Using Herpetofauna and Culture to Determine Conservation Priority Habitats on Malaita Island, Solomon Islands

Home , Pothos (plant), Prasinohaema virens

EVALUATING TROPICAL FOREST ECOSYSTEMS: USING HERPETOFAUNA AND CULTURE TO DETERMINE CONSERVATION PRIORITY HABITATS ON MALAITA ISLAND, SOLOMON ISLANDS

Edgar John Maeniuta Pollard

A thesis submitted in fulfilment of the requirements for the degree of Masters in Environmental Science

School of Geography, Earth Science and Environmental Sciences

Faculty of Science, Technology and Environment

The University of the South Pacific

July, 2013

Declaration

Statement by Author I, Edgar John Maeniuta Pollard, 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 other degree at any institution, except where due acknowledgment is made in the text.

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Statement by Supervisor The research in this thesis was performed under my supervision and to my knowledge is the sole work of Mr Edgar John Maeniuta Pollard.

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Acknowledgements Mauriha e hitarana rapena noni. Apu ana noni e hitarana kahu ni mauriha, susurina noni e hitarana okiraha ana mauriha, manina noni e hitarana mako e watea mauriha, ihihu ana pau na noni e hitarana ai ma maasu e watea mauriha

This thesis is dedicated to the people of Are’Are, Malaita. For captured within this work is a glimpse of the richness their lands and people hold. Raemanoha Rikaa.

I would like to thank the stewards, landowners and tribes of the Arata’s (land) and villages that this study was conducted in, for allowing access onto their land. I would also like to thank my field guides and assistants – John Mahane, Pauro Horipeo, Wencis Rohoia, Francis Aniratana Jr and Peter Aitai. For the mountains we climbed, the rivers we crossed, for being soaking wet in the middle of the forest in the middle of the night and for the friendship and knowledge shared along the way, I will be forever thankful. Special thanks also to; Myknee Sirikolo for knowledge imparted on the identification of plants, Mike McCoy on the identification of reptiles and Patrick Pikacha on the identification of frogs.

A very big tagio tumas for my supervisors, mentors and advisors – Dr. Gilianne Brodie, Dr. Clare Morrison and Prof. Randy Thaman, who’s tireless hours and constructive feedback help shape this entire project from start to finish. A special thanks to Patrick Pikacha for teaching me hands on skills and knowledge regarding frogs and bush fieldwork and Marika Tuiwawa who’s advice and encouragement and wise counsel was valued.

I would also like to thank the University of the South Pacific, for enabling me to carry out this research and for the support rendered, especially the Departments of Biology and Geography. This work would also not be possible without the help of my sponsors, the Solomon Islands government and the USP research office.

Last but not the least I would like to thank my ever supportive family, mum, dad and wife Patricia who have all stuck by me and supported me through it all.

Abstract Within the context of the global biodiversity crisis there is a need to identify conservation priority habitat types. This study aims to identify important forest habitats for conservation priority setting on the island of Malaita, Solomon Islands. To achieve this five different forest habitat types were sampled to quantify richness of biodiversity based on richness and abundance of frogs and lizards (herpetofauna) as biological indicators. In addition, interviews with local community members were conducted to gather associated local cultural knowledge on frogs, lizards and forest habitats. The study focused on unlogged coastal, unlogged lowland and unlogged upland forests, logged lowland forests and plantation teak forests, with the two latter having significant human influence resulting in reduced herpetofaunal richness. Prioritisation methods used to identify important forest habitat types were based on: 1) species richness and abundances, 2) ‘important’ (threatened, totem, rare and indicator) species presence, 3) cultural importance of the forest habitat and 4) the threatened status of the forest habitat. It was found that: 1) lowland forests contained the greatest species richness and the greatest number of important species, 2) lowland forests also had the highest cultural value based on locally described uses, and 3) coastal forests were under the greatest threat from anthropogenic activities. The overall results show the importance of biological sampling being coupled with cultural knowledge to improve our understanding of forest habitat value for conservation action.

Abbreviations CBD Convention on Biological Diversity CBSI Central Bank of Solomon Islands CI Conservation International CRV Combined Rank Value CV Cultural Values DQS Diurnal Quadrat Sampling FAO Food and Agriculture Organization FTV Forest Threat Value GDP Gross Domestic Product IBA Important Bird Area IFA Important Forest Area IHA Important Herpetofaunal Area IUCN International Union for Conservation of Nature ISV Important Species Value ITCZ Inter Tropical Convergence Zone MDG Millennium Development Goals MPA Marine Protected Area MoFR Ministry of Forestry and Research (Solomon Islands) NVES Nocturnal Visual Encounter Survey OJP Ontong Java Plateau PHCG Pacific Horizons Consultancy Group PNG Papua New Guinea SINSO Solomon Islands National Statistics Office SPC Secretariat of the Pacific Community SPRH South Pacific Regional Herbarium SRAV Species Richness and Abundance Value SVL Snout Vent Length TEK Traditional Ecological Knowledge TK Traditional Knowledge UNEP United Nations Environment Programme WCMC World Conservation Monitoring Centre iii

Table of Contents Acknowledgements ...... i Abstract ...... ii Abbreviations ...... iii Table of Contents ...... iv List of Tables ...... viii List of Figures ...... x CHAPTER 1: INTRODUCTION ...... 1 1.1 Introduction ...... 1 1.2 Rationale and Justification for Study ...... 2 1.3 Objectives/Aims and Hypotheses ...... 4 1.4 Structure and Outline of Thesis ...... 5 CHAPTER 2: BACKGROUND ...... 6 2.1 Tropical Biodiversity ...... 6 2.1.1 Importance of tropical biodiversity ...... 6 2.2 Tropical Forest Ecosystems ...... 7 2.2.1 Status and importance of tropical forests ...... 7 2.3 Herpetofauna ...... 8 2.3.1 Status and Importance of herpetofauna ...... 8 2.3.2 Indicators of ecosystem health, the use of herpetofauna...... 9 2.4 Threats and Decline of Biodiversity ...... 10 2.4.1 Specific threats to tropical forests ...... 12 2.4.2 Specific threats to tropical herpetofauna ...... 15 2.5 Conservation of Biodiversity...... 16 2.5.1 What is conservation? ...... 16 2.5.2 How do we conserve biological diversity ...... 18 2.5.3 Conservation and traditional ecological knowledge (TEK) ...... 21 CHAPTER 3: STUDY LOCATION AND GENERAL METHODOLOGY ...... 23 3.1 Study location ...... 23 3.1.1 Solomon Islands ...... 23 3.1.2 Malaita ...... 25 3.1.3 Are`Are study site ...... 30 3.2 Pilot study and General Methodology ...... 31 iv

3.2.1 Pilot Study ...... 31 3.2.2 Major Fieldwork ...... 33 CHAPTER 4: RICHNESS AND ABUNDANCE OF FROGS, GECKOS AND SKINKS ON MALAITA ...... 37 4.1 Introduction ...... 37 4.2 Specific Methodology ...... 37 4.3 Results ...... 39 4.3.1 Summary of results ...... 39 4.3.2 Nocturnal herpetofauna ...... 39 4.3.3 Diurnal herpetofauna ...... 49 4.3.4 Additional species ...... 55 4.3.5 Species behaviour and Indicator species ...... 55 4.4 Discussion of Results ...... 56 4.4.1 Indicator Species ...... 56 4.4.2 Herpetofaunal richness comparisons to other studies ...... 57 4.4.3 Herpetofaunal richness comparisons to other Solomon Island islands ...... 57 4.4.4 Malaitan Herpetofaunal richness compared to McCoy and Pikacha ...... 60 4.4.5 Evaluation of methods used ...... 62 4.5 Summary of herpetofaunal richness and abundance ...... 62 CHAPTER 5: FOREST HABITAT AND HERPETOFAUNAL RICHNESS ...... 64 5.1 Introduction ...... 64 5.2 Specific Methodology ...... 67 5.3 Results ...... 69 5.3.1 Unlogged Coastal Forest ...... 69 5.3.2 Unlogged Lowland Forest ...... 70 5.3.3 Unlogged Upland Forest ...... 72 5.3.4 Logged Lowland Forest...... 73 5.3.5 Teak Plantation Forest ...... 75 5.3.6 Comparison of herpetofauna richness in the different habitat types ...... 76 5.3.7 Priority forest habitat based on herpetofauna species richness ...... 78 5.3.8 Impact of habitat degradation and modification ...... 80 5.4 Discussion ...... 83 5.5 Summary ...... 86

CHAPTER 6: TRADITIONAL KNOWLWEDGE OF HERPETOFAUNAL BIODIVERSITY AND FORESTS IN ARE`ARE, MALAITA ...... 87 6.1 Introduction ...... 87 6.2 Specific Methodology ...... 88 6.3 Results ...... 89 6.3.1 Herpetofauna...... 89 6.3.2 Forests ...... 101 6.3.3 Informants knowledge of frogs and lizards by age and gender ...... 108 6.4 Discussion ...... 109 6.4.1 Traditional knowledge of herpetofauna ...... 109 6.4.2 Threatened forest habitats...... 109 6.4.3 Loss of cultural practises and traditional knowledge ...... 110 6.4.4 Loss of traditional knowledge in the younger generation ...... 110 6.5 Summary ...... 111 CHAPTER 7: POTENTIAL PRIORITY HABITATS AND STRATEGIES FOR FOREST BIODIVERSITY CONSERVATION ...... 112 7.1 Introduction ...... 112 7.2 Methods for Prioritisation ...... 113 7.3 Results ...... 115 7.3.1 “Species richness and abundance value” (SRAV) ...... 115 7.3.2 “Important species value” (ISV) ...... 116 7.3.3 “Cultural value” (CV) ...... 116 7.3.4 “Forest threat value” (FTV) ...... 117 7.3.5 “Combined rank value” (CRV)...... 117 7.4 Discussion ...... 119 7.4.1 Species richness and abundance ...... 119 7.4.2 Important species ...... 119 7.4.3 Culture ...... 120 7.4.4 Forest threat ...... 120 7.4.5 Combined ...... 120 7.6 Conclusion ...... 121 CHAPTER 8: OVERALL SUMMARY OF RECOMMENDATIONS FOR FUTURE CONSERVATION WORK ON MALAITA ...... 122 8.1 Introduction ...... 122 vi

8.2 Important Recommendations for Future Conservation work on Malaita based on Literature ...... 122 8.2.1 The importance of culture ...... 123 8.2.2 The importance of conservation science ...... 123 8.2.3 The importance of policy...... 124 8.3 Important Recommendations for Conservation work on Malaita based on this Study ...... 124 8.4 Conclusion ...... 125 LITERATURE CITED ...... 126 Appendix A: Ethnological Questionnaire ...... 136 Appendix B: Species Descriptions with Field Photographs...... 145 Frogs ...... 145 Lizards (Geckos) ...... 153 Lizards (Skinks) ...... 156

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List of Tables Table 2.1 Categories of goods and services provided by biodiversity 6 Table 2.2 Consequences of deforestation 13 Table 2.3 Logging yield, Solomon Islands 14 Table 2.4 Criteria used in the prioritisation of biodiversity conservation 18 Table 2.5 Percentage Terrestrial and Marine Protected Areas Cover 20 Table 3.1 Comparison of population density among Solomon Island 27 Provinces Table 3.2 Soils of Malaita 29 Table 3.3 Total no. of transects and quadrats carried out and in each 34 habitat Table 4.1 Summary of nocturnal results 39 Table 4.2 Summary of diurnal results 50 Table 4.3 A comparison of species behaviour and habitat preferences of 55 the 21 herpetofaunal species encountered during sampling Table 4.4 A selection of global tropical herpetofaunal studies similar to 58 the current study Table 4.5 A comparison of the recorded richness of frogs, geckos and 60 skinks of the 6 major islands of the Solomon Islands archipelago Table 4.6 Species lists according to McCoy (2006) and Pikacha et al. 61 (2008) vs species actually observed in this study Table 5.1 The 8 Major categories of forests found in the Solomon Islands 65 Table 5.2 Descriptions based on personal observations of the five habitat 68 types used in this research study Table 5.3 The dominant species of plants from the four floral groups 69 found in unlogged coastal forests Table 5.4 The dominant species of plants from the four floral groups 71 found in unlogged lowland forests Table 5.5 The dominant species of plants from the four floral groups 72 found in unlogged upland forests Table 5.6 The dominant species of plants from the four floral groups 74 found in logged lowland forests viii

Table 5.7 The dominant species of plants from the four floral groups 75 found in teak plantation forests Table 5.8 Difference in average encounter rates and species presence in 82 logged lowland forest compared with unlogged lowland forests Table 5.9 Difference in average encounter rates and species presence in 83 teak plantation forest compared with unlogged lowland forests Table 6.1 Vernacular and likely scientific nomenclature of frogs based on 90 questionnaire surveys Table 6.2 Summarised associated uses of different frog species as 95 described by informants Table 6.3 Vernacular and likely scientific nomenclature of lizards based 97 on questionnaire surveys Table 6.4 Summarised associated uses of different lizard species as 100 described by informants Table 6.5 Coastal forest uses, changes and perceived impact on 102 herpetofauna Table 6.6 Lowland forest uses, changes and perceived impact on 103 herpetofauna Table 6.7 Upland forest uses, changes and perceived impact on 104 herpetofauna Table 6.8 Logged forest uses, changes and perceived impact on 105 herpetofauna Table 6.9 Plantation forest uses, changes and perceived impact on 106 herpetofauna Table 6.10 Forest threat values calculated from uses described by 107 informants Table 7.1 Summary of four categories for conservation prioritisation used 113 in this study with descriptions Table 7.2 Species richness and abundance values 115 Table 7.3 Important species values 116 Table 7.4 Cultural values 117 Table 7.5 Forest threat values 117 Table 7.6 Combined ranked values 118

List of Figures Figure 1.1 Examples of (a) a frog and (b) a lizard 4 Figure 3.1 Map of the Solomon Islands archipelago 24 Figure 3.2 The island of Malaita and surrounding islands. 25 Figure 3.3 a) 13 Lingual groupings of Malaita. b) 30 Political wards of 26 Malaita Figure 3.4 Malaita Island with central peaks and rivers 27 Figure 3.5 The Solomon Islands archipelago in relation to the Ontong Java 29 Plateau and Greater Bukida Island Figure 3.6 The age and gender demographics of the Tai ward 30 Figure 3.7 Transect distance-species curve constructed using data from 32 pilot study Figure 3.8 Quadrat area-species curve constructed using data from pilot 33 study Figure 4.1 Batrachylodes vertebralis nocturnal (transect) mean encounter 41 rate for each habitat type Figure 4.2 Bufo marinus nocturnal (transect) mean encounter rate for each 42 habitat type Figure 4.3 Ceratobatrachus guentheri nocturnal (transect) mean encounter 43 rate for each habitat type Figure 4.4 Discodeles guppyi nocturnal (transect) mean encounter rate for 44 each habitat type Figure 4.5 Platymantis guppyi nocturnal (transect) mean encounter rate for 45 each habitat type Figure 4.6 Platymantis solomonis nocturnal (transect) mean encounter rate 45 for each habitat type Figure 4.7 Platymantis weberi nocturnal (transect) mean encounter rate for 46 each habitat type Figure 4.8 Cyrtodactylus salomonensis nocturnal (transect) mean 48 encounter rate for each habitat type Figure 4.9 Nactus multicarinatus nocturnal (transect) mean encounter rate 49 for each habitat type Figure 4.10 Emoia cyanogaster diurnal (quadrat) mean encounter rate for 51 x

each habitat type Figure 4.11 Emoia pseudocyanura diurnal (quadrat) mean encounter rate 52 for each habitat type Figure 4.12 Sphenomorphus concinnatus diurnal (quadrat) mean encounter 54 rate for each habitat type Figure 5.1 Encounter rates of herpetofaunal species found in coastal forest 70 Figure 5.2 Encounter rates of herpetofaunal species found in lowland 71 forest Figure 5.3 Encounter rates of herpetofaunal species found in upland forest 73 Figure 5.4 Encounter rates of herpetofaunal species found in logged forest 74 Figure 5.5 Encounter rates of herpetofaunal species found in teak forest 76 Figure 5.6 Comparison of average herpetofauna species richness in the 77 different habitat types based on nocturnal surveys (transects) Figure 5.7 Comparison of average herpetofauna species richness in the 78 different habitat types based on diurnal surveys (quadrats) Figure 5.8 Comparison of total combined nocturnal and diurnal 79 herpetofaunal species richness Figure 5.9 Average abundances per transect/quadrat 80 (nocturnal/transects=blue and diurnal/quadrats=red) Figure 5.10 A comparison of total herpetofauna species richness in 81 unlogged lowland, logged lowland and teak plantation forests Figure 6.1 Graph of informant’s age and gender against average number of 108 frogs and lizards described Figure 7.1 Graphic representation of priority habitat types based on Table 118 7.6

CHAPTER 1: INTRODUCTION

1.1 Introduction During the Earth Summit in Rio in 1992, the United Nations Convention on Biological Diversity (CBD) (UN 1992a) was ratified by 168 nations, including the Solomon Islands and several other Pacific countries (CBD 2012). Sections a) and b) of Article 8 in the CDB (UN 1992a) state that contracting parties shall a) “establish a system of protected areas… to conserve biological diversity” and b) “develop guidelines… for the selection, establishment and management of protected areas”. The Solomon Islands signed and ratified the CBD in 1995 but is yet to establish a recognised protected areas system. As signatories to this global agreement, there is an urgent need to establish protected areas in the Solomon Islands to conserve its unique biological diversity.

Biological diversity refers to the diversity of life, which ranges in scale from molecules to ecosystems, encompassing genes, species and taxa, populations and communities (UN 1992b, Margules et al. 2002, Spray and McGlothlin 2003). Also included are the interactions and ecosystem processes within and between these entities such as nutrient and energy cycling, predation, competition, mutation, adaptation and migration (UN 1992b, Margules et al. 2002, Spray and McGlothlin 2003). Thaman (pers. comm.) also stresses that this diversity includes human diversity and “ethnobiodiversity”, which is defined as “the knowledge, uses, beliefs, management systems, language and taxonomy that a given human society or group has for their biodiversity”.

Tropical forests are not only important as the richest habitat for terrestrial biological diversity but also represent natural capital or renewable wealth for the people of the Pacific (Montagnini and Jordan 2005, Pauku 2009). Forests have provided and continue to provide many goods and ecosystem services, including goods such as, timber, fuel, medicine, insecticides, rubber, resins, ornamental plants, oils, waxes, tannin, canes, bamboos, fibre, fruit, spices and honey. Ecosystem services provided by forests including shade, shelter, habitat for diverse biota,

1 watershed preservation, erosion control, soil fertility, nutrient cycling, climate regulation, pollution reduction and carbon sequestration, as well as providing the basis for activities such as ecotourism (Khan 2001, Montagnini and Jordan 2005, Pauku 2009).

Dinerstein and Wikramanayake (1993) and Fa et al. (2004) both identified the Solomon Islands forest eco-region as a global biological diversity hotspot due to its high species richness and endemism. However, large-scale, unregulated and illegal logging operations have seriously threatened forest biodiversity, resulting in an ecological and cultural disaster reducing the subsistence ability of the people and their standard of living (Crocombe 2001, McCoy 2006, PHCG 2008, Pikacha 2008). The rate of this logging harvest is unsustainable and environmentally degrading, with predictions that commercially viable forest stocks of the Solomon Islands will be exhausted by 2015 (PHCG 2008).

In a perfect world, all biodiversity should be conserved. There are, however, many competing demands on natural resources as well as limited financial, technical, physical, institutional and human resources available for conservation. Therefore to achieve success, efforts need to be focused, prioritized and strategic (Singh et al. 2000, Spector 2002, Allison 2003, Lindenmayer et al. 2007, Wilson et al. 2009).

1.2 Rationale and Justification for Study The Secretariat of the Pacific Community (SPC) identified eleven priority research and development themes for Pacific Island forests that included; germplasm, food security, reforestation, climate change, traditional knowledge, environmental services, invasive species, forest market products, community agroforestry, endangered species and sustainable forest management (SPC 2009). The current study will relate primarily to the themes highlighted in bold and the following information will explain and highlight the need and rationale for this biodiversity research on the island of Malaita, in the Solomon Islands.

In the process of prioritising areas for conservation, there is a need for quality baseline data on biodiversity as a basis for informed decision making (Gascon et al. 2004). In this context, biodiversity assessments are required before selecting areas for protection to increase the chances of successful conservation interventions 2

(Lindenmayer and Franklin 2002, Allison 2003). Information on patterns of diversity, distribution, endemism, rarity and endangerment provide important information to help in the formation of conservation priorities and plans (Allison 2003, Gascon et al. 2004). In addition, due to the complexity of biodiversity, surrogates are required, and these can be subsets of species, species assemblages or habitat types such as “vertebrates” or “vegetation” (Margules and Pressey 2000, Margules et al. 2002, Allison 2003).

Unfortunately, to date such baseline biodiversity information is not available for much of the Solomon Islands, there are only rough estimates concerning the diversity and richness of most taxa (Morrison et al. 2007). Of particular importance is information on species richness, species interactions and ecological process and patterns (Purvis and Hector 2000).

On the island of Malaita, there are currently no officially recognised protected areas and little biological research has been carried out. The work that has been undertaken is mostly in the form of species inventories (McCoy 2006, Pikacha et al. 2008). Filardi et al. (2007) proposed the “Central Malaitan Highlands” and “Maramasike- Are’Are of Malaita” as Birdlife International, Important Bird Areas (IBAs), however these areas still have no official protection. Thus the current project will be the first of its type on the island of Malaita and will in addition to biological surveys also try to analyse relationships between forest areas, species richness and inter-related cultural values.

Within this context the use of frogs and lizards (herpetofauna) as surrogates is seen as having great potential for conservation prioritisation (Lewandowski et al. 2010). This is due to their susceptibility and fragility, particularly in the case of amphibians, in the face of habitat modification (Pough et al. 1998, Wells 2007). Additionally, these faunal groups are abundant in forests and are generally easy to identify (Pough et al. 1998, Wells 2007). The Solomon Islands is home to 86 currently described species of reptiles (McCoy 2006) and 21 species of frogs (Pikacha et al. 2008). These numbers are, however, incomplete and new species are being found and made known to science through both natural and genetic discovery (Brown 2012).

Globally, many studies looking at herpetofaunal diversity and habitat types have selected and compared forest fragments with continuous forests (Bell and Donnelly 2006, Hillers et al. 2008) and compared fragments among themselves (Bickford et al. 2010). Other studies compared different secondary, primary and plantation forest habitats (Ernst et al. 2006, Gardner et al. 2007, Herrera-Montes and Brokaw 2010) or have compared herpetofaunal diversity between disturbed and undisturbed sites (Pineda and Halffter 2004, Garner et al. 2008). These studies have helped lay the foundation for this research study, as similar methods will be used.

To date most of the data and studies on habitat modification, fragmentation and herpetofauna in the tropics are currently based on Amazonian experiments (Bell and Donnelly 2006). However a need for similar comparative research in the Pacific Islands is recognized (Kingsford et al. 2009), to address the lack of Pacific Island case studies, because of the seriousness of threats to biodiversity in the region.

1.3 Objectives/Aims and Hypotheses The overall aim of this study is to identify priority forest habitats for conservation on the island of Malaita using a combination of biological and ethnological data. It will use selected herpetofauna groups: frogs, lizards (Figure 1.1a-b) – as surrogates for overall habitat health and conservation value.

a b

Figure 1.1a-b Examples of (a) a frog (Ceratobatrachus guentheri) and (b) a lizard (Corucia zebrata) © Edgar Pollard

The specific objectives of this current research study are therefore as follows;

1. To survey different forest habitats on Malaita to determine the abundance, richness and local conservation status of native frogs and lizards (herpetofauna). 2. To define relationships between herpetofaunal incidence, forest habitat type and degree of habitat degradation. 3. To carry out community-based ethnobiological surveys to examine local perceptions, knowledge and cultural uses of herpetofauna and including perceptions of the conservation status of forests and associated herpetofauna. 4. To identify potential priority forest habitats and strategies for forest biodiversity conservation based on the results of objectives 1, 2 and 3.

1.4 Structure and Outline of Thesis The structure of this thesis follows the objectives described above and is divided into eight chapters. Chapter 1 describes the context of the research problem including a brief background, objectives of the study and the research rationale. Chapter 2 focuses on reviewing. Chapter 3 focuses on the general research methodology including a description of the study area, field techniques and a summary of the pilot study. Chapter 4 presents the results of the herpetofauna field surveys on Malaita and addresses the first objective – determining the abundance, richness and conservation status of herpetofauna. Chapter 5 addresses the second objective – the relationships between herpetofaunal incidence, forest habitat type and degree of habitat degradation. Chapter 6 focuses on the third objective – obtaining the local perceptions, knowledge and cultural uses of herpetofauna plus community perceptions of the conservation status of forests and associated herpetofauna. The seventh chapter addresses the final objective and identifies and discusses the conservation priority forest habitats on Malaita. The final chapter discusses the implications of the overall results of this research study for forest conservation on Malaita and provides recommendations for future research and resource management.

CHAPTER 2: BACKGROUND

2.1 Tropical Biodiversity

2.1.1 Importance of tropical biodiversity Biodiversity is the ‘biological wealth’ of the Planet; it provides many beneficial goods and services which can be grouped into four categories (provisioning, cultural, regulating and supporting) (Table 2.1). These services are essential to human livelihoods and therefore link people with the environment (Khan 2001, Spray and McGlothlin 2003, Pauku and Lapo 2009, Kareiva and Marvier 2011). Provision of these beneficial goods and services on our planet is dependent on the overall health of this biodiversity, which is formally defined as the diversity of genes, species, populations and ecosystems (UN 1992b). This biodiversity also affects a community’s ability to recover after disturbances, environmental change and will be especially important for adaptation for survival during long-term global climate change (Kareiva and Marvier 2011).

Table 2.1 Categories of goods and services provided by biodiversity, collated from Khan (2001), Spray and McGlothlin (2003), Pauku and Lapo (2009) and Kareiva and Marvier (2011)

Categories Goods and Services Provided

Provisioning Food, water, fuel, medicines, materials, shelter and shade

Cultural Aesthetic, spiritual, recreational and educational services Climate and weather, flood and disease/pest regulation, Regulating erosion control and the filtration and purification of water and wastes Nutrient cycling, soil formation, oxygen production, Supporting carbon sequestration, primary productivity and the maintenance of gases and ecosystem function

2.2 Tropical Forest Ecosystems

2.2.1 Status and importance of tropical forests Forests are defined as “land spanning more than 0.5 hectares with trees higher than 5 meters and a canopy cover of more than 10 percent” (FAO 2010). Tropical forests fall between the 30° north and south latitudes and are characterised by warm humid conditions year round (Moran 2006). These tropical forests cover around 6- 8% of the earth’s surface but are believed to hold over 50% of the earth’s biodiversity (Moran 2006) with somewhere between 10-50 million species (Dauvergne 2001). Of the 25 global biodiversity hotspots identified by Myers et al. (2000), 17 contain tropical forests which clearly indicates the importance that this habitat type plays in global biodiversity.

Tropical forests as components of biodiversity provide four main functions; (1) productive (timber, fibre, fuel wood and non-timber products), (2) environmental (climate regulation, carbon sequestration and storage, biodiversity reserve and soil and water conservation), (3) social (recreation and subsistence for local populations and cultures) and (4) aesthetic, scientific and spiritual values (Bennet 2000, Montagnini and Jordan 2005, Lindenmayer 2009, Pauku 2009).

