A Comparative Study of Bird Abundance and Diversity in Mature

Home , Agathis, Dacrydium, Fiji goshawk, Fiji parrotfinch, Forest honeyeater, Parrotfinch

A COMPARATIVE STUDY OF BIRD ABUNDANCE AND DIVERSITY IN MATURE SECONDARY AND LOGGED MID-ALTITUDE RAINFORESTS IN NAKOBALEVU AND SAVURA, SOUTHERN VITI LEVU HIGHLANDS, FIJI ISLANDS.

Nitassha Kavita Shankaran

A thesis submitted in fulfillment of the requirements for the degree of Master of Science in Biology

School of Biological and Chemical Sciences Faculty of Science, Technology and Environment The University of the South Pacific

June, 2010

DECLARATION

Statement by Author I, Nitassha Shankaran declare that this thesis is my own work and that, to the best of my knowledge, it contains no material previously published, or substantially overlapping with material submitted for the award of any degree at any institution, except where due acknowledgment is made in the text.

ACKNOWLEDGEMENTS

I would like to thank the following people in particular for their efforts in helping make this study a success:

5 The Darwin Initiative, BirdLife International Pacific and the University of the South Pacific who provided funding to complete this study 5 Dr. Timothy Markwell, Dr. Linton Winder, Dr. Uma Khurma, James Millett and Dr. Clare Morrison for their technical support and help with the logistics of the study 5 Vilikesa Masibalavu for introducing birding to me as an exciting activity and for teaching me a significant amount about bird identification 5 Avinesh Prasad and Dinesh Kumar for driving us to the study locations every morning 5 Staff of BirdLife International Pacific for assistance during training and research site establishment 5 Vido and Neno for keeping us safe in the forest 5 Janesh Kumar and Preetika Prasad for their support, proof-reading assistance and being my confidants in the tussles faced during this study 5 And to all other people who I have missed who deserve my due respect and gratitude

I dedicate this thesis to my mother, Late Mrs. Sangita Sharma, who taught me the value of education, my brother, Anvil Shankaran, for his love and support and especially to my husband, Janesh Janendra Kumar, for his support and inspiration in whatever I do.

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ABSTRACT

To establish the impact of logging on the diversity and abundance of forest birds in the mid-altitudinal rainforests of the Southern Viti Levu Highlands of Fiji, random point counts were carried out in mature secondary as well as recently logged forests. An overall total of 120 point counts were carried out from August to November, 2007 and January to February, 2008.

This study showed that there were more individuals in recently logged forests at the two locations, Nakobalevu and Savura (ANOVA: F4, 116 = 31.66, P treatment = 0.000). In addition to this, there were significantly more bird species found in recently logged forests as well (ANOVA: F4, 116 = 19.61, P treatment = 0.000). These results were also reflected with results from the Shannon - Weiner Index where the species richness (S) was higher in recently logged forests compared to the mature secondary forests. The species richness was S=27 and S=24 in recently logged forests at Nakobalevu and Savura respectively. The evenness measure was higher in mature secondary forests than in recently logged forests indicating that in recently logged forests a few species have larger numbers but most species have low numbers. This study also established indicator species for the two forest types. Even though there were bird species with high associations with the mature secondary forest, they were extensively present in recently logged forests as well. Therefore, no indicator species were found specifically for the mature secondary forests. Barking pigeons (Ducula latrans), the Fiji bush-warbler (Cettia ruficapilla) and the Fiji white-eye (Zosterops exploratory) were found in higher numbers and had stronger associations with the recently logged forests at both Nakobalevu and Savura. The Fiji parrotfinch (Erythrura pealii) and the White-rumped swiftlets (Aerodramus spodiopygius) were the only two bird species which were only present in the recently logged forests. Other species that have been recorded in this study have shown to have flexible habitats, that is, they were found in both mature secondary and recently logged forests at both the study locations.

This study indicates that logging does affect the bird abundance and diversity. Increases in species richness found at recently logged forests do not directly relate to

iv conservation priorities as the additional species observed were associated with disturbed habitats and were of lower conservation concern than species that were associated with mature secondary forests. Results from this study show that even though species richness is higher in recently logged forests compared to mature secondary forests, the mature secondary forests have a more stable resident native breeding land bird population and therefore bird populations with a higher conservation priority. This study shows that Fijian forests and Fijian birds have some adaptability towards habitat disturbance. The avian ecosystem is able to recover after logging provided the forest is permitted to recover and where source populations survive.

CONTENTS

Declaration ii Acknowledgements iii Abstract iv List of figures ix List of tables x

CHAPTERS Chapter 1 1 Introduction 1 1.1 General study of Fiji 1 1.1.1 Location and topography of Fiji Islands 1 1.1.2 Climate 2 1.2 Fiji’s flora 3 1.3 Biodiversity conservation in Fiji 4 1.3.1 Biodiversity and conservation of wildlife in Fiji 4 1.3.2 Origin and distribution of birds in Fiji 6 1.3.3 Understanding of bird biodiversity 7 1.4 Loss of biodiversity caused by logging 7 1.4.1 Logging and its effects on bird populations 7 1.4.1.1 Treefall gaps 9 1.4.1.2 Fragmentation and habitat loss 10 1.4.1.3 Edge effects on bird population 12 1.4.1.4 Effects of logging on bird nesting 13 1.4.1.5 Invasive alien plants and animals 14 1.4.1.6 Unfavourable succession pathways: A 15 consequence of logging 1.5 The Study 17 1.5.1 Location, vegetation and habitat types 17 1.5.2 Logging disturbance history 20 1.6 Aims and objectives of the study 21

Chapter 2 22 Methodology 22 2.1 Sampling design and method 22 2.2 Statistical analysis 24 2.2.1 Two-way ANOVA 24 2.2.2 Shannon - Weiner index and Pielou’s species evenness 25 2.2.3 Dufrêne and Legendre’s Indicator Value Method (IVM) 25

Chapter 3 27 Results 27 3.1 Number of individuals 27 3.2 Number of species 28 3.3 Shannon - Weiner index and Pielou’s species evenness 31 3.4 Dufrêne and Legendre’s Indicator Value Method (IVM) 35 3.4.1 Nakobalevu 35 3.4.2 Savura 36 3.5.Diet type and habitat of bird species found in mature 39 secondary forest and recently logged forest

Chapter 4 40 Discussion 40 4.1 Bird abundance in mature secondary forests and recently 40 logged forests 4.2 Species richness and diversity in mature secondary forests 41 and recently logged forests 4.3 Indicator species’ for mature secondary forests and 44 recently logged forests 4.4 Conclusion and recommendations 45

Bibliography 47

Appendix 1: Establishment of number of points required 62

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Appendix 2:Diet, feeding habit and status of bird species recorded 64 at study locations

viii LIST OF FIGURES

Figure Description Page 1 Map of Fiji in the Eastern Melanesian island region 2 2 Location of Important Bird Areas in Fiji 6 3 Map of Important Bird Area FJ10 situated in the Southern 17 Highlands of Viti Levu, Fiji Islands 4 Map of study locations showing vegetation types from 19 forest survey conducted in 1992 5 Map of study plots at Nakobalevu and Savura 24 6 Mean number of individuals found in MSF and RLF at 28 Nakobalevu (n=30) 7 Mean number of individuals found in MSF and RLF at 28 Savura (n=30) 8 Mean number of bird species found in MSF and RLF at 30 Nakobalevu (n=30) 9 Mean number of bird species found in MSF and RLF at 31 Savura (n=30)

LIST OF TABLES

Table Description Page 1 Forest cover on larger islands within the Fiji group based 9 on National Forest Inventory 1990-1993 2 Fijian vernacular name and its scientific equivalent for 21 major tree species’ harvested from the recently logged forest in Nakobalevu and Savura Forest Reserve from 2000-2005 3 Sampling design used for the study 23 4 Results from two-way ANOVA for number of individuals 27 5 Total number of bird species found in the two forest types 29 at Nakobalevu and Savura 6 Results from two-way ANOVA for number of species 30 7 Relative abundance of bird species recorded in mature 32 secondary forest (MSF) and recently logged forest (RLF) at study locations 8 Bird species richness (S), absolute number of birds 35 observed (N) Shannon’s indices (H1) and Pielou’s species evenness (E) at each study location 9 Total number of individual birds and the total number of 35 families [ ] recorded at the study locations, Nakobalevu and Savura 10 Species established as being indicative of forest type at 37 Nakobalevu using the Indicator Value Method 11 Species established as being indicative of forest type at 38 Savura using the Indicator Value Method

CHAPTER 1

INTRODUCTION

1.1 General study of Fiji 1.1.1 Location and topography of Fiji Islands The Fiji Islands, located approximately 850 km east of Vanuatu, comprises of over 300 named islands. They are located at 16-20◦S latitude and 178◦E - 178◦W longitude (Fig 1). The cluster of more than 300 islands (97 of which are inhabited (Mueller- Dombois & Fosberg 1998) occupies an ocean territorial area of about 650,000km2, of which less then 3 percent (18333km2) is the total land area. The largest islands, Viti Levu and Vanua Levu, have a land area of 10, 390km2 and 5,538km2 respectively (Watling 2004). In addition to this, Fiji also has of two midsized islands: Taveuni (434km2) and Kadavu (408km2) and many smaller islands (Mueller-Dombois & Fosberg 1998).

The highest elevations of the main islands in Fiji are 1324m (Tomanivi or Mt. Victoria) (Watling 2004) on Viti Levu, 1032m (Mt. Batini) on Vanua Levu, 1241m (Mt. Uluiqalau) on Taveuni and 838m (Mt. Washington) on Kadavu. The Fijian high islands of Viti Levu can be classified into three topographical types (Mueller- Dombois & Fosberg 1998). These are as follows:

1. Flatlands: Makes up 16% of the Viti Levu surface. These are the coastal plains, many of which are subject to periodic inundation during heavy rains and traversed by meandering delta forming rivers. 2. Rolling and hilly lands: Comprises 17% of Viti Levu’s surface. Often found adjacent to flatlands and a majority of this land type occurs as interior plateaus among the steeplands. 3. Steeplands: Generally supports more natural vegetation and makes up 67% of Viti Levu’s surface.

