ECOLOGY, IMPACT AND TRADITIONAL KNOWLEDGE OF RESIN

HARVEST ON THE WILD DAMMER TREE-CANARIUM STRICTUM ROXB. IN

THE NILGIRI BIOSPHERE RESERVE, WESTERN GHATS, INDIA

A DISSERTATION SUBMITTED TO THE GRADUATE DIVISION OF THE UNIVERSITY OF HAWAI‘I AT MᾹNOA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF

DOCTOR OF PHILOSOPHY

IN

BOTANY

AUGUST 2014

By

Anita Varghese

Dissertation Committee:

Tamara Ticktin, Chairperson David C. Duffy Jefferson Fox Orou Gaoue Christopher A. Lepcyzk

Keywords: non timber forest produce, resin, harvest, traditional ecological knowledge, mixed effects models, phenology, Canarium strictum, Nilgiri biosphere reserve

ACKNOWLEDGEMENTS

This intense journey of the past years would not have been possible without grace from the Almighty and support, and encouragement of family, friends, teachers, colleagues and institutions. To all of them I remain deeply grateful and wish to express my heartfelt gratitude.

My committee members – Dr Tamara Ticktin (Chairperson) for her unflinching faith in her student’s abilities and for being the wonderful person she is.

Drs David Duffy, Jefferson Fox, Christopher Lepcyzk and Orou Gaoue for comments, suggestions, ideas and encouragement.

My family and friends- Parents, daughter, sister, nieces, brother in law and cousins for their support and faith in what I do. My daughter, Merab who helped me set out on this journey and was there to help me take it to the end point. Friends and neighbours in Kotagiri – Pratim, Sneh, Mathew and Annie, my motivators and my local guardians. Pastor and members of the Union Church in Kotagiri, who have been a source of strength. Chamanlalji and Shipradi, for their blessings and countless lessons about life. Mari and Stan, from Gudalur for their encouragement. Anu, Sunita,

Kochu, Durga – friends for a long time. Teachers at Rishi Valley School, who adopted Merab as their own. Hawaii my home away from home and so much like my home. Tamara, Gustavo and Ylang, my family away from family. Vandana, Lisa,

Isabel, Alesandre, Jennifer, Daniela, Georgia, Natalie, Katie, Gioconda, Shimona – for the learning times and fun times. A special word for Jennifer my constant ‘J-seek’.

The Botany department at UHM - a great place to be and Dr. Tom Ranker, Dr.

Alison Sherwood and Ms Patty Bedoya have been especially supportive.

My organisation – Keystone Foundation- a small NGO with big ideas - an inspiring place to be. Sneh, Pratim and Mathew – founders who shared their dreams

ii with me. My team members especially Sumin, colleagues and well-wishers who have made the place so vibrant. For help in field work that I received from Aradukuttan,

Mahadesh, Sudhakar, L. Rajendran, Senthil Prasad, Maya,Vasu, Punit, Rajan, Karian,

Rangasamy, Rangan, and Lingan.

My funding agencies – The East West Center, PEO International Scholarship,

East West Center Summer Research Grant, Jean E Rolles Scholarship, UH

Foundation Grants, Ruffords Small Grants, Botany in Action-Phipps Conservatory,

Mohammed bin Zayed species conservation fund, and Keystone Foundation.

Many people knew from a long time that I should get out and do my PhD and

I am grateful to them for their advice – Tony Cunningham who started this idea,

Pratim Roy who nurtured it, Tamara Ticktin who welcomed it and Patricia Shanley who helped me take that final decision.

I deeply miss my good friends, T.Kunhimohammed and S.S.Manoharan who were there when I started out but are not here on earth to see this journey reach its destination.

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ABSTRACT

Harvest of plant products from the wild are an important source of livelihood to millions of people who live near forests. The impact of harvesting occurs at many levels to the ecology of the species. Harvesters are guided by their traditional ecological knowledge that also aids in reducing the impact of the harvest. I use the case study of Canarium strictum Roxb, (Burseraceae) a semi-evergreen tree harvested for resin by several indigenous communities in the Indian subcontinent. Resin is used locally for rituals and healing purposes and traded widely for industrial uses. I investigated the effects of resin harvesting on the ecology and phenology of the species on 89 trees in three regions of the Nilgiri Biopshere Reserve, Western Ghats, for two years. Seed germination experiments with seeds from harvested and not harvested trees were also undertaken. Through focus group discussions with harvesters, and using a fuzzy logic approach I documented their perceptions on the ecology of resin harvest and trees. I found that harvesting practices, size of the tree along with the characteristic of the tree flush colour were significant predictors of resin harvest. My results show that harvesting of resin has no negative effect on the growth rate, and fruit production. However harvested trees flowered at different times from not harvested trees and showed increased fruit production and seed germination rates. I found that resin harvesting was a prevalent practice among the indigenous people of the region and many of the factors perceived by the harvesters to influence resin quality and status of resin tree numbers in the forest coincided with factors observed in the ecological studies. Overall my results suggest that harvesting of resin has relatively low impact on the ecology of C. strictum and harvesters of resin make decisions on resin harvest based on a number of ecological factors. My results illustrate some of the detailed knowledge that harvesters have with regard to a lesser

iv studied species like C. strictum, and community based monitoring programs that build on this knowledge can ensure strategies that allow for sustainable use while meeting the goals of conservation.

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TABLE OF CONTENTS

ACKNOWLEDGEMENTS ...... II

ABSTRACT ...... IV

LIST OF TABLES ...... VIII

LIST OF FIGURES ...... X

CHAPTER 1. INTRODUCTION ...... 1

Overview ...... 1

Nilgiri Biosphere Reserve (NBR), Western Ghats India ...... 2

Impacts of harvesting from the wild ...... 3

People and forests in India ...... 5

Community based ecological monitoring ...... 6

Research questions and dissertation outline ...... 8

CHAPTER 2 - IMPACT OF RESIN HARVEST ON THE BIOLOGY OF CANARIUM STRICTUM ROXB ...... 10

Introduction ...... 10

Materials and Methods ...... 13

Results ...... 18

Discussion & Conclusion ...... 20

CHAPTER 3 - PHENOLOGICAL PATTERNS IN CANARIUM STRICTUM ROXB. (BURSERACEAE), A RESIN HARVESTED SPECIES OF THE NILGIRI BIOSPHERE RESERVE, WESTERN GHATS, INDIA ...... 29

Introduction ...... 29

Materials and Methods ...... 31

Results ...... 35

Discussion ...... 39

CHAPTER 4. TRADITIONAL ECOLOGICAL KNOWLEDGE OF RESIN GATHERERS OF THE NILGIRI BIOSPHERE RESERVE, WESTERN GHATS, INDIA ...... 55 vi

Introduction ...... 55

Materials & Methods ...... 58

Results ...... 63

Discussion ...... 67

CHAPTER 5. CONCLUSIONS ...... 85

Main findings ...... 85

Contributions to scientific literature ...... 89

Future directions ...... 92

APPENDIX.A. MAPS OF STUDY REGIONS WITH APPROXIMATE LOCATION OF SITES ...... 95

APPENDIX B. PHOTOS OF STUDY SITE, RESIN, RESIN TREE, AND HARVEST METHODS ...... 96

APPENDIX C. DETAILS OF METHODS AND RESULTS FROM THE LINEAR MIXED EFFECTS MODELS ON PREDICTORS OF RESIN HARVEST AND EFFECT OF RESIN HARVEST ON GROWTH, FRUIT PRODUCTION AND SEED GERMINATION ON THE WILD DAMMAR TREE (CANARIUM STRICTUM) ...... 100

APPENDIX D UNIVERSITY OF HAWAIʻI COMMITTEE ON HUMAN SUBJECTS RESEARCH EXEMPTION ...... 102

BIBLIOGRAPHY ...... 103

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LIST OF TABLES

Table.2.1. Characteristic of 89 Canarium strictum trees in relation to harvest methods, percentage harvested, quantity or resin yielded and size of tree across three regions of the Nilgiri Biosphere Reserve ...... 24

Table.2.2. Predictors of harvest status (yes/no) of C. strictum trees from a binomial generalized linear mixed effects model...... 25

Table.2.3. Predictors of quantity of resin harvested/tree in C. strictum trees over a two year period, from a general linear mixed effects model with a negative binomial error structure...... 25

Table.2.4. Predictors of growth of C. strictum trees from a linear mixed effects model...... 26

Table.2.5. Predictors of quantity of fruit produced in C. strictum trees from a generalized linear mixed effects model with a negative binomial error structure...... 27

Table.2.6. Predictors of the probability of C. strictum seed germination from a binomial linear mixed effects model...... 27

Table.2.7. Summary table of predictors of resin harvest and effects of harvest on growth, germination, and fruit production in C. strictum ...... 28

Table.3.1. Characteristics of Canarium strictum trees – Percent of trees in different conditions during study period...... 45

Table.3.2. Effects of size as determined by diameter at breast height, openness of canopy and location on the colour of flush in Canarium strictum trees from a binomial generalized linear mixed effects model...... 46

Table.3.3. Observations of animals dependant on the Canarium strictum tree, flush, flower or fruit ...... 47

Table.4.1. Details of harvester villages, communities, access, quality of resin and market price of resin across four regions in the NBR ...... 75

Table.4.2. Factors affecting resin quality as perceived by resin harvesters from Chamrajnagar region, arranged in fuzzy logic sets ...... 76

Table.4.3. Factors affecting resin quality as perceived by resin harvesters from Chamrajnagar region, arranged in fuzzy logic sets ...... 77

Table.4.4. Factors affecting resin quality as perceived by resin harvesters from Kotagiri region, arranged in fuzzy logic sets ...... 78

Table.4.5. Factors affecting resin quality as perceived by resin harvesters from Nilambur region, arranged in fuzzy logic sets ...... 79

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Table.4.6. Factors affecting number of resin trees as perceived by resin harvesters from Coonoor region, arranged in fuzzy logic sets...... 80

Table.4.7. Factors affecting number of resin trees as perceived by resin harvesters from Kotagiri region, arranged in fuzzy logic sets ...... 81

Table.4.8. Recent changes in resin harvest methods across the four regions in the NBR ...... 82

Table.4.9. Summary of factors perceived by resin harvesters important to resin quality ...... 83

Table.4.10. Summary of factors perceived by resin harvesters important to number of trees...... 83

Table.4.11. Comparison of perceptions in Traditional Ecological Knowledge and findings in Ecological studies in relation to aspects of resin harvest and resin trees .. 84

Table C.1. Specifications of full linear and generalised linear mixed effects models (LMM and GLMM) used to predict resin harvest and effect of resin harvest on growth fruit production and seed germination in C. strictum ...... 100

Table C.2. Estimates and standard errors for main effects and interactions that were non-significant (p>0.05) by likelihood ratio test during model reduction in Chapter 2. All terms were significant in Growth model...... 101

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LIST OF FIGURES

Figure. 3.1. Rainfall (mm) over 2011-12 in three study regions located in the Nilgiris district of the Nilgiri Biosphere Reserve ...... 48