Most of the world’s tropical forests are located in South America, Asia and Africa, the Oceania region holds a very small proportion, however this portion is very unique in its diversity and isolation that has led to a very high level of narrow distribution-ranged, endemic species (Smith et al. 2007, Woinarski 2010). Tropical forests with their trees and genetic resources are also recognised as the base of cultural, economic and ecologically sustainable development within the Pacific Islands (SPC 2009). For example, the forestry sector in the Solomon Islands employs around 3% of the labour force which earns roughly US$ 57 million per year, which is approximately 17% of the country’s GDP (FAO 2010). The forests of the Solomon’s are also estimated to hold around 182 million tonnes of carbon stock in the living forest biomass at an average of 82 tonnes per hectare, this is significant for global carbon cycles and storage (FAO 2011).

2.3 Herpetofauna

2.3.1 Status and Importance of herpetofauna The word herpetology is based on the Greek word herpes, meaning creepy thing (Pough et al. 1998). Herpetofauna (amphibians and reptiles) are ectotherms, they also share common lineage and have therefore been placed under the study of herpetology (Pough et al. 1998). There are currently 7104 described species of amphibians and 9766 described species of reptiles globally (AmphibiaWeb 2013, Uetz 2013). Of these the number listed under some form of threat by the IUCN Red- List is 3324 with 635 critically endangered (IUCN 2012) However, still 2196 species remain under the data deficient category. Herpetofauna are found on all major land masses of the world (including most oceanic islands) except for the continent of Antarctica and the island of Greenland (Uetz 2013). The majority of herpetofaunal species are forest dwellers as forests provide a rich array of microhabitats (Heyer et al. 1994, Khan 2001).

Amphibians and reptiles both play an important role in the energy flow and nutrient cycling of ecosystems (Pough et al. 1998). As ectotherms they require little energy for body maintenance and therefore act as reserves of energy (Cloudsley- Thompson 1999). Also due to their ectothermy, the proportion of energy consumed that is used to generate new tissue is high at close to 50% (Heyer et al. 1994). This is around 25 times greater than birds and mammals, indicating the importance that herpetofauna play in overall forest biomass (Pough et al. 1998).

Since amphibians and reptiles play important roles in ecosystems it is important to understand the impacts of land-use practices on these animals (Bell and Donnelly 2006). One clear example of this linkage on islands is the plant-lizard interactions that have co-evolved to produce unique interactions such as the mutualistic flower- visiting and fruit-consuming species of lizard (Olesen 2003).

Additionally the skin permeability in amphibians is an evolved adaption that enables gas and water exchange through the skin, this however adds to the sensitivity of amphibians to environmental changes especially in the water and air (Pough et al. 1998, Wells 2007).

2.3.2 Indicators of ecosystem health, the use of herpetofauna. We cannot survey everything everywhere. To address questions regarding the health and integrity of ecological landscapes a particular species or taxa is selected to act as a surrogate for the whole ecological community and the ecosystem, these are called ecological indicators (Hilty and Merenlender 2000). Such indicators have parameters, such as density, absence/presence, and infant survivorship that can be used to indicate ecosystem conditions (Hilty and Merenlender 2000).

To help select suitable indicators for this study the following sampling considerations as outlined by Feinsinger (2001) were used.

i. Objective sampling – the indicator should be able to be effectively and objectively sampled through direct observation with limited biases. ii. Efficient sampling – the indicator should be able to be efficiently sampled producing good data quickly without too much need for setup. iii. Sample size – the indicator should be able to provide a large number of replications. iv. Sampling expense – the indicator should involve minimised costs in equipment and procedure. v. Familiarity – the natural history and taxonomy of the indicator should be well known. vi. Scale – the scale at which the indicator operates should be the same as the scale of the ecological conservation concern, e.g. species and habitats. vii. Sensitivity – the indicator should be sensitive to factors related to the ecological conservation concern. viii. Aptness as a surrogate – the indicator should respond consistently to environmental change over time. ix. Consistency – the indicator should be equally accessible and active at all times when sampling occurs. x. General interest – the indicator should also respond to factors that concern local communities.

Herpetofauna, the taxa involved in this study meet all the above categories relatively well. In addition, herpetofauna (especially amphibians) are also often cited

9 as an ideal indicator group for ecological studies, because of their sensitivity to changes in moisture and temperature regimes, two-part life cycle, diversity in reproductive methods and weak dispersal abilities (Pineda and Halffter 2004, Smith and Rissler 2010). Amphibians are abundant and functionally important in many ecosystems around the world and most are easily identified and are of global conservation concern because of their well-documented, widespread decline (Bennett 1999, Stuart et al. 2004, Smith and Rissler 2010).

To further support the use of herpetofauna, a review of surrogate studies by Lewandowski et al. (2010) showed herpetofauna to be the most effective surrogate taxa, in comparison to arthropods, birds, fungi, mammals, plants, molluscs and all vertebrates. An example is the leaf litter frogs of West Africa that showed strong negative response to minor degradation of their habitat (Hillers et al. 2008). Estrada et al. (2010) also found that no taxon is a good “umbrella group” (representation group for other taxa) but that reptiles were the most appropriate as their results match most closely with other vertebrate taxa. Hilty & Merenlender (2000) further suggested that multiple indicator taxa be used as single taxa cannot accurately reflect system health. Allison (2003) also found that species richness patterns for different taxonomic groups show little overlap, emphasizing the importance of surveying all taxa if possible. Therefore, biological indicators are not the answer to everything but are a useful way of surveying biodiversity, and the combination of amphibians and reptiles together is among the most useful (Lewandowski et al. 2010).

2.4 Threats and Decline of Biodiversity We are in the midst of our planet’s sixth mass extinction event (Gascon et al. 2004, Kingsford et al. 2009), biodiversity loss is 1000 to 10000 times the expected background extinction rate (Khan 2001). This sixth event is considered a crisis because for the first time such a mass extinction is anthropogenicly driven (Brodie et al. 2013). A high rate of species extinctions can change the dynamics of ecosystems by altering: energy flows, the composition and structure of plant and animal communities, behaviour in organisms and cause an overall disruption of ecological and environmental processes (Kareiva and Marvier 2011, Tuomainen and Candolin 2011). The loss of even a few important “keystone species” can cause a trophic cascade and the structural collapse of entire ecosystems (Hairston et al. 1960). 10

Direct causes of this huge global biodiversity loss include destruction of natural habitats via activities such as agriculture, deforestation, mining, urbanization, over- fishing, intensive agriculture, invasive species, environmental and industrial pollution (Khan 2001, Spray and McGlothlin 2003, Kareiva and Marvier 2011). Plus a lack of regulation with poor government policies that are inconsistent and disregard the value of biodiversity (Khan 2001, Spray and McGlothlin 2003, Kareiva and Marvier 2011). Less direct or cryptic underlying causes of biodiversity loss also include: international trade, globalisation, shifting cultural attitudes, lack of knowledge of sustainable resource use, a lack of economic valuing for biodiversity, the use of inappropriate technology and increased economic growth (Khan 2001, Spray and McGlothlin 2003, Kareiva and Marvier 2011). Most of these latter causes are creating an increasing and unsustainable demand for natural resources and energy (Wilson and Peter 1988).

Globally, exponential population growth has increased the pressure placed on natural biological resources, poverty leads to encroachment on marginal lands and protected areas and the unsustainable harvesting of resources such as mangrove wetlands (Kareiva and Marvier 2011, Brodie et al. 2013). In addition, the introduction of exotic species (which in many cases has led to the extinction of native species) and an increasing discovery for uses of biodiversity has put further pressure on previously non-targeted organisms in both the terrestrial and marine environments (Wilson and Peter 1988, Khan 2001, Spray and McGlothlin 2003).

Tropical ecosystems are also threatened by human-induced changes in biogeochemical cycles of carbon, nitrogen, phosphorus and also global climate change with its associated increases in temperatures, sea-level rise and altered weather patterns (Gascon et al. 2004, Pauku 2009, Becker et al. 2010). Kingsford et al. (2009) described the six major causes of biodiversity decline in the Oceania region as habitat loss and degradation, invasive species and disease, climate change, overexploitation and pollution all of which are further exacerbated by a lack of political capacity. As in many other countries of the region, the biodiversity of the Solomon islands is threatened, mainly by intensive logging, inappropriate land use practises in agriculture and mining and over-exploitation of natural resources, all of which are exacerbated by natural disasters, climate change, pollution, invasive 11 species and population increase (PHCG 2008, Pauku and Lapo 2009). Highly threatened ecosystems in the Solomon’s include mangrove forests, wetlands and coastal forests as these are habitats that interact more frequently with people and have highly sought after resources (Pauku 2009).

2.4.1 Specific threats to tropical forests Tropical forests are one of the most essential ecosystems on the planet but are also considered one of the most threatened (Pineda and Halffter 2004, Hillers et al. 2008). Tropical forests are particularly vulnerable, firstly because they keep most of their nutrients in living organisms and therefore rely on the work of decomposers to recycle nutrients (Moran 2006). Secondly tropical forest flora and fauna tend to have smaller populations and ranges and are therefore more susceptible to environmental changes (Moran 2006, Woinarski 2010). Thirdly tropical forests play an important role in weather and climatic processes especially through the action of evapo- transpiration, so if more forests are lost then rainfall will diminish in many areas (Schwartzman et al. 2000, Moran 2006, FAO 2011). Degradation of tropical forests therefore threatens the existence of many birds, reptiles and mammals especially significant keystone species that play vital roles in ecosystems, such as dispersal of forest seeds (Pineda and Halffter 2004, Pauku 2009). Forest degradation will also impact on the resilience ability of forests to recover from disturbances and degradation (Pauku 2009, Woinarski 2010).

A major process that degrades forests is deforestation otherwise known as logging (Lindenmayer 2009). Deforestation is the “removal of forest and the subsequent conversion of land to other uses” (Moran 2006). Logging activity has many consequences (Table 2.2) and usually results in an extremely fragmented forest landscape especially through the construction of logging roads deep into natural forest areas (Moran 2006, Dutson 2011). Forest fragmentation is when forests are cleared in an unsystematic, unplanned way and this leads to a totally changed forest community structure which leads to the eventual loss of certain species and the introduction of invasive species (Hill and Curran 2001, Moran 2006, Filgueiras et al. 2011, Brodie et al. 2013). Fragmentation also creates edge habitats in forests and these areas are more exposed than natural forests and are thus unsuitable habitats for many native species (Hill and Curran 2001, Pineda and Halffter 2004, Moran 2006). 12

Table 2.2 Consequences of deforestation (Dauvergne 2001, Khan 2001, Gascon et al. 2004, Morrison et al. 2007, FAO 2011)

Consequences of Deforestation loss of habitat for wildlife On biota species extinctions

On ecosystems reduced ecosystem productivity

On the release of carbon dioxide into the atmosphere atmosphere

loss of topsoil and a decline in soil fertility decreased microbial activity On the soil increased landslides severe wind and water erosion

siltation of waterways and reefs disruption to local hydrological cycles lower stream flow, On the water lowered water table, lower water quality more widespread and frequent flooding

Timber utilization or logging is the most lucrative and common form for usage of forest resources in the world and is also recognized as one of the main threats to vertebrate diversity globally (Moran 2006). The forestry industry in the Solomon Islands is a major player in the export revenue sector bringing in around 70% of total export revenue in 2008 (MoFR 2009). Of the 598,000 hectares of harvestable forest in the Solomon’s 288,000 hectares has already been logged with remaining stocks estimated to be depleted by 2015 (PHCG 2008). Annual estimated sustainable yield from natural forests since 1994 is displayed in Table 2.3. In relation to actual yield, it clearly shows volumes almost five times the sustainable level.

Table 2.3 Solomon Islands logging yield. Adapted from Dauvergne (2001), PHCG (2008) and CBSI (2010)

Year Estimated sustainable yield Actual yield (m³) (m³) (Dauvergne 2001 and (Dauvergne 2001 and PHCG 2008) CBSI 2010) 1994 276 000 826 000 1998 223 000 650 000 2007 320 000 1 444 003 2008 320 000 1 523 000 2009 320 000 1 064 445 2010 320 000 1 428 211

An indirect effect of deforestation is the expansion and creation of degraded forests, secondary forests and exotic species plantation forests referred to as monocultures (Gardner et al. 2007, Herrera-Montes and Brokaw 2010). Plantation forests are forests predominantly composed of trees established through planting and/or deliberate seeding and may be composed of native or introduced species which are established usually for timber production (FAO 2010). A similar threat is the extensive conversion of lowland forests into oil-palm plantations as seen on the islands of Guadalcanal, New Ireland and proposed also on Malaita (Dutson 2011).

Contributing factors that also threaten the nature of tropical forests in Oceania (similar to causes threatening biodiversity as a whole described earlier) include increasing population numbers, poverty, commercial exploitation, corrupt governance, breakdown of cultural values, pressure to get cash from resources and a lack of economic incentives to conserve biodiversity (Moran 2006, Pauku 2009, Woinarski 2010). The State of the Environment report for the Solomon’s (PHCG 2008) stated that the forests and soils of the Solomon’s are “running out”, they are losing the ability to sustain people and also to sustain themselves.

Overall, the future for the “tropical forests of Oceania is bleak” mainly due to a host of factors including direct exploitation and modification of natural ecosystems (Woinarski 2010). The Oceania region (including Australia and New Zealand) holds 14 an estimated 191 million hectares of forested areas and this has decreased slowly over the past 20 years (FAO 2011). The Solomon Islands holds therefore some of the last remaining untouched forests of the tropical world but the islands are under increasing threat to large scale degradation and habitat loss at alarming rates (PHCG 2008, Pikacha 2008). This has great effects on the local biodiversity as complex ecosystems are broken and the biota that depends on these forests begin to disappear (Pikacha 2008).

2.4.2 Specific threats to tropical herpetofauna Of all vertebrates, the amphibians have the highest proportion of species threatened with extinction and are facing a significant global decline (Blaustein and Kiesecker 2002, Stuart et al. 2004, Cushman 2006, Gardner et al. 2007, Garner et al. 2008, Bombi 2009). Amphibians are particularly sensitive to habitat degradation and fragmentation and these factors are viewed as major contributors to the global amphibian decline (Pineda and Halffter 2004, Ernst et al. 2006, Hillers et al. 2008). Amphibians are exceptionally vulnerable to habitat degradation compared to other terrestrial vertebrates because of their relatively low tolerance to environmental extremes and pollution, high susceptibility to pathogens, specific breeding-habitat requirements, and competition and predation from invasive species (Pough et al. 1998, Cushman 2006, Bickford et al. 2010). Reptiles are also facing a similar fate but is not as well documented (Bombi 2009).

Habitat degradation is the biggest threat to herpetofauna especially in the tropics where more than 80% of all amphibians and reptiles are found (Pough et al. 1998). The opening of the tree canopy through selective logging results in microclimate alterations which place constraints on certain frog species (Hillers et al. 2008). Also the degradation of forests creates changes in canopy structure, leaf-litter environment and loss of microhabitats, all necessary for healthy herpetofaunal populations (Gardner et al. 2007).

Invasive alien species also pose a great threat to native amphibians as they modify habitats, affect reproductive success and directly impact amphibian species through predation and competition (Christy et al. 2007, Martin and Murray 2011).

Illegal exporting of herpetofauna by collectors supplying the North American and European pet trades are also threatening the long-term survival of the many more charismatic species in the Solomon’s such as the lizards Corucia zebrata, Varinus indicus and the Candoia snakes (McCoy 2006). The trade in reptiles and amphibians caught in the wild is mostly unregulated with only a limited number of species being monitored on CITES (Schlaepfer et al. 2005). However, even the figures show that there is a significant amount of species being traded (Schlaepfer et al. 2005) with the potential to contribute to global herpetofaunal declines.

Global warming is also believed to be contributing to the decline in amphibians and lizards (Wake 2007). Climate change may also have severe impacts on amphibians, as temperature and moisture, two important variables that define their distribution are also two components that will be directly impacted by forecasted global climate change (Wells 2007). Such changes will in particularly impact those restricted range species that are adapted to cooler, wetter conditions on mountain tops and ridges (Pikacha et al. 2008).

In addition, the perceptions of humans towards some species are also a potential threat for herpetofauna (Pough et al. 1998). In New Caledonia for example children are warned not to kill lizards as they may be killing their own ancestors, whereas in some parts of Iran lizards are killed because they are believed to carry the devil’s soul (Pough et al. 1998).

2.5 Conservation of Biodiversity

2.5.1 What is conservation? Understanding that our biodiversity is threatened creates a need for conservation actions. A goal of biodiversity conservation is to maintain variety of life, all that is known and unknown, measured and unmeasured, the variety of life on earth (Margules et al. 2002). The three main objectives of the Convention on Biological Diversity (CBD) (UN 1992a) are: (1) the conservation of biological diversity, (2) the sustainable use of its components and the fair and (3) equitable sharing of the benefits arising out of the utilization of genetic resources.

“Protection of biodiversity” as a goal is too general and naive and needs to be placed in context, focused and specific (Kati et al. 2004). Conservation targets can take the form of habitats, communities, species, ecological processes and services but a conservation plan should be focused on a “subset”. A “subset” can be a single species, a subset of species or by focusing on the threat status of a species (Kati et al. 2004). As discussed by Kati et al. (2004) different types of species used in conservation are: keystone (linked to many other species), umbrella (covers other species), flagship (charismatic or culturally important species), indicator (reflects the health of the environment or the effectiveness of conservation interventions) and focal species (those sensitive to dominant threats). However, Kareiva and Marvier (2011) found that recommendations from studies based on single particular taxa generally failed to provide protection for other taxa. It is therefore important to consider the conservation of functional traits (eg. pollinating insects or carnivorous birds) and diversity among forest species may serve community populations better, in the form of ecosystem functioning and community robustness, than just the focusing of conservation efforts on specific species (Ernst et al. 2006).

Three global strategies (that are not mutually exclusive) for conservation have to date been utilized. One is the “hotspot” approach favoured by Conservation International (CI) that focuses on areas with the most “threatened and distinctive” biota (Olson and Dinerstein 1998, Myers et al. 2000). Myers et al. (2000) has defined 25 global hotspots based on species endemism and degree of threat, these hotspots are thought to contain 44% and 35% of all plant and vertebrate diversity respectively. Biodiversity hotspots are areas with a large number of species or large number of threatened, rare and/or endemic species (Kati et al. 2004) and the conservation of these hotspots is described as a ‘silver bullet’ strategy in cost- effectiveness for biodiversity conservation. The second is the representative ecoregion approach that focuses on conserving sites in major ecosystems and habitat types (Gascon et al. 2004). Thirdly, Kati et al. (2004) identified the complementary network (where conservation areas complement one another through-out a network of protected areas) and richness hotspot approaches combined as the best possible approaches for conserving the entire biological diversity of an area.

2.5.2 How do we conserve biological diversity

2.5.2.1 Conservation priorities An important challenge facing tropical biodiversity conservation is determining methods for prioritisation and then to implement effective conservation in these identified priority areas (Becker et al. 2010). Priority setting in conservation identifies “where, how, on what, and when” we should act first, knowing well that we cannot do everything, everywhere at once (Wilson et al. 2009). The purpose of priority setting is to limit and minimize the current loss of biodiversity and to “effectively and efficiently” achieve at least some preservation of biodiversity within the limited resources and funds available (Margules et al. 2002, Wilson et al. 2009).

Prioritisation of biodiversity conservation is dependent on many criteria (Table 2.4) (Lewandowski et al. 2010). The current research study will try to incorporate as many of these criteria as possible in its decision-making.

Table 2.4 Criteria used in the prioritisation of biodiversity conservation collated from Ratcliffe (1977), Sanderson et al. (2002), Fa et al. (2004) and Hill et al. (2005)

Species Level Criteria Site Level Criteria

species abundance, richness and size diversity naturalness and the intrinsic rarity and endemicity appeal of an area evolutionary uniqueness and representativeness or typicalness diversity of phenotypic traits fragility of a site protection status land use change and human role in the ecosystem and species influence involved in multi-species sites with recorded histories, interactions (e.g. keystone potential monetary value, species) importance in geographical or ecological processes

According to Wilson et al. (2009), in conservation prioritisation certain variables need to be identified, we need to know; what the assets are? (e.g. details of biodiversity such as frog species richness) What the threats are? (e.g. logging and invasive species) What possible actions can be taken? (e.g. protected areas) How much those actions will cost? (e.g. human and financial resources) The current research will attempt to capture as many of these influencing variables as possible. However, based on the work of Kareiva and Marvier (2011) an additional question that needs to be answered is: should “limited conservation funds be spent on saving near-extinct species or should it be invested into the prevention of more abundant species becoming rare”? Also based on the work of Brooks et al. (2006) another question is: should “environmental services which are also threatened be incorporated into conservation planning”? (services such as carbon sequestration, climate stabilization, maintenance of water quality, minimization of pest and disease outbreaks).

2.5.2.2 Conservation types Conservation actions can be separated into two types: ex situ and in situ. Ex situ conservation means “the conservation of components of biological diversity outside their natural habitats” (UN 1992a). This helps conserve wild and domesticated biodiversity through “aquaria, botanical gardens, herbaria, seed banks, cloned collections, microbial collections, field gene banks, forest nurseries, tissue and cell cultures, zoological gardens and museums” (Khan 2001).

In situ conservation means “the conservation of ecosystems and natural habitats and the maintenance and recovery of viable populations of species in their natural surroundings and in the case of domesticated or cultivated species, in the surroundings where they have developed distinctive properties” (UN 1992a). This includes the “legal protection of endangered species, preparation or implementation of species habitat recovery or management plans and the establishment of protected areas to conserve either individual species and/or habitats” (Khan 2001). According to Khan (2001) both in situ and ex situ actions will need to go hand in hand to achieve successful conservation of biodiversity.

Protected areas are currently considered one of the most effective methods for in situ species conservation (Brooks et al. 2004, Bombi 2009). The Food and Agriculture Organization (FAO 2010) defined protected areas as “areas dedicated to the protection and maintenance of biological diversity, natural and associated cultural resources and managed through legal or other effective means”. Protected areas are also defined by the CBD as “a geographically defined area which is designated or regulated and managed to achieve specific conservation objectives” (UN 1992a). Protected areas are considered as an effective means of protecting ecosystems and species in the tropics (Bruner et al. 2001). This occurs on land mainly by preventing land clearance, and should continue to be an important part of long-term terrestrial biodiversity conservation (Bruner et al. 2001). Both terrestrial and marine protected areas (MPA)s not only protect the biological diversity but can also help maintain long-term, sustainable industries such as fisheries, timber harvesting and ecotourism (Kareiva and Marvier 2011).

The IUCN and UNEP-WCMC (2011) created a world database of nationally designated and known protected areas of all countries showing that globally nearly 13% of the planet’s area is under some form of protection. Goal 7 of the Millennium Development Goals (MDG)(developed by the United Nations) is to ensure environmental sustainability and with that recommendations for national protected area cover by 2020 (UN 2011). A summary of global, regional and the national figures for recognised protected areas show the Solomon Islands to be highly unlikely to achieve the MDG targets for protected area systems (Table 2.4).

Table 2.5 Percentage Terrestrial and Marine Protected Areas Cover and MDG recommendations collated from IUCN and UNEP-WCMC (2011)

Level Percentage (%) of Percentage (%) of terrestrial area under marine area under some some form of protection form of protection MDG targets by 2020 17 10 Global 12.7 7.2 Oceania region 4.9 2.8 Solomon Islands 0.09 0.12

2.5.3 Conservation and traditional ecological knowledge (TEK)

2.5.3.1 How has TEK been used in conservation Huntington (2000) defines traditional ecological knowledge (TEK) as “the knowledge and insights acquired through extensive observation of an area or species” which is usually shared orally. For thousands of years indigenous peoples have used TEK to survive, build and maintain their unique cultures (Bennet 2000, Thaman et al. 2010, FAO 2011). TEK is the basis for people’s livelihoods and maintains the cultural, economic and traditional practises (Bennet 2000, Thaman et al. 2010, FAO 2011).

Some of the richest areas of biodiversity globally are controlled by local indigenous people (Painemilla et al. 2010), as in the Pacific region where a vast majority of land and natural resources are traditionally owned (Brodie et al. 2013). Indigenous people use customary laws and traditional practices that have kept their rich resources intact (Painemilla et al. 2010). These traditional practises, skills and wisdom have been used by local villagers to help them adapt to change (Lauer and Aswani 2010, Painemilla et al. 2010), and can still offer useful guidelines for many communities in the Pacific for biodiversity protection and conservation management (Huntington 2000, Berkes 2004, Painemilla et al. 2010).

It is known that governments and natural resource managers still have a lot to learn from indigenous communities (Painemilla et al. 2010, Woinarski 2010). Cinner & Aswani (2007) recommend that ‘hybrid institutions’ be formed from the merging of customary management systems and contemporary conservation initiatives. These ‘hybrid institutions’ would use traditional ecological knowledge and also scientific knowledge to conserve and further improve respect for traditions and local acceptance of conservation values (Cinner and Aswani 2007).

2.5.3.2 How have the Pacific people practiced conservation The Pacific people hold traditional beliefs and practised a close relationship with the environment; concepts such as species recovery, conservation and sustainability are not new and may even inadequately define such relationships (Read 2002, Bayliss-Smith et al. 2003). Forests, the focus of this research have a sacredness and ancestral significance that are an integral part of Melanesian culture and the respect 21 and appeasement to the spirits and the forest is of an utmost importance (Bennet 2000). This is a stark contrast to the degradation that Melanesian forests are facing today.

Early traditional conservation methods revolved around restriction of access to resources by time, place or from certain people (Bennet 2000, Crocombe 2001). In the Solomon’s basic customary conservation practices are in the form of: a) sacred sites, that restrict access to certain areas for certain members of the community, b) social prohibitions which is the restriction on certain species by certain groups which could also be limited to certain times of the year and c) sequential prohibitions which rotate areas limiting certain groups to harvesting some resources in the form of temporary closures (Caillaud et al. 2004). However, many of these practices are breaking down and being lost (Thaman 2002).

CHAPTER 3: STUDY LOCATION AND GENERAL METHODOLOGY

3.1 Study location

3.1.1 Solomon Islands The Solomon Islands, the third largest archipelago in the South Pacific, is located between 6-12ºS and 155-168ºE and composed of a double chain of approximately one thousand islands (Figure 3.1) extending over 1450 km in a south-eastern direction (Mueller-Dombois and Fosberg 1998). The political Solomon Islands (made up of 10 provinces; Choiseul, Western, Isabel, Central, Guadalcanal, Malaita, Makira, Temotu, Rennell & Bellona and Honiara City) consist of the Solomon’s archipelago minus Bougainville the largest geological island which is politically part of Papua New Guinea (PNG). As a nation state the Solomon Islands is located 1,800 km north east of Australia and around 5,800 km south west of the Hawaiian Islands. Most of the Solomon’s islands are of a volcanic origin as the archipelago is situated along the “Pacific ring of fire”, in the subduction zone between the Pacific and Indo- Australian plates (PHCG 2008). Many of the larger islands are still very much geologically active (Pikacha et al. 2008, PHCG 2008). Though the total land area of the Solomon’s is around 28,785 km² (Mueller-Dombois and Fosberg 1998) the country has rich marine resources and a total marine area of around 1.3 million km² (Gough et al. 2010).

The Solomon Islands are regarded as one of the wettest places in the tropics with the climate described as tropical maritime (Whitmore 1969). Air temperature (coastal ranges from 25º to 32ºC) is relatively uniform all year round with major climate seasonality due to wind direction and rainfall (Mueller-Dombois and Fosberg 1998). Annual rainfall is in the range of 3000 to 5000 mm at sea level stations but in wind exposed higher altitudes, 7000 mm to 9000 mm can be expected per annum (Mueller-Dombois and Fosberg 1998, Pauku 2009). Due to the proximity of the islands to the equator, solar radiation reaching the islands is high and relatively uniform all year round. Also the effect of the “Inter Tropical Convergence Zone”

(ITCZ) is strongly felt especially during November to February, in the form of “monsoon-like” weather patterns (Ross 1973).