Figure 1: Map of Fiji in the Eastern Melanesian island region taken from Mueller- Dombois & Fosberg (1998)

1.1.2 Climate Fiji has a pronounced windward-leeward effect and orographic rainfall or relief precipitation due to the combining of the southeast trade winds with the mountainous topography. The southeastern sides of the islands receive approximately 3000mm of rainfall annually whereas the leeward sides receive 1800mm of rainfall per year (Mueller-Dombois & Fosberg 1998).

Fiji’s position in the subequatorial tropics of the Southern Hemisphere results in a slightly warmer and distinctly wet season from November through April and a slightly cooler season from June through to October. The period from mid-November

2 to mid-April is known as the cyclone or hurricane season in Fiji (Mueller-Dombois & Fosberg 1998). The relative humidity is usually over 85% in the wet zone and lowers in the dry zones.

Rainfall records from 2001 to 2005 indicated that Mt. Nakobalevu received a mean annual rainfall of approximately 2872mm with an average monthly rainfall of 239mm. The mean monthly rainfall recorded for the years 2006 and 2007 was 247.3mm and 275.7mm respectively while 615.9mm of rainfall was recorded for the month of January and 185.3mm for February, 2008 (Swastika Devi, pers comm, 2008). Both study locations fall in the leeward side of Fiji Islands thus receive a large amount of rainfall.

1.2 Fiji’s flora The Fiji Islands are often considered to have a floral foundation composed of Australian and New Guinean (or Papuan) elements (Mueller-Dombois & Fosberg 1998). According to Brownlie (1977) and Watkins (1995) a total of approximately 2600 vascular plants are present in Fiji. These include at least 310 Pteridophytes or non seed vascular plants (Brownlie 1977) and at least 2225 seed plants (Watkins 1995). Of these 1692 species (63%) are considered native of which 892 species (56%) are endemic to Fiji. The remaining 936 species (37%) are introduced exotics (Allison 2003).

The distribution of principal vegetation types in Fiji can be determined by four environmental parameters: climate, topography, critical substrate variations, and human influences. On this basis Mueller-Dombois & Fosberg (1998) described nine principal vegetation types in Fiji. These are:

1. Lowland rainforest: Extends from near the sea level to an altitude of 600m. It is found in the wet zone of Viti Levu and Vanua Levu receiving a mean rainfall of 2000-3000 mm annually. 2. Upland rainforest: Covers areas above an altitude of 600m. It has a cool, wet climate, the mean air temperature of about 5◦ to 6◦ C, cooler than in the lowland rainforest. These areas receive a mean annual rainfall of 2000-3000 mm.

3. Cloud forest: The mountaintops and ridges above 600m near the coast and those above 900mm inland, mainly cloaked in clouds. At such sites, moist clouds and horizontally driven rain adds large amounts of precipitation of more than 9000 mm of rainfall annually. 4. Dry forest: Present in the leeward lowlands of Fiji’s high islands, with a mean rainfall of 1750-2250mm annually. 5. Talasiqa vegetation: Meaning “sunburnt” land, it spreads from sea level to 1000m. It receives a mean annual rainfall from 1500-2500 mm. 6. Freshwater wetland vegetation: Occurs in Fiji only in the wet zone of Viti Levu, where it is restricted to poorly drained alluvial sites. 7. Mangrove forest and scrub: Seen along coastlines and river deltas of Fiji. 8. Coastal strand vegetation: Found on northeastern coast where there is little or no impact of anthropogenic factors. 9. Smaller island vegetation: Vegetation of small islands overall mean annual rainfall is near 2000 mm.

1.3 Biodiversity conservation in Fiji 1.3.1 Biodiversity and conservation of wildlife in Fiji All the Micronesian islands, tropical Polynesia and Fiji are part of the Polynesia- Micronesia Hotspot according to Conservation International. Fiji has one of the richest biota, with the highest degree of taxonomic endemism of all other Pacific island nations (Anonymous 2006). Areas containing an exceptional concentration of biodiversity are known as hotspots and can be measured by species richness, species endemism or both (Prendergast et al. 1993; Lumaret & Lobo 1996; Hobohm 2003). The total population of hotspots for organisms is approximately 3,235,250 out of which 65% is found in Hawaii and Fiji.

Fiji currently has 87 species of breeding birds widespread across Fiji. These breeding birds nest in Fiji. These can be divided into 57 native land species that are mostly forest birds; 19 native seabirds which mostly nest on small offshore islands and some on mountains of large islands; 11 introduced and alien species of birds as well as three extinct breeding birds. There are also 29 non-breeding seabird species, 22 species of non-breeding migrant waders and one species of non-breeding migrant

4 land bird in Fiji (Masibalavu & Dutson 2006). Non-breeding bird species do not nest in Fiji.

The 2004 IUCN (Anonymous 2006) Red List of threatened Species indicates that Fiji is home to 170 globally threatened species. These are: 17 bird species 3 turtles, 2 iguanas, 3 snakes, 3 frog, 4 bats, 10 geckos, 12 skinks 50 marine species such as whales, sharks and tuna 89 freshwater fish species of which 69 are native, 10 endemic and 10 are introduced 90 plants Little is known about the invertebrate fauna of Fiji (Anonymous 2005).

Conservation in Fiji began with the first nature reserves set up in Nandarivatu by the Colonial Government in 1958. Currently, there are several of conservation plans and organizations striving towards the conservation of Fiji’s biodiversity including BirdLife International, Nature Fiji and Wildlife Conservation Society to name a few. Several community based conservation projects have been set up with the help of conservation NGOs and the Fiji government combining the objectives of the NGOs and that of the Fiji National Biodiversity Strategy and Action Plan (NBSAP).

To date there is one declared Protected Forest, namely the Batiwai Forest (central division), with a Gulubia microcarpa (threatened species) palm population, of approximately 15,749 ha. There are 17 Forest Reserves covering an area of 26,203 ha and 7 Nature Reserves in Fiji covering an area of approximately 5,740 ha. In addition, the Sigatoka Sand Dunes, Bouma Heritage Park and Koroyanitu Heritage Park are protected under MOUs between the Government of Fiji and foreign partners.

BirdLife International has been identifying Important Bird Areas (IBAs) around Fiji. Important Bird Areas are internationally recognized sites which play a vital role for the conservation of birds (Masibalavu & Dutson 2006). For an area to be eligible as an IBA, sites must meet or exceed one or more global criteria and thresholds (Dutson & Masibalavu 2006). Studies by Masibalavu & Dutson (2006) have identified 14

IBAs (Fig 2) which cover 17% of the land area of Fiji (Masibalavu & Dutson 2006). All IBAs contain large areas of forest and range in size from about 28km2 to 720km2 with the exception of the tiny island of Vatu-i-Ra (FJ 05).

Figure 2: Location of Important Bird Areas in Fiji (Masibalavu & Dutson 2006).

1.3.2 Origin and distribution of birds in Fiji Paucity is a characteristic commonly seen in the birds of the oceanic islands (Watling 2004). The more remote and smaller the island, the more this characteristic is accentuated. According to Watling (2004) conditions for island hopping were better in the last two million years than at present, when the sea level was about 130m lower than today and most islands in the region were much larger than today. Due to a high degree of endemism, the exact origin and route of colonization of most bird species cannot be tracked with absolute certainty. However, a Papua New Guinean or Australian, or both, origin is thought to have supplied the ancestral stock of virtually all species (Watling 2004). Also according to Watling (2004) “Vanuatu may have been an essential stepping-stone during the invasion and colonization of the region

6 by bird populations, behaving as an apex of a funnel drawing species from both major source regions (Papua New Guinean and Australian), and directing them on to Fiji.”

1.3.3 Understanding of bird biodiversity In many respects bird biodiversity, both past and present, is better understood than for any other major group of organisms. This is largely due to the fact that birds inspire more extreme interest in humans than all other animals as they are often magnificent, are relatively easily observed and are neither too conspicuous nor too cryptic to identify and study (Norris & Pain 2002).

Knowledge of the geographical distribution of many species of birds and how this has changed in the last 100-200 years is relatively good. As a result, birds are used as a model species to investigate the location of biodiversity ‘hotspots’ of species- richness and endemism and the potential establishment of a network of globally - based conservation priority areas recently refined by BirdLife International into Endemic Bird Areas or EBAs (Bibby et al. 2000). BirdLife International has mapped every restricted range bird species to find avian centres of endemism. These areas are termed EBA and serve as excellent indicators of general biodiversity.

In Fiji, very little is known about the population trends of tropical birds and very few studies such as those carried out by BirdLife International have been carried out to assess the impacts of human activities on bird diversity and abundance and therefore establishing the overall ecological status of the forests of Fiji.

1.4 Loss of biodiversity caused by logging 1.4.1 Logging and its effects on bird populations Forests support about 65% of the world’s terrestrial taxa (World Commission on Forests and Sustainable Development 1999) and have the highest species diversity for many taxonomic groups, including birds (Gill 2004), invertebrates (Majer & Delabie 1994) and microbes (Crozier et al. 1999). Tropical forests are being logged or converted to agricultural land at an ever increasing rate (Chapman & Chapman 1997). A study by Pimm & Raven (2000) suggests that about 1 million square

7 kilometers of forest worldwide is eliminated every 5 to 10 years. Conserving forest diversity is therefore a critical task (Lindenmayer et al. 2006). Tropical birds are generally understood to be less resilient to habitat disturbances such as logging, compared to their counterparts in the temperate regions (Warkentin et al. 1995; Marsden 1998; Hughes et al. 2002; Sodhi et al. 2005).

Logging has several consequences for bird populations such as affecting the foraging and nesting behaviour of different bird species, and increasing access for invasive alien predators and competitors of the bird population. Other indirect factors of logging also affect the bird biodiversity. For example, seedling suppression affecting forest cover and composition and making pathways for unfavourable succession trends in logged forests in turn affecting bird populations (Paul et al. 2004).

