05 ” ” ” 9:35am 06 35 05.17443 039.83011 13 Antiaris toxicaria 36 55 06 ” ” ” 9:53am 07 46 05.17448 039.83051 14 Antiaris toxicaria 20 26.5 08 ” ” ” 10:16am 08 25 05.17448 039.83049 15 Antiaris toxicaria 19 16 07 ” ” ” 10:34am 09 13 05.17391 039.83057 15 Borassus aethiopum 15 40.4 05 ” ”

65 ” 10:44am 10 50 05.17341 039.83063 14 Antiaris toxicaria 14 53 06 ” ” ” 11:01am 11 20 05.17322 039.83056 14 Antiaris toxicaria 30 65 07 ” ” ” 11:20am 12 35 05.17321 039.83059 15 Antiaris toxicaria 35 63.7 10 ” ” ” 11:35am 13 37 05.17324 039.83060 15 Antiaris toxicaria 36 59 05 ” ” ” 11:47am 14 250 05.17311 039.83077 12 Antiaris toxicaria 30 64.6 12 ” ” ” 11:57am 15 40 05.17311 039.83073 13 Antiaris toxicaria 42 53.7 08 ” ” ” 12:13pm 16 200 05.17328 039.83006 14 Antiaris toxicaria 32 60 13 ” ” ” 12:30pm 17 20 05.17329 039.82994 15 Mangifera indica 15 65 22 ” ” ” 12:49pm 18 40 05.17394 039.82967 16 Antiaris toxicaria 25 55 10 ” ”

66 Appendix 10: Number of species counted during dry season at Wete roost site. Assessment on spatial-temporal variation of Pteropus voeltzkowi population. Site, Wete Port GPS Location, S 05.06100, E 039.71848 Elevation 17m Date Time Roost Number GPS location Elevati Plant species Height D bh Canopy Seaso Types of from site ID of species (Coordinate) on of the (cm) diameter nality habitat counted plant Latitude Longitude

18/2/20 4:05pm 01 150 05.06176 039.71996 06 Terminalia catappa 25 60 20 Dry Resident 4:18pm 02 480 05.06187 039.71951 03 Terminalia catappa 25 61.8 19 ” ” 4:34pm 03 150 05.06187 039.71936 02 Terminalia catappa 15 55.6 11 ” ” 4:51pm 04 70 05.06181 039.71939 04 Calophyllum inophyllum 10 35 10 ” ” 5:09pm 05 880 05.06182 039.71947 09 Terminalia catappa 28 65 15 ” ” 5:26pm 06 50 05.06169 039.71952 12 Terminalia catappa 20 20.7 20 ” ” 5:39pm 07 1500 05.06169 039.71952 08 Terminalia catappa 27 37.5 33 ” ” 5:58pm 08 750 05.06161 039.71945 11 Terminalia catappa 25 40.3 30 ” ” 6:15pm 09 500 05.06166 039.71940 10 Terminalia catappa 27 54.7 25 ” ” 6:28pm 10 550 05.06162 039.71939 10 Terminalia catappa 29 49.6 16 ” ”

67 19/2/20 9:15am 11 300 05.06139 039.71951 17 Mangifera indica 16 30 11 ” ” 9:34am 12 450 05.06132 039.71929 18 Mangifera indica 28 47.8 18 ” ” 9:51am 13 200 05.06127 039.71926 17 Syzygium cumini 18 30.3 10 ” ” 10:07am 14 850 05.06156 039.71921 12 Terminalia catappa 17 45.6 35 ” ” 10:26am 15 30 05.06137 039.71912 19 Terminalia catappa 12 50.2 9 ” ” 10:45am 16 650 05.06149 039.71897 05 Terminalia catappa 30 43.8 13 ” ” 10:58am 17 30 05.06147 039.71889 04 Vitex doniana 26 20.5 9 ” ” 11:11am `18 250 05.06143 039.71851 06 Mangifera indica 28 29.7 11 ” ” 11:30am 19 250 05.06208 039.71880 04 Pterocarpus angolensis 30 20.5 14 ” ” 11:49am 20 550 05.06203 039.71931 09 Terminalia catappa 32 45.8 19 ” ”

68 Appendix 11: Number of species counted during wet season at Wete roost site. Assessment on spatial-temporal variation of Pteropus voeltzkowi population. Site: Wete roost site GPS Location, S05.06206, E039.71877 Elevation: 13m Date Time Roost Number GPS location Elevati Plant species Height Dbh Canopy Season Types of from site ID of species (Coordinate) on of the diamete ality habitat counted plant r Latitude Longitude