Figure 3.1 Map of the Solomon Islands archipelago including provinces.

Tropical cyclones are a factor periodically affecting the Solomon Islands, with storm events creating wind-fall gaps in the forests (Whitmore 1969, Mueller- Dombois and Fosberg 1998). It is believed that many organisms of these islands have adapted to cyclone disturbed and modified habitats (Ross 1973, Filardi et al. 2007), and that cyclones help maintain species diversity and composition within and between forest types (Burslem 1999).

Most islands are covered in dense tropical forest with the majority of flora of Malesian (southeast Asia) affinity (Mueller-Dombois and Fosberg 1998). The source of biota dispersal is believed to flow through New Guinea from southeast Asia into the west, therefore islands with closest proximity to the mainland source area in the west (Western Is. and Choiseul) have greater diversity and species richness than islands further eastward (Malaita and Makira) (McCoy 2006, Hamilton et al. 2009). Whitmore (1969) believed the vegetation of the Solomon Islands to be an incomplete subset due to incomplete migration of biota from Malesia and possibly also due to the comparatively young volcanic age of some islands. 24

The Solomon Islands has claims of possessing greater terrestrial biodiversity than any other Pacific island nation except PNG (PHCG 2008). The Solomon Islands rainforest eco-region is included in the world’s “Global 200” (a list of ecoregions identified by WWF as priorities for conservation WWF 2012) and categorised as “globally outstanding.” In terms of species richness and uniqueness, the Solomon Islands host more “restricted range” and recorded endemic birds than any area in the world, and also has the largest skink in the world, the largest insect-eating bat and some of the largest native rats (Pauku and Lapo 2009). The overall richness of the Solomon Islands is not only in its natural biodiversity but also in its cultural diversity with over 70 surviving indigenous languages and many more dialects (PHCG 2008).

3.1.2 Malaita

3.1.2.1 Malaita Province The province of Malaita approximately 80 km east of Honiara (Figure 3.1) is made up of the main Malaitan Island and the southern adjacent island, Sa’a, which is separated from the main island by a narrow passage plus other much smaller surrounding islands (Figure 3.2). Malaita has a high population density and is home to roughly a third of the total Solomon Islands population (Table 3.1). Since most Malaitan’s still depend on subsistence agriculture the high population has impacted the natural Figure 3.2 The island of Malaita forest vegetation in many areas in and surrounding islands. addition to the timber industries (Filardi et al. 2007). The province is culturally divided into 13 lingual groupings (Figure 3.3a) and also politically divided into 30 wards (Figure 3.3b).

Figure 3.3a-b a) 13 Lingual groupings of Malaita. b) 30 Political wards of Malaita

3.1.2.2 Malaita Island The island of Malaita lies in a northwest to southeast direction and measures around 190 km in length. The centre of the large main island is located at 9° S and 161° E (Polhemus et al. 2008). Its width ranges from 10-40 km in the widest parts with a total land area of approximately 4200 km² (Moore 2007). It is the third largest and fourth highest island in the Solomon Islands, with a central mountain range that includes a number of peaks reaching over 1000 m (Figure 3.4). The highest peak, Mount Kolovrat (Alasa’a) has an elevation of 1433 m.a.s.l. (Filardi et al. 2007, Polhemus et al. 2008). Dominant landforms include “steep, narrow ridges, fluvial plains, karst mountains, valleys, swamps and coastal landforms” (Moore 2007, PHCG 2008). Geologically this rugged topography is relatively young with much “folding, thrusting and deformation” and many crystal clear fast-flowing streams, which account for the relative absence of long coastal estuaries on Malaita (Petterson et al. 1999, PHCG 2008, Polhemus et al. 2008). Lagoons are also a common feature of the island with the lagoons of the Lau (renowned for its artificial islands), Langa 26

Langa (renowned for its shell money making people) and Are’Are (known for its expansive mangrove forests) constituting some of the most widely known features of Malaita province internationally. There are also extensive coastal swampy areas indicating past existence as lagoons (Moore 2007).

Figure 3.4 Malaita Island with central peaks and rivers

Table 3.1 Comparison of population density among Solomon Island Provinces (adapted from Law 2011 and SINSO 2011) .

Province Total land area Total population Population density (km²) (2009) (SINSO (persons/ km²) 2011) Choiseul 3,837 26,372 6.9 Western 5,475 76,649 14.0 Isabel 4,136 26,158 6.3 Central 615 26,051 42.4 Rennell-Bellona 671 3,041 4.5 Guadalcanal 5,336 93,613 17.5 Malaita 4,225 137,596 32.6 Makira-Ulawa 3,188 40,419 12.7 Temotu 895 21,362 23.9 Honiara 22 64,609 2936.8 Total 28,400 515,870 18.2

As on others oceanic islands, the sea plays a tempering effect on the climate of Malaita island, with daily temperature ranges from 25°C to 32°C and high levels of wetness and humidity. In common with many islands in the Pacific is the diurnal weather pattern of “clear sunny mornings and afternoon showers” (Ross 1973, Pauku 2009) and a “windward-leeward effect”, with the western coast of Malaita receiving an annual average rainfall of 3750 mm whereas the eastern mountains in the direct path of the prevailing south easterly’s receive more than 7500 mm/year (Moore 2007).

Malaita Island is the highest point on the Ontong Java Plateau (OJP) (Figure 3.5) originating from volcanic activity around 125-121 Ma (Miura et al. 2004, Polhemus et al. 2008). This volcanic intrusive core with pelagic sedimentary overlaying gives the geology of Malaita a unique dual volcanic and sedimentary base (Ross 1973, Petterson et al. 1997). The OJP is world’s largest oceanic plateau covering an area of approximately 1600 km x 800 km with an average crustal thickness of 33 km (Miura et al. 2004). According to Petterson et al. (1997) the OJP situated on the Pacific plate collided with the Indo-Australian plate along the Solomon Islands arc subduction zone around 25-20 Ma. Due to the large mass of the OJP a subduction flip occurred whereby the Pacific plate ceased total subduction and the India-Australian plate began subduction under the Pacific Plate (Petterson et al. 1997, Ishikawa et al. 2004). Due to stress and folding of the subsequent crusts Malaita emerged above sea- level at about 5-2 Ma (Petterson et al. 1997, Ishikawa et al. 2004) and this relatively recent emergence has implications in regard to the arrival of biota to the island (Polhemus et al. 2008).

During the Pleistocene glacial episodes (the last being around 12,000 years ago) where sea-levels dropped to around 120 m below present a ‘Greater Bukida’ island (Figure 3.5) was formed joining the islands from Bougainville in the north right down through Choiseul and Isabel down to the Florida group in the central Solomon Islands and possibly including Guadalcanal. The islands of Malaita and Makira were never part of this ‘Bukida’ island indicating that direct contact with a greater species pool was limited and that greater endemism may have occurred on these two islands compared to other islands in the archipelago, due to their relative isolation (Jameson and Ratcliffe 2009). 28

Figure 3.5 The Solomon Islands archipelago in relation to the Ontong Java Plateau and Greater Bukida Island

The soil type of Malaita consists of strongly to slightly moderate weathered leached soils with low base status, organic and decomposed peat (PHCG 2008). Soil type and soil use relationships are important to the people of Malaita, as described by Ross (1973) for the people of northern Malaita (Baegu) who have simplistically identified four soil types: sandy, inland, dry black/brown and red (Table 3.2).

Table 3.2 Soil types of Malaita, adapted from Ross (1973)

Soil Type Comment

“Sandy soil” found on the coasts and useful for coconuts and yams

“Inland soils” of wet black/brown sediments which are too heavy and not well drained and is therefore used only for some Taro

“Dry is well drained and is best used for gardens, supporting rich Black/brown” variety of agricultural crops

“Red soil” does not absorb water well and forms a hard crust and is commonly used as a location for settlements and hamlets due to its firmness

3.1.3 AreÀre study site The AreÀre lingual group in the south of Malaita Island has the largest land area on the island (Figure 3.3a). The Tai ward (Figure 3.3b) found in the AreÀre lingual group area was where the fieldwork for this research was focused. Tai ward was selected because with a relatively low population density, the native vegetation has remained relatively intact until the commencement of logging operations in the area in 2002 which then caused subsequent heavy degradation throughout the region (pers. obs.). There are still however fragments of pristine forest remaining, especially further inland and at higher altitudes that house the most pristine representations of Malaita’s native flora and fauna.

The age and gender demographics of the Tai ward based on the 2009 census (SINSO 2011) are shown in Figure 3.6. The ward shows a young population with 52.9% of the 85+ population below the 80-84 age of 20, which is 75-79 Female 70-74 typical for most of the 65-69 Male 60-64 Solomon’s. There is a 55-59 significant change in 50-54 45-49 population between 40-44

Age class Age 35-39 the ages of 15-24, 30-34 which is probably 25-29 20-24 accountable to 15-19 temporary migration 10-14 5-9 due to education or 0-4 work. For example, in -10 -5 0 5 10 % the Tai ward, there are only four schools that Figure 3.6 The age and gender demographics of cater for students up to the Tai ward based on the 2009 census (SINSO form three level and 2011) students wishing to continue must therefore leave the area. There is also very low numbers of older citizens with only 5.8% of persons over the age of 60. These are usually the persons with greater in-depth traditional knowledge and their low numbers indicate the potential threatened nature of this knowledge. 30

3.2 Pilot study and General Methodology This research study has included: a pilot study reconnaissance survey followed by major field surveys comprised of formal transect and quadrat sampling and questionnaire surveys to gather local indigenous knowledge.

3.2.1 Pilot Study A pilot or reconnaissance study was conducted in the month of August 2011 to principally pre-test and identify any problems with the already designed field methodology as well as determine optimal quantities, locations and exact methodologies for quadrats and transects.

3.2.1.1 Transects Nocturnal Visual Encounter Survey (NVES) transects were carried out in each of four selected forest habitat types: unlogged lowland forest, unlogged upland forest, teak plantation forest and logged forest (during the pilot stage unlogged coastal forest was yet to be included). Two 600 m transects were surveyed in each habitat type, recording cumulative herpetofaunal species abundance and richness at the 300m, 400m, 500m and 600m marks along each transect. The collected combined data for all habitat types show the cumulative mean number of species observed at each distance mark (Figure 3.7). This graph was then used to determine the optimum length for transects. Only 43% (2.00) of species were encountered within the first 300 m. Seventy-three per cent (3.12) of species were encountered within the first 400 m and 94% (4.00) of species were encountered within the first 500 m. Speed along a transect depended on terrain, undercover vegetation thickness and herpetofaunal abundance (as more species led to more time taken to record results). The average speed per 100m was 20min. This led to the decision to select 500 m as the optimum transect length in terms of time efficiency and species encounter rates, so that two transects could be completed each evening.

5 4.5 4 3.5 3 2.5 2

species observed observed species 1.5 1 0.5 Cumulative mean of herpetofaunal of mean herpetofaunal Cumulative 0 300 350 400 450 500 550 600 Distance travelled along transect (m)

Figure 3.7 Distance-species curve constructed using data from pilot study. Data from all habitat types were combined (2 transect replicates x 4 habitat types).

3.2.1.2 Quadrats Diurnal quadrat sampling (DQS) was carried out in each of the four habitat types: unlogged lowland forest, unlogged upland forest, teak plantation forest and logged forest (during the pilot stage unlogged coastal forest was yet to be included). Four different sized quadrats were carried out in each habitat type, recording herpetofaunal species abundance and richness at the 4 m², 6 m², 8 m² and 10 m² area scales. The combined data for all habitat types generated a line graph representing the cumulative mean of species observed for each quadrat type to determine optimum quadrat area (Figure 3.8). At 4 m² a mean of 0.75 species were observed, at 6 m² a mean of 1.25 species were observed, at 8 m² a mean of 2.13 species were observed and at 10 m² a mean of 2.63 species were observed. The time taken to sample a 10x10m quadrat was 1 person/hour (2 persons x 30 min) enabling 3 quadrats to be completed per morning within the optimum time for herpetofaunal activity which was 9am to 12pm (Heyer et al. 1994). So with the aim to observe maximum species diversity within the optimum time and area, 10 m² was used in the actual sampling.

3.5

2.5

1.5

species observed observed species 1

0.5 Cumulative mean of herpetofaunal of mean herpetofaunal Cumulative 0 45678910 Size of quadrat (m²)

Figure 3.8 Area-species curve constructed using data from pilot study. Data from all habitat types were combined (4 quadrat replicates x 4 habitat types).

3.2.2 Major Fieldwork The research methods for measuring variables and obtaining data follow standard methods for amphibian studies outlined in Heyer et al. (1994) and were also adapted from a study by Gardner et al. (2001). This current research undertook a non- manipulative experimental design of passive observation to discern the relationship between herpetofauna richness and forest habitat type.

D’Cruze & Kumar (2011) recommended that when dealing with herpetofauna a variety of sampling methods should be used to provide a greater comprehensive evaluation. Frogs belonging to the order Anura will be sampled and lizards belonging to the family Gekkonidae (geckos) and family Scincidae (skinks). Frogs and geckos are predominantly nocturnally active animals and most skinks are diurnally active, therefore the two different sampling methods used were Nocturnal Visual Encounter Surveys (NVES) for frogs and geckos and Diurnal Quadrat Sampling (DQS) for skinks following Heyer et al. (1994) and Gardner et al. (2007). The herpetofauna observed were identified to species level using Frogs of the Solomon Islands (Pikacha et al. 2008) for frogs and Reptiles of the Solomon Islands (McCoy 2006) for geckos and skinks.

Three field trips were conducted over an 8 month period (September, January and April) to take into account the effects of seasonality on species abundance. In each field trip each of the 5 selected habitat types was surveyed with 6 transects and 9 quadrats amounting to a total of 90 transects (3 trips x 5 habitats x 6 transects) and 135 quadrats (3 trips x 5 habitats x 9 quadrats). However due to limited availability of teak, coastal and upland forests fewer transects and quadrats were carried out in these habitat types (Table 3.3). It is important to note that these five forest habitat types are mutually exclusive.

Table 3.3 Total no. of transects and quadrats carried out in each forest habitat type.

Habitat type Transects (NVES) Quadrats (DQS) Feasible Desired Feasible Desired Unlogged coastal forest 12 18 18 27 Unlogged lowland forest 24 18 36 27 Logged lowland forest 16 18 24 27 Unlogged upland forest 18 18 27 27 Teak plantation forest 10 18 15 27 Total 80 90 120 135

3.2.2.1 Transects (NVES) Transects can effectively track “species diversity, abundances and density”, this is a useful method for sampling along gradients and also within and across habitat types along a straight line with a fixed length and direction (Bennett 1999). Also at night frogs are more mobile and can be encountered at higher rates using torches (Hill et al. 2005). Nocturnal geckos are best found by night time torchlight searching as some species give off “eye-shine” and many are paralysed by the torch beam to make capture easier (Heyer et al. 1994).

Visual Encounter Surveys helps to determine species richness of an area, species assemblage of the area and relative abundances upon a certain time period expressed in person-hours (Bennett 1999). Four basic assumptions of the VES are; i) every individual of every species has the same chance of being observed during a survey, 34 ii) each species is likely to be observed during each sampling session, iii) an individual is recorded only once in each survey, iv) results from two or more observers surveying the same area simultaneously are identical (Bennett 1999). These basic assumptions were accepted and applied in this study.

In the NVES, a 500 m x 6 m belt transect was placed in each habitat type. Sampling began around sunset at 1830 hrs and covered 2 line transects of the same habitat type per evening at a fixed effort of 2 man hours per transect (2 persons x 1 hour). All transects were located at least 100 m from the forest edge. Transects within the same forest habitat types were separated by a minimum distance of 200 m and separated by a minimum of 500 m for transects between different forest habitat types. This was done to minimize the problems of edge effects and pseudo- replication as discussed in Heyer et al. (1994).

3.2.2.2 Quadrats (DQS) Quadrat sampling involved the random placement of quadrats within a habitat to thoroughly search visually and by hand for herpetofauna. Quadrats can record species presence and absence, abundances and densities (Bennett 1999). This method is usually used for sampling in leaf litter and on stream sides where species densities can be high. Skinks are primarily diurnal but some species are also active during the night and therefore many skinks are found in and amongst leaf litter (Heyer et al. 1994). Assumptions of the technique were that all animals are equally available to the observer to be observed and that observers should not be changed as this may add a bias. According to Bennett (1999) strengths of this technique are “hands on experience, the observation of cryptic species and juveniles” and this is a good efficient technique for sampling multiple species in “heterogeneous habitats”.

For the DQS, surveys were timed to coincide with the temperature window occurring between 0900hrs and 1100hrs where reptiles are likely to bask in the sun (Hill et al. 2005). 10 m² randomly placed quadrats were used during 0900 hrs and 1100hrs covering 3 quadrats at a fixed effort of 1 person-hour per quadrat (2 persons x 0.5 hours). Randomization for the placement of the quadrats is carried out using the ‘randomized walk’ method where the observer uses pre-determined compass directions and distances for the placement of the quadrats (Heyer et al. 1994). So

35 beginning at a random location a randomized walk of a set distance and direction will lead to the placement of the north corner of the quadrat.

All quadrats were located at least 100 m from the habitat type edge to avoid edge effects that may be unrepresentative of a given habitat. Quadrats within the same forest habitat types were separated by a minimum distance of 50m and separated by a minimum of 500 m from quadrats between different forest habitat types.

3.2.2.3 Ethnological Questionnaires Documenting patterns of human use and local knowledge of biodiversity is an important aspect of any conservation research project and rich species specific data can be collected through systematic surveying of local community members (Heyer et al. 1994). Therefore, questionnaire surveys were designed and carried out to record the perceptions and knowledge that local people have of herpetofauna and forest habitats. Villages within the Tai political ward were selected for questionnaire surveys, with villages located adjacent to surveyed forests. A total of 30 questionnaires were administered which included 10 questionnaires to persons over the age of 60, 10 between the ages of 30 and 60 and 10 to person under the age of 30, with a gender ratio of 15 females and 15 males. The open ended questionnaire used can be found in Appendix 1.

CHAPTER 4: RICHNESS AND ABUNDANCE OF FROGS, GECKOS AND SKINKS ON MALAITA

4.1 Introduction Although ecologically important, the herpetofauna of Malaita, specifically the frogs geckos and skinks have been poorly studied. This probably results from the lack of funding for such research, limited human-resource capacity and a decreasing amount of natural habitat essential for such biodiversity studies. However, McCoy (2006) and Pikacha et al. (2008) have produced useful field guides with species lists for Malaita. Pikacha et al. (2008) describes eight species of frogs for Malaita although more recent genetic work by Pikacha (unpub. data) may result in the identification of additional single island endemic frog species. McCoy (2006) describes six species of geckos and 14 species of skinks for Malaita island, although as in the case of P. Pikacha, R. Fisher (unpub. data) also suggests new genetic species. This study will test and build on these species lists by describing local abundances and identifying possible species additions. None of the species encountered are currently classified as endemic to Malaita. All species except the introduced B. marinus are regional endemics. All species encountered are classified by IUCN as Least Concern, except two that are Near Threatened.

This chapter addresses objective 1: To survey forest habitats on Malaita to determine the abundance, richness and local conservation status of frogs, skinks and geckos. The results will begin with a summary of total herpetofaunal richness and abundance encountered during the surveys followed by individual species analysis. Field photographs (in situ) and species that were found during this study are provided in Appendix B.

4.2 Specific Methodology Species were observed and recorded using the standard techniques for herpetofauna sampling described in Chapter 3. Night-time sampling (transects) targeted the nocturnally active herpetofauna (all frogs, geckos and 1 skink – Corucia zebrata) and the day-time sampling (quadrats) targeted the diurnally active herpetofauna (all skinks except for C. zebrata). For each individual animal

37 encountered the following details were recorded: species name, specific habitat and microhabitat, whether vocalizing or not (in the case of frogs) and if collected or not. Following on Bennett (1999) collected animals were placed in sealed bags for closer inspection or photography. In situ species identifications were then later confirmed using identification keys and species descriptions in McCoy (2006) and Pikacha et al. (2008). Once identified all collected specimens were then released at the site of capture on the following morning.

Encounter rates of different species were compared among habitat types using Kruskal Wallis tests. Significant results were those with P < 0.05 (critical value H ≥ 9.49, df = 4), if results were found to be significant then a bar graph was produced to display this. Species commonality is simply defined as rare, (<4 total encounters), uncommon (4-16 total encounters), common (17-64 total encounters) and very common (>64 total encounters), number cut offs were determined using an exponential gradient multiplied by 4.

Of the species that were not rare, species were split into generalist (encountered in greater than two habitats) or specialist (encountered in only 1 or 2 habitats) behaviour based on the number of habitats they were encountered in. A species preferred forest habitat type is based on the habitat type with the highest mean abundance for that particular species regardless of whether it displays generalist or specialist behaviour.

From the specialist species that exhibited forest habitat preferences, it is possible to deduce possible indicator species for forest health. Indicator species of forest health will be defined in this study as 1) showing high population abundances in relatively undisturbed areas, 2) showing low population abundances in areas of habitat degradation and 3) cannot be naturally rare so that they are difficult to encounter.

Species richness and abundance were chosen metrics for the study mainly because of simplicity both for data collection and data analysis. Richness provides information that is both easy to understand and data that provides direct information on the diversity of an area. Abundance also provides information on the relative “health” of populations for communities and individual species. 38

4.3 Results

4.3.1 Summary of results A total of 21 herpetofaunal species were encountered during both nocturnal and diurnal sampling: 8 frogs, 3 geckos and 10 skinks (Appendix B). For each species commonality based on encounter rates and microhabitat preference based on observations in the field will be listed followed by a short description with supporting statistical tests and figures.

4.3.2 Nocturnal herpetofauna A comparison of the results for nocturnal herpetofaunal commonality (encounter rate), micro-habitat preference, and total encounters show the introduced species Bufo marinus to be dominant (Table 4.1). The IUCN Red-list status (IUCN 2012) and endemic status (McCoy 2006, Pikacha et al. 2008) of each species is also listed in Table 4.1.

Table 4.1 Summary of nocturnal results. Commonality = rare, (<4 total encounters), uncommon (4-16 total encounters), common (17-64 total encounters) and very common (>64 total encounters). SI = Solomon Islands. Endemic status taken from McCoy (2006) and Pikacha et al. (2008) and Red- list status taken from IUCN (2012) Species Endemic status status IUCN Malaita Commonality on preference habitat micro- Observed individuals encountered Total sum of

Red-list

Frogs Batrachylode S.I. Least Very 1-2 above 210 s vertebralis national Concern common ground on endemic epiphytes, ferns and tree trunks Bufo marinus Introduced Least Very On the ground, 326 (invasive) Concern common along tracks species and still-water bodies 39

Ceratobatra- S.I. Least Very On or under 86 chus national Concern common leaf litter guentheri endemic Discodeles S.I. Least Very On rocks 111 guppyi national Concern common beside moving endemic waterways Platymantis S.I. Least Very Arboreal, 2- 73 guppyi national Concern common 10m above the endemic ground on broad-leafed shrubs, trees and palms Platymantis S.I. Least Common On the ground, 21 solomonis national Concern in caves and endemic holes Platymantis S.I. Least Very On the ground, 104 weberi national Concern common in holes and endemic dead logs Rana kreffti S.I. Least Common On the ground, 51 national Concern close to still endemic waters Geckos

Cyrtodactyl- S.I. Near Uncommon Large trees 14 us national Threatened salomonensis endemic G. oceanica Regional Least Uncommon Tree trunks, 14 endemic Concern Pandanus spp. Nactus Regional Least Common Tree trunks, 64 multicarina- endemic Concern tree hollows tus Skinks

Corucia S.I. Near Rare Tree trunks 3 zebrata national Threatened with dense endemic epiphytes

Batrachylodes vertebralis Boulenger, 1887 Based on the total sum of encountered individuals (210) in all habitat types Batrachylodes vertebralis (Appendix B) is classed as very common on Malaita Island (Table 4.1). Observed microhabitat preference especially for vocalising males is between 1-2 m above ground on tree trunks, epiphytes (eg. Asplenium nidus), ferns, the tree fern (Cyathea vittata) and occasionally found on the ground.

There is a significant difference between habitat types with more B. vertebralis found in upland forest (145) than any other habitat type (KW test transects H = 13.75, df = 4, P < 0.05, Figure 4.1).

Figure 4.1 Batrachylodes vertebralis nocturnal (transect) mean encounter rate for each habitat type, Error Bars: 95% Confidence Interval.

Bufo marinus Linnaeus, 1758 Based on the total sum of encountered individuals (326) in all habitat types Bufo marinus (Appendix B) is classed as very common and had the greatest sum of encounter of all species (Table 4.1). Observed microhabitat preference is on the ground especially in cleared or semi-cleared areas such as along bush tracks and aggregations have been observed around still water bodies. This species seems to favour drier conditions and areas of high anthropogenic activity.

There were significant differences in encounter rates between habitat types, with significantly less B. marinus found in upland forest, especially compared to logged and teak forests but there were no differences in frog abundance between the other four habitats (KW test transects H = 27.00, df = 4, P < 0.05, Figure 4.2).

Figure 4.2 Bufo marinus nocturnal (transect) mean encounter rate for each habitat type, Error Bars: 95% Confidence Interval.

Ceratobatrachus guentheri Boulenger, 1887 Based on the total sum of encountered individuals (86) in all habitat types Ceratobatrachus guentheri (Appendix B) is classed as very common (Table 4.1). Observed microhabitat preference is on or under dead leaves with high preference for areas of thick leaf litter. Bamboo (Nastus obtusus) groves provide safe areas for eggs and juveniles.

There was a significant difference in the encounter rates of C. guentheri in the different habitats (KW test transects H = 38.15, df = 4, P < 0.05). Individuals were most common in the upland forest habitats followed by lowland and logged habitats. No individuals were found in the coastal and teak plantations (Figure 4.3).

Figure 4.3 Ceratobatrachus guentheri nocturnal (transect) mean encounter rate for each habitat type, Error Bars: 95% Confidence Interval.

Discodeles guppyi Boulenger, 1884 Based on the total sum of encountered specimens (111) in all habitat types Discodeles guppyi (Appendix B) is classed as very common (Table 4.1). Observed microhabitat preference is on rocks besides streams especially waterfalls and on the ground in riparian forest but not far from waterways.

Almost all D. guppyi were encountered in lowland forest (110) habitats more than any other habitat (KW test transects H = 25.93, df = 4, P < 0.05, Figure 4.4). No individuals were found in teak, upland or coastal habitats.

Platymantis guppyi Boulenger, 1887 Based on the total sum of encountered specimens (73) in all habitat types Platymantis guppyi (Appendix B) is classed as very common (Table 4.1). Observed microhabitat preference is between 2-10 m above the ground on broad-leafed trees, shrubs (Elatostema sp., Alpinia oceanica and Cominsia guppyi), ferns, palms and the epiphyte Asplenium nidus.

There was a significant difference between the encounter rates of P. guppyi in lowland, upland and logged forests with teak forests, (KW test transects H = 14.76, df = 4, P < 0.05, Figure 4.5). Platymantis guppyi individuals were most commonly found in the lowland (38) and upland (21) forest habitats followed by logged and coastal forest habitats. No individuals were found in the teak plantations.

Figure 4.4 Discodeles guppyi nocturnal (transect) mean encounter rate for each habitat type, Error Bars: 95% Confidence Interval.

Platymantis solomonis Boulenger, 1887 Based on the total sum of encountered specimens (21) in all habitat types Platymantis solomonis (Appendix B) species is classed as common (Table 4.1). Observed microhabitat preference is on the ground, in caves, holes and under ledges.