In Fiji, approximately half of the forests have been lost as consequence of land clearance for agriculture and through repeated fires (Masibalavu & Dutson 2006). Many remaining areas of tropical rainforests in Fiji have been heavily degraded. An accurate account of forest cover was given by the 1990-1993 National Forest Inventory satellite images (Table 1) (Masibalavu & Dutson 2006). A study by Claasen (1991) states that deforestation in Fiji is moderate but continuing, occurring at a nominal rate of between 0.5-0.8 percent per year. The mean deforestation rate estimated by the Food and Agriculture Organization of the United Nations (0.2% per year) between 1990 and 2000 is believed to be less accurate than Claasen’s (1991) estimate (Masibalavu & Dutson 2006). However, the new forest inventory to be released soon shows an increased rate of forest loss (James Millett, pers comm, 2008). Watling (2004) states that the most considerable forest loss caused by disturbance recorded in the past 30 years was the conversion of about 50,000 ha of good tropical rainforest to mahogany plantation in areas such as Nukurua and Tailevu. According to Masibalavu & Dutson (2006), in addition to the overall reduction in the extent of the forest, many of Fiji’s forests suffer degradation from logging. Studies by Kretzschmar et al. (2003), Richner & Kretzschmar (2001) and Watling (1988) state that birds in Fiji are less tolerant to the poor environmental standards practiced by some logging companies which harvest too many trees which in turn allows pioneer vegetation such as vines to dominate and repress forest regeneration.

Table 1: Forest cover on larger islands within the Fiji group based on National Forest Inventory 1990-1993 (Masibalavu & Dutson 2006). Areas Dense Medium Scattered Pine & Total Dense & (km2) natural dense natural mahogany land medium forest natural forest plantation area dense forest natural forest (% of total land area) Viti 1273 2684 1074 975 10378 3958 Levu (38%) Vanua 1403 1250 349 368 5535 2653 Levu (48%) Taveuni No data: cloud obscured aerial photograph Kadavu 124 101 83 4 408 225 (55%) Gau 2 49 12 3 140 51 (37%)

1.4.1.1 Treefall gaps Tree fall gaps play a significant role in the determining the distribution, abundance and diversity of forest organisms (Wunderle et al. 2006b). When a tree is cut down and falls, it opens the forest canopy. Tree fall gaps are also created when trees fall naturally. This initiates a dynamic succession pathway commencing with little or no canopy, and terminates in a mature phase with closed canopy (Wunderle et al. 2006b). The gap phase is represented by a period of rapid plant growth and colonization (Wunderle et al. 2006b).

Both positive and negative results for bird populations have been documented in the case of treefall gaps created by logging. Treefall gaps allow more light to reach the forest floor of logged forests resulting in a hotter, drier environment with a dense thicket of gap understory vegetation which can result in the reduction of bird abundance and diversity in logged forests (Thiollay 1992). When many treefall gaps occur in a forest area understory bird species of unlogged forests disappear as the conditions of gap understory sharply contrasts the conditions of the surrounding forest understory with respect to microclimate, detritus, productivity and plant species composition (Wunderle et al. 2006a). Another negative response to treefall

9 gaps documented is the hesitance of bird species to travel across gaps and the influence of this on foraging and nesting behaviour (Robichaud et al. 2002).

In contrast, certain species of birds may increase in numbers in response to an increase in gaps especially nectarivores and frugivores (Thiollay 1992). Plants in the gap understory and the area immediately surrounding the gap respond to increase in light positively through increased flower and fruit production (Smith 1991). This increase in fruits and flowers would thus draw bird species relying on fruits and flowers for food. More light, and the presence of fruits and flowers also makes the gaps more attractive to some gap species than would the darker understory of unlogged forests (Wunderle et al. 2006a).

1.4.1.2 Fragmentation and habitat loss Habitat fragmentation can be defined as the conversion of a large continuous patch of habitat into smaller, isolated or remotely connected patches surrounded by a matrix of other habitat types (Wiens 1989). The size and spatial distribution of tropical forest patches and connectivity of forest patches across the landscape are severely affected by logging (Putz et al. 2001). Logging roads and activities associated with the construction of logging routes can greatly influence the permanence of forest fragments created. Fragmentation of the tropical forest has been described as the single greatest threat to global biological diversity (Laurance et al. 2002). Forest fragmentation has many problems for the population dynamics of birds (Trine 1998; Cornelius et al. 2000) and may be the major contributing factor in the extinction of various bird species (Fahrig 1997).

Studies on birds in logged tropical forests have documented a variety of changes to the bird community after logging (Johns 1996; Alexio 1999; Styring & Hussin 2004). Understory insectivores are considered to be particularly sensitive to forest disturbance and fragmentation (Thiollay 1992; Johns 1996; Laurance et al. 2002). There are few bird species that occupy all ecological niches and thus have a high capacity for environmental adaptation (Arroyo et al. 1996). A great percentage of the birds in tropical forests develop in several environments and therefore have less specific requirements for microenvironments or inter-specific relationships for

10 development than non-tropical forest birds. However, some species are present that have special requirements for water environments, food availability, open places for foraging and defined conditions for nesting (Deferrari et al. 2001).

Research by Hill & Curan (2003) suggests that forest fragmentation decreases species number and alters community composition of birds accounted for by reduction in forest area, change in forest shape and an increase in isolation of the remaining forest fragments. Although there seems to be no single pattern of change after the creation of forest fragments, generally a homogenizing effect between the fragmented forests and the surrounding habitats occurs as species not normally found in closed unlogged forests penetrate the forest interior while sensitive forest species become low in abundance or disappear (Bawa & Seidler 1998; Lambert 1992).

Forest spatial geometry controls fundamental ecological processes such as speciation, dispersal, migration, competition and extinction. It thus influences both the abundance and diversity of birds within a given area (Laurance et al. 2002; Hill & Curran 2003). Fragmentation reduces the area of the patches, decreasing the suitability of these fragments for sensitive species that have large home ranges (Freemark & Merriam 1986; Estades & Temple 1999). Like fragmentation, habitat loss is also a primary environmental cause of biodiversity decline at local, regional as well as global scales (Dirzo & Raven 2003). A review of birds and mammals in Australia suggests that habitat patch isolation only becomes important in terms of species richness or abundance when the percentage of habitat decreases below a 20- 30% threshold (Westphal et al. 2003).

Dispersal is one of the key processes determining the probability of persistence of bird populations in fragmented landscapes (Merriam 1991; Wiens et al. 1993). Connectivity is defined by Merriam (1991) as the “degree to which absolute isolation is prevented by landscape elements which allow organisms to move among patches.” After logging, the patches of forest become isolated from each other, affecting dispersal movements of isolation sensitive species which become confined to each fragment (Temple & Cary 1988; Villard et al. 1996). A high level of connectivity is considered essential to the long term persistence of metapopulations in fragments

11 because it increases the probability of recolonisation after extinction caused by logging (Merriam & Saunders 1993).

The consequences of fragmentation for bird populations vary, depending on factors such as the time since fragmentation, fragment size, distance between fragments, and the shape of fragment and features of a species’ life history (Hansen & Urban 1992). As fragmentation continues in many parts of the tropics, conservation efforts must focus on understanding the effects of forest spatial geometry on species diversity as only then can biodiversity be influenced significantly through the management of forest area, shape and isolation (Hill & Curan 2003).

1.4.1.3 Edge effects on bird population Many biologists consider edges between adjacent forest fragments a positive feature of the landscape for wildlife, in particular for birds (Paton 1994). This view is based on Leopold’s (1933) “law of interspersion” which states that increases in the amount of edge habitat results in higher population densities. This effect on bird populations has been reflected in many studies.

Forest edge effects are characterized by changes in the abiotic conditions some distance into the forest. This includes features such as increased amounts of sunlight, higher wind speeds penetrating up to 300m (Pimm et al. 2003), and larger fluctuations in the temperature and humidity (Saunders et al. 1991; Murcia 1995). Therefore, forest edges are hotter, drier and have more tangled vegetation such as vines, shrubs and creepers (Pimm et al. 2003). These changes in abiotic conditions thus affect plant community assemblages and structure, in turn affecting bird populations.

There have been several studies conducted on edge effects on bird abundance and diversity, varying results from positive to negative effects. One of the most commonly cited explanations for bird populations declines is the increased nest predation and parasitism rates near the edges (Paton 1994). Another frequently cited reason for a decrease in bird population is the increased negative interactions with species from adjacent habitats (Estades & Temple 1999).

In contrast, some studies have shown evidence of positive effects of edge effects on bird abundance and diversity. The amount of sunlight reaching the forest floor at the edges of fragmented forests increases due to treefall gaps favoring frugivores, nectarivores and insectivores (Thiollay 1992; Plumptre 1997; Owiunji & Plumptre 1998; Dale et al. 2000). On the other hand, Dale and colleagues (2000) have also reported a decrease in the abundance of bark-gleaning, leaf-gleaning, and ground insectivores near the forest edges. In general, several species showed significant preferences for “the edge”, but few also showed significant avoidance of the edge (Dale et al. 2000).

1.4.1.4 Effects of logging on bird nesting Breeding is one of the vital ecological aspects of bird population dynamics and yet is the most difficult to evaluate (Dranzoa 2001). The commencement of breeding is determined by a number of interacting environmental and physiological factors, whereas breeding success is influenced by both ecological resources and climatic factors (Thompson et al. 1992). The most dramatic disturbances to breeding and nesting of bird populations involve the widespread clearing of forests which occurs during harvesting of all trees in a given area for timber (Robinson & Robinson 2001). Dranzoa (2001) found that breeding success in selectively logged forests was significantly lower which resulted in lower abundance and diversity of birds in logged forests. This was accredited to reduced nest availability, fewer nest materials, increased competition for nest sites and significant increase in nest predation and parasitism. The long term persistence of bird populations requires successful reproduction, which is rare in highly disturbed landscapes (Donovan et al. 1995; Trine et al. 1998; Robinson & Robinson 2001). According to Dranzoa (2001) the influence of past logging on bird breeding potential and success differ from species to species. Low levels of breeding after logging appear to be a phenomenon of tropical forest birds (Dranzoa 2001). “Open crevice” nesters and “tree hollow” nesters are known to commonly breed in pristine forests rather than the logged forests (Dranzoa 2001). The pattern of low numbers of these birds clearly corresponds to the nest resource declines in logged and regenerating forests. In a study by Dranzoa (1995), results indicated that the available nest sites for this particular group of “open crevice” nesters were reduced by over 50% by logging.