Figure.3.2 Phenology of Canarium strictum trees located in 5 sites in 3 regions of the NBR across a two year period ...... 49

Figure.3.3. Percent of harvested and not harvested trees in flush across three regions of the NBR in 2012-13 ...... 50

Figure.3.4. Percent of harvested and not harvested trees in flower across three regions of the NBR in 2012-13 ...... 51

Figure.3.5. Percent of harvested and not harvested trees in fruit across three regions of the NBR in 2012-13 ...... 52

Figure.3.6. Total amount of resin harvested across two years from Canarium strictum trees in three regions across the NBR ...... 53

Figure.3.7. Distribution of Canarium strictum trees with green and red flush in relation to diameter at breast height ...... 54

B.1. Vegetation characteristics of study sites ...... 96

B.2. Grades of resin found in the region ...... 97

B.3. Characteristics of resin tree, Canarium strictum ...... 98

B.4. Resin harvesting tools and practises...... 99

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CHAPTER 1. INTRODUCTION

Overview

Balancing the goals of human development with conservation of biodiversity gained momentum in the past three decades with the release of the Brundtland

Commission report of the United Nations which described conservation and development as “opposite sides of the same coin” (WCED, 1987). The need to strike this balance by reducing poverty and stemming biodiversity loss became global challenges echoed in the Convention on Biological Diversity and the Millenium

Development Goals (Barrett et al, 2011). Poverty was seen as the cause for environmental degradation though this view has increasingly been discounted in recent times when all the social, political, economic and ecological factors are taken into account (Barrett, et al, 2011; Duraiappah, 1998; Wunder, 2001).

An estimated 1.5 billion people world over are dependent on forests for their livelihoods (Chao, 2012) and forest incomes contribute between 20-27% of the household incomes (Vedeld, et al, 2004; Wunder, 2014). In many forest based livelihood interventions, development goals have been achieved to some level, but not so with conservation goals (Belcher & Schreckenberg, 2007; Morsello,et al, 2012).

High dependence on forests does not necessarily mean high levels of conservation especially in relation to species that are of high conservation priority but not of immediate use (Freese, 1997; Madhusudan & Shankar Raman, 2003). A relook at conservation by defining it in a more robust way to include approach, management, incentive, and knowledge can be seen as a way forward in improving community based conservation (Berkes, 2004).

Associating forest dependence to incomes has meant that cultural practises, healing traditions, wild food consumption, sacred spaces, cultural identity and

1 traditional knowledge benefits from the forest has received less attention. These benefits have a direct link to the quality of forest dependant people’s lives and support a system of knowledge, practise and belief also defined as their traditional ecological knowledge (Berkes, 1999). This knowledge not only helps indigenous communities survive in the forests around them but also defines their cultural beliefs and practices

( Gadgil, 1992).

In my dissertation I seek to understand the impact of harvests and traditional ecological knowledge of, a wild harvested species of cultural and economic importance. I use the case study of the dammar tree (Canarium strictum Roxb.) from which resin (also known as ‘dhupa’ locally and frankincense commonly) is collected for local consumption and trade. I study the effects of harvesting on the ecology and phenology of the species, with specific observations on the timing of leaf flush, flowering, fruiting, and on growth rates, seed germination, fruit production and survival. I explore the traditional ecological knowledge with regard to resin harvest of the indigenous people who harvest the resin. My study seeks to establish linkages between science and traditional ecological knowledge in order to create a foundation for a long term community based ecological monitoring program for a biosphere reserve located within a biodiversity hotspot.

Nilgiri Biosphere Reserve (NBR), Western Ghats India

The Western Ghats, also known as the Sahyadri Hills are a 1600 km long mountain range along the west coast of India and cover approximately 1,40,000 sq km. The region was declared a biodiversity hotspot in 1988 and is under threat from developmental activities related to agriculture and urbanization and has a density of

267 people/sq. kms of area, highest for the biodiversity hotspots of the world

(Cincotta, et al, 2000; Das et al., 2006). A recent report commissioned by the

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Government of India on conservation measures for the Western Ghats has become controversial, as vested interests have portrayed the report as anti- development leading to much debate and civil unrest (Gadgil, 2014)

The Nilgiri Biosphere Reserve (NBR) is part of the Western Ghats and lies between 100 45’N to 12 0 N and 760 E to 770 15’ E with a total area of 5520 sq.km spread across the three southern states of Karnataka, Kerala and Tamil Nadu. The region was designated as a biosphere in 1986 by the Man and biosphere program of the United Nations. Both the Western Ghats and the NBR are home to mega fauna like the Asian Elephant (Elephas maximus) and the tiger (Panthera tigris) and are known to have a long history of human land-use, more than 12,000 years (Subash

Chandran, 1997). In the region of the NBR there are over 20 distinct indigenous (‘ adivasi’) people groups with a total population size of nearly 2,00,000 (Keystone

Foundation, 2007). Many communities live close to the forests and are well recognised for their forest related skills, especially of honey hunting (Demps, et al,

2012; Roy et al., 2011).

Impacts of harvesting from the wild

Plant products from the wild are harvested extensively and intensively

(Shackleton, 2010). NTFP are defined here as goods, other than timber and firewood, of plant origin and derived from forests which may be used directly, such as food, fibre, medicine and construction materials or processed further to yield oils, soap substitutes and other commodities; and these products may be traded for money or bartered for other goods and services (De Beer and McDermott, 1996). Management of non-timber forest products (NTFP) gathered momentum world over as a means of poverty alleviation through localized value addition initiatives, catalysed by research from the Peruvian Amazon by Peters et.al (1989), which showed that the income from

3 sustainable NTFP collection was more than returns from timber harvesting or any other alternative land uses. Increasingly traditional sustainable harvest practises of

NTFP as practised by many communities across the world is being documented and management recommendations have sought to bring those practises on board (Ticktin et al, 2010).

NTFP harvest systems can have ecological impacts at multiple ecological scales, from individuals to ecosystems and consequences of harvest depend on many factors, including the part(s) of the plant harvested, the life history characteristics of the species, the nature and intensity of harvest and/or management practices, and the larger socioeconomic, political and ecological context from which the products are harvested (Cunningham 2001, Ticktin et al, 2012). NTFP harvest sustainability requires not only that NTFP populations are able to persist over the long-term, that but also that harvest does not negatively affect community and ecosystem functions.

In reality, both the management and ecological impacts of harvesting any given species can vary enormously across space and time (e.g. Nantel et al. 1996; Siebert

2000; Gaoue & Ticktin 2007). High dependence of local communities on NTFP has been reported to be a cause for loss of forest cover and forest degradation in the

Western Ghats (Davidar 2008, Karanth et al. 2006). However recent studies in the region have also shown that harvest of NTFP has less detrimental effects on the population dynamics of the species than previously thought (Ticktin et al, 2012) and there is a need to consider the multiple processes that may be affecting harvested populations (Mandle et al, 2013).

Plant exudates like gums and resins are important NTFPs that have a number of uses in medicine, ritual, industry and early records of its use are reported from

Arabia-1000 BC and Mayan civilisation - 600 BC (Langenheim,2003). Trees produce

4 resin for a number of reasons, primarily to avoid auto toxicity, defend themselves from parasites, and prevent herbivory (Langenheim, 2003). Resins also play a role in attracting pollinators and seed dispersers (Langenheim, 2003). Research has shown that harvesting resin impacts conduction of nutrients and hormones involved in the production of flower buds (Primack 1987); reduces number of fruits and viable seeds

(Rjikers 2006) and keeps populations in a vegetative state (Cunningham & Mbenkum,

1993; Cocks & Dold, 2004). In contrast some studies show that removal of resin or other exudates including latex, did not affect the flower or fruit production (Bitariho et al 2006; Schumann et.al 2010; Bladauf et al 2014).

Canarium strictum Roxb., is a heavily harvested Indian resin tree, which produces the resin known as ‘black dammar’. C. strictum is a semi evergreen tree, found in the wild and rarely in cultivation, distributed unevenly along the Western

Ghats and other parts of India. Information about its conservation status nationally or globally is lacking, though at the level of the region C. strictum has been reported to be a species of conservation concern (Ravikumar et al, 2000). Typically, resin is either collected opportunistically from trees where is has gathered, or by making incisions and returning to collect resin after a period of time. In some regions, harvesters set fire at the base of the tree to increase resin production before making incisions (Varghese et al 2008).

People and forests in India

India has only 2% of the world’s forests and nearly 21% of the country’s total land cover is under forests (Varadarajan, 2014). An estimated 100 million people are forest dependent in India (Kabra, 2009) of which close to 89 million are indigenous or adivasi people who have close cultural and economic links with the forests (Census of

India, 2010). The National Medicinal Plant Board of India estimates about 960

5 species of medicinal plants are used in trade, of which 178 species have annual consumption levels in excess of 100 metric tonnes and exports in the range of INR 10 billion (http://nmpb.nic.in/index.php). The bulk of the published research on wild plant harvests in the Western Ghats, show that the harvested species are on the decline

(Ravikumar & Ved 2000).

In the NBR studies have shown that upto 40% of incomes of indigenous people come from the forests (Pain, 2009). This high use of the forests linked to conservation has been identified as a priority research area in tropical conservation

(Bawa et al, 2004). Participation of forest-dwelling indigenous communities in the conservation process is undergoing a rapid change in the Indian context due to the recently implemented Scheduled Tribes and Other Traditional Forest Dwellers

(Recognition of Forest Rights) Act, 2006 also referred to as the Forest Rights Act of

2006. The Act broadly provides for two sets of rights – land rights, both private and common, and community rights over forest resources. Community rights over forests include rights over grazing lands, NTFP collection areas and other common property resources. The Act also provides for declaration of critical wildlife habitats and the preservation of threatened species or habitats. This law recognises the local communities rights to manage natural resources and biodiversity and provides opportunities to jointly manage biodiversity along with the state (Bawa, et al, 2011).

Community based ecological monitoring

Indigenous users of natural resources are guided by their traditional ecological knowledge which deals with complexity through rules of thumb and indicators which are used to simplify the questions, consistent with fuzzy logic thinking (Berkes 2009).

Fuzzy logic models allow management of complex non-linear systems traditionally with industrial process and more recently with ecological, economic and geophysical

6 systems by offering more opportunity to use human judgment (Zadeh 2008). Fuzzy logic uses language variables ‘And If Then’ rules to bring precision into imprecise information (Zadeh 2008).

Monitoring has been defined as “the systematic measurement of variables and processes over time” and “assumes that there is a specific reason for that collection of data, such as ensuring that standards are being met” (Spellerberg, 2005). Monitoring is one way that conservationists rely on traditional wild produce harvester communities to be partners in sustainability, although this alone may not be enough and we need to find a complementarity between science and the traditional ecological knowledge of the harvesters (Moller et.al 2004).