There was a significant difference in the encounter rates of P. solomonis in lowland and upland forests compared to other forests, however no significant difference existed between lowland and upland forests (KW test transects H = 13.58, df = 4, P < 0.05, Figure 4.6). Platymantis solomonis individuals were found in the lowland (15) and upland (6) forest habitats. No individuals were found in the logged, coastal or teak plantation forests. 44

Figure 4.5 Platymantis guppyi nocturnal (transect) mean encounter rate for each habitat type, Error Bars: 95% Confidence Interval.

Figure 4.6 Platymantis solomonis nocturnal (transect) mean encounter rate for each habitat type, Error Bars: 95% Confidence Interval. 45

Platymantis weberi Schmidt, 1932 Based on the total sum of encountered specimens (104) in all habitat types Platymantis weberi (Appendix B) is classed as very common (Table 4.1). Observed microhabitat preference is on the ground and in holes with calling males usually found in slightly elevated positions such as fallen logs or tree stumps.

There was a significant difference in the encounter rates of P. weberi in lowland and upland forests with coastal forests (KW test transects H = 15.39, df = 4, P < 0.05), however there was no significant differences between other habitats (Figure 4.7). Platymantis weberi individuals were commonly found in the lowland (45) and upland (30) forest habitats, followed by logged and teak plantation habitats. The lowest number of individuals was found in the coastal forests.

Figure 4.7 Platymantis weberi nocturnal (transect) mean encounter rate for each habitat type, Error Bars: 95% Confidence Interval.

Rana kreffti Boulenger, 1884, also known as Hylarana kreffti Based on the total sum of encountered specimens (51) in all habitat types Rana kreffti (Appendix B) is classed as common (Table 4.1). Observed microhabitat

46 preference is on the ground and close to still/stagnant water where males call with loud distinctive notes.

There was no significant difference in the encounter rates of H. kreffti in the different forest habitat types (KW test transects H = 5.98, df = 4, P < 0.05).

Cyrtodactylus salomonensis Rösler, Richards & Günther, 2007, formally known as C. louisiadensis De Vis, 1892 Based on the total sum of encountered specimens (14) in all habitat types Cyrtodactylus salomonensis (Appendix B) is classed as uncommon (Table 4.1). Observed microhabitat preference is on large tree trunks especially those without climbing epiphytes such as P. pinnata, Canarium sp. and Ficus sp. trees.

There was a significant difference in the encounter rates of C. salomonensis in lowland and logged forests with all other forests, however there is no significant difference between the two (KW test transects H = 14.08, df = 4, P < 0.05, Figure 4.8). Cyrtodactylus salomonensis individuals were commonly found in the lowland forests (12) followed by logged forest (2) habitats, no individuals were found in the upland, teak plantation and coastal forests.

Gehyra oceanica Lesson, 1830 Based on the total sum of encountered specimens (14) in all habitat types Gehyra oceanica (Appendix B) is classed as uncommon (Table 4.1). Observed microhabitat preference is on tree trunks but especially on broad-leafed shrubs such as Pandanus sp. and the sago palm Metroxylon salomonense.

There was no significant difference in the encounter rates of G. oceanica in the different forest habitat types (KW test transects H = 2.88, df = 4, P < 0.01).

Nactus multicarinatus Günther, 1872 Based on the total sum of encountered specimens (64) in all habitat types Nactus multicarinatus (Appendix B) is classed as common (Table 4.1). Observed microhabitat preference is on the ground on tree trunks and especially in and around tree hollows. This species seems to favour drier conditions and areas of high anthropogenic activity.

There was a significant difference in the encounter rates of N. multicarinatus individuals in the different forest habitat types (KW test transects H = 10.68, df = 4, P < 0.05, Figure 4.9). Nactus multicarinatus individuals were found in all forests habitat types. Transects in logged forests had significantly lower values than lowland and teak forests.

Figure 4.8 Cyrtodactylus salomonensis nocturnal (transect) mean encounter rate for each habitat type, Error Bars: 95% Confidence Interval.

Corucia zebrata Gray, 1855 Based on the total sum of encountered specimens (3) in all habitat types Corucia zebrata (Appendix B, plate 12) is classed as rare (Table 4.1). Observed microhabitat preference is on tree trunks, especially tree trunks with thick climbing epiphytes such as P. pinnata, Canarium sp. and Ficus sp. trees. Due to its rarity and lack of recordings no statistical tests were carried out in relation to habitat preference.

Figure 4.9 Nactus multicarinatus nocturnal (transect) mean encounter rate for each habitat type, Error Bars: 95% Confidence Interval.

4.3.3 Diurnal herpetofauna A comparison of the results for diurnal herpetofaunal commonality (encounter rate), micro-habitat preference, and total encounters show the native species Emoia pseudocyanura to be dominant (Table 4.2). The IUCN Red-list status (IUCN 2012) and endemic status (McCoy 2006, Pikacha et al. 2008) of each species is also listed in Table 4.2.

Emoia atrocostata freycineti Duméril & Bibron, 1839, Solomon Islands subspecies Based on the sum total of encountered specimens (1) in all habitat types Emoia atrocostata freycineti (Appendix B) is classed as rare (Table 4.2). Observed microhabitat preference is on the ground and on rocks within or beside the intertidal zone, this indicates its non-preference for forested areas. Due to its rarity and lack of recordings no statistical tests were carried out in relation to habitat preference.

Table 4.2 Summary of diurnal results. Commonality = rare, (<4 total encounters), uncommon (4-16 total encounters), common (17-64 total encounters) and very common (>64 total encounters). SI = Solomon Islands. Endemic status taken from McCoy (2006) and Pikacha et al. (2008) and Red- list status taken from IUCN (2012) status preference Species Endemic status IUCN on Malaita Commonality micro Observed individuals encountered Total sum of - habitat

Red

-list

Skinks

E. atrocostrata S.I. national Least Rare Rocks and 1 endemic Concern close to inter-tidal zone E. cyanogaster Regional Least Uncommon Arboreal, on 6 endemic Concern tree trunks and shrubs 1-5m. E. nigra Regional Least Common Leaf litter, 35 endemic Concern clearings, tree trunks E. S.I. national Least Very Leaf litter, 189 pseudocyanura endemic Concern Common clearings, tree trunks, shrubs and ferns P. virens Regional Least Rare Arboreal 5m 1 endemic Concern S. bignelli S.I. national Least Uncommon On the 6 endemic Concern ground, under debris S. concinnatus S.I. national Least Very On the 94 endemic Concern common ground, leaf litter S. cranei S.I. national Least Rare Within tree- 1 endemic Concern fern trunks S. solomonis S.I. national Least Uncommon In the 6 endemic Concern ground, rotting wood

Emoia cyanogaster Lesson, 1826 Based on the total sum of encountered specimens (6) in all habitat types Emoia cyanogaster (Appendix B) is classed as uncommon (Table 4.2). Observed microhabitat preference is between 1-5 m above the ground on tree trunks and branches, especially those with think epiphytic cover.

There was a significant difference in the encounter rates of E. cyanogaster in logged and lowland forests with all other forests (KW test quadrats H = 16.06, df = 4, P < 0.05, Figure 4.10). Emoia cyanogaster individuals were found in logged and lowland forests and no individuals were encountered in all other habitat types.

Figure 4.10 Emoia cyanogaster diurnal (quadrat) mean encounter rate for each habitat type, Error Bars: 95% Confidence Interval.

Emoia nigra Jacquinot & Guichenot, 1853 Based on the total sum of encountered specimens (35) in all habitat types Emoia nigra (Appendix B) is classed as common (Table 4.2). Observed microhabitat preference is on the ground especially along bush paths and amongst leaf litter, with some specimens also found around 1m above the ground on tree trunks.

There was no significant difference in the encounter rates of E. nigra individuals in the different forest habitat types (KW test transects H = 9.13, df = 4, P < 0.05).

Emoia pseudocyanura Brown, 1991 Based on the total sum of encountered specimens (189) in all habitat types Emoia pseudocyanura (Appendix B) is classed as very common (Table 4.2). Observed micro-habitat preference is on the ground especially along bush paths and amongst leaf litter; specimens are also found around 1m above the ground on tree trunks, shrubs and ferns (eg. Alpinia oceanica), it is commonly found basking in direct sunlight.

There was a significant difference in the encounter rates (H = 36.99, df = 4, P < 0.05) with teak plantation and logged forests showing higher encounter rates compared with the other forest types (Figure 4.11). Although individuals were found in all forest habitats they were more common in logged and teak forests, followed by coastal then lowland and upland forests.

Figure 4.11 Emoia pseudocyanura diurnal (quadrat) mean encounter rate for each habitat type, Error Bars: 95% Confidence Interval.

Prasinohaema virens Boulenger, 1883 Based on the total sum of encountered specimens (1) in all habitat types Prasinohaema virens (Appendix B) is classed as rare (Table 4.2). Observed microhabitat preference is arboreal at around 5 m above the ground along ends of branches. Due to its rarity and lack of recordings no statistical test were carried out in relation to habitat preference.

Sphenomorphus bignelli Schmidt, 1932 Based on the total sum of encountered specimens (6) in all habitat types Sphenomorphus bignelli (Appendix B) is classed as uncommon (Table 4.2). Observed microhabitat preference is on the ground and often burrowing under dead debris or leaf litter.

There was no significant difference in the encounter rates of S. bignelli individuals in the different forest habitat types (KW test transects H = 3.87, df = 4, P < 0.05).

Sphenomorphus concinnatus Boulenger, 1887 Based on the total sum of encountered specimens (94) in all habitat types Sphenomorphus concinnatus (Appendix B) is classed as very common (Table 4.2). Observed microhabitat preference is on the ground especially along bush paths and amongst leaf litter; some specimens also burrow into rotting debris.

There was a significant difference in the encounter rates of S. concinnatus in teak plantation and lowland forests with all other forests (KW test quadrats H = 12.61, df = 4, P < 0.05, Figure 4.12). Sphenomorphus concinnatus individuals were found in all forest habitats except coastal forest and were most common in teak plantation and lowland forest habitats in transects.

Figure 4.12 Sphenomorphus concinnatus diurnal (quadrat) mean encounter rate for each habitat type, Error Bars: 95% Confidence Interval.

Sphenomorphus cranei Schmidt, 1932 Based on the total sum of encountered specimens (1) in all habitat types Sphenomorphus cranei (Appendix B) is classed as rare (Table 4.2). Observed microhabitat preference is within the trunks of tree ferns (Cyathea vittata). Due to its rarity and lack of recordings no statistical tests were carried out in relation to habitat preference.

Sphenomorphus solomonis Boulenger, 1887 Based on the total sum of encountered specimens (6) in all habitat types Sphenomorphus solomonis (Appendix B, plate 21) is classed as uncommon (Table 4.2). Observed microhabitat preference is on the ground, often burrowing under debris, rotting wood or leaf litter.

There was no significant difference in the encounter rates of S. solomonis individuals in the different forest habitat types (KW test transects H = 5.33, df = 4, P < 0.05).

4.3.4 Additional species In addition to the 21 species encountered during the sampling surveys there were also five other species observed outside of sampling areas. These included one frog (Litoria thesaurensis Peters, 1877), two geckos (Lepidodactylus lugubris Duméril and Bibron, 1836 and Hemidactylus frenatus Duméril and Bibron, 1836) and two skinks (Eugongylus albofasciolatus Shaw, 1802 and Lamprolepis smaragdina Lesson, 1830). Litoria thesaurensis and E. albofasciolatus were observed only once at night in lowland forests and appear to be rare on the island of Malaita. Lepidodactylus lugubris, H. frenatus and L. smaragdina were found abundantly around homes and appear to be strong human commensals. Therefore, in total 26 species of frogs, geckos and skinks were observed, during field trip periods on Malaita.

4.3.5 Species behaviour and Indicator species Twelve species showed significant forest habitat preferences (7 frogs, 2 geckos and 3 skinks)(Table 4.3). All ‘very common’ species except for D. guppyi appeared to be generalist species and were found in more than two habitats, indicating that the majority of forest herpetofaunal biomass consists of such species. All species that did not exhibit habitat preference were ‘common’ and ‘generalists’.

Table 4.3 A comparison of species behaviour and habitat preferences of the 21 herpetofaunal species encountered during sampling. Possible indicator species are shaded, blank cells indicate insufficient data available to make allocations.

Species Commonality on Generalist or Preferred habitat, Malaita specialist forest with highest mean abundance Frogs B. marinus Very common Generalist Logged B. vertebralis Very common Generalist Upland C. guentheri Very common Generalist Upland D. guppyi Very common Specialist Lowland H. kreffti Common Generalist No preference

P. weberi Very common Generalist Lowland P. solomonis Common Specialist Lowland P. guppyi Very common Generalist Lowland Geckos C. salomonensis Uncommon Specialist Lowland G. oceanica Uncommon Generalist Coastal N. multicarinatus Common Generalist No preference Skinks C. zebrata Rare - No preference E. pseudocyanura Very common Generalist Teak E. atrocostata Rare - No preference E. cyanogaster Uncommon Specialist Logged E. nigra Common Generalist No preference P. virens Rare - No preference S. bignelli Uncommon Specialist No preference S. concinnatus Very common Generalist Teak S. solomonis Uncommon Generalist No preference S. cranei Rare - No preference

Three possible indicator species of healthy lowland forest are: the frogs: D. guppyi and P. solomonis and the gecko C. salomonensis. All are relatively common specialist species preferring lowland forests. These three species all have pros and cons as indicator species but from all herpetofauna observed during sampling they appear to be the best indicator species candidates for healthy lowland forest on Malaita.

4.4 Discussion of Results

4.4.1 Indicator Species Discodeles guppyi is a large and unique frog found in high numbers close to waterways (Table 4.1) and is easily distinguished from other species. It is also a charismatic species and well known by local inhabitants (see Ch. 6). However, being

56 an aquatic frog (as opposed to arboreal or ground frog) and found only in areas close to waterways, it may not be a useful indicator for general forest habitat, though use as an indicator of riparian or waterway health may be more applicable.

Platymantis solomonis is a medium sized frog found in good numbers in relatively healthy forest, although it is difficult to distinguish from P. weberi at a distance and usually grouped together with P. weberi (a smaller wider ranging frog) by locals (pers. obs.). However, due to its wide distribution in healthy forest P. solomonis is a good candidate for both lowland and upland forest health.

Cyrtodactylus salomonensis is a large, unique gecko found in relative low densities in lowland forests. It is easily distinguished from other lizards and is also a charismatic species to locals (see Ch. 6). Being an arboreal species with low densities, the encounter rate of this species is relatively low which may result in biased indications of forest health.

4.4.2 Herpetofaunal richness comparisons to other studies Many studies have sampled herpetofaunal richness in different habitat types globally, and several of these studies have been undertaken in tropical areas and have used similar methods to the current study (Table 4.4). For example, this study recorded a total of 8 frogs during sampling which is in the range of 7 and 23 species of frogs that were recorded in these previous studies. Also a total of 13 lizards (skinks + geckos) were recorded which also falls within the range of 3 to 30 lizards that were recorded in the previous studies. This suggests that the species richness found in this study corresponded well to similar studies around the world.

4.4.3 Herpetofaunal richness comparisons to other Solomon Island islands A comparison of the herpetofaunal richness found in this current study with the 5 other major islands in the Solomon’s archipelago according to McCoy (2006) and Pikacha et al. (2008) shows that Malaita has the second lowest herpetofaunal richness per square kilometre (behind Makira) of all the major Solomon Island’s islands (Table 4.5). The approximate island sizes and distances to nearest continental

57 landmass (biodiversity source area), in this case mainland PNG, is also listed in Table 4.5.

Table 4.4 A selection of global tropical herpetofaunal studies similar to the current study. Place used sampling methods Herpetofaunal sampled habitats Different richness herpetofauna Results year St udy authors and

of study of total of

Biological Station Costa Rica, 75 x 25m² plots primary andplantation forest Abandoned cacao gecko 18 frogs, 2 skinks and 1 Heinen (1992) undisturbed La Selva La

forest National Kibale Uganda, 50 x 25m² plots forest pine plantation and logged forest Undisturbed forest, lizards and10 frogs 5 Vonesh

(2001) Park

and Buton Islands, Sulawesi Indonesia, Indonesia, stream point and 20mindiurnal nocturnal traps,Pitfall driftnets, 1hr estuarine caves. habitats and coastaland towns, and villages farmland, forest, plantationsecondary forest, forest,moderately disturbed Minimally forest, disturbed geckos 13 frogs, 15 skinks and 10 Gillespie counts and 200m transects transects et al. Kabaena, Mu Kabaena, (2005) na

boards Brazil, transects traps P secondary rainforest Mature primary lizards and23 frogs 30 Gardner fence plantation forest and mature area of Amazonia of area (2007 itfall traps, itfall , , funnel , ) sticky trap and

Jari River

et al. ,

forest drift

upland rainforest Solomon Islands, Choiseul 12 x 50x100m plots mid secondary lowland forest, lowland swamp forest, plantation/gardens, rainforest, lowland palm forest, lowland coastal forest,Lowland skinks 15 frogs, 5 geckos and 7 Morrison - altitude rainforest and altitude rainforest et al. (2007)

orchards forests and Francais Montagne des Madagascar, 9 forest, clear Undisturbed skink 7 geckos and 1 2011)Kumar and (D'Cruze - 0.8ha plots

- cut traps pitfall with fences drift forest and native secondary Hong Kong 40 0.66² cover 100m transects, nocturnal and158 diurnal 116 E and7 frogs 5 lizards Sung xotic plantation

(2011)

- boards and boards forest

Islands, Malaita Solomon quadrats 120 x 10² and transects 80 x 500m teak plantations lowland and logged upland, lowland and Coastal, Unlogged geckos and 10 8 frogs, 3 (this study) Pollard, 2012

Isabel island, which is most similar in island size to Malaita has 10 more frog species, but two less gecko and four less skink species (McCoy 2006, Pikacha et al. 2008). Makira which is slightly smaller in size than Malaita is the island with the

59 most similar approximate distance from PNG to Malaita and has six less frogs, one less gecko and two less skink species (McCoy 2006, Pikacha et al. 2008). Therefore based on Table 4.5, island size seems to be a better determinant for lizard richness whereas distance from source a better determinant for frog richness.

Table 4.5 A comparison of the recorded richness of frogs, geckos and skinks based on Pikacha et al.(2008) and McCoy (2006) of the six major islands of the Solomon Island’s archipelago, with island size (UNEP 1998) and distance from mainland PNG (daftlogic.com 2012)

Island name No. of No. of No. of Total of Approx. Approx. Frogs Geckos Skinks the three island distance groups size from (km2) mainland PNG (km) Choiseul 19 4 15 38 2971 740 New Georgia 8 7 12 27 2037 760 Isabel 18 4 10 32 3665 870 Guadalcanal 12 12 15 39 5352 970 Makira 2 5 12 19 3190 1150 Malaita 8 6 14 28 Malaita (this 9 3 14 26 3836 1100 study)

4.4.4 Malaitan Herpetofaunal richness compared to McCoy and Pikacha One frog (Discodeles bufoniformis), one gecko (Lepidodactylus guppyi) and two skinks (Emoia cyanura and Emoia flavigularis) were recorded in the Malaitan herpetofaunal species lists of McCoy (2006) and Pikacha et.al (2010) but were not found during the current study sampling (Table 4.5). In addition, two frogs (R. kreffti and B. marinus) were found which had not been recorded in the species list of McCoy and Pikacha and are new records for the island of Malaita

Possible reasons why species were not observed or why some other species might have recorded minimal sightings fall into two categories. Either the species in question is rare/absent or the methods used were not suitable for accurate observation of the species. As sampling for herpetofauna was focused on forest habitats, gecko species such as L. lugubris, H. frenatus and L. smaragdina that are regarded as human commensals (McCoy 2006) were not expected to be found. In addition, the skink Emoia cyanura was not expected to be found on mainland Malaita as McCoy (2006) states that it is found only on nearby smaller islands in the Langa-Langa lagoon.

Table 4.6 Overall Malaitan herpetofaunal (frogs, geckos and skinks) species lists according to McCoy (2006) (M) and Pikacha et al. (2008) (P) and species observed in this study. (X = during sampling and x = outside of sampling but seen during field trip periods).

Species names Species names Recorded in lists species Encountered study in this Recorded in lists species Encountered study in this B. vertebralis P X H. frenatus M x B. marinus - X L. guppyi M - C. guentheri P X L. lugubris M x C. salomonensis M X L. smaragdina M x C. zebrata M X L. thesaurensis P x D. bufoniformis P - N. multicarinatus M X D. guppyi P X P. guppyi P X E. albofasciolatus M x P. solomonis P X E. atrocostrata M X P. virens M X E. cyanogaster M X P. weberi P X E. cyanura M - R. kreffti - X E. flavigularis M - S. bignelli M X E. nigra M X S. concinnatus M X E. pseudocyanura M X S. cranei M X G. oceanica M X S. solomonis M X

4.4.5 Evaluation of methods used With reference to ease of use and practicality in the field, the current methods were satisfactory with reference to physical demands on samplers and time availability to carry out sampling. The methods excelled in the encounter rates for frogs with the nocturnal VES recording a high rate for frog abundances and richness. However, there was a weakness in the lizard surveys because of a high chance of error in the identification of the fast moving lizards. Comparatively the relatively stationary nature of frogs, made the accuracy of identification higher. The methods used also had a weakness for under detection of arboreal species and therefore such species were probably under recorded. More exhaustive sampling including more sites and covering more seasons may result in an increased abundance of species such as C. zebrata, S. solomonis, S. cranei, E. atrocostata and P. virens. The observation of unobserved species such as L. thesaurensis, D. bufoniformis, L. guppyi, E. flavigularis and E. albofasciolatus, might then also be recorded.

Possible additional method to help improve the accuracy of visual identifications and increase the chance of capture for lizards especially the more cryptic species is the use of glue/sticky traps (Fisher 2011), cover-boards and funnel and pitfall traps with drift fences (Greenberg et al. 1994, Ryan et al. 2002). Traps can capture more difficult species however different techniques are better for different taxa and a combination of methods is recommended to achieve maximum species detections (Ryan et al. 2002).

4.5 Summary of herpetofaunal richness and abundance In summary nine frogs, five geckos and twelve skinks were observed on the island of Malaita. Of these 26 species, 12 indicated habitat preference based on the five different forest habitats sampled. Two previously unrecorded frog species (R. kreffti and B. marinus) were also found on the island although 3 species (D. bufoniformis, L. guppyi and E. flavigularis) previously recorded were not encountered. Bufo marinus is a well-known serious invasive pest species (GISD 2013) and its potential impacts on native fauna needs to be investigated further.

Most species encountered were of relatively high abundance and described as very common or common. However, four skinks were low in abundance having less

62 than four recordings during all sampling periods. No species are currently of a global conservation concern as measured by the IUCN Red-listing process, however specialist species such as D. guppyi and C. salomonensis are of local conservation concern due to habitat degradation.

Three species were selected as good possible indicator species for the health and intactness of lowland forest, the most highly threatened forest type in Solomon Islands (see Ch. 7). However, D. guppyi, C. salomonensis and P. solomonis were not suitable indicators for other forest types. More studies are needed especially with genetic work to identify and record and further herpetofaunal species on the island. The methods used provided excellent and expected results for Malaita, however the use of additional methods along with more exhaustive sampling may improve any further surveys. Methods to cater for arboreal species and also small, fast-moving skinks would also improve the accuracy of herpetofaunal sampling in any tropical forest.

CHAPTER 5: FOREST HABITAT AND HERPETOFAUNAL RICHNESS

5.1 Introduction The Solomon Islands has just over 2.2 million ha of forested areas which is approximately 79% of the total land area (FAO 2011). The composition of the forests are greatly dependant on disturbance levels and this disturbance results in a unique changing landscape (Burslem and Whitmore 1999). Gap formation (either man-made or natural) results in a “diverse fluctuating composition of climax species and pioneer species” in tropical forests (Bennet 2000). Forest ecosystems have become adapted to natural disturbances (eg. tropical cyclones) and species have adapted to take advantage of such disturbances resulting in a very resilient communities (Bennet 2000, Filardi et al. 2007).

Burslem et al. (2000) in a 30-year study on Kolombangara, Western Solomon Islands showed that cyclones only produce short-term impacts on intact forests and that major forest composition differences are caused by anthropogenic activities. Anthropogenic activities are the major factor influencing changes to the composition and distribution of Solomon forests and soil condition (Bennet 2000, Burslem et al. 2000). Forests are therefore not only communities of biological entities but are also a product of strong inter-relations with the resident human population. The first colonists cleared land and cut trees for agriculture, timber and fuel and also cultivated species of value such as the Canarium nut trees (Rolett and Diamond 2004). As quoted in Bayliss-Smith and Hviding (2003) “forests are in fact cultural artifacts exhibiting remarkable resilience in the face of both natural disturbance and human use over very long periods of time.”

Collation of the research of Ross (1973), Mueller-Dombois and Fosberg (1998), Bennet (2000), Pikacha et al. (2008) Pauku (2009) and FAO (2010) allows classification of the forests of the Solomon Islands into eight major categories (Table 5.1). Though these categories are useful guides, variations do occur within the different categories based on local topography, soil type and species composition. It is also important to note that distinct boundaries between the described forest types

are not easily defined and in many cases, a continuum of transformation may be more clearly observed (pers. obs.).

Table 5.1 The eight Major categories of forests found in the Solomon Islands as collated from Ross (1973), Mueller-Dombois and Fosberg (1998), Bennet (2000), Pikacha et al. (2008), Pauku (2009) and FAO (2010).

Comment Topographical Dominant botanical Forest location genera category also act marine ecosystems and ecological role Plays an important Inter Bruguiera Rhizophora forests 1. zone against high seas zone against high seas Saline swamp -tidal areas areas -tidal

s as a buffer

and for seasonrainy theinundated during usually that are areas draining poor Very Calophyllum Terminalia Metroxylon Cyrtosperma and riverine forests marshes, swamps 2. Freshwater , and

found on slopesfound and well sites drained Campnosperma, S T Flat inland areas Strongylocaryum Tapeinosperma Schizomeria Maranthes, Gmelina, Pometia,Parinari, Dillenia, such as trees Large 3. hort trees such as hort he most commercially exploited Lowland rainforest (also includes hill , Terminalia palms such as

Eleocarpus , Pandanus Barringtonia Leea Pterocarpus, Calophyllum , Canarium , and bamboos Endospermum Areca forest categoryforest and , Licuala ) and Vitex forest , , .

Is the sea Coastal close areas to Cerbera Callophyllum Ipomoea lowland beach forest) (also known as the 4. sea protection from the Casuarina Vigna Intsia Barringtonia a strong a strong Coastal forests

, ,

Terminalia Wollastonia , , Heritiera Canavalia

barrier of , , and , , , as montane the lower forest orchids, and orchids, Ardi wet and often cloudy,wet and often windy sites. tops and mountainridge peaks at commercial exploitation commercial This usually observed above 600 F Scheff Metrosideros trees Tall 5. ound on well-draining soils Upland ( rainforests sia, Rhododendron,sia, category has minimal lera, Podocarpus, Dacrydium Cyathea , Ficus , , tree-ferns , Psychotria, Psychotria, Eugenia, also known

bamboo, ) m asl on

climate Has a hospitable conditions. less there are where areas 1000masl U and Characterized by mosses moss forest 6. sually found above sually found Montane cloud or lichens lichens very cool, wet very

or in or other

isolated trees canopy and sparse with open Cleared and vines and invasive speciespioneer Dominated by 7. forests (also degraded L ogged forest

)

teak. teak. below onSee paragraph native and exotic species. around 28,000around ha Solomon Islands forest growth past lowland and coastal Mostly in areas of found Swietenia Acacia Terminalia,Eucalyptus, Campnosperma,Gmelina, 8. Plantation forests , Tectona .

and have of both

Teak (Tectona grandis) is an important timber plantation species especially on Malaita Island. For example, in 2009 a total of 363 kg of teak seeds were given by the Ministry of Forestry and Research (MoFR) to communities throughout the Solomon Islands to plant, of which 128 kg went to Malaita (MoFR 2009). Thus, since 2000 around 4,000 ha of teak have been planted throughout the country. During 2009 more than 103 ha of teak was planted in the country with over 12 ha of that on Malaita Island (MoFR 2009). These figures are probably an underestimate because they are only from areas under forestry observation and therefore don’t include all planted areas.