This drastic decrease in nesting sites also increases competition amongst the bird species requiring the same breeding resources, in turn, reducing the chances of successful breeding.

Breeding selection pressure acts most on those bird species with special requirements, for example, hollow nesters, crevice nesters, specific materials and so on (Pogue & Schnell 1994; Dranzoa 2001). The quality of nest sites and the nests themselves play an important part in facilitating breeding success of individual birds as most birds require specific characteristics of nests (Dranzoa 2001; Conner et al. 1994). Logging therefore, lowers the value of forests to birds with highly specialized breeding requirements.

Several studies have shown that predator and brood parasitic species, which can greatly reduce the breeding success of birds, increase in abundance in logged forests near forest edges (Robinson & Robinson 2001). Predator species such as rats, mongooses and cats also significantly increase in logged forests (Masibalavu & Dutson 2006). This is due to the increased access for invasive predator species by logging trails. High predation rates of ground and near ground nesting birds in most fragmented landscapes may reflect the abundance of such predators (Villard et al. 1996).

1.4.1.5 Invasive alien plants and animals The introduction of exotic pests and pathogens is an increasingly important ecological occurrence that is altering natural ecosystems worldwide by displacing native species, altering habitat, and modifying key ecological processes (Vitousek et al. 1996; Enserink 1999; Everett 2000; Mack et al. 2000). Logging is an important secondary disturbance leading to forest pest outbreaks that frequently lead to a more rapid removal of the existing bird species and more severe disruptions in the microenvironmental conditions and ecosystem processes (Kizlinski et al. 2002). In addition to damage caused directly by the extraction of trees, logging also enables increased accessibility for invasive alien predators. These disruptions in the microenvironmental conditions affect the bird populations more severely than the pest itself (Kizlinski et al. 2002).

Disturbance caused by logging of native tree species has been shown to facilitate colonization and establishment of invasive, non-native plants (Brown & Gurevitch 2004). The effects of colonization by non native plant species may affect foraging and nesting behaviour of some bird species. For example, studies have shown that logging facilitates the establishment of Psidium cattleianum Sabine (Myrtaceae), a shrub commonly known as the strawberry guava, in logged forests in Hawaii (Brown & Gurevitch 2004). P. cattleianum was seen not to invade closed, unlogged mature forests but rather the selectively logged, fragmented forests of Hawaii. Once established P. cattleianum can form monospecific stands that exclude the establishment of other plant species and therefore affect the composition of the forest and in turn influence the abundance and diversity of birds (Brown & Gurevitch 2004). Other less pervasive or abundant non-native invasive plants having similar effects on bird populations indicated in the study by Brown & Gurevitch (2004) were Eucalyptus robusta, a native Australian tree, and the Syzygium jambos, tree native to the Malay archipelago, Clidemia hirta (Melastomacaceae) which is a perennial shrub, Lantana camara (Verbenaceae) , a flowering plant, which are widely distributed throughout eastern Madagascar. Aggressive herbs, vines and shrubs also invade the forest floor after logging and can have immense consequences for the succession pattern of logged forests (Chapman et al. 1999).

1.4.1.6 Unfavourable succession pathways: A consequence of logging Stand maturation and plant species composition of forest communities’ changes over time (Toriola et al. 1998). This process, known as succession, is mediated by the competitive replacement of dominant plant species due to the availability of resources such as light and soil nutrients (Toriola et al. 1998). The impact of anthropogenic factors such as logging on the process of succession has been recognized as an important factor influencing bird abundance and diversity (Roberts & Gilliam 1995; Kubota et al. 2005). Some of the changes in forest plant species composition in logged areas have been observed to be due to succession but the nature of the change appears to be influenced by the intensity of timber extraction (Kubota et al. 2005). Acanthus pubescens Engl. (Acanthaceae) was found to dominate an area of high intensity timber extraction (harvest averaging 21m3 ha-1) compared to an area where relatively light harvesting (14m3 ha-1) of timber had taken place (Paul et al. 2004). 15

Following logging, in many cases, gaps become infested with aggressive pioneering herbs, vines and shrubs (Chapman & Chapman 1997). These aggressive herbs, strangling vines and shrubs have been shown to repress tree regeneration and halt succession towards a mature forest community (Paul et al. 2004). In tropical rainforests, logging is followed by an increase in lianas, ferns and accumulated debris which also impedes favorable succession pathways (Johns 1991; Cannon et al. 1998; Peres et al. 1999; Cleary et al. 2005). Subsequent regeneration and succession may be inhibited for extended periods of time by liana growth (Schnitzer & Bongers 2002). For example, in a study by Paul et al. (2004) in Kibale National Park, the large gaps created by mechanized logging 30 years ago were dominated by the sub- woody shrub, Acanthus pubescens Engl. (Acanthaceae). A. pubescens is commonly found in swamp forests and in disturbed forests throughout East Africa. Seedling and saplings in A. pubescens dominated areas are unable to germinate and persist in the presence of this invasive plant species. Another important factor preventing tree regeneration in these gaps may be the periodic physical collapse of A. pubescens. Collapses are enhanced by arching, vine-like growth forms of A. pubescens, which results in large networks of stems collapsing, often during the rainy season (Paul et al. 2004). This therefore affects the dynamics of the forested landscape and in turn having massive consequences on the bird populations (Paul et al. 2004).

A study by Pimm et al. (2003) provides evidence in support for unfavourable pathways after logging which involves the poor establishment of forest species. For instance, the mortality of transplanted seedlings of Copaifera multijuga was high in logged areas. C. multijuga survival was observed to be better in large old growth areas. It is important that a threshold of harvesting intensity is determined through research and followed to enable proper succession pathways which enhance the establishment of the native pioneer plants necessary for restoring important ecological processes of bird populations (Paul et al. 2004).

1.5 The Study 1.5.1 Location, vegetation and habitat types Mt. Nakobalevu is situated in the Viti Levu Southern Highlands and has an altitude of 464m at the top of mountain (Masibalavu &Dutson 2006). The vegetation and ecology of Mt. Nakobalevu has not been well researched except for studies by Waqa- Sakiti (2006) on the taxonomy, diversity and distribution of canopy Coleoptera beetles and recently by Tabudravu (2009). Mt. Nakobalevu is located in BirdLife International’s Important Bird Area (IBA) FJ10, home to globally threatened and restricted range forest bird species (Fig 3). The IBA (670 km2) is mostly lowland rainforests along a chain of hills and low mountains. The vegetation is comparatively stunted as a result of the shallowness of the soil and exposure to strong winds (Masibalavu & Dutson 2006). Field observations by Tabudravu (2009) of the same study sites indicated that the sizes of native trees in the two forest types did not differ much. However, there were more creepers, vines and shrubs in recently logged forests than in mature secondary forests at both Nakobalevu and Savura. The vegetation was more scattered and there were more open areas within the recently logged forests whereas the mature secondary forests were denser with less open areas (Fig 4).

Figure 3: Map of Important Bird Area FJ10 situated in the southern Highlands of Viti Levu, Fiji Islands (Masibalavu & Dutson 2006).

The protected forest area was primarily dominated by native trees such as the Agathis sp., Podocarpus sp., Hydriastele vitiensis, Balaka sp., and the ground vegetation comprising of Pandanus sp., ferns and shrubs (Tabudravu 2009). The recently logged forest area was at a lower elevation than the mature secondary forest, approximately 300m, was also lowland rainforest. The recently logged, located adjacent to the mature secondary forest, comprised of shrubs, dense ferns, shrubs, wild ginger, Pandanus sp. and native trees such as Agathis sp., Calophyllum sp., Palaquium sp., Myristica sp., Alphitonia sp., and Parinari sp. (Tabudravu 2009).

The comparative accessibility of the Savura Forest Reserve (Fig 4) and its protected status has resulted in previous ecological studies and botanical collections by several scientists. No plants have been harvested from the reserve since the establishment of the reserve. These include studies by Doyle (1998), and Keppel et al. (2005) on the botanical diversity at Savura. Savura, located in the southeastern Viti Levu, is one of the seven focal sites of the Fiji Pacific-Asia Biodiversity Transect (PABITRA) wet- zone transect is part of the province of Naitasiri. The Savura Forest Reserve is lowland rainforest and its adjacent forest reserve, Vago, acts as catchment areas for the Savura creek (Fig 4). Established in 1963, the Savura Forest Reserve comprises of 396.5 ha (Keppel et al. 2005). A recorded total of 560 native species of vascular plants out of which 52% of the plants are endemic have been recorded in the Savura Forest Reserve (Keppel et al. 2005). Of these, 345 (71% endemic) are dicotyledons, 93 (14% endemic) are monocotyledons, 117 (28% endemic) are ferns and 5 (none endemic) are gymnosperms. Besides these, 27 introduced species were also recorded.

The Savura Forest Reserve is also located in the BirdLife International’s Important Bird Area FJ10. The vegetation of the mature secondary forest at Savura was like that of the mature secondary forest at Nakobalevu with tall emergents and streams running through both. The recently logged forest was in close proximity to the mature secondary forest (Fig 4). The two forest types were separated by the Savura Valley Road. Reeds, vines, African Tulip were dominant at the forest edges of the recently logged forest in Savura.

Figure 4: Map of study locations showing vegetation types from forest survey conducted in 1992 (Management Services Division, Forestry Department, Colo-i- Suva Forestry Station).