To identify the effects of anthropogenic factors on ecological process it is important to monitor them over the long term. Phenology of plants, pollinator preferences and migratory patterns in birds are some of the more common processes that have been monitored. Biologists have relied a lot on participation of amateur naturalists, citizen groups and interested individuals from all walks of life

(Greenwood, 2007; Ledneva, Miller-Rushing, 2004). There is a need to link monitoring with local decision making processes, which in turn will make the monitoring sustainable and effective (Danielsen, 2005) . In developing countries fewer resources are available for these voluntary monitoring schemes and therein lies the challenge in designing effective, cost efficient methods (Danielsen et al 2005).

Locally based monitoring may be the ideal way for communities to report to external agencies on progress with managing habitats, species, and flows of ecosystem services in community or co-managed conservation areas (Danielsen et. al 2008). An opportunity in conservation-linked field studies lies in the use of local and indigenous knowledge, ensuring benefits to the people while meeting the goals of conservation.

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In the past, ecological monitoring was an integral part of traditional resource management practices in indigenous communities, many of which involved adaptive management and were highly effective at conservation (Moller et al, 2004).

Research questions and dissertation outline

In order to investigate the effects of harvest on the ecology and phenology of a wild resource and explore how the harvesters of the region understand the ecology of the species, I present a case study of C. strictum to ask :

1. What affects the probability that a tree (C. strictum) will be harvested for resin

or not? How does harvest impact the vital rates (growth, reproduction and

survival) of C. strictum?

2. What are the regional variations in phenology of the C. strictum, and does

harvest impact the phenology?

3. What is the traditional knowledge that exists about C. strictum and how can it

form part of ecological monitoring?

Chapter 2 of this dissertation deals with the impact of harvest on the vital rates of the wild dammar tree. I present the results of a two year study on drivers of resin production, resin harvest, growth, fruit production and germination in 89 C. strictum trees located in five locations across the NBR.

In Chapter 3, I present the results of observations done on the phenology (leaf flush, flowering, fruiting) of C. strictum trees and compare patterns between harvested and not harvested species.

In Chapter 4, I use the fuzzy logic approach to present the results of discussion that were held with resin harvesters on their knowledge in relation to resin quality and trees. This chapter also examines how this knowledge is changing and adapting

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In Chapter 5, I conclude by synthesizing my findings from the preceding chapters and discuss their implications for the conservation of a lesser studied species by including harvester communities in the monitoring of the species and its habitat. I summarise the contributions of my research to the fields of conservation biology and human ecology, discuss the limitations of my study and suggest future lines of research for integrating traditional knowledge in biodiversity conservation.

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CHAPTER 2 - IMPACT OF RESIN HARVEST ON THE BIOLOGY OF

CANARIUM STRICTUM ROXB

Introduction

Plant products from the wild have been harvested extensively and intensively across the world (Cunningham, 2011). The practice of collecting non timber forest products (NTFP) is an important part of the lives of some of the most marginalized communities of the world (Shackleton et al 2011). NTFP are defined here as goods, other than timber and firewood, of plant origin, derived from forests which may be used directly, such as food, fibre, medicine and construction materials or processed further to yield oils, soap substitutes and other commodities and these products may be traded for money or bartered for other goods and services (De Beer and

McDermott, 1996). Management through localized value addition of NTFP gathered momentum world over as a means of poverty alleviation, catalysed by research from the Peruvian Amazon by Peters et.al (1989), which showed that the income from sustainable NTFP collection was more than returns from timber harvesting or any other alternative land uses. Harvest of NTFP is reported to be the cause of population decline in a number of plants globally (Brummit and Bachmann, 2010). However, many NTFP, including those harvested for national and international sales, can be harvested sustainably (Ticktin & Johns, 2001; Schmidt & Ticktin, 2012; Schmidt et al. 2011).

Sustainable harvest of NTFPs can only be possible if the impacts of harvest at multiple ecological scales, from individuals to ecosystems, part(s) of the plant harvested to its life-history characteristics are taken into account (Ticktin, 2004).

Management practices, socio-economic, political and ecological context from which the products are harvested need to be considered when assessing the impacts of NTFP

10 harvest (Cunningham, 2001; Ticktin and Shackleton, 2011). For NTFP harvests to be sustainable it is critical that targeted plant populations are able to persist over the long-term, but also that harvest does not negatively affect community and ecosystem functions. In reality, both the management and ecological impacts of harvesting any given species can vary enormously across space and time (e.g. Nantel et al. 1996;

Siebert 2000; Gaoue & Ticktin 2010).

Plant resin, as an NTFP, has an important societal value and has been part of human cultures for a long time. Plant resin is used as such in medicine, and rituals, and processed for industrial uses related to the paint and varnish industry

(Langenheim, 2003). Trees produce resin for a number of reasons, primarily to avoid auto toxicity, defense from parasites and herbivores and also to attract pollinators and seed dispersers (Langenheim, 2003). The high cost of production to the plant, of this primary metabolite is offset by the benefits of pollination, dispersal and defense from pathogens for the tree. Harvesting or removal of resin, can impact conduction of nutrients and hormones involved in the production of flower buds (Primack 1987); reduce number of fruits and viable seeds (Rjikers 2006) and keep populations in a vegetative state (Cunningham & Mbenkum 1993, Cocks & Dold 2004). In contrast some studies show that removal of resin or other exudates including latex does not affect the flower or fruit production (Bitariho et al 2006; Schumann et.al 2010;

Bladauf et al 2014). However, this has not been widely studied and the general patterns remain unclear. Collection of plant exudates, if harvested in prescribed ways

(either by traditional knowledge or scientific practices), has the least impact on the species and its populations as compared to harvest of other vegetative or reproductive parts (Peters, 2001).

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Worldwide, resin has traditionally been harvested by tapping the tree which is a controlled wounding of the tree (Langenheim, 2003). Resin flow in lodge pole pine

(Pinus contorta) is influenced by wound size, diameter of the tree and tree health

(Nebeker et al, 1995). Studies that have looked at traditional practices of resin collection from South East Asia, Amazon and Africa show that resin production is highly dependent on tree size and crown diameter (Ella and Tongacan, 1987; Newton et al, 2010; Eshete et al, 2011). Traditional methods of resin harvest in South East

Asia involve the use of fire and tapping (Ibrahim et. al., 1987; Ella and Tongacan

1987). In the Phillipines, resin of Canarium luzonicum is harvested commercially and though it can be collected throughout the year from the rain forest, it is preferably harvested in the wet season when there is a greater resin flow (Langenheim, 2003).

Canarium strictum Roxb., is an important and heavily harvested Indian resin tree, which produces the resin known as ‘black dammar’. Resin from C. strictum is harvested for local consumption as well as for sale and the species is subjected to different types of harvest methods and frequency across the regions where it is found

(Varghese & Ticktin 2008). Typically, resin is either collected opportunistically from trees where it has collected naturally, or by making incisions and returning to collect resin after a period of time. In some regions, harvesters set fire at the base of the tree to increase resin production before making incisions (Varghese and Ticktin, 2008).

The goal of our study is to identify factors that predict patterns and quantities of C. strictum resin harvest, and assess the effects of resin harvest on vital rates

(growth, survival, reproduction) of the tree, across three regions of the our study area.

We address the following questions: 1) What affects the probability that a tree will be harvested for resin or not? 2) What are the predictors of resin quantity – that is, why do some trees yield more resin than others? 3) What effect does resin harvest, along

12 with tree size, flush colour, fruiting levels, canopy openness and location have on survival, growth, fruit production and seed germination?

We expected to find that location affected the status (harvested or not harvested) of the trees in our study area, since in two regions harvest involves incision making, while in the third it involves only gathering resin that has formed naturally.

We also expected that resin harvest has negative effects on the growth, reproductive output (number of fruits produced) and germination rates of seeds, owing to competition for resources between these processes and resin production.

Materials and Methods

Study species and site

Canarium strictum, Roxb is a lesser studied species in the wild and is found in semi evergreen, moist deciduous and riparian forests. C. strictum populations are distributed across the Western Ghats, parts of the Eastern Ghats, North East India into

Burma (Shashidharan, 2006). The tree is reported to be present up to an elevation of

1300m (Shashidharan, 2006). The flowers are polygamous (Gamble, 1915), though local indigenous people maintain that there is a male and a female tree. Individual trees can grow upto 50m (per.obs) and flush, flower and fruit gregariously. Trees are located in clusters, and all along the Western Ghats, the tree has been harvested for its highly valued resin which is sold as frankincense in the local markets.

Our study was conducted in the Nilgiri Biosphere Reserve (NBR) of the

Western Ghats biodiversity hotspot (Mittermerer et al 2005). We selected three regions, Coonoor, Kotagiri and Gudalur, based on prior knowledge on location of trees and distinct resin harvesting practices. All the regions selected were in semi evergreen and riparian forest patches with an elevation ranging from 900-1200m above sea level. The three regions had slight rainfall variations- Coonoor and Kotagiri

13 regions receive their maximum rainfall during the October-November period while

Gudalur region receives peak rainfall in July- August (Chapter 3). Trees in Gudalur region were located in a coffee plantation which was more than 50 years old and has evergreen forests adjoining the area. Within the plantation, C. strictum trees were left to grow as shade trees and as support trees for pepper vines. In Coonoor and Kotagiri,

C. strictum was found as a secondary pioneer, often occupying tree fall gaps and natural clearings in the forest. A fair amount of seedlings and saplings were found in the under storey around the adult trees. All these regions were relatively closed to disturbances from fire, grazing (domestic cattle), invasive species like Lantana camara and could be considered relatively undisturbed forests. Other dominant species found growing along with C. strictum are Elaeocarpus spp., Schelichera oleosa, Mesua ferrea, Persea macarantha, and Garcinnia gummi-gutta.

Resin harvesting practices

Harvesting of resin is an ancient practice that has been part of the ritual and traditions of the indigenous people of the NBR. The three main communities who were observed harvesting the resin belonged to the Irula, Kurumba and Kattunayaka people. In Kotagiri and Gudalur harvesting is done using tools like small axes and sharp curved knives. In Gudalur, it was the practice to set a low intensity fire at the base of the tree using dry leaf litter and burning the bark before making the incisions.

This practise has stopped since 30 years now, because of strict rules about starting fires imposed by the forest authorities. In Kotagiri and Gudalur incisions for tapping resin were observed to be made nearly three feet above the root and were left open for the resin to form. In Coonoor harvesting is practiced without tapping or causing any damage to the tree bark. Natural deposits of resin on very large trees collect at the forks of branches or natural cracks in the bark and these were collected as soon as

14 they had hardened and were not sticky, by which time they were black in colour and varied from 5cm to 15cm length (per.obs).

Selection of sites

We selected C. strictum trees above 25cm dbh in two sites each within a region with the exception of Gudalur region where we were able to locate only one site. We used 25 cm dbh as a cut-off because according to harvesters, only trees above this approximate girth are harvested. Sites were selected based on harvester’s prior knowledge of the area - the trees are not found evenly distributed in the forests so it was important to locate them with the help of the harvesters. In each site we looked for a minimum of 10 trees of which at least five were subjected to regular harvest and others were not harvested. In the Gudalur region we were able to locate 20 harvested and not harvested trees in total. Trees were also selected based on access for monitoring, and this was a challenge in the Coonoor region where the access was difficult even for the local people because of the terrain.