This chapter will define relationships between herpetofaunal incidence, forest habitat type and the degree of habitat degradation. It will begin with floral and herpetofaunal descriptions of each sampled forest habitat type. And then add a herpetofaunal richness analysis of all forest habitat types followed by results related to habitat degradation and modification.

5.2 Specific Methodology This current research study focused on five mutually exclusive forest types: 3) unlogged lowland, 4) unlogged coastal, 5) unlogged upland, 7) logged lowland and 8) teak plantation forests. These five categories were selected from the eight categories in Table 5.1 because they included the largest land cover area and were generally easily accessible. On the contrast, 1) saline swamp, 2) freshwater swamp and 6) montane forests were excluded from sampling due to great difficulty to access and because of minimal sampling area. The basic descriptions of these five forest habitats based on observations in the field are provided in Table 5.2.

Botanical lists (sorted into 4 floral group categories: canopy, understory, shrubs and epiphytes) were also generated to provide a brief botanical description of each forest habitat type. Photos of dominant plant species were taken to Honiara and identified by local botanist Myknee Sirikolo associated with the South Pacific Regional Herbarium (SPRH). Herpetofaunal abundance and richness was recorded for each forest habitat type using the sampling methods previously described in Chapter 3. Important floral plants will also be listed, these are plants that have any observed association with herpetofauna.

Table 5.2 Descriptions based on personal observations of the five habitat types used in this research study

Comments Description of General vegetation Forest forest habitat description habitat study sites type and sand gravel, or substrate, usually evident influence W the coast cases to adjacent and in m Flat land close to large trees coconuts and Abundance of Coastal forest 1. Unlogged ell- human- draining draining

any is

dark in colourdark in humusrich and Soils relatively are less than 300 m.a.s.l. slopes at elevations waterways and along adjacent to Along valleys and 20 m over many large trees thick canopy with by a characterized V forestLowland 2. egetation is Unlogged

habitation is evident pastevidence of human low temperatures, precipitationunder and this is frequently area M m.a.s.l usually above areas 500 A canopy < 15 m tall cate lichen Abundant moss and 3. forest long ridge tops,long ridge

oist substrate Unlogged Upland gorized by a gorized

. species and and

scale logging exposed areas are dry dry exposed are areas gardens intohave been turned remnants and areas that habitat hasThis forest have forestsLowland that species and new shrubs dominated by invasive usually undergrowth and thicktall trees canopy with very few L 4. forest the imp duetransformation to ack of aack of closed

Logged Lowland

undergone massundergone

acts of largeacts of , soils, in - forest lowlandFormally rows uniform homoge in relatively species established M undergrowth. sparse there was generally thick leaf litter and thick canopy cover, provided Teak 5.

ono-cultured ono-cultured Plantation forest

nous

5.3 Results

5.3.1 Unlogged Coastal Forest

5.3.1.1 Coastal plants A list of plants found in coastal forests revealed the dominant species to be canopy trees Calophyllum inophyllum, Barringtonia asiatica, and Rhus taitensis (Table 5.3). With shrubs such as Pandanus sp. were observed to provide important shelter for geckos.

Table 5.3 The dominant species of plants determined via photographic identification from the four floral groups found in unlogged coastal forests on Malaita in January 2012.

Floral Dominant Species group

Canopy Calophyllum inophyllum, Barringtonia asiatica, Ficus tinctoria, Alstonia spectabilis, Rhus taitensis, Scaevola taccada, Premna corymbosa, Rhus taitensis, Calophyllum vitiense, Cocos nucifera

Understory Erythroxylon ecarinatum, Medinilla rubescens, Inocarpus fagifer, Fagraea sp., Garcinia sp.

Shrub and Pandanus compressus, Nephrolepis sp., Crinum asiaticum, Piper sp., forest floor Pandanus sp., Acrostichum aureum, Discocalyx sp.

Epiphytes Asplenium nidus, Davalia solida, Dendrobium sp.

5.3.1.2 Coastal herpetofauna In the coastal forests a total of nine herpetofaunal species were encountered (Figure 5.1). The most abundant species found at night (transects) were Bufo marinus and Nactus multicarinatus. During the day (quadrats), the most common species was Emoia pseudocyanura.

3.5 Transects 3 Quadrats 2.5

1.5

0.5

Encounter rate rate per transect/quadrat Encounter 0 E. nigra E. R. kreffti R. P. guppyi P. P. weberi P. G. oceania G. B. marinus B. E. atrocostata N. multicarinatus N. E. pseudocyanura E. Species

Figure 5.1 Encounter rates of herpetofaunal species found in coastal forest

5.3.2 Unlogged Lowland Forest

5.3.2.1 Lowland plants A list of plants found in lowland forests revealed the dominant species to be canopy trees Vitex cofassus, Pometia pinnata, Canarium sp. and Ficus benjamina (Table 5.4). Plants that were observed to provide important shelter for herpetofauna include the canopy trees (P. pinnata, Canarium sp. and Ficus sp.), the epiphyte (Asplenium nidus), the shrub (Cominsia guppyi), the understory palms (Areca macrocalyx and Metroxylon salomonense) and the tree fern (Cyathea vittata).

5.3.2.2 Lowland herpetofauna In the lowland forests a total of 19 herpetofaunal species were encountered (Figure 5.2). The most abundant species found at night (transects) were Discodeles guppyi, Bufo marinus and Platymantis weberi, although D. guppyi were only encountered in transects in riparian forests beside streams. During the day (quadrats), the most common species were Emoia pseudocyanura and Sphenomorphus concinnatus. 70

Table 5.4 The dominant species of plants determined via photographic identification from the four floral groups found in unlogged lowland forests on Malaita in January 2012.

Floral Dominant Species group

Canopy Vitex cofassus, Pometia pinnata, Ficus benjamina, Ficus sp., Canarium sp., Gmelina moluccana

Understory Heterospathe minor, Areca macrocalyx, Calamus hollrungii, Metroxylon salomonense, Caryota rumphiana, Schizostachyum tessellatum, Heterospathe sp., Calamus hollrungii

Shrub and Selaginella rechingerii, Dennstaedtia sp., Elatostema sp., Cominsia forest floor guppyi, Cyathea vittata

Epiphytes Scindapsus salomoniensis, Pothos rumphii, Pothos hellwigii, Asplenium nidus

5 4.5 Transects 4 Quadrats 3.5 3 2.5 2 1.5 1 0.5 Encounter rate per transect/quadrat per rate transect/quadrat Encounter 0 E. nigra E. R. kreffti R. P. guppyi P. P. weberi P. D. guppyi S. bignelli C. zebrata G. oceania B. marinus B. C. guentheri C. S. solomonis S. P. solomonis P. B. vertebralis B. S. concinnatus S. E. cyanogaster C. salomonensis N. multicarinatus N. E. pseudocyanura E. Species

Figure 5.2 Encounter rates of herpetofaunal species found in lowland forest.

5.3.3 Unlogged Upland Forest

5.3.3.1 Upland plants A list of plants found in upland forests revealed the dominant species to be canopy trees Schefflera sp. and Calophyllum vitiense and understory plants Cyathea vittata (tree fern) and Nastus obtusus (bamboo) (Table 5.5). Plants that were observed to provide important shelter for herpetofauna include the epiphyte (Asplenium nidus) shrubs (Elatostema sp., Alpinia oceanica) and understory plants (C. vittata and Nastus obtusus).

Table 5.5 The dominant species of plants determined via photographic identification from the four floral groups found in unlogged upland forests on Malaita in January 2012.

Floral Dominant Species group

Canopy Schefflera sp., Trichospermum sp., Ficus variegata, Calophyllum vitiense, Neonauclea orientalis

Understory Cyathea vittata, Litsea sp., Areca macrocalyx, Garcinia sp., Nastus obtusus, Saurauia sp., Gulubia macrospadix

Shrub and Calanthe triplicate, Elatostema sp., Scindapsus sp., Goodyera sp., forest floor Pleomele angustifolia, Dennstaedtia sp., Cyathea vittata, Tmesipteris sp., Piper sp., Selaginella rechingeri, Leea indica, Crinum asiaticum, Pleomele angustifolia, Begonia sp., Alpinia oceanica, Freycinetia sp., Nephrolepis sp.

Epiphytes Asplenium nidus, Myrmecodia salo, Piper sp., Psychotria sp., Pothos sp., Coelogyne sp., Lycopodium phlegmarioides

5.3.3.2 Upland herpetofauna In the upland forests a total of 14 herpetofaunal species were encountered (Figure 5.3). The most abundant species found at night (transects) were Batrachylodes vertebralis and Ceratobatrachus guentheri. During the day (quadrats), the most common species was also C. guentheri, though all were juveniles.

8 Transects 7 Quadrats 6

Encounter rate per transect/quadrat per rate transect/quadrat Encounter 1

0 E. nigra E. S.cranei R. kreffti R. P. guppyi P. P. weberi P. S. bignelli B. marinus B. C. guentheri C. S. solomonis S. P. solomonis P. B. vertebralis B. S. concinnatus S. N. multicarinatus N. E. pseudocyanura E. Species

Figure 5.3 Encounter rates of herpetofaunal species found in upland forest

5.3.4 Logged Lowland Forest

5.3.4.1 Logged plants A list of plants found in upland forests revealed the dominant species to be invasive alien epiphytes, Merremia peltata and Mikania micrantha, undergrowth trees Timonius timon, Rhus taitensis and Ficus sp. (Table 5.6). There is an obvious lack of upper canopy, with the a few remnant species that included Vitex cofassus, Pometia pinnata, and Canarium sp.. Shrubs such as Alpinia oceanica were observed to provide important shelter for skinks.

5.3.4.2 Logged herpetofauna In logged forest a total of 15 herpetofaunal species were encountered (Figure 5.4). Logging usually occurs in lowland forests and therefore similarities in species composition were found between the two habitat types. The most abundant species

73 found along at night (transects) was Bufo marinus. During the day (quadrats), the most common species was Emoia pseudocyanura.

Table 5.6 The dominant species of plants determined via photographic identification from the four floral groups found in logged lowland forests on Malaita in January 2012.

Floral Dominant Species group

Canopy Vitex cofassus, Pometia pinnata, Canarium spp.

Understory Timonius timon, Rhus taitensis, Areca macrocalyx, Macaranga tanarius, Ficus copiosa, Ficus septica, Paraserianthes falcata

Shrub and Nephrolepis biserrata, Alpinia oceanica, Nephrolepis hirsutula, forest floor Spathoglottis plicata

Epiphytes Merremia peltata, Mikania micrantha

Transects 2.5 Quadrats 2

1.5

0.5

Encounter rate per transect/quadrat per rate transect/quadrat Encounter 0 E. nigra E. P. virens P. R. kreffti R. P. guppyi P. P. weberi P. D. guppyi G. oceania B. marinus B. C. guentheri C. B. vertebralis B. S. concinnatus S. E. cyanogaster C. salomonensis C. N. multicarinatus N. E. pseudocyanura E. Species

Figure 5.4 Encounter rates of herpetofaunal species found in logged forest.

5.3.5 Teak Plantation Forest

5.3.5.1 Teak plantation plants A list of plants found in upland forests revealed the dominant species to be the cultivated species Tectona grandis (Table 5.7). There are however, a few shade tolerant shrub and understory plants, which provide ground cover such as Nephrolepis, Ficus. and Piper sp. and Selaginella rechingeri. The thick leaf litter created by T. grandis was observed to provide an important shelter for skinks.

Table 5.7 The dominant species of plants determined via photographic identification from the four floral groups found in teak plantation forests on Malaita in January 2012.

Floral group Dominant Species

Canopy Tectona grandis

Understory Ficus septica, Ficus chrysochaete, Ficus variegata, Euodia triphylla, Ficus wassa, Tarrena sp.

Shrub and Nephrolepis biserrata, Selaginella rechingeri, Alpinia oceanica, forest floor Pteris sp., Dendrocnide salomonensis, Costus speciosus, Nephrolepis hirsutula, Cyrtosperma johnstonii

Epiphytes Piper betel, Piper sp.

5.3.5.2 Teak plantation herpetofauna In the teak plantation forest a total of 10 herpetofaunal species were encountered (Figure 5.5). The most abundant species found at night (transects) was Bufo marinus. During the day (quadrats) the most common species were Emoia pseudocyanura and Sphenomorphus concinnatus.

3.5 Transects 3 Quadrats 2.5

1.5

0.5 Encounter rate per transect/quadrat per rate transect/quadrat Encounter 0 E. nigra E. R. kreffti R. P. weberi P. C. zebrata G. oceania B. marinus B. S. solomonis S. S. concinnatus S. N. multicarinatus N. E. pseudocyanura E. Species

Figure 5.5 Encounter rates of herpetofaunal species found in teak forest.

5.3.6 Comparison of herpetofauna richness in the different habitat types

5.3.6.1 Nocturnal (Transects) A comparison of results based on the nocturnal visual encounter surveys’ (transects) for all habitats focusing on herpetofaunal species active at night shows there were obvious differences in species richness on transects between the different habitat types (KW test H = 28.010, df = 4, p < 0.05, Figure 5.6). At night (transects), lowland and upland forests show significantly higher herpetofauna species richness than coastal habitats. Lowland forests also had significantly higher species richness than teak forests.

Figure 5.6 Comparison of average herpetofauna species richness in the different habitat types based on nocturnal surveys (transects) conducted on August 2011 to April 2012, Error Bars: 95% Confidence Interval.

5.3.6.2 Diurnal (Quadrats) A comparison of results based on diurnal quadrat sampling (quadrats) for all forest habitats focusing of herpetofauna species active during the day shows there were no obvious differences in species richness in quadrats in the different habitat types (KW test H = 8.583, df = 4, p > 0.05, Figure 5.7). During the day (quadrats) all forest habitat types recorded similar values for species richness per quadrat, however species assemblages differed.

Figure 5.7 Comparison of average herpetofaunal species richness in the different habitat types based on diurnal surveys (quadrats) conducted on August 2011 to April 2012, Error Bars: 95% Confidence Interval.

5.3.7 Priority forest habitat based on herpetofauna species richness Looking at the average nocturnal herpetofaunal species richness per transect (Figure 5.6), lowland forest recorded the highest average value of 5.2 species. When looking at average diurnal herpetofaunal species richness per quadrat (Figure 5.7), teak forest recorded the highest average value of 1.9 species. Overall, lowland forest has the highest total herpetofaunal species richness value with 18 species recorded (Figure 5.8). Logged forest (15) is second followed by upland forest (14). In teak forest and coastal forest only 10 and 9 species were recorded, respectively. From these, only coastal forest, upland forest and logged forest recorded a single species that was not found in any other forest habitat type.

20 18 16 14 Skinks

12 Geckos 10 Frogs 8

Species richness richness Species 6 4 2 0 Coastal Lowland Upland Logged Teak Forest Habitat Type

Figure 5.8 Comparison of total combined nocturnal and diurnal herpetofaunal species richness.

When comparing nocturnal species abundances (transects), upland forest clearly has the greatest abundance per species value with an average of 1.8 individuals encountered per species per transect (Figure 5.9a). Comparing diurnal abundances (quadrats), teak forest displays the greatest abundance per species with an average of 1.4 individuals encountered per species per quadrat.

If we remove the introduced, invasive cane toad Bufo marinus (Figure 5.9b) from the comparisons to only include native fauna, there is a significant decrease for nocturnal (transect) results in the average species abundance in logged (0.7) and teak (0.4) forests and a slight decrease in coastal forests (0.9). This shows that the presence of B. marinus in degraded landscapes is important and can bias data representation.

a) Including b) Excluding Bufo marinus Bufo marinus

2 2 1.8 1.8 1.6 1.6 1.4 1.4 1.2 1.2 1 1 0.8 0.8 0.6 0.6 transect/quadrat 0.4 transect/quadrat 0.4 0.2 0.2 0 0 Average species abundance per per abundance species Average Average species abundance per per abundance species Average

Figure 5.9a & b Average abundances per transect/quadrat (nocturnal/transects=blue and diurnal/quadrats=red), both including and excluding the introduced, invasive Bufo marinus.

5.3.8 Impact of habitat degradation and modification It was estimated due to the location and surrounding habitat that the majority of logged and teak plantation forests were formerly lowland forests. Therefore, comparisons will be drawn between these three habitat types (unlogged lowland forest, logged lowland forest and teak plantation forest) to try to quantify the impact of degradation and modification on lowland forests solely based on differences in herpetofaunal abundance and richness (Figure 5.10). Unlogged lowland forests have the highest total number of species with 18 followed closely by logged lowland forests with 15 then teak forests with only 10 species.

20 18 16 14 12 10 8 6 Species Richness Richness Species 4 2 0 Unlogged Lowland Logged Lowland Teak Plantation (previously lowland) Forest Habitat Types

Figure 5.10 A comparison of total herpetofauna species richness in unlogged lowland, logged lowland and teak plantation forests (previously lowland).

There are differences in herpetofaunal species average encounter rates and presence in logged lowland forest communities in comparison to those of unlogged lowland forest communities. The difference in average encounter rates = logged lowland forest average encounter rates minus unlogged lowland forest average encounter rates (Table 5.8). Six species recorded higher encounter rates in logged forested habitats than in unlogged lowland forested habitats (positive encounter rate difference value). Eight species recorded lower encounter rates (negative encounter rate difference value) and four species were not encountered at all in logged forested habitats compared with unlogged lowland forested habitats. Therefore, with a net loss of four species and a change in abundance for all 14 other species it is reasonable to suggest that logging results in a change in the species composition of herpetofaunal species.

Table 5.8 Difference in average encounter rates and species presence in logged lowland forest compared with unlogged lowland forests, the difference in average encounter rates = logged lowland forest average encounter rates - unlogged lowland forest average encounter rates.

Species Difference in average encounter rates B. marinus + 4.8 E. pseudocyanura + 1.6 H. kreffti + 0.7 B. vertebralis + 0.4 E. nigra + 0.2 E. cyanogaster + 0.1 G. oceanica - 0.1 C. guentheri - 0.3 C. salomonensis - 0.4 N. multicarinatus - 0.5 S. concinnatus - 0.5 P. weberi - 0.8 P. guppyi - 1.0 D. guppyi - 4.5 P .solomonis Absent C .zebrata Absent S. solomonis Absent S. bignelli Absent

There are differences in herpetofaunal species average encounter rates and presence in teak plantation forest communities in comparison to those of unlogged lowland forest communities. The difference in average encounter rates = teak plantation forest average encounter rates minus unlogged lowland forest average encounter rates (Table 5.9). Six species recorded higher encounter rates in logged forested habitats than in unlogged lowland forested habitats (positive encounter rate difference value). One species of skink recorded no significant difference in abundance (zero encounter rate difference value). Two species recorded lower encounter rates (negative encounter rate difference value) and nine species were not encountered at all in teak forested habitats compared with lowland forested habitats. Therefore, with a net loss of nine species and a change in abundance for eight other species it is reasonable to suggest that replacing lowland forest with teak plantation forest results in a net change in the species composition of herpetofaunal species.

Table 5.9 Difference in average encounter rates and species presence in teak plantation forest compared with unlogged lowland forests, the difference in average encounter rates = teak plantation forest average encounter rates - unlogged lowland forest average encounter rates.

Species Difference in average encounter rates B. marinus + 5.6 E. pseudocyanura + 2.1 S. concinnatus + 1.7 E. nigra + 0.5 N. multicarinatus + 0.3 H. kreffti + 0.2 C .zebrata 0.0 G. oceanica - 0.2 P. weberi - 1.1 B. vertebralis Absent E. cyanogaster Absent C. guentheri Absent C. salomonensis Absent P. guppyi Absent D. guppyi Absent P .solomonis Absent S. solomonis Absent S. bignelli Absent

5.4 Discussion In the current study, coastal forest on Malaita was found to be generally species poor for herpetofauna this is possibly due the saline and associated physiological drought conditions to which coastal forest are adapted (pers. obs.). Most herpetofauna, especially frogs are saline intolerant and freshwater dependent and are therefore absent from coastal areas that are in close proximity to saltwater (Balinsky 1981, Pough et al. 1998). Thus, most coastal forests in this study were also either fragmented due to the establishment of coconut plantations or degraded due to human activities such as pig farming and timber extraction and the abundance of invasive species (eg. rats, cats and dogs) (pers. obs.). Degraded and fragmented forests show decreased species diversity and richness (Hillers et al. 2008), which is evident for herpetofauna in coastal forests on Malaita. Two species (B. marinus and N. multicarinatus) that seem to favour drier conditions and areas of high anthropogenic activity were in high abundances in coastal forests. 83

Lowland forest was generally rich in herpetofaunal richness, which may be due to a high diversity of microhabitats. As Ernst et al. (2006) and Hillers et al. (2008) found that greater microhabitat diversity of breeding sites, vegetation structure and leaf litter cover act as influential variables and best explain frog abundance and species diversity in many cases. For example, the current study only encountered D. guppyi besides clean, small, fast flowing streams, a micro-habitat that was absent in upland, coastal, teak and logged forests. Also in support C. salomonensis and C. zebrata two of the largest lizards of the forest were found on P. pinnata, Canarium sp. and Ficus sp., large trees that were rare or absent in coastal, upland, teak and logged forests.

Upland forest was also found to be generally high in herpetofaunal richness and this may be due to its relatively undisturbed state and unique climatic conditions (pers. obs.). The conditions of the upland forests are cool and moist (Mueller- Dombois and Fosberg 1998) and these parameters are preferred by frog species (Wells 2007, Kohler et al. 2011). In this study, the relatively undisturbed nature of upland forests also resulted in high abundances in this habitat type.

In the present study, logged forest was generally high in herpetofaunal richness, possibly due residual microhabitat diversity and the adaptability of some species to modified habitats. However, P. solomonis, C. zebrata, S. solomonis and S. bignelli, all species found in unlogged lowland forests were absent in logged lowland forests. This is supported by Ernst et al. (2006) and Barlow et al. (2007) who found that logged forests only contained 60% of primary forest species in relation to lizards and “leaf-litter” frogs. The difference in species composition, abundance and richness between unlogged lowland and logged lowland forests is significant in this study as found in other similar studies (Vonesh 2001, Ernst et al. 2006, Thinh et al. 2012).

The strong adaptability of certain herpetofaunal species to habitat disturbance and degradation may also results in high species richness for logged forests. Ficetola and De Bernardi (2004) discuss that generalist species which are particularly mobile are able to adapt and exploit disturbed environments. Generalist frog species such as Bufo fowleri (Green 2005) and Osteopilus septentrionalis (Meshaka 2005) are known

84 to benefit from human altered landscapes (Wells 2007). This seems evident in the case of the frog Bufo marinus on Malaita.

It is also important to note that specialist species are the ones that are most affected by logging activities (Thinh et al. 2012), so we would expect to see species of greater conservation concern strongly impacted. However, intermediate levels of disturbance can also lead to higher species richness with a high number of both pioneer and climax species (Connell 1978). Geckos for example seem to favour disturbed habitats that provide an abundance of artificial shelter and egg-laying sites (Ineich 2010). This is supported with the presence of all geckos in logged forest habitat on Malaita.

In the current study, teak plantation forests were generally poor in herpetofaunal richness and this is most likely due to its modified state and homogeneity. According to Kanowski et al. (2005) and Barlow et al. (2007) the uniformity of plantations results in low species richness due to the lack of micro-habitats for herpetofauna. Another factor is the modification of forests through plantations creates changes in canopy structure, leaf-litter environment and loss of microhabitats, all necessary for herpetofauna (Gardner et al. 2007). Therefore, establishment of plantations is expected to result in a loss of certain forest herpetofaunal species and changes in forest community assemblages (Hillers et al. 2008). This supports the results of this study which found that herpetofaunal species such as P. solomonis, P. guppyi, D. guppyi, B. vertebralis, C. guentheri, C. salomonensis, S. solomonis, E. cyanogaster and S. bignelli were absent in teak forests as compared to lowland forests. However, teak forests are not expected to be biologically dead and can have certain conservation value (Lindenmayer et al. 2003, Carnus et al. 2006, Bremer and Farley 2010), as found in the current study by the presence of an IUCN red-listed species (C. zebrata).

Deforestation and degradation are the primary causes of species extinctions worldwide (Morgan 1987, Brooks et al. 2002, Brook et al. 2003, Hanski et al. 2007). An indirect effect of this deforestation is the expansion and creation of degraded forests, secondary forests and exotic plantation forests (Gardner et al. 2007, Herrera- Montes and Brokaw 2010). The formation of these new forest habitat types does not

85 suit most amphibians as indicated by the absence or reduction of their presence (Bell and Donnelly 2006, Gardner et al. 2007) and as seen on Malaita. Both teak plantation and logged forests demonstrated overall species assemblage change for herpetofauna. Logged forests displayed a reduction in abundances for eight species and an apparent loss of four herpetofaunal species. Teak forests displayed a reduction in in abundances for two species but displayed an apparent loss of nine herpetofaunal species. Reasons for this may include an increase or decrease in predator-prey relationships, a decrease or increase in suitable microhabitats plus alterations in the ecological conditions of the forest such as in temperature and moisture regimes (Bell and Donnelly 2006, Cushman 2006, Hillers et al. 2008).

5.5 Summary In summary, Malaitan unlogged lowland forests were found to have the highest herpetofauna species richness but unlogged upland forests had the highest average species abundance. Coastal forests have relatively low herpetofauna richness and abundance. Lowland forests have high species richness and moderate species abundances. Upland forests have moderate species richness and high species abundances. Logged forests have moderate species richness and moderate species abundances. Teak forests have relatively low species richness but high species abundances. There was a significant difference in the richness between forest habitats types for results based on nocturnal (transect) sampling but not for diurnal (quadrat) sampling. B. marinus had a significant impact on the abundance of herpetofauna in degraded habitats. Logging and the formation of teak plantations have resulted in a net herpetofaunal loss of 3 species for logged forests and 8 species for teak forests out of a total of 18 species found in unlogged lowland forest.

The “healthiest” habitat type using herpetofauna richness as a bio-indicator is lowland forest as expected and based on herpetofauna abundance, upland forest also as expected. The decreased richness evident in the modified habitats of logged forest and teak plantation forest further signify the impact that habitat degradation has on biodiversity loss. It is therefore important from a biodiversity conversation perspective that the degradation of forest habitats be minimized.

CHAPTER 6: TRADITIONAL KNOWLWEDGE OF HERPETOFAUNAL BIODIVERSITY AND FORESTS IN ARE`ARE, MALAITA

6.1 Introduction Traditional knowledge (TK) provides a foundation for successful living in natural environments; and this knowledge with its beliefs and customs form the ‘glue’ that creates social cohesiveness and cultural identity (Bennet 2000, Dutfield 2006, Thaman et al. 2010, FAO 2011). In Melanesia TK and cultural practices have developed and evolved over time resulting in interactions and relationships with the environment that are based on qualitative, holistic, oral approaches (Merculieff 2000, Caillaud et al. 2004, Painemilla et al. 2010). TK is wisdom, knowledge and information learned through experience, passed on from generation to generation and used in decision making, planning and the management of biodiversity among other things that are critical and beneficial to life in subsistence communities (Merculieff 2000). To further highlight TK’s importance Article 8 in the CBD tells contracting parties to “respect, preserve and maintain” the traditional knowledge, practises and innovations of local indigenous communities (UN 1992a). The value of TK in modern societies cannot be overlooked as many of these practises and beliefs may hold the key to sustainability in the Pacific islands.