Key: MSF 1- Nakobalevu Mature Secondary Forest MUF- Multiple Use Forest MSF 2- Savura Mature Secondary Forest PTF- Protection Forest RLF 1- Nakobalevu Recently Logged Forest PRF- Preserved Forest RLF 2- Savura Recently Logged Forest

1.5.2 Logging disturbance history Land ownership is the major factor determining the allocation of timber harvesting rights in the South Pacific. This is also the basis of timber harvesting in Fiji. The National Code of Logging Practice was launched in Fiji in 1990 and all logging licenses are subject to compliance with the Fiji National Code of Logging Practice. There are four categories of tenure for timber cutting rights in the natural forests in Fiji (FAO 2005). These are: Timber concessions (15-30 year periods) Long term license (10 years) Annual licenses Other licenses and prepayment licenses

The disturbance caused through tree harvesting (Table 2) for logged forests in both Nakobalevu and the Savura Forest Reserve was researched through personal communication with the Forestry Department officers and local villagers, Neno and Vido. A total of 87.627m3 of trees was harvested from Nakobalevu and the Savura Forest Reserve between 2000 and 2001 while a total of 449.503m3 was harvested between 2004 and 2005 (Kububola R., pers comm, March 7, 2008).

Table 2: Fijian vernacular name and its scientific equivalent for major tree species’ harvested from the recently logged forests in Nakobalevu and Savura Forest Reserve from 2000-2001 and 2004-2005. Family Fiji Vernacular Name Scientific Name Chrysobalanaceae Sa Parinari insularum

Gnetaceae Sukau Gnetum gnemon

Sapotaceae Bauvudi Palaquium porphyreum

Thymeleaceae Mavota Gonystylus punctatus

Euphorbiaceae Kauvula Endospermum macrophyllum

Myristicaceae Kaudamu Myristica spp.

Podocarpaceae Kuasi Podocarpus neriifolius

Araucariaceae Dakua makadre Agathis vitiensis

Podocarpaceae Dakua salusalu Decusscicarpus vitiensis

Podocarpaceae Yaka Dacrydium nidulum

1.6 Aims and objectives of the study The aim of this study was to investigate the impact of logging on the diversity and abundance of mid-altitudinal forest birds in the Southern Viti Levu Highlands. The relative density of individual bird species was estimated in adjacently located recently logged and mature secondary forests in two different locations within the Southern Viti Levu Highlands. The study evaluated the impact of logging on the abundance and diversity of forest bird species and using this information, indicator species for forest quality were identified.

CHAPTER 2

METHODOLOGY

2.1 Sampling Design and method Censusing tropical birds is notoriously difficult as most techniques were originally developed for temperate regions (Bibby & Etheridge 1993). Tropical bird species can have several calls which make them difficult to identify in dense tropical forests where chances of sighting birds is low. The bird community in this study was sampled using point count method (Bibby et al. 2000) to estimate bird species richness and abundance. This method is widely used in both temperate and tropical forest surveys. Studies were carried out at two locations, namely Nakobalevu and Savura in the southern Highlands of Viti Levu. Two forest types (treatments) within each location were studied; mature secondary forest where there has been no logging for over 20years and recently logged forest, logged 2-8 years ago. Three 100 ha plots for each forest type within each study location were allocated depending primarily on the characteristics of the habitat such as vegetation type and secondarily on accessibility to the plots (Fig 5). Following the method of ten stratified random point counts were carried out in each 100 ha plot (Bibby et al. 2000). Census points were generated using random number generation. Cumulative means were calculated for two variables (number of individuals and number of species) and plotted against the “point” numbers. The cumulative mean (Appendix 1) became constant after 30 “points” which indicated that 30 “points” per forest type in the two locations were adequate as per Naikatini (2005).

After arriving at each predetermined “point”, I spent a total of 10 minutes at each point, 3 minutes to let birds settle down after disturbance and 7 minutes of identifying and recording bird species within a 25-m fixed radius as in studies carried out by Sodhi et al. 2005 and Bibby et al. 2000. Points were identified using landmarks and with the help of guides who were very familiar with the area. Birds encountered or heard were not recorded for the first three minutes to ensure that there were no observer effects and to minimize the likelihood of double counting. Individuals arriving into the plot during point count were ignored during analysis

22 however individuals flushed out during arrival of the observer were considered (O’Dea et al. 2004). At each plot census points were spaced at least 100m apart to minimize the risk of counting the same individual twice. A distance of 100m between “points” was chosen as the terrain was rugged and due to time limitations. A total of 120 points were carried out for both locations during this study (Table 3).

Prior to the commencement of the actual survey, many trips were made to the study locations and other forests in the central division to learn the calls of bird species and learn to identify the different bird species. Help was sort from Vilikesa Masibalavu of BirdLife International in developing these skills. Preliminary studies were carried out in August 2007 to establish the survey area, plots and survey points. All surveys were carried out from 0600 – 1000 hrs on days with fine weather from August to November, 2007 and in January to February, 2008. The study was carried out during these months as it is believed to be the breeding season of some bird species and most tree flower and produce fruits during these months. Each plot within the two habitats was censused only once. Movement from census point to point (wherever possible) was randomized.

Table 3: Sampling design used for the study. Each plot was 100 ha and each point was at least 100 m apart. Observations were recorded within a 25 m radius of each point. Nakobalevu Mature Plot 1 Plot 2 Plot 3 secondary forest 10 points 10 points 10 points Recently Plot 1 Plot 2 Plot 3 logged 10 points forest 10 points 10 points Savura Mature Plot 1 Plot 1 Plot 3 secondary 10 points 10 points 10 points forest Recently Plot 1 Plot 2 Plot 3 logged 10 points 10 points 10 points forest

Figure 5: Map of study plots at Nakobalevu and Savura (Lands and Survey Department, Suva Fiji).

2.2 Statistical Analysis 2.2.1 Two-way ANOVA Data was checked for normality and homogeneity of variance using Kolmogorov- Smirnov test and Levene’s test respectively (SPSS 12.0 version). Data was not normally distributed therefore were square root transformed. Parametric tests were therefore justified on the basis that the data was normally distributed after square- root transformation. Two-way analysis of variance (ANOVA) was used to test for significant interactions between the two variables of interest; bird abundance and

24 diversity and the habitat type. The statistical significance level used in all tests was P< 0.05 (Dale et al. 2000; Dranzoa 2001; Kizlinski et al. 2002; Adekunle & Olagoke 2008). The R squared value indicated how closely the method and results of this study fit the model. A higher R squared value showed that the method and results of this study fit the model better.

2.2.2 Shannon - Weiner Index and Pielou’s Species Evenness Avian diversity in the two forest types within the two sites was obtained using a mathematical formula that takes into account the species richness and abundance of each species in the ecological community. The Shannon - Weiner diversity index provides a good learning tool for comparing two distinct habitats and combines the quantifiable measures, that is, species richness and species equitability. Pielou’s equitability on the other hand was used to compare the observed Shannon - Weiner diversity index against the distribution of individuals between the observed species which could maximize diversity. The equation for Shannon - Weiner diversity index (Pielou 1993; Krebs 2002; Kent & Coker 1992) that was used is:

Where H1 is the Shannon diversity index, S is the total number of species in the community, pi is the proportion of a species to the total number of birds in the community and Ln is the natural logarithm. Species evenness in each community was determined using Pielou’s equitability (EH) as stated by Kent and Coker (1992):

2.2.3 Dufrêne and Legendre’s Indicator Value Method (IVM) Dufrêne and Legendre’s (1997) Indicator Value Method provides an unequivocal method to detect indicator species by identifying similarities (groups) in sample composites (sites) (Dai et al. 2006). Besides establishing indicator species using absolute bird abundance, IVM is the only method available. The IVM method

25 produced Indicator Values (IV) by combining species abundance in a particular group and the faithfulness of occurrence of a species in a particular group (Carignan & Villard 2002). The Dufrêne and Legendre’s (1997) Indicator Value Method was used to establish which bird species were indicative of the habitat studied. The values were calculated as follows:

For each species i in each site group j, the relative abundance RAij, and the relative frequency RFij, was computed as follows: RAij = Aij / Ai.

Where Aij = the mean abundance of species i across sites of the group j, Ai. = the sum of the mean abundance of species i over all groups.

RFij = Sij / S.j where Sij = the number of sites in group j where species i is present, S.j.= the total number of sites in that group. Then the Indicator value (IV) of species i in the group j is:

IVij = RAij x RFij x 100

IV can range from 0 (no indication) to 100 (perfect indication) (Dai et al. 2006). Higher IV values indicate species are more representative for a given habitat. All species with a minimum total abundance of ten individuals were tested for associations with mature secondary forest and logged forest in Nakobalevu and Savura as per Cleary et al. 2005.

CHAPTER 3

RESULTS

3.1 Number of individuals A total of 30 point counts were carried out for each forest type at each location (Fig. 6 and 7). In total, 120 point counts were carried out for both location combined. There were more individual birds in recently logged forests in both Nakobalevu and

Savura (Table 4) than in mature secondary forests (ANOVA: F4, 116 = 31.66, P treatment = 0.000). The number of individual birds were not significantly (P>0.05) different between the two locations (ANOVA: F4, 116 = 1.023, P site = 0.314). Five hundred and seventy-eight individuals were recorded at Nakobalevu for both forest types combined while 501 individuals were recorded at Savura. There was no interaction found between the locations, Nakobalevu and Savura, and the forest types that were studied, mature secondary forest and recently logged forest (Table 4).