Data collection

In January 2012, a total of 89 trees across the three regions (Table.2.1) were tagged and measured for dbh at the beginning of the study and subsequently every six months for two years. We recorded the openness of the canopy (ranked as open, partially open, and closed), height of the tree, and history of harvest of each tree.

Every 15 days, presence of absence of leaf, flush, flower, and fruit were recorded, as well as flush colour. The bark of the tree was closely observed for fresh incisions made for harvest of resin. The quantity of resin harvested was estimated based on information from field assistants, who were also local resin harvesters in each region.

We undertook resin harvesting visits in all our regions to make independent assessments of resin weights and amounts per tree, this helped in validating the

15 estimates that were being made as part of our fortnightly data collection. In 2012,

May and June we counted the number of fruits produced by all fruiting trees. An estimate of fruit counts was done by portioning the tree crown into eight sections, counting the fruit on five random branches in each section and estimating number of fruiting branches for each section. Total fruit produced for the tree was then estimated as a multiple of number of fruits/branch times number of branches times number of sections. This estimate was always done by the principal investigator in order to standardise counts.

Germination experiments

In 2013 between July and October, seeds were collected from all harvested

(n=27) and not harvested (n=21) fruiting trees of the five sites in three regions. Seed collection was opportunistic and we collected a minimum of 50 seeds from each individual tree from the forest floor for the purposes of our experiments. A total of

253 seeds from Kotagiri, 101 seeds from Coonoor and 142 seeds from Gudalur were used as part of the experiment. Of these 239 seeds were from harvested trees and 257 from not harvested trees. Seeds were collected from directly under the crown and closer to the tree so as to rule out the possibility of a seed being from the neighbouring tree. We used a seed germination method which has been tried and perfected at a local organisation that undertakes native species planting. Seeds were treated by soaking in cold water for 7 days and left to dry in the sun for up to 3 hours, by which time the stony exocarp has split, after which the seeds are buried in nursery beds with a 1:3 ratio of soil to sand (L. Rajendran, unpublished nursery manual).

Germination tents were made with UV treated plastic sheets to maintain temperature and moisture conditions. A total of 490 seeds were planted and the seeds were watered every second day, and the number of seeds germinated were recorded at the

16 same time. When the radicle emerged from the soil, that was taken as the first day of germination and emerging seedlings were monitored for survival over a period of 6-8 weeks.

Data analysis

We used linear and generalised linear mixed effects model (LMER and

GLMER) to model resin production, harvesting status (yes/no), growth, germination, and fruit production of our study species (Table.C.1.) We first tested for multicollinearity between predictors and found none of our variables to be significantly correlated. We considered site within region as a random effect and since this was based on spatial configuration of our study design, this was retained in all our models. To model the probability of germination, we considered tree within site within region as a random effect. Fixed explanatory variables included region, total number of incisions made on the tree, total number of times a tree was harvested, canopy openness, dbh (log transformed), flush colour (green or red) and ranked fruiting intensity. Two way interactions of the main effects, between region, dbh and flush colour, were also included in the full model. Ranked fruiting intensity included none (trees produced flowers but did not fruit at all), low (trees produced very low numbers of fruits not more than 20 per tree, and only on one or two branches) and normal (trees produced more than 100 fruit). Fruit counts were found to be less than

20 or more than 100. To model the probability of tree survival, seed germination and tree being harvested for resin, we used binomial GLMMs. For resin quantity and fruit production, we used GLMMs with a negative binomial error distribution (Table C.1.).

For growth, we used a Gaussian error distribution. Fruit production data was modelled using the data from fruit producing trees (n=48). Resin quantity data was modelled using the data from resin harvested trees (n=26). All other models were based on data

17 from 89 trees. With the exception of fruit production, which was recorded only for

2012, and seed germination recorded only for 2013, the rest of the data were modelled over a two year period. Full models were reduced in a backwards stepwise process , sequentially dropping the least significant fixed effect term in the model, and looking at the AIC values. All analyses were completed in R.2.13.1(R Development Core

Team 2011) using the ‘lmer’ function in the lme4 package (Pinheiro et al 2011).

Results

Factors affecting resin harvest status and quantity

Nearly 50% (n=36) of the trees in the Kotagiri, 40% (n=20) in Gudalur and

20% (n=33) in Coonoor regions were harvested for resin in two years (Table.2.1). We found that region, total number of incisions, and flush colour were significant predictors of harvest status (Table.2.2). Region, in our study represented harvesting method and we found trees in Kotagiri and Gudalur, where harvest was done with incisions, were harvested significantly more than those in Coonoor. Trees that produce green flush were more likely to be harvested than those with red flush but when they were in a higher size class. There was a moderately significant interaction between size of the tree and flush colour (Table.2.2.). Fruiting intensity varied widely across regions and was not significantly correlated with harvest status (details in

Appendix C, Table.C.2).

In the two years of our study, we observed high variation in total resin quantities that were harvested, ranging from 10kg in Kotagiri region to 44kg in

Coonoor region (Table.2.1.). At the individual tree level, annual resin yields ranged from 0.1 to 18kg across regions. Size of the tree and number of new incisions made were significant predictors of the quantity of resin obtained (Table.2.3.). More incisions and larger trees yielded higher amounts of resin. Flush colour, fruiting

18 intensity and size of the tree were not significant predictors of quantity of resin harvested (Appendix C, Table.C.2.).

Effect of resin harvest on vital rates of C. strictum

The size of resin producing trees in our study sites ranged from 25cm to 142 cm dbh across the five sites in three regions. Average dbh (cms) sizes of the resin trees were 65±23.5 in Coonoor, 73±23.6 in Gudalur and 46±16 in Kotagiri

(Table.2.1). The final model for growth included all of our predictor variables even though their effect sizes were very small (Table.2.4). The variable with the biggest effect sizes was initial size (dbh of year one). Trees that produced lesser fruits and were in open canopy conditions were found to have lower growth rates. Tree size and red colour flush showed a negative interactive effect on growth of the tree.

Across the 48 trees that produced fruit in 2012, the mean (±SD) number of fruits counted was 8 ±7 in trees that fruited at a ‘low’ level and 470±296 in trees that fruited ‘normally’. Tree size and resin harvest status (harvested and not harvested) significantly predicted the probability of germination. Trees with red flush also produced significantly more fruit than trees with green colour flush. Finally the harvested trees produced significantly more fruit than not harvested trees. There was no significant interactive effect of flush colour and tree size on fruit production.

Region and canopy status were not significant predictors of number of fruits produced

(Appendix C, Table.C.2).

Seed germination rate, mean percentage (±SE) was significantly higher in harvested trees, 48% (±6.9) than in not harvested trees, 23% (±4). We found that size of the tree, and harvest status (harvested and not harvested) were significant predictors of the probability of germination. Seeds that were collected from trees that were harvested for resin showed higher germination rates than seeds collected from

19 trees that were not harvested for resin. Seeds collected from larger trees had a lower germination probability than those from smaller trees.

Over our two year observation period, six out of 89 trees died, all of which were harvested for resin. These dead trees were larger trees with dbh ranging from

40-140 cm. While two out of the six died because of other trees falling on them, four of them started to dry up and fell in the rain period. Even as we finished our monitoring in January 2014 we noticed that one more tree which was near a stream but was also a harvested tree, had started to show signs of drying up. However, none of our explanatory variables were significant predictors of the probability of survival.

Discussion & Conclusion

Our study on C. strictum harvests in the NBR show some variation across regions in terms of resin harvest but harvesting of resin did not have significant negative effects on the vital rates of the tree (Table.2.6).

Selection of trees to harvest resin

Tree characteristics may be an important determinant to the decisions governing harvest and these are based on observations and experiences of the harvesters who live close to the trees. In this context it is interesting that our results indicate that when the method of harvest is intensive as in the case of Kotagiri and

Gudalur where harvesters use incisions, trees were more likely to get harvested than those in Coonoor, where only resin naturally produced on the tree is removed. In the case of harvest of an oleo resin from Copaifera sp, environmental and morphological factors restricted productivity and not all trees produced amounts of oleo resin with similar regularity (Newton et al, 2010). However frequency of tapping, and size of the tree were shown to be significant predictors of resin production in Boswellia papyrifera in dryland forests of Africa (Eshete et al, 2011). This is likely to explain

20 some of the variation in our results. Size of the tree alone does not influence whether the tree will get harvested or not (Table2.6.), but size of the tree matters when the quantity of resin is taken into account; significantly larger volumes of resin were harvested from larger trees. This is the strategy that the resin harvesters of Coonoor employed, to wait for the tree to grow big and yield resin naturally. While Kotagiri and Gudalur harvesters tapped more trees, they did not actually obtain more resin at the stand level than did harvesters in Coonoor (Table 1). Although our models showed that at the individual level there was no significant difference in resin yield/tree across regions, this may be a result of the low sample size of trees harvested in Coonoor within the two year study period (Table 1).

The explanation for our findings that trees with green flush were significantly more likely to be harvested than those with red flush is not clear, since red flush apparently does not require more resources and is a protection mechanism against herbivores by keeping young leaves virtually devoid of nutritive value (Dominy et al

2013). This merits further investigation and has also been discussed further in Chapter

3.

Effects of harvest on vital rates

The trade-offs between growth, reproduction and defense has been a subject of much interest to evolutionary biologists. When resources like resin are removed from trees, the stress caused may result in poorer reproductive output and changes in their biology (Cunningham & Mbenkum 1998; Rjikers, 2006). In the case of resin extraction from Boswellia papyrifera Rjikers et al (2006) showed that trees that were subjected to experimental tapping produced fewer flowers, fruits and seeds. In their study on resin producing species Boswellia papyrifera, Groenendijk et al (2011), showed that poor regeneration in their study species was unlikely due to harvesting

21 and was driven by fire, grazing and beetle attacks. Eshete et al (2011) showed that seeds from trees that were untapped for resin in Boswellia papyrifera had a significantly higher percentage of germination that the tapped trees. On the other hand, Baldauf et al (2014), in their study on latex extraction (which includes bark defoliation), show that these harvests have a significant positive effect on fruit production of their study species in Brazil. Our results show that resin harvest did not have any significant negative effect on the growth of the trees, and that harvested trees had significantly higher fruit production and germination than non-harvested trees (Table.2.6). One explanation could be that trees that are able to produce resin have more resources than those that do not produce resin, and therefore have higher levels of fruit production and produce seeds that have higher probability of germination. Zuidema et al. (2009) and Jansen et al. (2011) have shown that for populations of some trees and palms, vital rates of some individuals are significantly greater than those of others, and that these ‘super performers’ therefore contribute disproportionately to population growth. This could also be the case with C. strictum and merits further research.