Thaman (2002) identifies the loss of traditional knowledge as a major threat to biodiversity itself and its preservation. He argues that, if the traditional names, uses and management systems of biodiversity are lost, the impetus for the conservation of these natural resources at a community level is also lost.

A main element of the focus of this thesis is to marry scientific and traditional information on the ecology, ethnobiology and conservation status of herpetofauna and forests on Malaita. This chapter will aim therefore carry out community-based ethnobiological studies to examine local perceptions, knowledge and cultural uses of herpetofauna and include perceptions of the conservation status of forests and associated herpetofauna. Questionnaires on the herpetofaunal species of Malaita (Appendix A) will then be discussed followed by perceived anthropogenic impacts

87 on and uses for the sampled forest habitat types. This chapter will therefore capture a glimpse of Malaitan TK and classification systems with the added purpose of documenting the learned observations that have occurred over generations on the island.

6.2 Specific Methodology Questionnaire surveys followed methods described previously in chapter 3. Aspects of traditional knowledge for which information was being sought included: the classification of frogs and lizards and the related uses associated with these animals and their different forest habitat types. In this context, focus is placed on the perceptions and knowledge that local people have regarding, skinks and geckos, their conservation status, forest habitat preferences, and conservation. The questionnaires were undertaken with the use of a local interpreter as many words and terms were unfamiliar to the primary researcher.

All informants belonged to the Are’Are dialect and all ten villages (Uwaisiwa, Swit point, Nahu, Tawaimare, Kopo, Mananawai, Komhauru, Tawaihuro, Hunanapuru and Ohanimeno) to which the informants belonged were found in the Tai ward. Informants were taken to quiet locations to be interviewed and interviews lasted between half an hour to an hour depending of amount of information shared by the informant. Thirty questionnaires were conducted to cover a sufficient number of age groups and villages but to allow completion within the study timeframe. The informants were selected across all age groups. The youngest informant was 16 years of age and the oldest was 99 years of age. Most age groups had one or two informants with the age group of 80 to 84 having the highest number of informants at five.

The questionnaire had two parts; in part 1 questions were formulated to specifically obtain information on herpetofaunal names, associated uses and perceived abundance. Part 1 questions were stated as:

What are the most important different frog and lizard species that you know? What are their names; where are they found; what is their abundance; and have they declined or increased in abundance and reasons for change in

abundance? What are their associated uses or other stories, tales or information on them?

Part 2 questions identified the important uses of the forests, the current status of these uses and the perceived impact on local herpetofauna. Only 21 informants answered part 2 questions as 9 of the older informants were not able to discuss the “current” uses and trends regarding forests due to their age. It is important to note that the questions under part two of the questionnaire were aimed at gaining a general overview of uses in coastal, upland, lowland, logged and plantation forests and were not an exhaustive description of specific uses. Forest threat values (FTV) were also calculated based on the sum of uses and the impact level of these said uses on the forests. The four threat impact levels were: 1) Destroys habitat, 2) degrades habitat, 3) disturbs habitat and 4) little or no impact To quantitatively compare between forest habitat types a simple formula (FTV = sum of (uses x use threat/impact level)) was created to estimate which habitat type is under the greatest stress or threat from humans. Part 2 questions were stated as:

What are up to 5 main uses associated with coastal, lowland, upland, logged and plantation forests? Have there been changes on this use and how do (if so) these impact frogs and lizards?

6.3 Results All information stated in this section was collected from the questionnaires and have been summarised according to species and forest type. Section 1 presents the results for herpetofauna (frogs then the lizards). Section 2 then presents results on forest types and their associated uses with regards to change in intensity of and perceived impacts on herpetofauna. Section 3 will then describe a short summary of the age and gender patterns of informants. Probable scientific classifications were based on descriptions of species by local informants and matched to species keys.

6.3.1 Herpetofauna

6.3.1.1 Frogs The surveys showed that there is rich and diverse local knowledge with different local names for taxonomically similar frog species in different sites (Table 6.1). A

89 total of 148 frogs were mentioned by informants with a total of 31 distinct names summarized or grouped into 9 described or identifiable species (Table 6.1).

Table 6.1 Vernacular and likely scientific nomenclature of frogs based on questionnaire surveys of people from Uwaisiwa, Swit point, Nahu, Tawaimare, Kopo, Mananawai, Komhauru, Tawaihuro, Hunanapuru and Ohanimeno villages in Tai Ward Malaita.

Most common Other names Frequency of Probable scientific local also known as times mentioned classification based on vernacular by informants Pikacha et al. (2008). See frog names x/30 also list in Appendix B Pari 26 Discodeles guppyi Hahaia haha’a, koe, kii 20 Batrachylodes vertebralis Oripasu, tarapasu, ma 19 Ceratobatrachus guentheri pau-pasu, pe’u, oripasu papa, tara iki Otohao otokao, 19 Platymantis guppyi oripapa Pina-iki iki-iki 17 Hylarana kreffti Puroko, ori niaoke 16 Bufo marinus Ten ten, 16 Hylarana kreffti Kori-niu 16 Platymantis weberi Ka`a-ka`a ka'a kaka, koen 6 Platymantis solomonis mako, ori Taramena koe rahuta, 5 Litoria thesaurensis nonoto, koe memea

Pari (Discodeles guppyi) is a well-known frog. It is the largest native frog found in the forest, having a dark smooth backside and yellowish underside. Fully grown it is said to resemble a new-born child and has very long legs giving it the ability to

90 jump very far. It is said to make many distinct sounds including a very loud whistling sound, a barking sound and a sound like the cry of a new-born child.

According to informants, it is common to rare depending on location and is found close to fresh water systems in inland lowland forests. It can be found along valleys, beside streams on rocks at night, in thick forested areas and at headwater systems. It is also able to dive and stay under water for long periods of time.

This species (pari) was an important protein component in bush diets. It was caught and eaten during feasts, usually cooked by roasting and is said to taste like freshwater fish. It is usually hunted when it is raining by listening for its call, and the month of March is said to be the best month to catch this frog. Informants believe that if a person snaps a twig when encountering this frog, it will render the frog immobile as it will think that one of its legs has just been broken. A whistle made from a stick can also be used to find the frog, which responds to the sound made by the whistle. It is also caught and eaten by dogs and cats.

This species is also a totem to certain tribes and believed to bring fertility to gardens if found in them (Table 6.2). The consumption of this frog by these tribes is prohibited. The bones of this frog were also used medicinally to rub against the body of children to prevent snake and centipede bites.

Hahaia (Batrachylodes vertebralis) also known as haha`a, koe or kii is a small dark coloured to yellowish frog with red, white and yellow stripes along the back, with smooth skin and long small legs. It has two distinct calls, a beeping sound and a soft haa haa sound. It is said to be found in valleys, forests, creeks and swamps, on trees and on ra leaves and wet areas, in particular upland forests, but is not found on the ground. It is also known to form aggregations around forest pools in the dead logs after big rains and is said to lay eggs in bubbles on trees. The best time to encounter this frog is between 8pm and 12 midnight and it is also known to urinate when jumping.

This frog was also eaten and can be cooked in bamboo; traditionally, it is an important food for feasts and when it is presented at a feast, an auapu (important woman) would eat it to signify that feasting can begin. It is also protected with

91 seasonal taboos for harvesting and was such an important food that it could be bought with pata-ni-hanua (traditional shell money). Folklore regarding the frog is that if its legs are broken when caught a tree will fall on you in the forest (Table 6.2).

Oripasu (Ceratobatrachus guentheri) also known as tarapasu, ma pau-pasu, pe`u, oripasu papa and tara-iki is also well-known. This frog is a very distinct medium-sized species that resembles a leaf, having a “sharp” nose and “sharp” eyelids. It has rough, camouflaged skin of many colours and patterns, with the underside usually a paler colour.

“Oripasu” is said to be common to rare depending on habitat type, with greater abundance in undisturbed upland forests. It is normally found on the ground, amongst leaf litter in forests and also inside deep holes. It prefers cooler habitats such as the upland forests and valleys and lays its eggs in damp places on the ground. If encountered the startled frog will flatten its body against the ground instead of jumping away and, if handled, will usually inflate its abdomen.

This frog is roasted after being gutted and was an important food for feasts (Table 6.2). Its bones are known to be used for mato’oha (sorcery) to bring luck and to increase garden fertility. This frog is also used as a medicine for opa-opo (swollen stomach) and for bedwetting in children by rubbing against the child’s stomach. Its urine is also drunk to heal stomach illnesses and its saliva can be used to treat snake and centipede bites. In a traditional historical story this frog was responsible for protecting an area in the mountains from being destroyed by black magic, the evidence is the presence of a distinctive hill, known as Hurakaia, which still juts up from the middle of the forest.

Otohao (Platymantis guppyi) also known as otokao and oripapa has an elongated body with smooth skin, its back varies in colour from red, brown to yellow, white and is also said to have camouflaged patterns. This frog is said to jump high and long and its abdomen will expand when handled. It is said to be a common arboreal species and is found in inland forests and swampy, wet areas, on common plants such as rao (Metroxylon salomonense), ra (Cominsia guppyi), papareo (Asplenium nidus) and kakake (Cyrtosperma chamissonis).

This frog was eaten and was roasted in bamboo and was also sold and used for trade and exchange (Table 6.2). It was commonly hunted using traps made in ra leaves. March is said to be an important month for catching these frogs and this frog is also preyed upon by snakes. It is better known for its medicinal use in preventing bed-wetting in children, which is done by rubbing the frog on the child’s stomach or making the frog to pee on the child’s head. This frog is also regarded as a koe maea (tabu frog) and is a totem to certain tribes with its call believed to signal death if found calling near a house.

Pina-iki (Rana kreffti) also known as iki-iki is well known by locals. It has a smooth slimy body, striped with black brown on its back, white-yellow on its underside and has very long legs. It makes a sharp “iki” call and is therefore named as such. It is a common species found in forested areas close to creeks, pools of water and swampy, muddy areas. It is said to be found around houses and is particularly abundant close to pig feeding sites.

This frog was also eaten but some tribes are not allowed to eat them as it is a totem and can signal death or sickness if it is heard calling or found in the house (Table 6.2). It was also used to determine the thoughts and feelings of ancestral spirits.

Puroko (Bufo marinus) or ori-niaoke is the introduced cane toad. This is a relatively large, “ugly” frog with rough skin and a warty appearance. It is said to be found everywhere, in all habitats, especially along drains, water pools and in gardens. It is thought to be increasing in numbers and can be very abundant in coastal areas where human modification is evident.

According to an informant, “Puroko” was introduced and deliberately spread throughout the island in the 1940s by a Commissioner Bell and Chief Alick Nonohimae, primarily to eat insects and kill snakes such as the poisonous ma- ara`ara (Salomonelaps par). It has poison in its skin which kills snakes. Locally, around the 1950s a man named Patere Wate introduced it to Rohinari village and to further show its perceived importance at the time, three men were fined for accidently killing one at Wairokai village.

Informants believe that this frog eats termites in houses and also eats human faeces. It is also said that the Chinese eat this species. Folklore regarding this species from Nahu village is that it used to have teeth but they were stolen by a shark (Table 6.2).

Ten-ten (Rana kreffti) is also a well-known frog. Two informants said that this frog resembles the Pina-iki (previous species) but is smaller and the author believes that both are the same species but ten-ten refers to the smaller vocal males of the species. This frog is a small to medium-sized, dark brown frog with black stripes on its back, yellow whitish under parts and a thin abdomen. It is said to be a relatively common but secretive species and found on the ground close to pools, creeks, still bodies of water and muddy areas.

There are no reported uses for adults of this species, but the juveniles are used as fishing bait for catching eels (Table 6.2).

Kori-niu (Platymantis weberi) is a small, very vocal frog named after its distinctive loud call, sounding like “körii.” It has a dark brown back with rough skin, a pale coloured underside and an elongated body.

This species is relatively common and is said to be found on the ground in forests, under dead logs, along waterways and around houses, it is also common in muddy areas and around pig feeding areas, but prefers upland forests and valleys. This frog was also eaten (Table 6.2).

Kaà kaà (Platymantis solomonis) also known as kaà kaka, koen mako and ori is a large frog with long legs and a long jump, but not as big as the pari (D. guppyi) and has a dark reddish colour. It is relatively uncommon and is said to be found under stones in inland forested areas close to water such as near streams and rivers and has a very loud call sounding like its name. It was also eaten and its meat is said to have a greasy texture (Table 6.2).

Taramena (Litoria thesaurensis) also known as koe rahuta, nonoto and koe memea is a small usually yellow pale frog, but can have shades of brown, green, yellow and white with smooth skin that is relatively uncommon. It has big eyes and round pads on its feet to help grip onto leaves. It is said to be predominately arboreal 94 and not too common but can be found on leaves such as the ra (Cominsia guppyi) and papareo (Asplenium nidus), It is found in coastal areas on rocks and is also found on large leaves such as those of kakake (Cyrtosperma chamissonis) and around freshwater pools. It is preyed upon by cats and snakes and is also eaten by humans (Table 6.2). This species is used to rub against a child’s stomach to help prevent bedwetting.

Table 6.2 Summarised associated uses of different frog species as described by informants

USES Food Trade Totem Medicinal Sorcery Folklore Fishing

Local frog names Pari X X X Hahaia X X X Oripasu X X X X X Otohao X X X X Pina-iki X X X Puroko X Ten-ten X Kori-niu X Ka`a-ka`a X Taramena X X

The following descriptions briefly define the uses stated above;

“Food” means that this particular species was eaten by humans. With reference to certain frog species as food the past tense is used as all respondents claim to no longer be eating these animals, mainly because most settlements are located in coastal areas now away from the main frog populations. Frogs were usually cooked by two methods either by direct roasting on hot coals or steamed in bamboo. 95

“Trade” is the use of frogs (usually cooked) as an item for barter and exchange, therefore giving the frog monetary value. Many frog species are “Totem” animals to certain tribes, informants always said other tribes and did not reveal the names of the tribes. The term koe maea when referring to these totem species gives certain powers that lead to reverence for the particular species. These frogs act as symbols to local villagers often with a negative connotation such as foreseeing death for a certain family or household. However, some of these species can also be viewed in a positive sense such as the promise of fertility to gardens. Species that are totems would not normally be eaten if the species was the totem for your tribe and therefore would be protected and revered by the tribe’s people. “Medicinal” use basically refers to the application of the frog or its parts externally to the sick person to heal them. Common ailments cured with frogs are of the abdominal or stomach area and these are still being practised in some parts of Are`Are. “Sorcery” refers to the single application of a certain frog’s bones for the use as good luck charms or to aid in the increase of garden fertility. “Folklore” refers to certain cultural stories or myths that are associated with a particular species. “Fishing” refers to the use of a frog specimen for the act of fishing.

6.3.1.2 Lizards The surveys show rich local knowledge that is diverse even in a localised area with different names for the same scientific species and possibly for different growth phases, colour forms or sexes. A total of 179 lizard mentions were provided by informants, with a total of 27 distinct names summarized into 12 described species. The frequency of times mentioned by informants for these 12 species varied and their probable scientific classifications were estimated (Table 6.3).

Ikiko asi (Emoia pseudocyanura) also known as iikiko niapa and iikiko ha`arirato is a small, smooth, light coloured skink with two dark lateral stripes along its body and a long greenish-blue tail. It is common and found everywhere, along paths, on tree trunks in the forest, around homes and enjoys basking in direct 96 sunlight. It is opportunistically hunted and can be roasted and eaten and is also used as fishing bait (Table 6.4). Ikiko asi is said to lay 1-2 eggs at a time and is also medicinally used by young boys to rub against their faces to prevent facial hair growth.

Table 6.3 Vernacular and likely scientific nomenclature of lizards based on surveys of people from Uwaisiwa, Swit point, Nahu, Tawaimare, Kopo, Mananawai, Komhauru, Tawaihuro, Hunanapuru and Ohanimeno villages in Tai Ward Malaita.

Most common Other names Frequency of Probable scientific local vernacular also known times mentioned classification based on lizard names as by informants x/30 McCoy (2006) Ikiko asi iikiko niapa, 29 Emoia pseudocyanura iikiko ha’arirato Unu 25 Corucia zebrata Paru paru 24 Emoia nigra Kuma kuma ni-iira, 21 Gehyra oceanica kuma nima’asu Rarani rarahuto, 17 Cyrtodactylus iikiko raran salomonensis Oru oru 17 Eugongylus albofasciolatus Kuma-ni-nima 17 Hemidactylus frenatus Iikiko ota iko warawa 14 Prasinohaema virens Iikiko mamatoru, iko wapu 13 Sphenomorphus bignelli Iikiko haho iikiko niasi 6 Emoia atrocostata Iikiko puru 5 Sphenomorphus concinnatus Iko ma 5 Emoia cyanogaster

Unu (Corucia zebrata) is the largest skink in the forest and can grow to be as large as an adult’s forearm. It has large eyes and a very long tail which can be used when climbing. It has a shiny body with visible scales and is usually greenish but can vary in colour and marking patterns which are usually well camouflaged to match its surroundings. The unu has very sharp teeth that can give a painful bite and which are said to protrude out the sides of the mouth when mature. It is a comparatively slow moving creature and is common to rare depending on forest habitat and is usually found in tree tops, tree hollows, trees with dense epiphytic growth on trunks and also in trees close to water ways, favoured plant species include large banyans (Ficus spp.) and Vitex cofassus trees.

Unu is found in cohabitation with both the native opossum (Phalenges orientalis) which is regarded as its enemy, and the native lizard rarani (C. salomonensis) which the unu is said to play tricks on. This animal is hunted and eaten and is said to have tasty greasy meat (Table 6.4). This was an important source of protein in the past however is becoming rare and was also used in sacrifices for ancestral worship.

Paru-paru (Emoia nigra) is a large skink with an all-black to brown back that has a reddish shine with a faded striped pattern; the under parts are a pale yellow. It is common along footpaths, on dead logs, rubbish heaps and tree trunks in forested areas and also coconut plantations. The paru-paru enjoys basking in direct sunlight and is said to lay 1-2 eggs at a time. This species is preyed upon by cats and dogs and also used as bait for fishing (Table 6.4).

Kuma (Gehyra oceanica) also known as kuma ni-iira or kuma ni-maasu is a widespread gecko and is well known. This species is a medium-sized, coloured white to light brown with dark specks and markings and has a fat body and big eyes with some specimens having a split tail. It is commonly found in lowland and upland forests, in tree hollows and trees such as the coconut and betel nut palms and also inside kakake leaves. Kuma is usually found in pairs. It is also found in an around homes and is known to eat moths and other insects inside houses. If handled the species has the ability to shed its skin as a defensive mechanism. This gecko can be eaten and is also used as a good-luck charm (sorcery) in gambling (Table 6.4).

Rarani (Cyrtodactylus salomonensis) also known as the rarahuto or iikiko raran. This is a very large lizard (but not as big as the unu) with big protruding eyes and rough dry skin that can be shed upon contact. It is predominately brown in colour with a striped pattern likened to an army camouflage design. It is most commonly found at night in lowland forests on trees and palms such as the sago palm, tree hollows and around rotting wood though it is rarely encountered. Its most active times are said to be between 11pm and 1am. The population of this species is thought to be declining and clearing of land and logging are believed to be predominant causes. It can also be eaten and was an important animal food for feasts (Table 6.4). Folklore regarding this species is that disobedient children will often be frightened by parents, saying that their eyes will turn into the eyes of a rarani if disobedient. Currently this lizard is valued in Honiara for around SBD$500 for the exotic pet trade and a few men have devised traps to catch this lizard. Some men also claim that this lizard has the ability to find gold.

Oru oru (Eugongylus albofasciolatus), named because of the “öru öru öru” sound that it makes, it is a very large dark coloured ground skink that has lighter orange coloured patterned stripes across its back as well as visible scales. It is rare, but found in caves, holes, under dead logs, rocks and rotting rubbish piles in forested areas. This nocturnally active species is very cryptic and escapes quickly making it hard to catch. It has sharp teeth and a painful bite which is poisonous and can, reportedly, be fatal. It is also a totem for some tribes signalling death if encountered (Table 6.4). Certain evil spirits are believed to take the form of this lizard, causing childbirth difficulties and insanity in victims. It can also be eaten and is usually cooked in bamboo.

Kuma-ninima (Hemidactylus frenatus) is a common small introduced whitish, light-coloured gecko found only in and around houses, especially close to light sources where it can be seen chasing insects. It was reportedly brought over from Guadalcanal by ancestors living in Marau and is believed to be good for the home by keeping insect numbers down.

Iikiko ota (Prasinohaema virens) also known as iko warawa is a small green skink, which is common on and around palms, such as the betel nut and in bamboo

99 thickets. It is difficult to catch and can be used for fishing bait and also be eaten (Table 6.4). Some specimens are said to have split tails.

Iikiko mamatoru (Sphenomorphus bignelli) also known as iko wapu is a small, smooth bodied skink that is dark coloured with a reddish sheen. It is found in inland forests, on the ground, amongst leaf litter, under rocks and dead logs, rubbish piles and is quiet commonly seen by people when digging mounds when gardening. It is used for fishing bait (Table 6.4) and is also known to be preyed upon by snakes.

Table 6.4 Summarised associated uses of different lizard species as described by informants

Uses Food Trade Totem Medicinal Sorcery Folklore Fishing

Iikiko asi X X X Unu X X X Paru-paru X Kuma X X Rarani X X X Oru-oru X X X Kuma-ni-nima Iikiko ota X X Iikiko mamatoru X Iikiko haho X X X Iikiko puru Iko ma X X

Iikiko haho (Emoia atrocostata) also known as iikiko niasi is a skink with a greenish grey body with many thin dark stripes across its back. It has a long tail and is a commonly seen in coastal areas on tree trunks, around houses, and especially on rocks in the inter-tidal zone. It enjoys basking in the sun, is also used as fishing bait (Table 6.4) and is preyed upon by cats. It is also a totem and tabu animal for certain 100 tribes and was used in traditional sacrifices, nowadays traditional sacrifices are no longer practised.

Iikiko puru (Sphenomorphus concinnatus) is a small to medium sized lizard with black to brown shiny skin and a striped pattern on its back with lighter under parts. This species is fairly common and can be found on walls of houses, in the forests along footpaths, in the grass and on tree trunks in the mornings. It is also preyed upon by cats. No mention of human use was provided by informants.

Iko ma (Emoia cyanogaster) is a medium to large sized, yellow- green lizard that is found climbing along trees and on dead logs in forested areas. It is uncommon and preyed upon by cats, birds and snakes and can also be eaten by humans (Table 6.4). Some have double tipped tails and its bones are also used as good-luck charms (sorcery) in gambling.

6.3.2 Forests The different uses associated with forests by local custodians do not only influence the physical nature of forests but also strengthens the perceived cultural value of these rich ecosystems. These custodians have authority over the forests and its inhabitants, authority which in many cases has been abused. An understanding of the relationship that local people have with their forests will help in any planning or prioritisation for any conservation activity. General uses associated with different habitat types have been compared simply to observe which habitats are most “useful” to locals. The most “useful” forests can also be regarded as the most threatened and under greater human related “stress”.

6.3.2.1 Coastal forests A total of 11 different uses were described by informants for coastal forests (Table 6.5). All of these uses are said to be currently increasing. Coastal plants useful to the local people include C. inophyllum used for timber, R. taitensis used for firewood and Pandanus sp. used for making mats and other traditional items. Overall this forest type is experiencing an increase in human activity with increasing impact on local herpetofauna and can therefore be classed as the forest habitat type under the greatest threat from anthropogenic impacts.

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Table 6.5 Coastal forest uses by locals, changes and perceived impact on herpetofauna, listed in order of number of mentions.

Perceived impact on x/21 No. Human uses Changes herpetofauna/forest 1 Food gathering Increase No impact 10 Harvesting building 9 2 Increase Disturbs habitat materials 3 Recreation Increase No impact 7 4 Feeding pigs (fenced) Increase Disturbs habitat 6 5 Timber extraction Increase Destroys habitat 5 6 Creating plantations Increase Destroys habitat 4 Collecting bush 4 7 Increase Disturbs habitat materials eg. firewood Destroys habitat and may 2 8 Creating settlements Increase accidently kill some species. Materials for 1 9 Increase Disturbs habitat traditional items 10 Hunting Increase Disturbs habitat 1 11 Gardening Increase Degrades habitat 1

6.3.2.2 Lowland forests A total of 13 different uses were described by informants for lowland forests (Table 6.6). Most of these uses are said to be currently increasing apart from traditional worship and other cultural related activities such as traditional burial sites and the harvesting of materials for cultural items. Hunting is listed as ‘no change’ by 16 out of 21 informants and this is due to the limited number of pigs found in this forest type, which is the primary target for hunting activities although possums, pigeons, bats and lizards are also caught. Lowland plants useful to the local people include V. cofassus and P. pinnata used for timber, M. salomonense and A. macrocalyx used for thatching, flooring, walling, and C. vittata as food. Overall like the coastal forest this forest type is experiencing an increase in human activity with increasing impact on local biodiversity including herpetofauna. 102

Table 6.6 Lowland forest uses by locals, changes and perceived impact on herpetofauna, listed in order of number of mentions.

Perceived impact on No. Human uses Changes x/21 herpetofauna/forest 1 Gardening Increase Degrades habitat and may accidently 13 kill some species. 2 Harvesting Increase Disturbs habitat 10 building materials 3 Feeding pigs Increase Disturbs habitat 9 (fenced) 4 Creating Increase Destroys habitat 8 settlements 5 Creating Increase Destroys habitat 5 plantations 6 Timber extraction Increase Destroys habitat 5 7 Food gathering Increase Disturbs habitat 5 8 Hunting No Disturbs habitat and some species may 4 change be targeted such as the Unu. 9 Traditional Decrease No impact but some species may be 3 worship used as sacrifices 10 Materials for No Disturbs habitat 1 traditional items change 11 Collecting of Increase Disturbs habitat 1 ornamental plants 12 Burial sites Decrease Disturbs habitat 1 13 Collecting water Increase No impact 1

6.3.2.3 Upland forests A total of 11 different uses were described by informants for upland forests (Table 6.7). Most of these uses are said to be currently decreasing in status apart from harvesting for building materials, settlement expansion and surveying of land

103 which is increasing. Overall this forest type is experiencing a decrease in human activity that results in a lower impact on local biodiversity including herpetofauna.

Table 6.7 Upland forest uses by locals, changes and perceived impact on herpetofauna, listed in order of number of mentions.

Perceived impact on x/21 No. Human uses Changes herpetofauna/forest No impact but some species may be 16 1 Hunting Decrease targeted such as the Unu. Harvesting building 14 2 Increase Disturbs habitat materials Feeding pigs 7 3 Decrease Disturbs habitat (unfenced) Creating 7 4 settlements Decrease Disturbs habitat expansion Destroys habitat and may accidently 5 5 Gardening Decrease kill some species. No impact but some species may be 5 6 Traditional worship Decrease used as sacrifices 7 Food gathering Decrease No impact; some species targeted 3 Surveying of tribal 2 8 Increase No impact land No impact; may create habitat for 1 9 Burial sites Decrease some species 10 Canoe building Decrease Disturbs habitat 1 11 Recreation Decrease No impact 1

6.3.2.4 Logged forests A total of 8 different uses were described by informants for logged forests (Table 6.8). Most of these uses are said to be currently increasing, particularly gardening. Food gathering, harvesting of building materials and hunting remain without change,

104 this is due to a limited supply of wild foods, building materials and pigs in this modified habitat type. Overall as with the previously described forests, this forest type is experiencing an increase in human activity (mostly gardening and plantation planting) with an increasing impact on local biodiversity including herpetofauna.

Table 6.8 Logged forest uses by locals, changes and perceived impact on herpetofauna, listed in order of number of mentions.