Table 4: Results from two-way ANOVA for number of individuals Source Type III df Mean Square F Sig. (P) Sum of Squares Model 951.024a 4 237.756 215.507 0.000 Forest type 34.929 1 34.929 31.661 0.000 Location 1.129 1 1.129 1.023 0.314 Forest type*Location .157 1 .157 .142 0.707 Error 127.976 116 1.103 Total 1079.000 120 a.R Squared = 0.881 (Adjusted R Squared = 0.877)

0 Mature secondary forest Recently logged forest

Figure 6: Mean number of individuals found in MSF and RLF at Nakobalevu (n=30)

Figure 7: Mean number of individuals found in MSF and RLF at Savura (n=30)

3.2 Number of species Bird species from eighteen different families were observed and recorded in the entire study. A higher number of species were noted in the recently logged forests of both study locations, Nakobalevu and Savura (Table 5). The number of bird species in mature secondary forests were significantly less (Fig. 8 and 9) compared to

28 recently logged forests (ANOVA: F4, 116 = 19.609, P treatment = 0.000). Individuals from the species Zosterops exploratory (Fiji white-eye) were most common in all forest types. The total number of bird species encountered in the two locations did not differ significantly (P= 0.597) (Table 6 & 7). As in the test for number of individuals, there was no interaction noted in the number of bird species recorded between the treatments and the sites (P= 0.279).

ANOVA results show that there is a difference between the response variables (abundance and diversity) according to the forest type, that is, mature secondary forest versus the recently logged forest, but there is no difference between the two locations on the abundance and diversity. Regardless of the level of treatment, there is no difference between the two locations. More individuals were recorded in recently logged forests in both locations. There was also more species present in logged forests in both Nakobalevu and Savura (Table 5). The interaction effect (forest type*location), or the lack thereof, shows that the differences shown between the forest types are consistent across the two locations.

Table 5: Total number of bird species found in the two forest types at Nakobalevu and Savura Mature Secondary Forest Recently Logged Forest

Nakobalevu 17 27

Savura 23 24

Table 6: Results from two-way ANOVA for number of species Source Type III df Mean Square F Sig. (P) Sum of Squares Model 522.493a 4 130.623 356.464 .000 Forest type 7.186 1 7.186 19.609 .000 Location .103 1 .103 .281 .597 Forest type*Location .434 1 .434 1.185 .279 Error 42.507 116 .366 Total 565.000 120 a.R Squared = 0.925 (Adjusted R Squared = 0.922)

Figure 8: Mean number of bird species found in MSF and RLF at Nakobalevu (n=30)

Figure 9: Mean number of bird species found in MSF and RLF at Savura (n=30)

3.3 Shannon - Weiner index and Pielou’s species evenness A total of 192 individual birds from eleven different families were recorded during the study of mature secondary forest at Nakobalevu whilst 386 individual birds from sixteen families were recorded in recently logged forest (Table 9). Species richness, calculated using Shannon - Weiner index, was higher in recently logged forest (S=27) at Nakobalevu compared to the 17 species recorded at mature secondary forest (Table 7). The Shannon - Weiner index (Table 8) for mature secondary forest was H1=2.57 which was higher than in recently logged forest (H1=2.43). Pielou’s species evenness which indicates the proportion of individuals among the species recorded was higher in mature secondary forest (E=0.91). Therefore, individuals were more evenly spread among species of birds recorded in mature secondary forests than in recently logged forest at Nakobalevu (E= 0.74). The values for evenness range from 0 to 1 where a sample of equal numbers of individuals of the same species has a value of 1.

A higher number of individual birds were recorded at the first study location, Nakobalevu (Table 9). In total, 166 individuals (fourteen families) were recorded in mature secondary forest and more than double (N=335 from 18 families) the number of individuals was recorded at the recently logged forest (Table 9). Twenty three different species were observed at mature secondary forest while 24 species were observed at recently logged forest at Savura (Table 5). Diversity indicated by the Shannon - Weiner index (Table 8) at mature secondary forest was H1=2.793. This was slightly higher compared to the diversity at recently logged forest at Savura which was H1=2.635. Species evenness was slightly higher at the mature secondary forest (E=0.891) than in recently logged forest (E=0.829) at Savura indicating that there were similar number of individuals of each species recorded in mature secondary forest.

The evenness measure was higher in mature secondary forests than in recently logged forests which means that in recently logged forests a few species have large numbers and most species have low numbers. On the other hand, the mature secondary forest examined had more species that had large numbers and there were just a few species with low numbers. Hence only a few species do well in recently logged forests but in mature secondary forests more species have higher numbers and hence do better.

Table 7: Relative abundance of bird species recorded in mature secondary forest (MSF) and recently logged forest (RLF) at study locations Family Scientific Name Common Nakobalevu Savura Name and MSF RLF MSF RLF species status Threskiornithidae Anas Pacific 0 0 0 5 superciliosa black duck

Phasianidae Accipiter Fiji 0 0 0 1 rufitorques goshawk

Columbidae Ducula latrans Barking 13 16 10 15 pigeon

Chrysoenas Golden 2 8 3 3 luteovirens dove Psittacidae Prosopeia Masked 8 18 13 9 personata shining parrot

Phigys Collared 0 8 5 0 solitarius lory

Cuculidae Cacomantis Fan -tailed 2 1 2 1 flabelliformis cuckoo

Apodidae Aerodramus White - 0 17 0 24 spodiopygius rumped swiflet

Alcedinidae Todirhamphus White - 1 7 5 1 chloris collared kingfisher

Sturnidae Acridotheres Common 0 4 2 3 tristis mynah

Pycnonotidae Pycnonotus Red -vented 0 3 1 11 cafer bulbul

Muscicapidae Turdus Island 11 2 11 8 poliocephalus thrush

Sylviidae Cettia Fiji bush – 22 45 29 42 ruficapilla warbler

Eopsaltriidae Petroica Scarlet 0 2 1 1 multicolor robin

Monarchidae Rhipidura Streaked 17 10 10 11 spilodera fantail

Mayrornis Slaty 15 15 7 16 lessoni monarch

Clytorhynchus Lesser 6 2 4 3 vitiensis shrikebill

Clytorhynchus Black - 0 3 1 0 nigrogularis faced shrikebill

Myiagra Vanikoro 3 7 5 12 vanikorensis broadbill

Myiagra Blue - 10 9 14 13 azureocapilla crested broadbill

Pachycephalidae Pachycephala Golden 6 1 6 4 pectoralis whistler

Campephagidae Lalage Polynesian 10 22 2 18 maculosa triller

Zosteropidae Zosterops Fiji white - 35 146 11 86 explorator eye

Zosterops Silvereye 0 1 0 0 lateralis

Ploceidae Erythrura pealii Fiji 0 5 0 9 parrotfinch

Erythrura Pink -billed 0 1 0 0 kleinschmidti parrotfinch

Meliphagidae Myzomela Orange - 18 12 9 22 jugularis breasted myzomela

Foulehaio Wattled 0 8 1 0 carunculata honeyeater

Gymnomyza Giant forest 13 13 14 17 viridis honeyeater

Table 8: Bird species richness (S), absolute number of individual birds observed (N) Shannon’s indices (H1) and Pielou’s species evenness (E) at each study location Study Location S N H1 E Nakobalevu Mature secondary forest 17 192 2.6 0.91 Recently logged forest 27 386 2.4 0.74

Savura Mature secondary forest 23 166 2.8 0.89 Recently logged forest 24 335 2.6 0.83

Table 9: Total number of individual birds and the total number of families [ ] recorded at the study locations, Nakobalevu and Savura Study Location Forest Type Total Nakobalevu MSF RLF 192 386 578 [11] [16]

Savura 166 335 501 [14] [18]

3.4 Dufrêne and Legendre’s Indicator Value Method (IVM) 3.4.1 Nakobalevu Twelve bird species (minimum total abundance of more than ten individuals) had significant associations for the designated forest types at Nakobalevu (Table 10). A large number of individuals (MSF=164, RLF=314) of these twelve species were observed at Nakobalevu. Of these, ten species were associated with mature secondary forests and ten species with recently logged forests. Eight species had significant associations with both forest types. These were the Barking pigeon, Fiji bush-warbler, Streaked fantail, Slaty monarch, Polynesian triller, Fiji white-eye,

Orange-breasted myzomela and the Giant forest honeyeater. The strongest associations were shown between the Orange Breasted Myzomela and mature secondary forest (IV=28) at Nakobalevu followed by Streaked Fantail and Giant Forest Honeyeater with an indicator value of 18.89 and 18.33 respectively. The Fiji Bush-warbler had the strongest association with recently logged forest (IV=53.73). The second strongest association was seen between recently logged forest and Fiji White-eye with an indicator value of 51.09. Polynesian Triller also showed association with the recently logged forest at Nakobalevu (IV= 34.38).

3.4.2 Savura A total of fourteen bird species showed significant interaction at the second study location, Savura (Table 11). Eight bird species had association with mature secondary forest and twelve with recently logged forest. Six species were recorded in both forest types had a minimum abundance of 10 individuals. These were the Barking pigeon, Fiji bush-warbler, Streaked fantail, Blue crested broadbill, Fiji white-eye and the Giant forest honeyeater. Fiji Bush-warbler had the strongest association with mature secondary forest with an indicator value of 25.87. Other species such as the Island Thrush and the Giant Forest Honeyeater also had association with this forest type. The indicator value for these two species was 19.3 and 18.06 respectively. The highest indicator value at Savura’s recently logged forest was shown by the Fiji Bush-warbler (IV=45.35). This was followed by the Polynesian Triller which had an indicator value of 36 with the recently logged forest at Savura. The Orange Breasted Myzomela which had an indicator value of 35.48 had the third strongest association with recently logged forest at Savura. The White- rumped swiftlet was only recorded in the recently logged forests at both study locations. No species were recorded that were exclusive to the mature secondary forests at the two study locations.