Implications for sustainable management

Our research suggests good potential for the sustainable harvest of C. strictum.

Resin harvest as practiced across the three study regions, and including both tapping and opportunistic harvest of resin naturally produced, does not appear to have negative effects on growth, fruit production or germination. However, although our models showed no significant effects of harvest on mortality, it is possible that this may be due to sample size and the relatively short duration (two years) of our study.

It is noteworthy that all six of the trees that died in our study were harvested, and that other authors have reported (but not tested) that harvested trees appear to fall easily in

22 the wind (pers comm – resin harvesters). Even if harvest leads to early mortality, management that includes protecting and out planting saplings, could alleviate potential negative effects. The potential for out planting efforts exists as local organizations in the region already grow C. strictum seedlings for distribution to communities.

Our study was limited in that it was not possible to test the effects of different harvest practices within the same region. Thus comparisons between tapping with incisions (Kotagiri and Gudalur) and natural harvest (Coonoor) are confounded by other potential differences between these regions. These kinds of tests would be possible at field stations and could yield further insight on optimal strategies for harvest. Ultimately though, decisions about harvesting practises are influenced by social factors too (Chapter 4). Overall our two year study provides a baseline for a lesser studied species, and a type of harvest (exudate harvest) for which very little is known. In order to understand fully the dynamics of harvested species under different abiotic and biotic conditions there is a need for long term monitoring of harvested populations which could contribute to conservation aims for the species and the ecosystem.

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Table.2.1. Characteristic of 89 Canarium strictum trees in relation to harvest methods, percentage harvested, quantity or resin yielded and size of tree across three regions of the Nilgiri Biosphere Reserve

Total resin Average Trees Average Total Trees quantity size of harvested size of all Region Harvest method trees harvested harvested harvested in both years trees dbh observed (%) in 2 years trees dbh (%) (cms) ±SD (kgs) (cms)±SD Coonoor Natural 36 12 5 44 65 (±23.5) 77±22 Incision Gudalur (fire 30 20 40 30 20 73(±23.6) 80±27 years ago) Kotagiri Incision 33 45 27 10 46(±16) 46±17

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Table.2.2. Predictors of harvest status (yes/no) of C. strictum trees from a binomial generalized linear mixed effects model.

Fixed effects Estimate SE Z value p-value Intercept -14.4565 5.6567 -2.556 0.010599 Region Gudalura 1.3859 0.6867 2.018 0.043570 Region Kotagiria 2.4153 0.7108 3.398 0.000679 Log dbh1 3.2590 1.3155 2.477 0.013234 Flush colour redb 11.0031 6.2814 1.752 0.079827 Logdbh1:Fl.cl.red -2.8153 1.5165 -1.856 0.063389

Random effects SD

Site 1.324e-05 aAs compared to Coonoor region bAs compared to flush colour green

Table.2.3. Predictors of quantity of resin harvested/tree in C. strictum trees over a two year period, from a general linear mixed effects model with a negative binomial error structure.

Fixed effects Estimate SE t value p-value Intercept 0.582786 0.327410 1.780 0.075078 Total new incisions 0.034091 0.008248 4.133 3.57e-05 Logdbh1 0.254241 0.073203 3.473 0.000514

Random effects SD site 1.255e-07

25

Table.2.4. Predictors of growth of C. strictum trees from a linear mixed effects model.

Fixed effects Estimate Std Error t-value (Intercept) -0.06944 0.06222 -1.12 Dbh (log transfrmd) 1.019602 0.014621 69.74 Region Gudalura 0.007031 0.0084 0.84 Region Kotagiria 0.003689 0.008764 0.43 Total resin quantity (log transformed) 0.000103 0.00064 0.16 Canopy openb -0.00655 0.011122 -0.59 Canopy partialb 0.00027 0.01119 0.02 Flush redc -0.05227 0.073534 0.71 Fruit level normald 0.003053 0.009742 0.31 Fruit level lowd -0.00736 0.008383 -0.88 Logdbh:flcolred -0.01356 0.017819 -0.76

Random effect SD Site 0.0 a Compared to Coonoor. bAs compared to closed canopy cAs compared to flush colour green dAs compared to fruit level none

26

Table.2.5. Predictors of quantity of fruit produced in C. strictum trees from a generalized linear mixed effects model with a negative binomial error structure.

Fixed effects Estimate SE Z-value p-value

Intercept -6.1152 1.6709 -3.66 0.000252

Log of dbh 2.5407 0.3919 6.483 8.98e -11 Harvested treesa 0.7377 0.2966 2.487 0.012867 Flush colour redb 1.0962 0.3316 3.305 0.000949

Random effects SD site 0.00E+00 aAs compared to not harvested trees bAs compared to flush colour green

Table.2.6. Predictors of the probability of C. strictum seed germination from a binomial linear mixed effects model.

Fixed effects Estimate SE Z-value p-value Intercept -5.4836 2.9640 -1.850 0.0643 Log of dbh 1.3515 0.7073 1.911 0.0560 a Harvested trees 0.9794 0.4089 2.395 0.0166

Random effects SD tree.id: site 5.957e-01 site 5.631e-05 aAs compared to not harvested trees

27

Table.2.7. Summary table of predictors of resin harvest and effects of harvest on growth, germination, and fruit production in C. strictum

Predictor Region Region Harvest Total Canopy Canopy Fruit Fruit Flush Total Total variables Size Gudalura Kotagiria (Y/N) resin openb Partialb level level colour number times Dbh Harvested Lowc Normalc Red d incisions harvest Response (cms) (gms) variables

Harvest status + * + + + * Resin quantity + + Growth + # + + + - + - + - # Fruit production + + + Germination - + aCompared to Coonoor region; bCompared to close canopy; cCompared to fruit level none; d Compared to flush colour green *size and flush colour are slightly correlated, the interaction between these terms has a negative effect on probability of a tree to get harvested #size and flush colour are correlated, the interaction between these terms has a negative effect on overall growth

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CHAPTER 3 - PHENOLOGICAL PATTERNS IN CANARIUM STRICTUM

ROXB. (BURSERACEAE), A RESIN HARVESTED SPECIES OF THE

NILGIRI BIOSPHERE RESERVE, WESTERN GHATS, INDIA

Introduction

For centuries human beings have used changes in plants to forecast seasons, predict fruit yields, and determine animal migrations (Lantz et al, 2003). The field of science dedicated to the study of seasonal biological activities and their responses to environmental variation is phenology (Rathcyke et al, 1985). For plants, phenological events and phases are an important part of their life cycle, from reproduction to growth to maturing of fruit to leaf senescence. Phenological events have to be synchronised as in the case of seed and pollen dispersal, which is vital to reproduction, and is linked to the arrival of pollinators or seed predators/dispersers

(Van Schlaik et al, 1993). Many long term observation studies have shown that the timing and intensity of these phenological events are sensitive to temperature, light, precipitation and snowmelt dynamics (Chapman et al 2005; Alexander et al 2011;

Clinger et.al 2013). The links between phenology, and climate, and more recently climate change exist at the global scale (Parmesan and Yohe, 2003; Root et al., 2003).

In the first attempt globally to correlate present day flowering records with flowering records from museums and herbaria, Primack et. al. (2004) were able to demonstrate a significant response of plant flowering time to changing spring temperatures over the past century, specifically, that plants are flowering earlier because of warmer spring temperatures.

Besides climate change, other anthropogenic factors affect the phenological patterns in species. Periodic harvesting, grazing and fires have been shown to affect the flowering, fruiting and production of new leaves in many harvested species the

29 world over (Ticktin, 2004; Gaoue et al, 2008; Schmidt. 2011; Mandle et al, 2013).

The harvest of non-timber forest products (NTFPs) such as fruit, bark, roots, and resins, was shown as a significant contributor to declines in plant populations in many tropical forests (Brummitt et al, 2010). In a review paper on non timber forest produce harvesting, Brites and Morsello (2012) showed that 67% of studies evaluating the reproductive rates of harvested species reported a negative impact of harvesting. In their study on the effect of tapping for frankincense from Boswellia papyrifera, Rjikers et al (2006) show, that in heavily harvested trees, flower and fruit production was limited and non-viable seeds were produced.

Plants produce leaves, flowers, fruits and seeds at particular times of the year for important functions in their lifecycle (Rathcyke et al, 1985). In the case of exudates like resin, which are produced by plants, they are important as defence for injured tissues and to prevent attacks from insects and fungi (Langenheim, 2003).

Many plants in the tropics keep their young leaves virtually devoid of nutritive value till they reach full size, also called the delayed greening strategy, in order to protect them from herbivores (Kursar and Coley, 1992; Van Schlaik 1993; Dominy 2013).

The anthocyanin pigments present in young leaves are acidic therefore unpalatable to herbivores (Karageorgou et al, 2006). These pigments cause the leaves to look red and thereby making them invisible to most invertebrate herbivores, who lack a visual receptor for red (Dominy, 2013).

While rates of production (of leaf, flower, fruits and seeds) have been studied in relation to harvest pressure, lesser number of studies have been undertaken on effects of harvesting practises on the timing of various phenological events (for example Baldauf, 2014). The first ecological study on black dammar (Canarium strictum Roxb., Burseraceae), an important resin-producing medicinal species in the

30

Western Ghats in India showed variations in population structure based on harvest types (Varghese and Ticktin, 2008). Several authors have expressed concerns that populations of C. strictum are disappearing due to tapping practices (Kannan, 1992 ;

Augustine and Krishnan, 2006). Fruits of C. strictum are reported to be dispersed by large birds (Ganesh and Davidar, 2001), including some species of endangered hornbills (Bucerotidae) (Kannan, 1992). Canarium fruits are the second most important species, next to figs, in the diet of more than 16 species of hornbills

(Kitamura, 2011).

Our study looks at the phenology C. strictum Roxb. and provides details of the phenology (flush, flower and fruit timings) and life history of C. strictum across three regions of the Nilgiri Biosphere Reserve (NBR). We further explore differences in phenology between harvested and not harvested C. strictum trees across the regions.

We do this by asking the following questions 1) What is the timing of flush, flower and fruit events and does this vary across regions; 2) How do these events vary between harvested and not harvested individuals; 3) What intraspecific variations exist in the flush colour of C. strictum and what factors can explain this variation?; 4)

What animals are dependent on the flush, fruit and flower of the tree?

Materials and Methods

Study site

The Nilgiri Biosphere Reserve (NBR) is part of the Western Ghats chain of mountains of the Indian peninsula, and lies between 100 45’N to 12 0 N and 760 E to

770 15’ E with a total area of 5520 km2 spread across the three southern states of

Karnataka, Kerala and Tamil Nadu (Prabhakar, 1994). Altitude varies from 250m to

2650m, and at least four of the major rivers of south India originate in this region - the

Bhavani, Moyar, Kabini and Chaliyar rivers. The region is home to endemic and

31 endangered fauna like the Lion Tailed Macaque (Macaca silenus), Bengal Tiger

(Panthera tigris), King Cobra(Ophiophagus Hannah), Asiatic Elephant (Elephas maximus), Great Indian Pied Hornbill (Buceros bicornis) (Daniels, 1995). The floral diversity is unique to the region especially in the forests of the high elevation which have several species and genera that are endemic and endangered (Keystone, 2007).