No. Human uses Changes Perceived impact on x/21 herpetofauna/forest 1 Gardening Increase Destroys habitat and may accidently 18 kill some species. 2 Creating plantations Increase No impact 7 3 Harvesting building No Disturbs habitat 3 materials change 4 Collecting bush Increase Disturbs habitat 3 materials, eg. firewood 5 Food gathering No No impact 3 change 6 Materials for Decrease Disturbs habitat 2 traditional items 7 Hunting No No impact but some species may be 1 change targeted such as the Unu. 8 Creating settlements Increase Disturbs habitat 1

6.3.2.5 Plantation forests A total of four different uses were described by informants for plantation forests (Table 6.9). This is the forest type with the least amount of human uses. All of these uses are currently increasing as most are directly linked to the plantation itself such as the maintenance and harvesting of the plantation. Overall this forest type is experiencing an increase in human activity with an increasing impact on local biodiversity and herpetofauna.

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Table 6.9 Plantation forest uses by locals, changes and perceived impact on herpetofauna, listed in order of number of mentions.

Perceived impact on x/21 No. Human uses Changes herpetofauna/forest Maintenance of Disturbs habitat and may 16 1 Increase Plantation accidently kill some species. Collecting and 5 2 harvesting plantation Increase Disturbs and degrades habitat crop 3 Recreation Increase No impact 3 4 Feeding pigs (fenced) Increase Disturbs habitat 2

6.3.2.6 Forest threat value (FTV) As stated by the informants the associated uses described by them for the five different forest habitat types can be divided into four threat levels namely: 1) destroys habitat, 2) degrades habitat, 3) disturbs habitat and 4) has little or no impact (Table 6.10). The uses of greatest concern are those that destroy habitat, are increasing and are also of a modern, commercial, unsustainable nature, specifically the creation of plantations and the extraction of timber. Uses that are present and/or increasing are calculated, with a level 1 impact having a value of 3, level 2 impacts having a value of 2, level 3 impacts having a value of 1 and level 4 impacts having a value of 0. Modern, commercial and unsustainable uses will have their value doubled to signify impacts. Based on the overall calculated forest threat value, coastal forests (46) have the highest value followed closely by lowland forests (42), third are logged forests (28) then upland forests (15) and plantation forests (10) have the lowest forest threat value.

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Table 6.10 Forest threat values calculated from uses described by informants. P = present and I = increasing. Threat level values are: Level 1 = 3, level 2 = 2, level 3 = 1 and level 4 = 0. Shaded uses reflect modern commercial unsustainable practises and will be multiplied by 2 to signify threat impact.

Coastal Lowland Upland Logged Plantation Uses P I P I P I P I P I 1) Destroys habitat (value = 3) Creating plantations (x2) 6 6 6 6 6 6 Timber extraction (x2) 6 6 6 6 Creating new settlements 3 3 3 3 3 3 3 2) Degrades habitat (value = 2) Canoe building 2 Gardening 2 2 2 2 2 2 2 3) Disturbs habitat (value = 1) Harvesting building materials 1 1 1 1 1 Materials for traditional 1 1 1 1 items Food gathering 1 1 1 1 1 1 Surveying of tribal land 1 1 Burial sites 1 Feeding pigs 1 1 1 1 1 1 1 Hunting 1 1 1 1 1 Collection of ornamental 1 1 plants Collecting bush materials eg. 1 1 1 1 firewood Harvesting plantation crop 2 2 (x2) Plantation work (x2) 2 2 4) Little or no impact on habitat (value = 0) Water collection 0 0

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Traditional worship 0 0 Recreation 0 0 0 0 Collection of medicine Forest threat value Values 46 42 15 28 10

6.3.3 Informants knowledge of frogs and lizards by age and gender The average number of frogs and lizards described varies across different age and gender classes (Figure 6.1). The group with the greatest knowledge of frogs was both the males and females over the age of sixty, both with an average of 6.4 species of frog mentioned. The group with the highest knowledge regarding lizards was the males below the age of thirty, with an average of 7.4 species of lizards mentioned.

7 Frogs 6 Lizards 5

3 No. of Species of Species No. 2

1 4.4 7.4 5.8 6.6 6.4 7 3.6 4.6 5 6.3 6.4 6.4 0 <30 30-60 >60 <30 30-60 >60 Male Female Respondant's Age & Sex Groups

Figure 6.1 Graph of informant’s age and gender against average number of frogs and lizards described.

General trends indicate that males are more knowledgeable than females regarding information on herpetofauna with males having higher averages for all age classes except above sixty where they are even. Male informants had a combined average knowledge of 12.5 frogs and lizards per questionnaire whereas females 108 averaged 10.8 frogs and lizards per questionnaire. The data therefore indicates that older the informant the greater the amount of information provided regarding herpetofauna and that males above the age of sixty have the richest traditional knowledge regarding herpetofauna. Informants ‘below thirty’ averaged 10 species descriptions per questionnaire, informants aged ‘thirty to sixty’ averaged 12 species descriptions per questionnaire and the informants ‘above sixty’ averaged 13 species descriptions per questionnaire.

6.4 Discussion

6.4.1 Traditional knowledge of herpetofauna A total of 58 distinct herpetofaunal names were recorded from informants and these were placed into 21 distinguishable species (Appendix B). Associated with the variety of names were seven categories of traditional use, which is similar to a study by Lohani (2011) in Nepal that found six categories of traditional uses for 49 animals, including three frogs. There was considerable overlap with Lohani (2011) with regards to use categories, however Lohani (2011) also mentioned the use of animals for weather forecasting but did not mention the use of animals in fishing and trade that were mentioned in the current study. Globally reptiles have been identified to be traditionally important for medicinal uses (Alves et al. 2008) as recorded for five species in this study. However there is a lack of published literature on traditional knowledge in relation to herpetofauna, which further adds to the importance of the information collected in this current study.

6.4.2 Threatened forest habitats Due to Christianity, education and a desire for participation in a cash economy people are known to have moved from upland areas to the coast of Malaita (Keesing 1967). This is evident in the abandoned stone wall remnants of settlements located in upland forests (pers. obs). As found in this study, upland forests have a low level of human associated threat, due to the distance from the majority of human settlement areas. The relocation of settlements in coastal areas has also led to an increase in the access of locals to the coastal and lowland forest habitats (Keesing 1967). Both forest types have a high record of use by locals which has resulted in a high level of human associated threats for both forest habitat types. 109

6.4.3 Loss of cultural practises and traditional knowledge In this study the forest practices of ‘traditional worship’ and ‘collecting materials for traditional items’ are decreasing as described by informants. This decrease indicates a loss of knowledge and culture and is brought about by factors such as: 1) a decrease in the supply of the traditional materials, 2) a decrease in the importance and need for these traditional items and 3) a shift in lifestyle toward “modern” alternatives. Traditional worship is now replaced mainly with Christianity, and this has also resulted in a decrease in traditional practices (Keesing 1967).

Globally cultural diversity including traditional ecological knowledge (TEK) are under threat due to a range of related processes including westernisation and a change in lifestyle (Caillaud et al. 2004, Brosius and Hitchner 2010, Painemilla et al. 2010). As stated by Caillaud et al. (2004) “the survival of traditional knowledge is vital to ensure sustainable conservation of [natural] resources in Melanesia”. Therefore traditional knowledge surrounding but not limited to herpetofauna and forest habitats needs to be preserved to help us achieve sustainable development and sustainable societies. There is a need for the conservation of both the biodiversity and its inter-related traditional information.

6.4.4 Loss of traditional knowledge in the younger generation In this study there seems to be a difference of traditional knowledge with the younger generation (below 30yrs) recording less knowledge than the eldest generation (above 60yrs). This is also supported by Lohani (2011) and Garcia (2006) who also found a lack of knowledge with younger people revealing less knowledge than elder people. The reasons for this include: 1) a decrease of knowledge transmitting events and interaction between the older and younger generation (Garcia 2006, Lohani 2011); 2) a decrease in availability in wild food plants and animals to allow interaction; 3) social stigmatization leading to a lack of interest in younger people; and 4) the attendance in school which limits time for traditional knowledge acquisition (Garcia 2006, Lohani 2011).

Since it is known that TEK persists, is developed and thrives while in application, if its application ceases to be practiced the TEK will be lost (Charnley et al. 2007). Likewise, if the traditional knowledge and practices surrounding herpetofauna cease

110 to be practiced and shared this information will also be threatened with extinction. For example methods for the capture and cooking of frogs will be lost along with traditional customs and stories associated with individual species.

Additional patterns observed include: that most of the information coming from older informants and especially from those that have spent a large amount of time living in the forest habitats. Where the informant grew up or spent their childhood was important in relation to the knowledge that they had, those that grew up in inland settlements as opposed to the coast had a higher level of understanding regarding herpetofauna and forests.

6.5 Summary In summary a total of 58 distinct herpetofaunal names were recorded from informants and these were placed into 21 distinguishable species, associated with seven categories of traditional use. Upland forests show the least amount of pressure from human activities with decreasing intensity for most uses due to an exodus of settlements to the coast. Therefore due to this weaker threat pressure upland forests would be a priority for conservation action. Lowland and coastal forests are under the greatest (and increasing) pressure from locals, this is mainly due to the close proximity of these habitat types to the human settlement areas. Logged and plantation forests are also under high pressure but due to their modified state with limited biological diversity they would not be priority candidates for conservation. However, it is also important to note that habitats faced with the greatest threats may also warrant a greater need for conservation actions.

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CHAPTER 7: POTENTIAL PRIORITY HABITATS AND STRATEGIES FOR FOREST BIODIVERSITY CONSERVATION

7.1 Introduction Conservation effort needs to be focused due to the limited financial and technical resources available (Myers et al. 2000, Bottrill et al. 2008, Wilson et al. 2009). Therefore, there is a pressing need to identify priority areas and strategies for conservation action (Margules et al. 2002, Wilson et al. 2009).

Conservation prioritisation is the process of identifying conservation priorities and making recommendations that will provide policy makers and donors with the necessary information to achieve the shared vision of biodiversity conservation (Collins and Storfer 2003). Conservation prioritisation is based on a number of inter- related principles including irreplaceability and vulnerability (Margules et al. 2002, Wilson et al. 2009). Irreplaceable areas contain unique species and habitats and are considered a high priority for conservation planning (Margules et al. 2002). Vulnerability is influenced by: the rarity of, the level of threat faced by, and the ecological importance of the species or habitats (Fa et al. 2004). Margules et al. (2002) believes that priority conservation areas should also have two roles, they should represent the biodiversity of the region and they should separate the biodiversity from the processes that threaten it.

Effective prioritisation requires sound information on the conservation status of species and ecosystems, including the vulnerabilities of, and threats to biodiversity (Beebee and Griffiths 2005, Wilson et al. 2005). Effective prioritisation also requires the effective combination of scientific methods, community engagement and traditional knowledge (Collins and Storfer 2003) a method that is being used in this study.

Key Biodiversity Areas (KBAs) represent global conservation prioritisation as they are designated areas of high biodiversity-conservation priority based on global standards and thresholds (Eken et al. 2004, Bass et al. 2011). The overall goal of KBAs is to apply standardised scientific methods for selecting globally significant 112 biodiversity sites for conservation actions (Eken et al. 2004). On a local scale important forest areas (IFAs) and important herpetofaunal areas (IHAs) once identified can also be included in the conservation prioritizing process.

Therefore an aim of this study is to identify important forest areas (IFAs) and important herpetofaunal areas (IHAs) on the island on Malaita that will help us prioritise conservation efforts at local scales. This chapter will address identification of potential priority forest habitats and strategies for forest biodiversity conservation based on the results of previous chapters. It will also discuss different methods of conservation prioritisation.

7.2 Methods for Prioritisation Singh et al. (2000) collated a list of 17 categories based on 47 global studies which focused on conservation prioritisation. Of the 17 categories, 4 categories (1) richness/diversity, 2) important species, 3) socio-cultural and 4) level of threat) were used in this study to identify conservation priority forest habitats (Table 7.1). The 4 categories were selected because of their relevance to this study and the opportunity to collect data to be used in these prioritisation categories.

Table 7.1 Summary of four categories for conservation prioritisation used in this study with descriptions based on Singh et al. (2000)

Categories General description Specific description of method used used for based on Singh et al. in the current study conservation (2000) prioritisation Refers to the number and “Species richness and abundance density of species in an value” (SRAV). This category refers area, with the greater to the total and mean species richness

1. Richness richness the higher per transect/quadrat and mean species priority. abundance per transect/quadrat as described in Chapter 5 (Figure 5.6 and 5.7).

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Refers to ecologically, “Important species value” (ISV). This economically and category refers to presence of near- symbolically important threatened, rare, totem and indicator species and can also refer species encountered in each habitat to endemic, threatened type. Near-threatened species are 2. Important and keystone species. classed as such by the IUCN Red-list species Areas with more such criteria, rare and indicator species are species having higher those defined in Chapter 4 (Table 4.1 priority. and 4.2) while totem species are those described by local communities in Chapter 6 (Table 6.2 and 6.4).

Refers to the non- “Cultural value” (CV). This category economic value of the refers to the perceived importance of site as part of culture, the forest habitat types to local 3. Socio-cultural aesthetics or history and communities and the general uses religion. recorded in Chapter 6 (Table 6.5, 6.6, 6.7, 6.8 and 6.9).

Refers to the level and “Forest threat value” (FTV). The type of pressures that the threat and the pressure that locals site is under. place on the forests as perceived by informants. The forest threat value is 4. Level of calculated based on the informant’s threat descriptions of the current status of the described uses and their impact on the relevant forest habitat types as per Chapter 6 (Table 6.10).

A fifth added approach the “combined rank value” (CRV) category will also be used. It will result in the combination of different prioritisation types so to achieve a holistic and inclusive approach to prioritisation setting for conservation areas. These 5 categories will then become the “methods” used for conservation prioritisation. 114

Under each of the five categories there are sub-categories and each habitat type will be assigned a value for each sub-category. These values were ranked and points assigned based on the rank (eg. 1st = 5 points, 2nd = 4 points, 3rd = 3 points, 4th = 2 points and 5th = 1 point). Each category will therefore have a final rank value for priority conservation habitat based on the sum of the ranks of each sub-category, with the lowest value having highest priority. This method was created for the purposes of this study alone and is not based on any other known studies. It is also important to note that for initial result purposes each category and subsequent sub- category has equal weighting.

7.3 Results

7.3.1 “Species richness and abundance value” (SRAV) Lowland forest is the highest priority forest habitat type based on the combination of species richness and species abundance whilst coastal forest is the least important (Table 7.2). With regards to the ranked sum of the total number of species observed, mean species richness (nocturnal and diurnal combined) and mean species abundance (nocturnal and diurnal combined), lowland forest can be said to have the highest SRAV and therefore be of high conservation priority.

Table 7.2 “Species richness and abundance values”, (total species + mean species richness and abundance) with higher rank values having greater conservation priority. (1st = 5 points, 2nd = 4 points, 3rd = 3 points, 4th = 2 points and 5th = 1 point) Forest Total Mean species richness Mean species SRAV habitat species per transect/quadrat abundances per value from type (TS) (nocturnal and transect/quadrat ranked diurnal combined) (minus B. marinus) sum of sub- categories Coastal 5th 9 = 5th 3.5 = 4th 1.5 = 2nd (1+2+4) = 7 1st Lowland 18 = 1st 6.7 = 1st 1.3 = 3rd (5+5+3) =

13 3rd Upland 14 = 3rd 5.4 = 3rd 2.0 = 1st (3+3+5) =

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2nd Logged 15 = 2nd 5.7 = 2nd 1.5 = 2nd (4+4+4) =

12 4th Teak 10 = 4th 5.4 = 3rd 2.0 = 1st (2+3+5) = plantation 10

7.3.2 “Important species value” (ISV) Lowland forest is the highest priority based on the combination of important species value sub-categories, whilst coastal and teak forests are equally least important (Table 7.3). With regards to the ranked sum of the number of near-threatened species, number of totem species, number of rare species and number of indicator species, lowland forest can be said to be a priority to be conserved based on its ISV.

Table 7.3 “Important species values”, (number of “near-threatened”, “totem”, “rare” and “indicator” species per habitat type) with higher rank values having greater conservation priority. (1st = 5 points, 2nd = 4 points, 3rd = 3 points, 4th = 2 points and 5th = 1 point)

Forest No. of near- No. of No. of No. of ISV value from habitat threatened totem rare indicator ranked sum of type species species species species (IS) sub-categories (NTS) (TS) (RS) Coastal 4th 0 = 3rd 3 = 2nd 1 = 3rd 0 = 4th (3+4+3+2) = 12 Lowland 1st 2 = 1st 5 = 1st 2 = 2nd 3 = 1st (5+5+4+5) = 19 Upland 3rd 0 = 3rd 3 = 2nd 3 = 1st 1 = 3rd (3+4+5+3) = 15 Logged 2nd 1 = 2nd 3 = 2nd 1 = 3rd 2 = 2nd (4+4+3+4) = 15 Teak 4th 1 = 2nd 1 = 3rd 1 =3rd 0 = 4th plantation (4+3+3+2) = 12

7.3.3 “Cultural value” (CV) Lowland forests are the highest priority and have the highest CV based on general uses of the forest as described by participants, whilst coastal and upland forests are also a priority (Table 7.4).

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Table 7.4 “Cultural values”, (number of general uses described by locals) with higher values having greater conservation priority. (1st = 5 points, 2nd = 4 points, 3rd = 3 points, 4th = 2 points and 5th = 1 point) Forest habitat type General uses CV value from ranked sum of sub-categories Coastal 11 =2nd 2nd (4) Lowland 13 = 1st 1st (5) Upland 11 = 2nd 2nd (4) Logged 8 = 3rd 3rd (3) Teak plantation 4 = 4th 4th (2)

7.3.4 “Forest threat value” (FTV) Coastal and lowland forests are the highest priority with the highest FTV, based on the perceived impacts that the general uses have on the forests described by participants (Table 7.5). These FTVs incorporate the current status, impact on the environment and the scale of the activities/uses into the analysis.

Table 7.5 “Forest threat values”, with higher values having greater conservation priority. (1st = 5 points, 2nd = 4 points, 3rd = 3 points, 4th = 2 points and 5th = 1 point) Forest habitat Forest threat values FTV rank from ranked sum of sub- type (FTV) categories Coastal 46 = 1st 1st (5) Lowland 42 = 2nd 2nd (4) Upland 15 = 4th 4th (2) Logged 28 = 3rd 3rd (3) Teak plantation 10 = 5th 5th (1)

7.3.5 “Combined rank value” (CRV) A combined rank value was determined by adding the SRAV, ISV, CV and FTV values. Combination of these values results in a final priority rank that clearly indicates lowland forests as the highest conservation priority (Table 7.6).

A visual representation of all priority methods shows that lowland forests are consistently of a high priority with all methods (Figure 7.1). Lowland forests score first in all but one prioritisation method making it clearly the forest type of highest conservation priority. Logged forest even with its evidently modified state is still of 117 high conservation priority with high SRAV and ISV and rates as the second forest of highest conservation priority. Coastal forests are third equal in conservation priority ranking first for FTV. Upland forests are also third equal with a high SRAV value. Teak plantation forests are of the least priority for biodiversity conservation and have low values for all prioritisation methods.

Table 7.6 “Combined rank value”, the combination of the four category values for conservation prioritisation. “Species richness and abundance values”, “important species values”, “cultural values” and “forest threat values”. (1st = 5 points, 2nd = 4 points, 3rd = 3 points, 4th = 2 points and 5th = 1 point) Forest habitat type SRAV ISV CV FTV Combined Rank Value (CRV)

Coastal 5th 3rd 2nd 1st (1+3+4+5=13) 3rd

Lowland 1st 1st 1st 2nd (5+5+5+4= 19) 1st

Upland 3rd 2nd 2nd 4th (3+4+4+2= 13) 3rd

Logged 2nd 2nd 3rd 3rd (4+4+3+3= 14) 2nd

Teak plantation 4th 3rd 4th 5th (2+3+2+1= 8) 4th

SRAV ISV CV FTV CRV

Coastal Coastal Coastal Coastal Coastal

Lowland Lowland Lowland Lowland Lowland

Upland Upland Upland Upland Upland

Logged Logged Logged Logged Logged

Teak Teak Teak Teak Teak

Figure 7.1 Graphic representation of priority habitat types based on Table 7.6 (the darker shade has the higher priority)

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7.4 Discussion

7.4.1 Species richness and abundance In this study the use of “species richness and abundance” to identify values of conservation priority has been useful. The use of species richness for habitat biodiversity comparison is very popular (Heinen 1992, Kerr 1997, Gascon et al. 1999, Vonesh 2001, Darwall and Vie 2005, Gillespie et al. 2005, Burgess et al. 2006, Gardner et al. 2007, Uehara-Prado et al. 2007, D'Cruze and Kumar 2011), however not so for abundance. The current study found that based on species richness, unlogged lowland forest is the priority habitat for conservation. This result is supported by Gardner et al. (2007) who found high species richness in similar “primary” forest. However it is contradictory to Vonesh (2001) in Uganda, who found greater species richness in “logged” forests. In addition, based on species richness Gascon et al. (1999) found a significant difference between sampled forest habitats, whereas Uehara-Prado et al. (2007) found no difference between forest habitats. Therefore it is safe to say that species richness alone is not generally a useful tool for conservation prioritisation because of its variable responses and exclusion of biologically important areas that are species poor (Kerr 1997, Eken et al. 2004).

7.4.2 Important species In the current study the use of “important species” to identify values of conservation priority has been useful. Previous studies of conservation prioritisation based on the IUCN Red Listed species (Eken et al. 2004, Darwall and Vie 2005, Pleguezuelosa et al. 2010) and indicator or keystone species (Darwall and Vie 2005) are common however the use of “culturally important” species as defined by Lohani (2011) is not so common. The current study has utilized all three individual species sub-categories (red-list, indicator and culturally important species) to identify lowland forest as the priority conservation habitat. “Important species” when combined with other criteria has therefore been shown to be especially useful for conservation prioritisation as also indicated by Eken et al. (2004) and Darwall and Vie (2005).

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7.4.3 Culture In this study the use of “culture” to identify values of conservation priority has been beneficial. The use of cultural, traditional or social values for conservation management and planning is infrequently found however it is also increasing as shown by the studies of Pedroso-Junior and Sato (2005), Chazdon et al. (2009), Raymond et al. (2009) and Bryan et al. (2011). The current study found that based on cultural values alone, unlogged lowland forest is the priority habitat for conservation (Figure 7.1). Since locals maintain strong ties with their surrounding biodiversity, their associated knowledge of the biodiversity is vital for conservation planning and prioritisation. The importance of using cultural values is also supported by the previous studies of Pedroso-Junior and Sato (2005) and Painemilla et al. (2010). However care must be taken with the use of cultural knowledge as shown by Bryan et al. (2011) working in Australia who found a negative correlation between social values of areas as defined by locals and the corresponding ecological values.

7.4.4 Forest threat The use of threats or the vulnerability of an area is commonly used in conservation prioritisation (Reyers 2004, Wilson et al. 2005, Brooks et al. 2006, Burgess et al. 2006, Cannon et al. 2007). The current study found that based on human threats to the forest the most vulnerable habitat type and therefore a priority conservation area is unlogged, coastal forest. In contrast, Cannon et al. (2007) found that lowland forests on alluvial soils to be under the most threat on Sulawesi, Indonesia and Burgess et al. (2006) found the mountainous regions on the African continent to be the most vulnerable. Site accessibility and close human habitation are two highly influential factors to the forest’s vulnerability (Burgess et al. 2006, Cannon et al. 2007), as is the case of coastal forests in this study. It is important to note that there are additional threats to forest habitats that locals may not know about, such as invasive species and global climate change.

7.4.5 Combined In this study the use of a “combined value” to identify forest conservation priority areas has been invaluable. The use of combined values for habitat biodiversity comparison is relatively common and strongly recommended (Eken et al. 2004, Burgess et al. 2006, Chazdon et al. 2009). For example, some studies 120 combined a measure of irreplaceability (e.g. endemic species) and vulnerability (e.g. threats) see Reyers (2004) and Burgess et al. (2006). Some studies combined scientific and local knowledge (Raymond et al. 2009, Raymond et al. 2010) while Wilson (2009) recommends that prioritisation decisions should include data on biodiversity, threat and cost. This study found that based on a combined rank value from species richness and abundance, important species, cultural values and forest vulnerability, unlogged lowland forest is the overall priority habitat for conservation. Similarly, Burgess et al. (2006) used the integration of biological values and threats for the entire continent of Africa and found lowland and montane forests as conservation priorities due to their globally significant biological values and high threats. The advantage of the combined values method is that it is more inclusive of a wide variety of inputs from science and society resulting in a more holistic approach.

A final result that stood out was that logged lowland forest emerged as being the second highest forest conservation priority. According to Gardner et al. (2007), Herrera-Montes and Brokaw (2010) and Gibson et al. (2011) logged or secondary forests do not provide an adequate substitute for primary forests, however some species may find these modified habitats favourable and therefore provide a valuable contribution to forest conservation

7.6 Conclusion Based on species richness and abundance, important species, cultural values and forest threats, lowland forests are the priority forest conservation habitat on Malaita. Logged forest is also of significant conservation value even in its disturbed state and also presents an additional opportunity for direct conservation action.

The current study has shown that it is possible to combine conservation biology science and traditional ecological knowledge to address the present conservation challenges in the Solomon Islands. With the baseline data provided here, the people of Malaita will have a vital starting point for discussion of future conservation action and steps that need to be taken.

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CHAPTER 8: OVERALL SUMMARY OF RECOMMENDATIONS FOR FUTURE CONSERVATION WORK ON MALAITA

8.1 Introduction The overall aim of this study was to identify priority forest conservation habitats on the island of Malaita using a combination of scientific and ethnological methods. The objectives included: 1) determining the abundance and richness of the Malaitan herpetofauna (frogs, geckos and skinks). 2) Defining the relationships between herpetofaunal occurrences, forest habitat type and forest habitat degradation. 3) Examining local perceptions on herpetofauna and forests and 4) identifying priority conservation forest habitats.

The methods used in this study were field surveys and questionnaires. Field surveys consisted of 40 days (and nights) sampling transects and quadrats in 5 different forest habitat types (coastal, lowland, upland, logged and teak plantation). Thirty individual questionnaires were completed in 10 villages on Malaita in close proximity to the forest study sites. Forest habitat prioritisation for conservation was then determined based on five values calculated from information gathered through the field surveys and questionnaires.

8.2 Important Recommendations for Future Conservation work on Malaita based on Literature Areas that should be prioritized are those with high species richness and diversity especially across different taxa, areas such as biogeographic crossroads where intersections of dominant habitat types create such areas (Spector 2002). The Solomon Islands lie at a biogeographic crossroads between the continental biota of Malesia or Australasia and the isolated, mostly oceanic, islands of the Pacific. No other primarily oceanic archipelago is considered to have a greater proportion of the planets living biodiversity, with exceptional patterns of endemism and richness also in culture and way of life (Filardi et al. 2007). Tropical regions are particularly vulnerable and their rich biodiversity and ever increasing threats make them a high priority for conservation effort (Gascon et al. 2004).

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8.2.1 The importance of culture Conservation is about people, our ability to address and deal with social, cultural and community issues and link this with the needs of biodiversity (SPBCP 2001, Chan et al. 2007, Brodie et al. 2013). Conservation assessments therefore need to incorporate cultural, social, economic and political factors (Gascon et al. 2004, Knight and Cowling 2007, Wilson et al. 2009, Tengberg et al. 2012). In order for conservation to achieve any degree of success, the local communities who “own” the biodiversity need to be able to make informed decisions about the sustainability and use of their natural resources (Pough et al. 1998, Schwartzman et al. 2000, Read 2002, Danielsen et al. 2009, Game et al. 2011). The conservation agenda and implementation plan must be set by these local groups (Smith et al. 2009) and planned and managed in its own individual context (Brosius and Hitchner 2010).