21/3/20 8:00am 01 90 05.06189 039.72015 6 Terminalia catappa 25 60 20 Wet Resident ” 8:18am 02 350 05.06194 039.71950 17 Terminalia catappa 25 61.8 19 ” ” ” 8:34am 03 85 05.06201 039.71942 16 Terminalia catappa 15 55.6 11 ” ” ” 8:51am 04 550 05.06196 039.71937 16 Calophyllum inophyllum 10 35 10 ” ” ” 9:09am 05 95 05.06181 039.71948 16 Terminalia catappa 28 63.6 26 ” ” ” 9:26am 06 150 05.06198 039.71941 16 Terminalia catappa 20 16 09 ” ” ” 9:44am 07 120 05.06206 039.71915 15 Terminalia catappa 29 50.8 17 ” ” ” 10:02am 08 40 05.06203 039.71904 14 Terminalia catappa 17 49 12 ” ” ” 10:19am 09 55 05.06208 039.71875 13 Pterocarpus angolensis 30 60.7 25 ” ” ” 10:34am 10 160 05.06143 039.71846 16 Mangifera indica 28 64.4 30 ” ”

69 ” 10:38am 11 25 05.06145 039.71884 13 Vitex doniana 26 38 13 ” ” ” 10:55am 12 250 05.06152 039.71894 11 Terminalia catappa 30 55 15 ” ” ” 11:14am 13 115 05.06156 039.71911 13 Terminalia catappa 27 43.7 16 ” ” ” 11:33am 14 420 05.06157 039.71937 17 Terminalia catappa 30 48 12 ” ” 22/3/20 2:30pm 15 1200 05.06163 039.71943 18 Terminalia catappa 29 64.7 13 ” ” ” 2:49pm 16 480 05.06131 039.71928 23 Mangifera indica 28 64.5 22 ” ” ” ” 3:05pm 17 110 05.06127 039.71930 24 Syzygium cumini 25 59 23 ” ” ” 3:25pm 18 37 05.06121 039.71941 26 Terminalia catappa 21 53.6 25 ” ” ” 3:45pm 19 700 05.06170 039.71948 22 Terminalia catappa 32 64.8 18 ” ” ” 4:16pm 20 17 05.06173 039.71954 20 Terminalia catappa 20 16 13 ” ”

70 Appendix 12: Habitat preference of P.voeltzkowi SN Plant species being eaten Part of the plant Preference GPS Location Elevation Season being consumed Latitude Longitude 01 Mangifera indica Fruits Frequency -05.06131 039.71930 23 Dry/wet 02 Ficus sur Fruits Frequency -05.00555 039.83448 20 Dry/wet 03 Psidium guajava Fruits Frequency -05.06555 039.83228 16 Dry 04 Terminalia catappa Fruits Frequency -05.06152 039.71894 11 Dry 05 Syzgium comini Fruits Frequency -05.06127 039.71930 24 Wet 06 Syzgium comini Flowers Moderate -05.06127 039.71930 24 Wet 07 Syzgium comini Seeds Rare -05.06127 039.71930 24 Dry 08 Litchi chinensis Fruits Frequency -05.06121 039.71941 26 Wet 09 Uapaca guineensis Fruits Frequency -05.06208 039.71875 30 Dry 10 Ceiba pentandara Pollen Moderate -05.06202 039.71907 8 Dry 11 Ceiba pentandara Flower Moderate -05.06202 039.71907 8 Dry 12 Vetex doniana Fruits Frequency -05.17391 039.85057 15 Wet 13 Anacardium occidental Fruits Rare -05.09863 039.75117 67 Dry 14 Celosia trigyna Fruits Moderate -05.00451 039.83243 16 Dry 15 Celosia trigyna Leaves Rare -05.00451 039.83243 16 Dry 16 Carica papaya Fruits Frequency -05.00555 039.83448 20 Dry 17 Musa paradisiaca Fruits Frequency -05.17394 039.82967 16 Dry 18 Artocarpus altilis Fruits Moderate -05.17329 039.82994 22 Wet 19 Ficus sur Pollen Moderate -05.00555 039.83448 20 Wet

71 Appendix 13: Mean number of days of seed germination between processed and unprocessed seeds from different plant species and number of seeds germinating when they planted in a plastic beaker. S/N Plant species Mean day of seeds that germinated Bats processed seeds Unprocessed seeds 1. Ficus sur 9 14 2. Psidium guajava 21 33 3. Syzygium comini 10 - 4. Vitex doniana 13 -

The number of seeds germinating when they planted in a plastic beaker and their percentages, between processed and unprocessed seeds from different plant species.

Number of seeds that was germinate S/N Plant species Processed Percentage % Unprocessed Percentage % 1. Ficus sur 33/50 66 10/50 20 2. Psidium guajava 21/50 42 8/50 16 3. Syzygium comini 13/50 26 Nil Nil 4. Vitex doniana 9/50 18 Nil Nil

72 Appendix 14: Result from Linear mixed-effects model. 1. Chi-square tests of independence Chi-square tests showed that there significant variation of number of Pemba Flying Fox between villages; χ2 =307.29, df = 264, p-value = 0.03449, but variation between seasons and habitat were insignificant; χ2 =85.31, df = 88, p-value = 0.5614 and χ2 = 95.523, df = 88, p-value = 0.2737 2. Linear mixed-effects model fit by maximum likelihood where Rost.ID is Random effects and fixed effects are Pemba flying fox, village and season.