The range of topography and climate has resulted in sharp gradients of vegetation composition, ranging from thorny scrub forest dominating the north-eastern region and intergrading westwards into dry and moist deciduous forests and wet evergreen forests towards the western parts of the reserve (Prabhakar, 1994).

Study species

Canarium strictum, Roxb. is a large tree, buttressed, up to 30 m tall with a clear bole trunk, brownish bark, with terete branchlets which are ferruginous tomentose. Leaves are compound, imparipinnate, alternate, spiral and clustered at twig ends upto 40 cm in length. Leaflets are in 3-9 pairs with odd one at the apex; increasing in size towards apex. The flowers are polygamous and the fruit is an ellipsoid drupe, upto 5 cm long with one to three seeds enclosed within its woody endocarp. It is an occasional canopy tree in the evergreen forests up to 1300 m and reported from the Western and Eastern Ghats, parts of North East India and Myanmar

(Gamble 1915; Ravikumar & Ved 2000; Shashidharan, 2006). C. strictum has not yet been assessed for it conservation status, though at the regional level it is listed as a species of conservation concern, one among the 195 medicinal plants identified for conservation action (Ravikumar et al, 2000).

Vegetation, rainfall and resin harvest practices in study sites

Our study sites were located in three regions Coonoor, Kotagiri and Gudalur,

(Appendix A) in semi evergreen forest types where Mesua ferrea, Elaeocarpus spp.,

32

Schelichera oleosa, and Persea macarantha were dominant tree species. In Coonoor and Kotagiri, two sites each were selected based on the availability of a minimum of

10 adult C. strictum trees (above 25cm dbh) with an equal number of harvested and not harvested trees. At Gudalur, we chose only one site since we were able to find 20 trees with an equal number of harvested and not harvested individuals. The trees in

Gudalur were located within an established coffee plantation bordering the evergreen forests. Gudalur gets the bulk of its monsoon from the southwest monsoons which come between June to September (Figure.3.1), while both Coonoor and Kotagiri get their rainfall mostly during the north east monsoon which is between October and

December (Figure.3.1).

Three distinct methods of resin harvest: tapping, tapping with fire and collection from natural fissures on the bark are practised in the NBR (Varghese and

Ticktin, 2008). In the Coonoor region resin harvest is more opportunistic and resin is collected from natural deposits on the bark of the tree. During our field study we observed that some harvesters in Coonoor have started to make incisions on the bark of the tree. In Kotagiri, resin is harvested by making incisions on the bark of the tree.

In the Gudalur region it was the practise, 30 years ago, to use fire to prepare the tree after which the tree was incised to yield resin. The use of fire has been discontinued owing to forest laws preventing the use of fire and only incisions are made on the tree bark for harvesting resin.

Phenology and resin harvest observations

Canarium strictum populations were located inside forest areas that were accessible by four wheel drive jeeps and then walking for another 30min to three hours. We conducted observations every fortnight over a period of 24 months, on presence or absence of leaf, flush (colour), flower, and fruit. We monitored the trees

33 for presence or absence of fresh incisions for resin harvest. We also recorded the number of times resin was harvested from each tree and estimated quantity harvested.

For the Coonoor region resin harvest was identified by signs of damage to the bark.

Field assistants from local villages, who were also resin harvesters, were able to tell us how much resin was harvested each time. In addition to this research assistants accompanied harvesters on resin collection visits to the forest and carried with them a standard hand held weighing scale to get accurate measurements of quantity.

Observations of animals found eating fruit or leaves, building nests or visiting the flowers were also recorded. The entire sample included 89 adult trees, 45 of which were never once harvested for resin during the entire study period and 44 trees that were harvested regularly (Table.3.1.).

To identify phenological patterns we calculated the percentage of trees that were in flush, flower and fruit each fortnight and compared across regions; between harvested and not harvested trees, across years, and against resin quantities harvested.

Measurements of diameter at breast height (dbh) were recorded for all individuals at six month intervals. We tested for predictors of flush colour (green versus red) using a binomial generalized linear mixed model, where flush colour was the response variable; region, dbh and openness of the canopy were independent variables and site was a random factor. We reduced the full models in a backwards stepwise process sequentially removing the least significant fixed effect term in the model and testing for significance with log likelihood ratio tests (Zuur et al. 2009). All analyses were completed in R.2.13.1 (R Development Core Team 2011) using the lmer function in the lme4 package (Bates et al 2011)

34

Results

There were variations in flush characteristics, resin harvests, fruiting and flowering in C. strictum adult trees across time and region (Table.3.1.). Nearly 50% of the trees that were selected for the study were harvested for resin in the recent past and depending on the region, up to 15-61% of trees were harvested successively over the two year period (Table.3.1). The percentage of harvested trees was lowest in

Coonoor and highest in Kotagiri (Table.3.1). Between 15-54% of the trees in our study sites produced fruit in successive years and up to 40% of the trees flowered but did not fruit, depending on the regions.

Phenological patterns in Canarium strictum

Canarium strictum trees begin to produce young leaves or flush towards the end of December, going into January sometimes continuing to March (Figure.3.2.,

3.3.). Trees or individual branches re-sprout quickly when they have been damaged, and flush may also be produced during that time. Throughout the year flush would be present in more than 80% of the trees especially in two regions, Kotagiri and

Coonoor. During the study period we recorded damage to trees due to impact of falling neighboring trees. Trees in Gudalur region, owing to their location in a coffee plantation, with lesser neighboring trees, seldom, faced this natural disturbance and as a result flushing was observed to be less (Figure.3.2., 3.3.). Peak flush period was slightly different for all three regions but similar for sites within a region. Most trees in Gudalur region started to produce new flush in December and January, whereas

Coonoor and Kotagiri had a slightly delayed flush period from January to March.

Finally at no point in the year was a tree leaf less and during peak flush times the entire tree had only new young leaves. C. strictum leaves were observed to be present in the forest litter all through the year

35

The flowering period was relatively short in C. strictum trees and flowers persisted not more than 10 days. No flowers were observed fallen on the forest floor.

Maximum percentage of trees in flower was observed in February, March and April at all the sites (Figure.3.2., 3.4.). Extended flowering periods were observed in site 2 in

Coonoor region and site 1 in Kotagiri region (Figure.3.2., 3.4.). All flowering occurred before the onset of the rain period which begins in June (Figure.3.1).

Flowers of C. strictum are very fragrant and while some of the trees flowered gregariously others produced flowers only on select branches.

Fruits of the C. strictum tree were very prominent and stood out over the canopy when they were full grown and mature. Fruiting began new in the months of

April and no new fruits were observed on the canopy after July. Fruits were bluish green in colour and matured by July at which time they turn purplish blue and were visible even from a distance. Fruits persisted on the trees till about November

(Figure.3.2., 3.5.). At any point of time in the forest C. strictum fruits were available on any one tree at all of the sites (Figure.3.2, 3.5.). In one year in Coonoor young fruits were found lying on the forest floor, aborted at an early stage, in the subsequent year that particular tree dried up and died.

Overall the presence of flush on the tree was followed closely by flowering and in turn by fruiting (Figure.3.2). There were periods when phenological conditions overlapped; most often with flush and flower timings and in some rare cases fruiting overlapped with flush but this was also because fruits persisted from the previous year. There were times when fruit, flower and flush timing overlapped especially in

Kotagiri region (Figure.3.2). All trees were observed to be in flower once during the period of study.

36

Variation in phenology in relation to harvest of resin

Harvested trees flowered more frequently than non-harvested trees

(Figure.3.4.). Harvested trees in all three regions showed two flowering periods,

January to March and the other after October. Only in one case, in the Coonoor region, not-harvested trees also flower after October (Figure.3.2, 3.4.). In terms of the timing of the flush (Figure.3.3.) and fruit (Figure.3.4.) there were no differences between harvested and not harvested trees.

Overall resin was harvested every month in all the three regions (Figure.3.7).

There were no clear peaks except in the Coonoor region where the harvest was more opportunistic and peaked before the flowering period, February (Figure.3.4.) and after the rain period which ends in November (Figure.3.1.). In the Gudalur and Kotagiri it is likely that resin is harvested based on market demand ( Chapter 4) and less on biology of the tree.

Flush colour

Canarium strictum trees, were easily visible in the forest canopy as also the saplings in the under storey, because of their distinct red velvety flush. Adult trees flushed profusely lasting for a period of two weeks at which time no mature green leaves were observed on the tree. In all the three regions we observed individual trees produced two distinct flush colours, green and red. The probability for a tree to flush green was significantly greater for bigger trees than small trees (Table.3.2,

Figure.3.7.). Openness of canopy, fruiting intensity (whether a tree fruited or not) and region were not significant predictors of flush colour.

Animal interactions with Canarium strictum trees

Flush of the C. strictum trees was found partially eaten and lying on the forest floor in Coonoor and Gudalur region (Table.3.3.). Local indigenous resin harvesters

37 informed us that the Langurs and Indian Giant Flying Squirrel (Petaurista philippensis) forage on the young leaves. Interestingly in Coonoor there were huge cavities in the trunk of C. strictum tree under which the young flush was found on the forest floor. Flying squirrels have been observed living in these cavities and also were reported by local harvesters.

The wood of the C. strictum tree seems to be easy to burrow and one tree in

Kotagiri was the nest of the Golden backed Woodpecker or Flameback (Dinopium benghalense). The bird started to build its nest at the start of 2012 and till the end of

2013 there were three holes made along the trunk of tree.

Fruits of the C. strictum trees were eaten by a number of frugivores. In the fruiting season, seeds with their fruit coats removed were found in abundance on the forest floor. These were most likely fruits that have passed through the gut of a bird and since the fruits are medium sized it could have only been a large bird. Local informants told us that it was a pigeon (unidentified) and researchers in other forest areas have observed doves eating the fruit (R.Ganesan per.com). In the Coonoor and

Gudalur regions local harvesters have seen the Great Pied Hornbill (Buceros bicornis

) feeding on the C. strictum fruits. One of the most common sights during the fruiting season was the Malabar Giant Squirrel (Ratufa indica) feasting on the fruits in the canopy. They would eat the fruit and drop the seeds; at any time on the forest floor one could see seeds completely devoid of the fruit coat and also partially eaten fruits.

The seeds that fell on the ground were observed to be depredated on by rodents. Local harvesters also report that the Flying Squirrel also feeds on the fruits of the C. strictum.

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Discussion

Our results from observational studies across two years in five sites located in three regions of a biosphere reserve help in establishing a baseline for conservation on a less studied and highly used species. Harvests of plant parts have been shown to have significant impacts on the biology of the species and at multiple scales (Ticktin

2004). Our results show that C. strictum trees have phenological patterns that vary with harvest of resin. We also observed variations in flush colour which have not been recorded as yet (except in observations of resin harvesters-discussed in Chapter 4).