To truly understand the relationship between culture and nature, conservation biology and Traditional Ecological Knowledge (TEK) must be combined (Drew and Henne 2006). A partnership between science and law both traditional and modern is needed where the government can recognize and re-empower traditional laws and management systems (see Sulu in Caillaud 2004), as many of these traditional mechanisms now are no longer effective or respected (Bennet 2000, Crocombe 2001).

8.2.2 The importance of conservation science For conservation problems to be answered effectively, a clear definition of goals and the identification of actions and their likely costs and benefits needs to be made (Wilson et al. 2009). An overall goal of biodiversity conservation should be the “long-term survival of species and inter-related natural processes whilst excluding their threats” (Margules and Pressey 2000). Data on species and threats, costs and benefits is needed, and the success of conservation action depends on the quantity and quality of the data used to plan and design it (Kati et al. 2004). Therefore to get good data, good quality monitoring and research is vitally needed to contribute to effective decision making in conservation and resource management (Danielsen et al. 2009).

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8.2.3 The importance of policy Though scientific research is important to help us understand biodiversity declines, the power to really address and reverse biodiversity degradation lies with politics, legislation and community socioeconomics (Beebee and Griffiths 2005). There seems to be a significant lack of recognition from policy makers and leaders of the importance of the environment and biodiversity, its hugely threatened state and the need for immediate action (PHCG 2008). Consequently, there is an urgent need for greater partnership and collaboration between governments, NGOs and local communities (SPBCP 2001).

Globally, there is an acknowledged research-implementation gap in conservation science (Knight et al. 2008). Prioritisation is about being efficient but without implementation such activities become totally inefficient (Game et al. 2011). Successful implementation of conservation policies depends on education, awareness, political will, committed and knowledgeable leadership, community aspirations, social and economic capacity and scientific understanding blended with cultural and political institutions (Kingsford et al. 2009, Gough et al. 2010).

8.3 Important Recommendations for Conservation work on Malaita based on this Study It is a core aspect of this study to integrate both the biological and cultural values of forests and herpetofauna for conservation decision making. Therefore, any resulting conservation action must result in the preservation of both biological diversity and cultural diversity. Conservation prioritisation must be a process that includes all stakeholders at all levels.

Research findings and environmental conservation knowledge must be made available and user-friendly to locals. It is important to communicate in values and units that are understood by local resource owners for example, the unit of habitat type may be less understood than traditional land boundaries and units of tribal lands. It is also important to communicate in a language that is understood by all locals, this will substantially strengthen the chances of common understanding and common expectations. To be successful conservation must be driven by locals and cannot be seen as being imposed from the outside. Locals must have a complete knowledge of

124 costs and benefits of any conservation actions in order to remove any misconceptions.

However, collaboration is important and external stakeholders including government, NGO’s and possible financial and technical institutions can be engaged to improve conservation effectiveness. Training, capacity building and knowledge sharing with locals are of utmost importance. It is therefore essential to include landowners in biodiversity monitoring, this will help to ensure the long-term sustainability of conservation projects and also result in knowledge sharing between locals and any external stakeholders.

Malaita is an island with a high human population, density and birth rate that in turn creates a greater threat on the island’s biodiversity. The ÀreÀre region, the focus of this study was found to be a priority for conservation action, the results however can be translated for the rest of Malaita island. ÀreÀre also holds some of the last remaining “untouched” forests of Malaita and offer a great opportunity for conservation work. Also fit for mention are the Kwaio and Kwarae highlands of Malaita that house the highest mountains and only montane forests of the island.

8.4 Conclusion Achieved in this study was a greater understanding of herpetofaunal incidence on the island of Malaita. Also important was the documenting of traditional knowledge and understanding the threats to and importance of traditional knowledge to local communities and the conservation story. Unlogged lowland was identified as the priority conservation forest habitat type. Not achieved in this study was any actual conservation action or outcome.

To identify unlogged lowland forests as the priority conservation habitat type on Malaita is only the first step. Beyond this step is the actual development and implementation of conservation actions. Recommended principles for conservation action include the importance of culture, science and policy for successful outcomes. A holistic approach to conservation action by including scientific knowledge and methods with cultural knowledge and practices is vital. A realistic collaborative partnership between government, non-government stakeholders and resource owners is therefore essential. 125

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Appendix A: Ethnological Questionnaire Questionnaire to determine local community perceptions and knowledge regarding frogs, skinks and geckos (herpetofauna) and their local forest habitats No.:____ Name: ______Age: ______Gender: ______Village:______Date:______Time:______Interviewer: ______

What are the most important different frog (Ko`e) species that you know? What are their names; where are they found; what is their abundance; and have they declined or increased in abundance and reasons for change in abundance? What are their associated uses or other stories, tales or information on them?

1. , Description:______. Habitat/place found: ______. Abundance: C U R . Change in abundance: _No Change, Increase, Decrease _and reasons: ______Uses and other information:______.

2. , Description:______. Habitat/place found: ______. Abundance: C U R . Change in abundance: _No Change, Increase, Decrease _and reasons: ______Uses and other information:______.

3. , Description:______.

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Habitat/place found: ______. Abundance: C U R . Change in abundance: _No Change, Increase, Decrease _and reasons: ______Uses and other information:______.

4. , Description:______. Habitat/place found: ______. Abundance: C U R . Change in abundance: _No Change, Increase, Decrease _and reasons: ______Uses and other information:______.

5. , Description:______. Habitat/place found: ______. Abundance: C U R . Change in abundance: _No Change, Increase, Decrease _and reasons: ______Uses and other information:______.

6. , Description:______. Habitat/place found: ______. Abundance: C U R . Change in abundance: _No Change, Increase, Decrease _and reasons: ______Uses and other information:______.

7. , Description:______. Habitat/place found: ______. Abundance: C U R . Change in abundance: _No Change, Increase, Decrease

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_and reasons: ______Uses and other information:______.

8. , Description:______. Habitat/place found: ______. Abundance: C U R . Change in abundance: _No Change, Increase, Decrease _and reasons: ______Uses and other information:______.

9. , Description:______. Habitat/place found: ______. Abundance: C U R . Change in abundance: _No Change, Increase, Decrease _and reasons: ______Uses and other information:______.

10. , Description:______. Habitat/place found: ______. Abundance: C U R . Change in abundance: _No Change, Increase, Decrease _and reasons: ______Uses and other information:______.

What are the most important different lizard (gecko (kuma) and skink (iikiko, unu)) species that you know? What are their names; where are they found; what is their abundance; and have they declined or increased in abundance and reasons for change in abundance? What are their associated uses or other stories, tales or information on them? 138

1. , Description:______. Habitat/place found: ______. Abundance: C U R . Change in abundance: _No Change, Increase, Decrease _and reasons: ______Uses and other information:______.

2. , Description:______. Habitat/place found: ______. Abundance: C U R . Change in abundance: _No Change, Increase, Decrease _and reasons: ______Uses and other information:______.

3. , Description:______. Habitat/place found: ______. Abundance: C U R . Change in abundance: _No Change, Increase, Decrease _and reasons: ______Uses and other information:______.

4. , Description:______. Habitat/place found: ______. Abundance: C U R . Change in abundance: _No Change, Increase, Decrease _and reasons: ______Uses and other information:______.

5. , Description:______.

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Habitat/place found: ______. Abundance: C U R . Change in abundance: _No Change, Increase, Decrease _and reasons: ______Uses and other information:______.

6. , Description:______. Habitat/place found: ______. Abundance: C U R . Change in abundance: _No Change, Increase, Decrease _and reasons: ______Uses and other information:______.

7. , Description:______. Habitat/place found: ______. Abundance: C U R . Change in abundance: _No Change, Increase, Decrease _and reasons: ______Uses and other information:______.

8. , Description:______. Habitat/place found: ______. Abundance: C U R . Change in abundance: _No Change, Increase, Decrease _and reasons: ______Uses and other information:______.

9. , Description:______. Habitat/place found: ______. Abundance: C U R . Change in abundance: _No Change, Increase, Decrease

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_and reasons: ______Uses and other information:______.

10. , Description:______. Habitat/place found: ______. Abundance: C U R . Change in abundance: _No Change, Increase, Decrease _and reasons: ______Uses and other information:______.

11. , Description:______. Habitat/place found: ______. Abundance: C U R . Change in abundance: _No Change, Increase, Decrease _and reasons: ______Uses and other information:______.

What are up to 5 main uses associated with Primary (Wapu) Upland Forests (eg. Ohumae)?? Have there been changes on this use and how do (if so) these impact frogs and lizards??

1. ______Changes:___ No, Dec, Inc, Impacts on herps :

______

2. ______Changes:___ No, Dec, Inc, Impacts on herps :

______

3. ______Changes:___ No, Dec, Inc, Impacts on herps :

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______

4. ______Changes:___ No, Dec, Inc, Impacts on herps :

______

5. ______Changes:___ No, Dec, Inc, Impacts on herps :

______

What are up to 5 main uses associated with Primary Lowland (Oote) forest (eg. Houhou)?? Have there been changes on this use and how do (if so) these impact frogs and lizards??

1. ______Changes:___ No, Dec, Inc, Impacts on herps :

______

2. ______Changes:___ No, Dec, Inc, Impacts on herps :

______

3. ______Changes:___ No, Dec, Inc, Impacts on herps :

______

4. ______Changes:___ No, Dec, Inc, Impacts on herps :

______

5. ______Changes:___ No, Dec, Inc, Impacts on herps :

______

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What are up to 5 main uses associated with Secondary/Logged (Aru) forest (eg. Aimera)?? Have there been changes on this use and how do (if so) these impact frogs and lizards??

1. ______Changes:___ No, Dec, Inc, Impacts on herps :

______

2. ______Changes:___ No, Dec, Inc, Impacts on herps :

______

3. ______Changes:___ No, Dec, Inc, Impacts on herps :

______

4. ______Changes:___ No, Dec, Inc, Impacts on herps :

______

5. ______Changes:___ No, Dec, Inc, Impacts on herps :

______

What are up to 5 main uses associated with Plantation forests (bariki/farm) (eg. Teak)?? Have there been changes on this use and how do (if so) these impact frogs and lizards??

1. ______Changes:___ No, Dec, Inc, Impacts on herps :

______

2. ______Changes:___ No, Dec, Inc, Impacts on herps :

______

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3. ______Changes:___ No, Dec, Inc, Impacts on herps :

______

4. ______Changes:___ No, Dec, Inc, Impacts on herps :

______

5. ______Changes:___ No, Dec, Inc, Impacts on herps :

______

1. What are up to 5 main uses associated with Coastal (Haho) forest (eg. Rapi roto)?? Have there been changes on this use and how do (if so) these impact frogs and lizards??

1. ______Changes:___ No, Dec, Inc, Impacts on herps :

______

2. ______Changes:___ No, Dec, Inc, Impacts on herps :

______

3. ______Changes:___ No, Dec, Inc, Impacts on herps :

______

4. ______Changes:___ No, Dec, Inc, Impacts on herps :

______

5. ______Changes:___ No, Dec, Inc, Impacts on herps :

______

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Appendix B: Species Descriptions with Field Photographs All photographs were taken on Malaita in the Solomon Islands by Edgar Pollard in situ.

Frogs

1) Batrachylodes vertebralis Boulenger, 1887 Batrachylodes vertebralis is a small frog with males reaching 28 mm Snout-Vent Length (SVL) and females 30 mm SVL (Pikacha et al. 2008). The back is a grey, brownish, cream to tan, a dark band runs along the side of the head from the snout and there are usually dark specks or markings on its back. Occasionally a thin white stripe can be observed down the middle of the back and the hind legs have light transverse bands, the underside is yellowish to white (Pikacha et al. 2008). This species has been recorded on Choiseul, New Georgia, Isabel, Malaita, Guadalcanal, Ugi and Santa Ana in the Solomon Islands and is a native and endemic to the Solomon’s bio-region including Bougainville (Pikacha et al. 2008). It is found in low to mid altitude forests, degraded forests and plantations, males are usually found calling in elevated, sheltered positions (Pikacha et al. 2008). This frog is common with a stable, large population that is widely distributed with a tolerance for a range of habitats and is therefore listed as a species of Least Concern by IUCN (2012). This species may be threatened by clear-cutting such as logging and also by invasive species (IUCN 2012).

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2) Bufo marinus Linnaeus, 1758 Bufo marinus is a large introduced frog with males reaching 110 mm and females 250 mm SVL (Pikacha et al. 2008). The back is a pale brown/olive, there are usually large warts and dark marks with markings more visible in juveniles and the underside is yellowish to white (Pikacha et al. 2008). This species originates from Central America but is found on most major islands of the Solomon Islands including Choiseul, New Georgia, Kolombangara, Guadalcanal, Makira and Banika (Pikacha et al. 2008).This frog is also listed on the Global Invasive Species Database in the top 100 worst invasive species (GISD 2013).This frog adapts well to almost all habitats from urban areas, agricultural areas and coastal to upland forests, although road and track edges are preferred as it does not climb and thus dense vegetation hinders movement (Pikacha et al. 2008).

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3) Ceratobatrachus guentheri Boulenger, 1887 Ceratobatrachus guentheri is a medium sized frog with males reaching 65 mm and females 80 mm (Pikacha et al. 2008). It has a unique triangular-shaped head with pointed triangular skin flaps on the upper eyelids, snout, limbs and jaws. Coloration in this species is extremely variable ranging from bright yellow/orange to light/dark brown with variable spots and markings; the underside is a pale brown (Pikacha et al. 2008). This species has been found on all major islands in the Solomon Islands except for Makira and is a native and endemic to the Solomon’s bio-region including Bougainville (Pikacha et al. 2008). It is found on the forest floor in low to mid- altitude forests, degraded forests and plantations (Pikacha et al. 2008). This frog is common with a stable, large population that is widely distributed and with a tolerance for habitat modification. It is therefore listed as a species of “Least Concern” by IUCN (2012). However, this species may be threatened by live export for the foreign pet trade, collection for food and logging (IUCN 2012).

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4) Discodeles guppyi Boulenger, 1887 Discodeles guppyi is a large frog with males reaching 110 mm and females 250 mm (Pikacha et al. 2008). It is reddish to blackish brown with darker splotches, the throat, belly is whitish to yellowish, and the lips can have distinct transverse bands present. It is found on all major islands in the Solomon Islands except for Makira and is a native and endemic to the Solomon’s bio-region including Bougainville. It is found along streams and small rivers in lowland forests, degraded forests and occasionally in caves, males are usually found calling beside waterfalls at night (Pikacha et al. 2008). This frog is common with a stable, large population that is widely distributed with tolerance for habitat modification and is therefore listed as a species of “Least Concern” by IUCN (2012). This species may be threatened by live exporting for the pet trade, collection for food and logging (IUCN 2012).

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5) Platymantis guppyi Boulenger, 1887 Platymantis guppyi is a medium-sized frog with males reaching 75 mm and females 90 mm (Pikacha et al. 2008). The back ranges from yellowish to darker brown usually with darker spots or markings, the hind legs have faint but distinct transverse bands. It is found on all major islands in the Solomon Islands and is a native and endemic to the Solomon’s bio-region including Bougainville. It is found in closed canopy and old-growth forests and is arboreal, preferring trees 2-20 m above the ground (Pikacha et al. 2008). This frog is common with a stable, large population that is widely distributed with a tolerance for habitat modification and is therefore listed as a species of “Least Concern” by IUCN, although it may be threatened by logging (IUCN 2012) and plantation forest (teak) development.

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6) Platymantis solomonis Boulenger, 1884 Platymantis solomonis is a medium-sized frog with males reaching 56 mm and females 71 mm (Pikacha et al. 2008). The back is reddish to dark brown with darker splotches, the hind limbs have dark transverse bands and the underside is whitish to cream (Pikacha et al. 2008). It is found on all major islands in the Solomon Islands except for Makira and is a native and endemic to the Solomon’s bio-region including Bougainville (Pikacha et al. 2008) and found in low to mid altitude forests, degraded forests, coconut plantations and rural gardens (Pikacha et al. 2008). This frog is common with a stable, large population that is widely distributed with a strong tolerance for habitat modification and is therefore listed as a species of “Least Concern” by IUCN (2012). However on Malaita it was found that this species was only found in the less modified habitats of lowland and upland forests (pers. obs).

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7) Platymantis weberi Schmidt, 1932 Platymantis weberi is a medium sized frog with males reaching 35 mm and females 56 mm. The back is dark reddish to dark brown with red stripes common where the back meets the side of the body, the hind limbs have dark transverse bands and the underside is whitish to cream. It is found on all major islands in the Solomon Islands except for Makira and is a native and endemic to the Solomon’s bio-region including Bougainville. It is found in low to mid altitude forests, degraded forests, and plantations, and males are usually found calling in elevated, sheltered positions (Pikacha et al. 2008). This frog is common with a stable, large population that is widely distributed with a strong tolerance for habitat modification and is therefore listed as a species of Least Concern by IUCN (2012).

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8) Rana kreffti Boulenger, 1884 Rana kreffti is a medium-sized frog with males reaching 52 mm and females 82 mm. The back is mid to dark brown with no dark splotches, a black band runs along the side of the body from snout through eye towards the hind limbs, the underside is creamy yellow to white (Pikacha et al. 2008). It is found on all major islands in the Solomon Islands and is a native and endemic to the Solomon’s bio-region including Bougainville (Pikacha et al. 2008) in low- to mid-elevation forests, degraded forests, plantations, grasslands and swamps. It lays eggs in small pools (Pikacha et al. 2008). This frog is common with a stable population and is listed as a species of Least Concern by IUCN (2012).

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Lizards (Geckos)

1) Cyrtodactylus salomonensis Rösler, Richards & Günther, 2007 Cyrtodactylus salomonensis is a large gecko with an average SVL of 130 mm. Dorsal coloration is light yellowish brown to medium dark brown with dark broad cross-bands, ventrally it is grey to yellowish white. When nocturnally active a third to a half of the tail becomes white (McCoy 2006). This species is endemic to the Solomon Islands (Rosler et al. 2007) and has been recorded in the Shortland Islands, New Georgia, Isabel, Guadalcanal, and Malaita (McCoy 2006). This arboreal gecko is found mostly on the larger forest trees especially preferring hollows and Ficus spp. (McCoy 2006). It has been assessed and listed as “Near Threatened” on the red list (IUCN 2012).

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2) Gehyra oceanica Lesson, 1830 Gehyra oceanica is a native medium sized gecko with an average SVL of 90 mm (McCoy 2006). Dorsal coloration is light to dark brown with irregular lighter and darker flecks; ventrally it is cream to yellow (McCoy 2006).It is a widely dispersed species throughout the Pacific Islands and the Indo-Australian archipelago. In the Solomon’s It has been recorded on the islands of Shorthand’s, Mono, Choiseul, Rob Roy, New Georgia, Tetepare, Vangunu, Isabel, Guadalcanal, Tulagi, Ngela, Malaita Ontong Java, Makira, Ugi, Olu Malau, Bellona, Santa Cruz, Taumako, Reef Islands and Utupua where it is found mainly on larger trees especially preferring coconut and sago palms and sometimes found around homes (McCoy 2006). This gecko was naturally dispersed to the Pacific islands before human arrival and has adapted an ecology and reproductive biology to support its ability for cross ocean dispersal (Fisher 1997). This may also be applicable to other native lizards in the Pacific region. The IUCN assessment for this species is “Least Concern” (IUCN 2012).

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3) Nactus multicarinatus Günther, 1872 Nactus multicarinatus is a bisexual, small to medium-sized gecko with an average SVL of 60 mm. Dorsal coloration in this species is grey-brown with darker wavy transverse bands, ventrally it is cream to yellow (McCoy 2006). It is a native to Vanuatu and the Solomon’s bio-region including Bougainville. It is found on all major islands in the Solomon Islands, mostly on the ground but also on tree trunks in forests, plantations, gardens and urban areas (McCoy 2006). This gecko is common with a stable, large population that is widely distributed with a tolerance for habitat modification and is therefore listed as a species of “Least Concern” by IUCN, although it may be threatened by invasive species (IUCN 2012).

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Lizards (Skinks)

1) Corucia zebrata (Gray, 1856) Corucia zebrata is a very large sized native skink, probably the largest in the world (McCoy 2006) and has an average SVL of 350 mm. Its dorsal coloration is highly variable ranging from olive green, grey-green to khaki with lighter and darker flecks present; ventrally it is yellow-green to grey-green (McCoy 2006). This species is endemic to the Solomon Islands archipelago including Bougainville and has been recorded on the Shorthand’s Islands, Vella Lavella, Choiseul, New Georgia, Tetepare, Vangunu, Isabel, Guadalcanal, Ngela, Malaita, Makira, Ugi and Santa Ana. It is a nocturnal, arboreal skink, which is found mostly on the larger forest trees amongst dense foliage especially preferring hollows and Ficus spp. with mean home range sizes of around 0.17ha (McCoy 2006, Hagen and Bull 2011). The IUCN assessment for this species is “Near Threatened” (IUCN 2012).

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2) Prasinohaema virens Boulenger, 1883 Prasinohaema virens is a small native skink with an average SVL of 50 mm. Dorsal coloration is pale green to light olive green; ventrally it is bright yellow to yellow-green (McCoy 2006). This diurnal skink is found in PNG and the Solomon’s, in the Solomon’s it is very widespread and has been recorded on the Shortland Islands, Mono, Choiseul, Vella Lavella, New Georgia, Tetepare, Vangunu, Isabel, Guadalcanal, Ngela, Malaita, Ontong Java, Makira, Ugi, Olu Malau, Santa Ana, Santa Cruz, Vanikoro, Taumako, Utupua, Tikopia and the Reef Islands, where it is a totally arboreal forest dweller preferring trees with vines and creepers (McCoy 2006). The IUCN assessment for this species is Least Concern (IUCN 2012).

No picture was taken of this species due to low encounters because of arboreal nature.

3) Emoia atrocostata freycineti Duméril & Bibron, 1839 Emoia atrocostata freycineti is a medium sized native skink with an average SVL of 75 mm. Its dorsal coloration is grey to grey-green to black with lighter flecks that appear to form transverse bands; ventrally it is white with greenish hue (McCoy 2006). This sub-species is widespread throughout the Solomon’s and is also found in Vanuatu. It has been recorded on the Shortland Islands, Mono, Choiseul, Rob Roy, Vella Lavella, Ranongga, Gizo, Kolombangara, New Georgia, Tetepare, Vangunu, Isabel, Russell Islands, Guadalcanal, Ngela, Malaita, Ontong Java, Makira, Ugi, Olu Malau, Rennell, Bellona, Santa Cruz, Vanikoro and the Reef Islands where it is a common active diurnal skink found in coastal areas and rocky foreshores (McCoy 2006). The IUCN assessment for this species is “Least Concern” (IUCN 2012).

No picture was taken of this species due to rarity and speed of lizard evading capture.

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4) Emoia cyanogaster Lesson, 1826 Emoia cyanogaster is a large native skink with an average SVL of 85 mm. Its dorsal coloration is golden to greenish bronze with darker flecks occasionally present; ventrally it is yellow-green to lime-green (McCoy 2006). It is very widespread in the Solomon’s and is also found on Vanuatu and PNG. In the Solomon’s it has been recorded on the Shortland Islands, Fauro, Mono, Choiseul, Vella Lavella, New Georgia, Tetepare, Vangunu, Isabel, Guadalcanal, Ngela, Malaita, Ontong Java, Makira, Ugi, Olu Malau, Santa Ana, Rennell, Santa Cruz, Vanikoro, Utupua, Tikopia and the Reef Islands (McCoy 2006). This diurnal, arboreal skink is found in forested areas including gardens and plantations preferring vine covered trees (McCoy 2006). The IUCN assessment for this species is “Least Concern” (IUCN 2012).

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5) Emoia nigra Jacquinot & Guichenot, 1853 Emoia nigra is a large native skink with an average SVL of 100 mm (McCoy 2006). Dorsal coloration is glossy black or brown and ventrally it is white to dull cream (McCoy 2006).In the Pacific it has been recorded in PNG, Solomon Islands, Vanuatu, Fiji, Samoa and Tonga. This skink is the most widespread lizard in the Solomon Islands and is found on all islands. This diurnal, active skink is found mostly on the ground in a wide range of habitats from forests to human settlements and agricultural areas (McCoy 2006). The IUCN assessment for this species is “Least Concern” (IUCN 2012). Fisher (pers. com. 2012) indicates that this species will probably be split into three new species following recent genetic analyses.

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6) Emoia pseudocyanura Brown, 1991 Emoia pseudocyanura is an endemic moderate sized skink with an average SVL of 55 mm. Dorsal coloration is brown to black head fading out into a copper coloured tail with a mid-dorsal stripe and two lateral stripes present; ventrally it is creamy white to dull yellow. It is very widespread and has been recorded on the Shortland Islands, Choiseul, Isabel, Russell Islands, Guadalcanal, Ngela and Malaita (McCoy 2006). The Malaita population is believed to be a separate un-described species due to its distinct coloration (McCoy 2012, pers. comm., Fisher 2013 pers. comm.). This diurnal, semi-arboreal skink is found in a wide variety of habitats but prefers forest edges and areas (McCoy 2006). The IUCN assessment for this species is “Least Concern” (IUCN 2012).

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7) Sphenomorphus bignelli Schmidt, 1932 Sphenomorphus bignelli is a small skink with an average SVL of 35 mm. Dorsal coloration is light brown to black with irregular lighter and darker flecks, ventrally is grey to cream (McCoy 2006).This diurnal skink is endemic to the Solomon Islands and is found on the islands of Kolombangara, New Georgia, Tetepare, Vangunu, Russell, Ngela, Malaita and Guadalcanal where it found mostly on the ground in open shady areas amongst leaf litter (McCoy 2006). The IUCN assessment for this species is “Least Concern” (IUCN 2012).

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8) Sphenomorphus concinnatus Boulenger, 1887 Sphenomorphus concinnatus is a medium sized skink with an average SVL of 65 mm. Dorsal coloration is golden brown with darker flecks, ventrally is yellowish to dull orange-brown (McCoy 2006). This diurnal species is endemic to the Solomon Islands and is found on the Shortland Islands, Fauro, Choiseul, Rob Roy, Vella Lavella, Ranongga, Gizo, Kolombangara, New Georgia, Tetepare, Vangunu, Isabel, Ngela, Malaita and Guadalcanal where it is found mostly in forests and semi-cleared areas foraging amongst leaf litter (McCoy 2006). The IUCN assessment for this species is “Least Concern” (IUCN 2012).

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9) Sphenomorphus cranei Schmidt, 1932 Sphenomorphus cranei is a medium sized skink with an average SVL of 60 mm. Dorsal coloration is light brown to black with light and dark flecking; ventrally it is yellowish to orange-red. This sometimes diurnal skink is endemic to the Solomon Islands and is found on the Shortland Islands, Vella Lavella, New Georgia, Tetepare, Vangunu, Isabel, Ngela and Malaita where it is uncommon and is fairly moisture dependent (McCoy 2006). The IUCN assessment for this species is least concern (IUCN 2012). McCoy (pers. comm. 2011) believes that this specimen to be an undescribed S. cranei sub-species for the island of Malaita.

10) Sphenomorphus solomonis (Boulenger, 1887) Sphenomorphus solomonis is a small skink with an average SVL of 50 mm, dorsal coloration is glossy black or brown; ventrally it is white to dull cream. This nocturnal skink is very widespread and has been recorded on the Shortland Islands, Fauro, Choiseul, New Georgia, Isabel, Guadalcanal, Savo, Ngela, Malaita, Makira, Ugi, Santa Cruz, Taumako and the Reef Islands where it is found in forests living on the ground and amongst rotting wood and leaf litter in moist conditions (McCoy 2006). The IUCN assessment for this species is least concern (IUCN 2012).

No picture was taken of this species due to rarity and speed of lizard evading capture.

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