Linear mixed-effects model fit by maximum likelihood Data: P1 AIC BIC logLik 2333.732 2355.925 -1159.866

Random effects: Formula: ~1 | Rost.ID (Intercept) Residual StdDev: 24.42241 174.5196

Fixed effects: NPFF ~ Village + Season Value Std.Error DF t-value p-value (Intercept) 150.88627 32.89191 141 4.587337 0.0000 VillageM/ Mkuu 26.62675 40.71256 141 0.654018 0.5142 VillageNgezi -47.08644 37.49833 141 -1.255694 0.2113 VillageWete 221.85679 40.72325 141 5.447914 0.0000 SeasonWet -61.45684 26.69447 141 -2.302231 0.0228 Correlation: (Intr) VllM/M VllgNg VllgWt VillageM/ Mkuu -0.653 VillageNgezi -0.716 0.572 VillageWete -0.653 0.527 0.572 SeasonWet -0.405 0.000 0.000 0.000

Standardized Within-Group Residuals: Min Q1 Med Q3 Max -2.00505100 -0.36057539 -0.08559302 0.19125269 6.37383086

Number of Observations: 176 Number of Groups: 31

73 3. Linear mixed-effects model fit by maximum likelihood where Rost.ID is Ran dom effects and fixed effects are Pemba Flying Fox and village and season a nd DBH and Canopy diameter and Tree Height

Linear mixed-effects model fit by maximum likelihood Data: P1 AIC BIC logLik 2318.258 2349.963 -1149.129

Random effects: Formula: ~1 | Rost.ID (Intercept) Residual StdDev: 19.62443 164.5855

Fixed effects: NPFF ~ Village + Season + Dbh.cm + Canopy.dm + Height .m Value Std.Error DF t-value p-value (Intercept) -61.26340 61.57427 138 -0.994951 0.3215 VillageM/ Mkuu 29.90845 41.28668 138 0.724409 0.4700 VillageNgezi -105.32894 46.06087 138 -2.286734 0.0237 VillageWete 142.12659 46.80272 138 3.036716 0.0029 SeasonWet -63.54058 25.47593 138 -2.494141 0.0138 Dbh.cm 0.03481 1.15095 138 0.030247 0.9759 Canopy.dm 10.46915 2.95584 138 3.541851 0.0005 Height.m 4.52979 1.95344 138 2.318878 0.0219 Correlation: (Intr) VllM/M VllgNg VllgWt SesnWt Dbh.cm Cnpy.d VillageM/ Mkuu -0.449 VillageNgezi -0.258 0.438 VillageWete -0.177 0.548 0.542 SeasonWet -0.161 -0.014 -0.036 -0.015 Dbh.cm -0.351 0.090 0.567 0.181 -0.074 Canopy.dm -0.302 -0.211 -0.307 -0.554 0.018 -0.249 Height.m -0.350 0.174 -0.508 -0.037 0.021 -0.573 0.133

Standardized Within-Group Residuals: Min Q1 Med Q3 Max -2.18053781 -0.42954033 -0.07405422 0.31305034 5.71558167 Number of Observations: 176 Number of Groups: 31

74 Table 1: Linear mixed-effects model between Pemba Flying Fox and village and season

Coef± SE Df P

Intercept 150.88627±32.89191 141 0.0000*

Msitu Mkuu 26.62675±40.71256 141 0.5142

Ngezi -47.08644±37.49833 141 0.2113

Wete 221.85679±40.72325 141 0.0000*

Wet Season -61.45684± 26.69447 141 0.0228 * Random effect: 24.42 Site AIC value 2333.732

Table 2: Linear mixed-effects model between Pemba Flying Fox and village, sea son DBH, Canopy diameter and Tree Height.

Coef. value Std Error Df t-value p-value Intercept -61.26340 ±61.57427 138 -0.994951 0.3215 Msitu Mkuu 29.90845 ±41.28668 138 0.724409 0.4700 Ngezi -105.32894 ±46.06087 138 -2.286734 0.0237 Wete 142.12659 ±46.80272 138 3.036716 0.0029 Wet season -63.54058 ±25.47593 138 -2.494141 0.0138 Dbh.cm 0.03481 ±1.15095 138 0.030247 0.9759 Canopy.dm 10.46915 ±2.95584 138 3.541851 0.0005 Height. M 4.52979 ±1.95344 138 2.318878 0.0219 Random effect: Site 19.62 AIC value 2318.258

75 Appendix 15: Research Clearance from the University of Dodoma

76