We discuss our results with a view to expanding the understanding on the ecology of a harvested species and suggesting implications this has for conservation especially in the light of climate change.

Flush an important phenological event

Production of young leaves or flush is an important event in the lifecycle of C. strictum trees since it is one of the most frequent events recorded in our two year observations in the NBR. While there were clear peaks when whole trees had only new flush, but all through the year at least one branch per tree would have young leaves. This raised an interesting question about investment that the trees were making to produce new young leaves. According to one general estimate, approximately one third of plant species in tropical forests delay the greening of their leaves until full expansion (Coley and Kursar, 1996). This delayed greening strategy is thought to provide young leaves with protection from herbivores and common in shade tolerant species (Kursar and Coley 1992). This form of protection is derived from keeping the young leaves devoid of nutrition and not by investment in expensive physicochemical defences as previously thought (Dominy et al 2013). In the forests of the NBR especially in the months of January to March many of the C. strictum trees

39 are distinct because of their bright red young leaves. Flush is also produced throughout the year when the tree or branches begin to re-sprout. C. strictum is a secondary species and young saplings shoot up through gaps in the canopy of the semi evergreen forests. A recent study conducted across different forest types in the tropics showed that anthocyanin pigments make the leaves acidic therefore unpalatable to herbivores and because they are red in colour, they are invisible to most invertebrate herbivores who lack a receptor for red (Dominy, 2013; Karageorgou and Manetas

2006). While it is interesting that the C. strictum trees may be doing this to protect their young growing leaves, it would be interesting to know if this is timed with the emergence of herbivores or if it varies with herbivore densities.

However, our results show that not all C. strictum trees produce red flush and 10 – 55% of trees, depending on site, produced green coloured flush. This feature of the trees has not been reported elsewhere. The occurrence of leaf colour forms within a single plant population is of both ecological and evolutionary interest.

Intra species difference in colour of young leaves has been reported by Karageorgou and Manetas (2006) in Quercus coccifera. Smith (1986) studied the distinct leaf colour morphs in Byttneria aculeta and concluded that the green colour morph was more heavily attacked by herbivores and this was related to the frequency of occurrence of that morph in a particular habitat. Interestingly in the case of the

Quercus sp. photosynthetic efficiency was higher in the red coloured young leaves and the young green leaves were more prone to herbivore attacks (Karageorgou and

Manetas, 2006). Our results indicate that there is a correlation between size of the tree and colour of flush. As part of this study germination experiments were conducted in shade house conditions with 490 seeds of C. strictum (Chapter 2), interestingly no sapling was recorded with young green leaves. It can be that at some point in their

40 lifecycle, saplings with red flush grow into larger trees that produce green flush. We observed that green flush trees flower gregariously and produce more resin and less fruit (Chapter 2). It is possible that larger trees invest less in protection against herbivores and more in resin production. Secondary dioecious species in tropical evergreen forests of the Western Ghats have been observed transitioning from female to male trees after they have established in the middle storey and after attaining a certain girth class (R.Ganesan pers.com). C. strictum trees are reported to be polygamous, and it is possible that the relative proportions of male and female flowers on a tree shifts as the tree gets bigger. We were unable to test this because of the difficulty of obtaining flowers from these very tall trees, but this warrants further investigation.

Resin harvest and phenology

Wood exudates like resin are produced by the tree as a defence from fungal and insect attacks (Langenheim, 2003). Plants allocate carbohydrates for resource capture and growth, reproduction, storage and protection of captured resources, therefore tapping of resin is likely to enhance the trade-offs between growth processes and resin production (Rjikers, 2006). When timed properly, removal of resin can have least impact on the vitality of the tree but this is often disregarded for short term economic gains (Coppen, 1995). In Boswellia papyrifera tapping of resin is always done in the leafless dry period making the synthesis of new resin dependant on stored carbohydrates, and this result in low production of non viable seeds (Rjikers, 2006).

Depletion of carbohydrate reserves in other Boswellia serrata resulted in low fruiting levels during the leafless period (Sunnichan et al 2005). Our results show that there was no particular season when resin was harvested in Kotagiri and Gudalur regions but in the case of Coonoor region there were peaks when more than 5000gms of resin

41

(Figure.3.6.) was harvested from a single tree. Interestingly this peak was just after the monsoon rain of November (Figure.3.1) and before the flowering in February.

Since the harvesting in Coonoor region was not induced by incisions but took place more opportunistically or when resin was observed on the tree, one could presume that more resin is produced during these periods. With regard to flush and fruit production, harvested and not harvested trees showed the same patterns and it was only in the case of flowering that our results suggest a difference –in that more harvested trees flowered in two periods. In their study on Himantanthus drasticus in

Brazil, Baldauf et al (2014) observed that flowering activity increased when harvesting of bark exudate was 100%, though this did not guarantee an increase in reproductive output. One particular tree in Gudalur region is worth mentioning which flushed, produced mass flowers, fruits and also gave a kilogram of resin every three months consistently for the two year observation period. This tree was not even subjected to tapping and resin was flowing out of a natural break in its bark.

Flowering and fruiting in relation to climate

The Indian monsoons, estimated to be about 15-20million years old, are a major annual climatic event (Sanyal2010) that can be expected to have shaped the life history strategies of many plants and animals of the Indian subcontinent. Aravind et al

(2013) in their study for a common set of wind dispersed species show that mean fruiting time is associated with arrival of the wet season, independent of region. High solar radiation has been reported to trigger flowering events in many plants in the tropics (Ashton, 1988). In the case of C. strictum trees of the NBR, flowering occurred before the monsoons, independent of region, and was synchronously timed.

Mass flowering gives trees an adaptive advantage in attracting pollinators (Marquis

1988). In C. strictum trees we observed that the initiated flower buds would be

42 dormant till the young leaves expanded and turned green. Many species have been shown to flower immediately after leaf flushing, especially in tropical dry forests

(Murali & Sukumar 1994; Gunarathne and Perera, 2014). C. strictum fruits were abundant in some trees and persisted for up to eight months in all our regions. Fruits were initiated in late March, matured by June and persisted in the mature state till

October-November.

The phenology of tropical woody plants is shaped by both abiotic and biotic factors and irradiance and water stress have been shown to be the most influencing abiotic factors (Van Schlaik 2011). Studies on phenology need to be designed to understand how plants integrate abiotic and biotic factors added to the pressures from harvest in the case of species that are harvested for their products. We would like to emphasise that C. strictum is an important and less studied tree species of the Western

Ghats. The harvest of resin makes it a very important economic species and warrants more studies to establish the impacts of harvesting on the biology of the species. The tree is also found in natural habitats that are under pressure from changing climate, for instance seasonal streams and springs. For this same reason investing in the long term monitoring of the biology of the species is very crucial to understanding the linkages to the environment.

In many tropical forests there seem to be few plants that regularly produce edible fruits, seeds, flowers or leaves and these have been termed as “keystone plant resources” (Terborgh, 1986). In the case of C. strictum previous studies have recorded the importance of fruits from as a food source to rare birds and primates (Kannan,

1989; Kitamura 2011). In our study sites we report additional findings of young leaves being foraged by primates and nesting spaces provided by the trees. Our results clearly illustrate the presence of fruit and flush through the year, and as such

43 make this species a good candidate for a “keystone plant resource”. Although our study was carried out over 24 months, longer term monitoring would shed more light on inter annual variation and on the relationships between phenological patterns and local climate. This would contribute greatly to the understanding and conservation of this economically and ecologically important species.

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Table.3.1. Characteristics of Canarium strictum trees – Percent of trees in different conditions during study period

Trees Trees Trees that Total count of Trees with harvested fruiting flowered but Trees with Region Site trees observed green successively successively did not fruit red flush (%) for 2 years flush(%) over two over two in successive years(%) years (%) years (%) Coonoor Coo1 20 35 55 15 15 25 Coonoor Coo2 16 56 44 19 43 6 Gudalur Gud1 20 55 45 35 30 40 Kotagiri Kot1 13 62 38 61 54 8 Kotagiri Kot2 20 90 10 25 30 0

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Table.3.2. Effects of size as determined by diameter at breast height, openness of canopy and location on the colour of flush in Canarium strictum trees from a binomial generalized linear mixed effects model.

Fixed effects Estimate SE Z value p-value Intercept 8.8204 3.1791 2.775 0.0553 logdbh -2.1710 0.7557 -2.873 0.00407 canopyopen 0.6865 0.8480 0.810 0.41819 canopypartial 0.1863 0.8625 0.216 0.82898

Random effects SD site:region 0.000 region 0.348

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Table.3.3. Observations of animals dependant on the Canarium strictum tree, flush, flower or fruit

Region Site Frugivores mentioned/ observed consuming C. Animals or birds using tree Animals observed eating flush strictum fruits for nest

Coonoor Coo1 Unidentified species of pigeon, Malabar Giant Indian Giant Flying squirrel Gray Langurs (Semnopithecus squirrel(Ratufa indica), Indian Giant Flying (Petaurista philippensis) sp.) squirrel (Petaurista philippensis), Coonoor Coo2 Unidentified species of pigeon, Malabar Giant Indian Giant Flying squirrel Nilgiri Langur (Trachypithecus squirrel(Ratufa indica), Indian Giant Flying (Petaurista philippensis) johnii) squirrel (Petaurista philippensis), Gudalur Gud1 Unidentified species of pigeon, Malabar Giant None Nilgiri Langur (Trachypithecus squirrel(Ratufa indica),Great Indian Hornbill johnii) (Buceros bicornis) Kotagiri Kot1 Unidentified species of pigeon, Malabar Giant Golden backed woodpecker None squirrel(Ratufa indica), or Black rumped flameback (Dinopium benghalense ) Kotagiri Kot2 Unidentified species of pigeon, Malabar Giant None None squirrel(Ratufa indica),

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Figure. 3.1. Rainfall (mm) over 2011-12 in three study regions located in the Nilgiris district of the Nilgiri Biosphere Reserve

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Figure.3.2 Phenology of Canarium strictum trees located in 5 sites in 3 regions of the NBR across a two year period

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Figure.3.3. Percent of harvested and not harvested trees in flush across three regions of the NBR in 2012-13

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Figure.3.4. Percent of harvested and not harvested trees in flower across three regions of the NBR in 2012-13

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Figure.3.5. Percent of harvested and not harvested trees in fruit across three regions of the NBR in 2012-13

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Figure.3.6. Total amount of resin harvested across two years from Canarium strictum trees in three regions across the NBR

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Figure.3.7. Distribution of Canarium strictum trees with green and red flush in relation to diameter at breast height

54

CHAPTER 4. TRADITIONAL ECOLOGICAL KNOWLEDGE OF RESIN

GATHERERS OF THE NILGIRI BIOSPHERE RESERVE, WESTERN

GHATS, INDIA

Introduction

Traditional ecological knowledge (TEK) is a body of knowledge that has been handed down over many generations through an oral tradition and comprises a way of life in terms of practice, knowledge and belief (Berkes 1999). TEK helps indigenous communities survive in the environment around them and define their cultural beliefs and practices, for instance from the tradition of sacred groves in India (Gadgil 1992).