Table 10: Species established as being indicative of forest type at Nakobalevu using the Indicator Value Method. Only species with a minimum total abundance of ten individuals were tested for associations. * Absolute abundance less than 10 individuals. Species Mature Secondary Forest Recently Logged Forest RAij RFij IVij RAij RFij IVij

Barking pigeon 0.45 0.27 11.95 0.55 0.37 20.23

Masked shining * * * 0.69 0.43 30 parrot

White-rumped * * * 1 0.17 16.67 swiftlet

Island thrush 0.85 0.2 16.92 * * *

Fiji bush- 0.33 0.47 15.32 0.67 0.8 53.73 warbler Streaked fantail 0.63 0.3 18.89 0.37 0.23 8.64

Slaty monarch 0.5 0.23 11.67 0.5 0.37 18.33

Blue crested 0.53 0.17 8.77 * * * broadbill Polynesian 0.31 0.23 7.29 0.69 0.5 34.38 triller Fiji white-eye 0.19 0.2 3.87 0.81 0.63 51.09

Orange- 0.6 0.47 28 0.4 0.37 14.67 breasted myzomela Giant forest 0.5 0.37 18.33 0.5 0.37 18.33 honeyeater

Table 11: Species established as being indicative of forest type at Savura using the Indicator Value Method. Only species with a minimum total abundance of ten individuals were tested for associations.* Absolute abundance less than 10 individuals. Species Mature Secondary Forest Recently Logged Forest RAij RFij IVij RAij RFij IVij

Barking pigeon 0.4 0.27 10.67 0.6 0.3 18

Masked shining 0.59 0.27 15.76 * * * parrot

White-rumped * * * 1 0.13 13.33 swiftlet

Red-Vented * * * 0.92 0.13 12.22 bulbul Island thrush 0.58 0.33 19.30 * * *

Fiji bush- 0.41 0.63 25.87 0.59 0.77 45.35 warbler Streaked fantail 0.48 0.2 9.52 0.52 0.2 10.48

Slaty monarch * * * 0.70 0.27 18.55

Vanikoro * * * 0.71 0.33 23.53 broadbill Blue crested 0.52 0.3 15.56 0.48 0.33 16.05 broadbill Polynesian * * * 0.9 0.4 36 triller Fiji white-eye 0.11 0.07 0.76 0.89 0.3 26.60

Orange- * * * 0.71 0.5 35.48 breasted myzomela Giant forest 0.45 0.4 18.06 0.55 0.4 21.94 honeyeater

3.5 Diet type and habitat of bird species found mature secondary forest and recently logged forest A total of 29 different species of birds were recorded during this study (Table 5). Majority of these were recorded in both forest types with the exception of the following: White-rumped swiftlet and the Fiji parrotfinch which were only observed in the recently logged forests at both study locations Silvereye and the Pink – billed parrotfinch found only in the recently logged forest at Nakobalevu Fiji goshawk and Pacific black duck which were found only in the recently logged forest at Savura

Bird species which were predominantly insectivorous were common in both forest types at both study locations (Appendix 2). Ten out of the 17 bird species were predominantly insectivorous in the mature secondary forest in Nakobalevu while 16 of the 27 bird species in the recently logged forest. The Fiji parrotfinch was the sole bird species in recently logged forests in both study locations which an insectivore and also feed on grass (graminivore). Two predominantly omnivorous bird species, 3 predominantly frugivores and 2 predominantly nectarivores were recorded in the mature secondary forest at Nakobalevu while 4 omnivorous, 3 frugivores and 4 nectarivores were recorded in the recently logged forest.

A total of 23 bird species were recorded in the mature secondary forest at Savura. Of these 13 were predominantly insectivorous, 3 omnivorous, 3 frugivorous and 4 nectarivorous. In the recently logged forest at Savura, 15 were predominantly insectivorous, 4 omnivorous, 3 frugivorous and 3 nectarivorous bird species were observed.

CHAPTER 4

DISCUSSION

4.1 Bird abundance in mature secondary forests and recently logged forests This study was carried out to investigate the effects of logging on the diversity and abundance of mid-altitudinal forest birds in the Southern Viti Levu Highlands of Fiji. The relative abundance of bird species was estimated and using this information, indicator species for forest quality was identified. This study does not show any negative impact of logging on the abundance of birds. There were more individual birds recorded in recently logged forests at both Nakobalevu and Savura than in mature secondary forests (Table 7). The Fiji white-eye was even more abundant in the recently logged forests than the mature secondary forest. This was also the case with the Fiji Bush-warbler, Barking pigeon, Polynesian Triller and the Vanikoro Broadbill.

Results from this current study agree with studies on bird diversity and abundance which have attributed an increase in abundance of birds to the increased sunlight reaching the forest floor of recently logged forests (Thiollay 1992). As a result, plants in the logged understory and the area immediately surrounding the gaps created by logging respond positively to this increase in sunlight by producing more flowers and fruits (Smith 1991). This increase in fruits and flowers would thus be likely to draw bird species relying on fruits and flowers for food such as the Fiji White-eye, Polynesian Triller and the Barking pigeon the latter which is predominantly a fruit feeding species (frugivore). The increased production of flowers and fruits also means more nectar and therefore more insects would be attracted to the nectar (Thiollay 1992; Plumptre 1997; Owiunji & Plumptre 1998; Dale et al. 2000). This in turn would be expected to draw in insectivores such as the Fiji White-eye, Fiji Bush-warbler, Polynesian Triller and the Vanikoro Broadbill. This is seen in the current study where there is a higher number of these bird species in recently logged forests at the two study locations. More light, and the presence of fruits and flowers also makes “gaps” more attractive to some species than would the

40 darker understory of unlogged forests (Wunderle et al. 2006a). These would be bird species which are predominantly insectivorous, nectarivorous and frugivorous.

Some studies conducted in neotropical logged forests have reported significant decreases in the bird abundance in logged forests compared to unlogged forests. For example major changes in the vegetation structure caused by logging were stated as the reason for this decrease in abundance (Johns 1991; Thiollay 1992). In addition, more recent studies show that a decrease in abundance can result from the lower breeding success in logged forests (Dranzoa 2001). This resulted from reduced nest availability, fewer nest materials, increased competition for nest sites and a significant increase in nest predation and parasitism. Negative effects of logging on bird abundance and diversity are well documented by Faaborg et al. (1995) and Robinson et al. (1995). The influence of past logging on breeding potential and success differ from species to species (Dranzoa 2001). Some species are affected in a positive way by logging while some species are affected negatively. As seen in this study, frugivores may have benefited from logging as the increased sunlight most likely stimulated plants to produce more fruits and flowers which were consumed by fruit eating birds such as Barking pigeons. However, birds such as the Masked Shinning Parrot may have been affected negatively probably because trees with holes and fissures that this bird species would use to build their nests in were destroyed or lessened by logging.

4.2 Species richness and diversity in mature secondary forests and recently logged forests Species richness and diversity indices were higher in the mature secondary forest (MSF) and the recently logged forest (RLF) for bird communities at both Nakobalevu and Savura Forest Reserve (Table 8). There was a considerable difference in the species richness between the forest types in Nakobalevu. A total of 17 species were recorded at the mature secondary forest whereas 27 species were recorded at the recently logged forest at Nakobalevu. There was only one more species at the recently logged forest at Savura compared to the mature secondary forest (S=24). Diversity of the mature secondary forests at both study locations was higher as there were more species present which were equally abundant than in

41 recently logged forests where individuals of some species were more abundant than other bird species. The Shannon - Weiner index takes into account both the species richness and the evenness thus is a very good indicator of the ecological status of the forest (Fowler 2008). Results from this study show that even though species richness is higher in recently logged forests compared to mature secondary forests, the mature secondary forests may have a more stable bird population, looking at the results of the Shannon - Weiner index and Pielou’s Species Evenness, therefore a bird population with a higher conservation priority. Majority of the bird species recorded in this current study are endemic to Fiji (Appendix 2).

The differences in diversity of bird communities coincide with the changes in vegetation structure observed between mature secondary forests and recently logged forests. The changes in vegetation structure, with the production of large areas of regenerating habitats, may have allowed an increase in species richness in the bird community of recently logged forests in this study. This pattern of disturbance allows the coexistence of many forest species and species adaptable to forest edges in the recently logged forests resulting in an increase in species richness (Schemske & Brokaw 1981; Wunderle et al. 2006a; Johns 1991; Aleixo 1999). Similar results in species richness were also found among numerous groups of organisms such as birds and butterflies in studies carried out by Lugo (1988). The same pattern of increased species richness was also observed in studies of several animal groups such as beetles, butterflies, tapirs, rodents and primates found in selectively logged forests in Latin America, Africa and Asian Tropics (Frumhoff 1995).

Another factor contributing to the increase in species richness in recently logged forests from studies by Stouffer and Bierregaard (1995) is the proximity of the recently logged forests to the mature secondary forests. This permits a continuous flow of individuals between the recently logged forests and the mature secondary forests thus preventing the extinction in recently logged forests even of species with low dispersal abilities. According to Aleixo (1999) an extensive protected area with mature secondary forest close to logged forests, with almost 500 km2, provides a steady influx of individuals searching for territories or nest sites at logged forests. Other studies with which results from this study can be corroborated are by Johns (1991) and Lambert (1992). The secondary mature forests in this current study were

42 in close proximity (Fig. 5) to the recently logged forests therefore, it is highly possible that individuals are able to travel between the two forest types. In addition to this, this close proximity may also explain the presence of bird species such as the Barking pigeon, Masked shinning parrot and the Fiji bush-warbler that were expected to only be found in mature secondary forest but were also recorded in the logged study sites.

Some studies have reported significant decreases in bird species richness following logging when compared to unlogged forests (Johns 1991; Thiollay 1992). These decreases were due to extensive damage in the forest structure resulting in loss of breeding and nesting habitats and materials for the bird species (Thompson et al. 1992; Dranzoa 2001; Robinson & Robinson 2001). Logging also opens roads with give access to predatory species such as rats (Peh et al. 2005). Studies show logging trails drastically increase nest and brood predation therefore resulting in the decrease in species richness and diversity (Villard et al. 1996; Robinson et al. 1995; Peh et al. 2005). Another deleterious impact of logging on bird community affecting the species richness and diversity would probably be an increase in distance between the recently logged forests and the continuous unlogged forest which result in fragmentation and isolation effects (Thiollay 1992; Stouffer & Bierregaard 1995). The greater the distance between these two forest types the harder it becomes for the reestablishment of bird populations in the logged forests. In this study, as the two forest types were adjacent to each other, isolations effects leading to decrease in bird abundance and diversity in the recently logged forests were not observed.

In this study, logging did not reduce the species richness of the bird community in the mid-altitudinal tropical rainforests of the Viti Levu Southern Highlands in Fiji. This may have been directly due to the close proximity of the two forest types at the study locations. This result does not translate directly into conservation priorities, mostly due to the fact that additional species observed in recently logged forests are associated with disturbance, regenerative and semi open habitats, and consequently are more widespread and of lower conservation value than species restricted to mature secondary forests (Appendix 2) (Stotz et al. 1996).

4.3 Indicator species’ for mature secondary forests and recently logged forests There were significant associations between some bird species and the mature secondary forest in Nakobalevu and Savura (Table 10 & 11). However, these bird species could be established as indicator species for mature secondary forests as they were also found to have significant associations with recently logged forests. No indicator species were specifically found in this study for the mature secondary forests (Table 7, 10 & 11). This was because no bird species recorded only in mature secondary forests. This may have been due to the close proximity of the two forest types at the two study locations. The close proximity would allow birds from the mature secondary forest to travel to the recently logged forest and vice versa.

Indicators for the recently logged forests have been identified in this study. Barking pigeons, the Fiji bush-warbler and the Fiji white-eye were found in higher numbers and had stronger associations with the recently logged forests at both Nakobalevu and Savura. According to Watling (2004), the Fiji White-eye is a mixed feeder, feeding on insects in the canopy foliage, also on fruits and berries and seldom on nectar. The recently logged forest which receives a large amount of sunlight through the gaps created is abundant in nectar producing flowers, fruits and insects are therefore excellent habitats for the Fiji White-eye. The Barking pigeons and Fiji bush-warbler would also be abundant in recently logged forests due to the abundance in insects and fruit production in this forest type. The Fiji parrotfinch and the White- rumped swiftlets were the only two bird species which were only present in the recently logged forests (Table 7). These two bird species are clear indicators of recently logged forests and they are graminivorous and insectivorous respectively. Recently logged forests had open patches areas and were dominated with vines, shrubs and grasses and due to the high production of flowers, nectar and fruits may result in an increase in insects favoring these two bird species. The Fiji parrotfinch is known to specialize in seeds at the ‘milk stage’, and a favored food of this bird species is the Guinea Grass (Panicum maximum), dominant in degraded forests. The White-rumped swiftlets which is also an indicator of recently logged forests feeds exclusively on flying insects.

Other species that have been recorded in this study have shown to have flexible habitats, that is, they were found in both mature secondary and recently logged forests at both the study locations. According to Watling (2004), niche broadening is commonly seen in bird species in Fiji, especially in the Slaty Monarch and the Lesser Shrikebill. Only true endemics have very restricted habitat use, for example the Long-legged Warbler (not recorded in this study) associated with mature forest which has dense undergrowth and fast flowing rivers or waterfalls (Vilikesa Masibalavu, pers comm, 2008). Exceptions to this as noted by Watling (2004), are the Collared Lory, Wattled Honeyeater, Polynesian Triller and the Vanikoro Broadbill which are very adaptable species. In addition to this, the study also noted that Mynahs and Bulbuls are unable to fully inhabit mature secondary forests thus were not recorded in mature secondary forest at Nakobalevu which was less accessible than the mature secondary forest at Savura (Table 7). They can easily venture along the roads and rivers of the mature secondary forests in search for their weed colonized recently logged forests.

4.4 Conclusion and recommendations It can be concluded from this study that logging does affect the bird abundance and diversity. Disturbances caused by logging did reduce the diversity of the bird community. However, the species richness increased as a result of logging. This increase in species richness does not directly relate to conservation priorities as the additional species observed are associated with disturbed habitats and are of lower conservation value than species that are associated with mature secondary forests. For this reason, it is vital to invest in the conservation of mature secondary forests. It is also important to use diversity indices such at the Shannon - Weiner Diversity Index as it provides more information than simply the number of species present, that is, they account for some species being rare and others being common. Fijian forests and Fijian birds have some adaptability for habitat disturbance and the avian community is able to recover after logging provided the forest is permitted to recover and where source populations survive.

Based on the findings of this study, the following procedures are proposed to minimize the adverse effects of logging on bird communities in the tropical rainforests in the two study locations: 1. Logging practices should be well planned and should be of low intensity to minimize the effects on the forest bird communities, 2. logged areas should be close enough to unfragmented mature secondary forest to allow the recolonisation of bird species with high ecological value, 3. the exploitation of the forest should be carried out using as few roads as possible to reduce fragmentation and edge effects and to decrease accessibility to invasive alien predators, 4. minimizing the use of equipment to fell and transport trees to reduce effects of fragmentation and isolation, 5. long-term rotation should be used in areas designated for logging to allow time for regeneration.

This study leaves scope for further research which would benefit greatly with a larger sample size carried out in broader area and studies over a long term to establish with certainty the abundance and diversity of mature secondary and logged forests.

CHAPTER 5

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APPENDICES

APPENDIX 1

DETERMINATION OF THE NUMBER OF POINTS REQUIRED

10 9 8 7 6 mean 5 4 3 2 1 0 0 5 10 15 20 25 30 35 Point number

Graph of cumulative mean of number of individual birds recorded in mature secondary forests at Nakobalevu. Cumulative mean becomes relatively constant upon reaching 30 points indicating that carrying out further points would not significantly improve results.

10 mean 8

0 0 5 10 15 20 25 30 35 number of points

Graph of cumulative mean of number of individual birds recorded in mature secondary forests at Savura. Cumulative mean becomes relatively constant upon reaching 30 points indicating that carrying out further points would not significantly improve results.

APPENDIX 2

Diet, feeding habit and status of bird species recorded at study locations (Watling 2001, 2004). Scientific name Common Diet Species Commonality Feeding Name endemicity/s habit tatus Anas Pacific Animal and Resident, Not recorded Not superciliosa black duck vegetable native recorded matter breeding land bird Accipiter Fiji Mainly Fiji endemic/ Not recorded Not rufitorques goshawk insectivore Resident, recorded also rats native and lizards breeding land bird Ducula latrans Barking Frugivore Fiji endemic/ Common Forages pigeon Resident, in canopy native breeding land bird

Chrysoenas Golden Frugivore Viti Levu Fairly common Forages at luteovirens dove endemic/ sub- Resident, canopy native level breeding land bird Prosopeia Masked Mainly Viti Levu Fairly common Forages personata shining fruits and endemic/ throughou parrot seeds; also Resident, t forest flowers, native strata buds, breeding shoots land bird

Phigys Collared Nectarivore Fiji endemic/ Not recorded Not solitarius lory ; also soft Resident, restricted fruit native to breeding any land bird specific habitat

Cacomantis Fan -tailed Insectivore Resident, Fairly common Not flabelliformis cuckoo native recorded breeding land bird Aerodramus White - Insectivore Resident, Fairly common Aerial spodiopygius rumped native feeder swiflet breeding land bird

Todirhamphus White - Insectivore; Resident, Not recorded Not chloris collared also lizards, native recorded kingfisher young breeding birds, crabs land bird Acridotheres Common Insectivore; Introduced Not recorded Ground tristis mynah also fruit, breeding feeding grains, species domestic waste Pycnonotus Red - Omnivorou Introduced Very common Not cafer vented s but breeding recorded bulbul mainly species fruits, berries

Turdus Island Snails, Resident, Not recorded Ground poliocephalus thrush worms, native feeder insects; breeding also fruit land bird

Cettia Fiji bush – Insectivore Fiji endemic/ Common Forages ruficapilla warbler Resident, in native thick breeding vegetation land bird

Petroica Scarlet Insectivore Resident, Not recorded Forages multicolor robin native from breeding ground to land bird lower canopy level

Rhipidura Streaked Insectivore Resident, Fairly common Forages spilodera fantail native at breeding lower land bird canopy level

Mayrornis Slaty Insectivore Fiji endemic/ Common Forages at lessoni monarch Resident, sub- native canopy breeding level land bird

Clytorhynchus Lesser Mainly Resident, Fairly common Feed at vitiensis shrikebill Insectivore; native any also small breeding height fruits land bird Clytorhynchus Black - Insectivore Fiji endemic/ Not recorded Feeds at nigrogularis faced Resident, any shrikebill native height breeding land bird

Myiagra Vanikoro Insectivore Resident, Fairly common Forages at vanikorensis broadbill native canopy breeding level land bird Myiagra Blue - Insectivore Fiji endemic/ Fairly common Forages at azureocapilla crested Resident, lower broadbill native canopy breeding level land bird Pachycephala Golden Mainly Resident, Fairly common Forages at pectoralis whistler Insectivore; native any level also fruits breeding land bird

Lalage Polynesian Mainly Resident, Common Not maculosa triller Insectivore; native recorded also fruits breeding land bird

Zosterops Fiji white - Insects also Fiji endemic/ Very common Varies explorator eye fruits Resident, from native canopy to breeding any land bird height from ground Zosterops Silvereye Insects also Resident, Not recorded Canopy to lateralis fruits native ground breeding level land bird

Erythrura pealii Fiji Graminivor Fiji endemic/ Not recorded Not parrotfinch e and Resident, recorded insectivore native breeding land bird Erythrura Pink -billed Insectivore Viti levu Not recorded Feeds at kleinschmidti parrotfinch endemic/ any Resident, height native breeding land bird Myzomela Orange - Nectarivore Fiji endemic/ Fairly common Feeds at jugularis breasted Resident, any myzomel native height a breeding from the land bird ground Foulehaio Wattled Nectarivore Resident, Not recorded Feeds at carunculata honeyeater ; also small native any insects such breeding height as spiders land bird from the ground

Gymnomyza Giant forest Nectarivore Resident, Very common Forages at viridis honeyeater ; also small native canopy insects such breeding level but as spiders, land bird maybe small observed berries and at lower soft fruits levels