The idea that TEK is capable of adapting to both external changes and internal pressures has been a mainstay of human ecology for some time (Berkes et al 2000).

Yet by analyzing change primarily in terms of lost knowledge, the dynamic nature of

TEK, and causes for TEK loss, effects these may have on the capacity of communities to generate, transmit and apply TEK is given little emphasis (Gómez-Baggethun et al

2013). The young people in indigenous communities are important role players in the practice and sustenance of TEK and few studies have documented the knowledge that the young have in relation to forest gathering (Uprety et al 2012; Demps 2012).

Gathering products from the wild requires skill and as in the case of many activities like scaling rocks to collect honey, setting traps for hunting, fishing etc. The skill is linked closely to knowledge about the environment, behavior of the animal, seasonal changes, phenology etc. Use of TEK has become more common in conservation planning and resource assessment for reasons of efficiency, additionality and community participation (Berkes et al, 2000; Sheil et al 2004; Rist et al 2010;

Sundaram et al, 2012) Often this is interpreted as using local people to undertake field

55 work to reduce the costs of data collection and less recognition is given to the TEK which is an integral part of traditional resource management (Moller 2004).

The practical application of TEK and knowledge of the consequences of management that TEK brings is of great value for conservation aims (Rist et al, 2010).

In recent years, fuzzy cognitive modeling has emerged as a method to document traditional ecological knowledge especially with indigenous users of natural resources

(Berkes, 2008). In many communities decision making is guided by their TEK which deals with complexity through rules of thumb and indicators which are used to simplify questions about natural resource use, consistent with fuzzy logic thinking

(Berkes 2009). Fuzzy logic models allow management of complex non-linear systems traditionally with industrial process and more recently with ecological, economic and geophysical systems by offering more opportunity to use human judgment (Zadeh 2008). The knowledge used for decision making is not quantitative but consists of fuzzy sets of information using rules in the form ‘IF a certain situation occurs, THEN a known outcome is likely and can have several conditions linked by

AND, OR, NOT (Mackinson 2001; Uricchio 2004). Such approaches have been used as a tool in documenting fisher knowledge since it allows description of their ecological and technological knowledge (Grant & Berkes 2007). This approach has also been used in understanding community perceptions on conservation issues like illegal bush-meat consumption in the Serengeti National Park, Tanzania (Nyaki,

2013).

The complementarity between traditional and scientific sources of information has given more room for the use of TEK in ecological research (Moller et al, 2004;

Setty et al, 2008; Rist et al 2010; Sundaram et al 2012). TEK can not only add to scientific knowledge but present a different picture of reality making it of great value

56 especially in the case of lesser studied species. TEK can provide the much needed historical baseline which is often lacking in short term studies. Therefore we studied the TEK of resin harvest, a traditional practice of the indigenous communities with a view to document the decision making process and understand changes that are taking place to the practice. Very often harvesting practices are documented with a view to suggest improvements to methods and increase yields, there remains little discussion on the ecological knowledge that is generated as a result of these practices.

Our study was undertaken in the Nilgiri Biosphere Reserve (NBR), part of the

Western Ghats hotspot region which has a density of 267 people/sq. kms of area, the highest reported among the 25 biodiversity hotspots of the world (Cincotta et al

2000). An estimated 20 distinct indigenous groups live in the NBR and depend on the forest products for their livelihood needs (Keystone Foundation, 2007). Upto 40% of family income of the indigenous people in some areas of the reserve is derived from sale of forest products (Pain 2009). Forest products like honey, resin, medicinal plants, fruits, and leaves are gathered for local consumption and sale (Keystone

Foundation, 2007).

In our study we explore the knowledge of harvesting a locally important forest produce - resin from the tree, Canarium strictum, Roxb. (Burseraceae). We use a fuzzy logic approach to examine if and how ecological and management factors and their interrelationships are used to make decisions about harvest. We present the context of resin harvest in the NBR and then ask: 1) what factors are perceived by forest produce harvesters as important to determine quality of resin and availability of resin trees in the forest? 2) How are practices of resin harvest changing today?

We expected that knowledge about resin harvests would vary across regions because of the different harvesting practices involved. We expected harvesters of

57 resin to have detailed knowledge on the factors that affect resin quality and that these would include both biotic and abiotic factors. Finally we expected that TEK of resin harvesters can be useful for integrating for conservation and wild product management decisions in order to achieve conservation goals in landscapes where people and biodiversity co-exist.

Materials & Methods

Region

The Nilgiri Biosphere Reserve (NBR) is part of the Western and Eastern Ghats chain of mountains of the Indian peninsula, and lies between 100 45’N to 12 0 N and

760 E to 770 15’ E with a total area of 5520 km2 spread across the three southern states of Karnataka, Kerala and Tamil Nadu (Prabhakar, 1994). Altitude varies from 250m to 2650m, and at least four of the major rivers of south India originate in this region - the Bhavani, Moyar, Kabini and Chaliyar rivers. The region is home to endemic and endangered fauna like the Lion Tailed Macaque (Macaca silenus), Bengal Tiger

(Panthera tigris), King Cobra (Ophiophagus Hannah), Asiatic Elephant (Elephas maximus), Great Indian Pied Hornbill (Buceros bicornis) (Daniels, 1995). The floral diversity especially in the forests of the high elevation has several species and genera that are endemic and endangered (Keystone, 2007). The range of topography and climate has resulted in sharp gradients of vegetation composition, ranging from thorny scrub forest dominating the north-eastern region and intergrading westwards into dry and moist deciduous forests and wet evergreen forests towards the western parts

(Prabhakar, 1994).

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People

Amongst the more than 20 distinct indigenous groups in the NBR, we interacted with five of them: the Cholanayaka, Irula, Kattunayaka, Kurumba, and

Soliga (Table.4.1.). The Cholanayaka, Kattunayaka and Kurumba continue to practice their hunter gatherer way of life and are known for their honey hunting skills and collection of resin among other forest products. The Irula and Soliga additionally practice shifting cultivation. All five communities have strong links with the forests and live in close proximity to it (Appendix A). In an inventory of the forest plants of the NBR, knowledge of more than 300 species of plants are documented for each community (Keystone, 2006, 2008).

Resin tree

C. strictum, is an important and heavily harvested tree, which produces the resin known as ‘black dammar’. C. strictum is a semi evergreen tree found in the wild, rarely in cultivation and is distributed unevenly along the Western Ghats and other parts of India (Shashidharan, 2006). Little is known about C. strictum’s conservation status nationally or globally, but regionally C. strictum has been reported to be a species of conservation concern because of the high pressure from harvest of resin

(Ravikumar and Ved 2000).

In India, resin harvesting is an ancient practise that has been recorded in ancient texts, and in the Western Ghats too resin is an important part of rituals and has been harvested for a long time. Resin from C. strictum is harvested for local consumption as well as for sale and the species is subjected to different types of harvest methods and frequencies across the region (Varghese & Ticktin 2008).

Typically in the Kotagiri region, resin is either collected opportunistically from trees where it has collected naturally, or by making incisions and returning to collect resin

59 after a period of time. In the case of resin harvest from Nilambur region, the common practise is to set fire at the base of the tree before making incisions.

Methods

Focus group discussions were held in 2012-13, in nine villages across five regions where C. strictum is harvested in the NBR (Table.4.1). Villages were selected based on their proximity to the forests where the resin trees were present and on their harvest methods (Appendix A).

Introductory meetings were held in all the nine villages to introduce the research questions and to request the community’s willingness to participate in the discussions. Focus group discussions, as well as informal conversations were held in the language of the state: Tamil for Tamil Nadu, Malayalam for Kerala and Kannada for Karnataka. We used the help of an interpreter for discussions held in Kannada. All the communities have their distinct languages but are also comfortable to speak in the state languages. We obtained informed consent from interviewees and Human

Subjects Review Board exemption (Appendix D).

Focus group (7-13 participants) discussions to understand the traditional knowledge of resin harvest were organised in three sessions over time in each of the nine villages. Each discussion session was planned for one hour, and questions and topics discussed were as following: Session 1) Main topic: Resin quality – We start with a general discussion on resin harvest practises –prevalence and legal issues. We then discussed tools used for harvest, how many grades of resin are collected, what is the resin used for (is there any animal or plant that needs the resin), which is the season for resin collection and preparation of the tree, and why are there many grades of resin? We also requested people, if they were willing, to show us resin that they had harvested recently from the forests to give us an idea of the quality of the resin.

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Session 2) Main topic: Number of resin trees in the forest. We start by asking how often do the harvesters go to the forest for resin collection, how far away are the resin trees, how much resin does one tree give, when does the tree give flower, fruit and young leaves, what other animals and plants need the resin tree, what conditions are important for there to be a high number of resin trees in the forest and if they have seen any changes to the numbers of resin trees in the past ten years. Session 3) The main points from earlier discussion are presented to the group. Each factor that was mentioned in the first two sessions is represented on a large sheet of paper and participants added more information if they felt anything was missing. Connecting lines were drawn between factors by the participants to indicate relationships between factors. Session three was organised at the level of the region (n=4) and harvesters from each of the villages came together for this session. The information from this session was later organised in sets using the ‘fuzzy logic’ principles for interpretation

(Berkes 2009) which relies on using the language variables ‘IF (situation) AND

(condition) THEN outcome’ (Mackinson 2001).

Resin of C. strictum was mostly harvested by the men of the community and only in the case of the Cholanayaka people in the Nilambur region were women part of the collection activity. Wage work is an important part of indigenous people’s daily lives and attending meetings at the cost of their wages was not desirable. Most of the meetings took place on holidays or at late evening sessions in the villages, after people had returned from work. In some cases when it was not possible to meet on holidays, compensatory wages were given for all who participated in meetings.

Informal discussions

As part of our ecological study on C. strictum harvest, we monitored 89 trees in five locations fortnightly (Chapter 3). On all these monitoring field trips I was

61 accompanied by a minimum of two field assistants, one of whom was from the local area and also a resin harvester.

Context of resin harvest in the NBR

Harvest legality, access, market demand, resin harvest practices and resin quality vary across the five study regions (Table.4.1). Collection of resin is not permitted for commercial sale by Tamil Nadu state forest authorities in Coonoor and

Kotagiri regions and therefore resin is used only for home consumption and local use.

In the regions of Nilambur which is in Kerala state and Chamrajnagar which is in the state of Karnataka, resin collection is permitted and traded in the open market and also through the government owned co-operative societies.

Access to the resin trees is mostly restricted to indigenous communities, except in the case of the Kattunayaka village in Nilambur (Table.4.1) where the trees were located in a privately owned coffee estate at which both indigenous and non- indigenous people were employed for daily work. The resin trees in Chamrajnagar were located within the protected area of a tiger sanctuary and only the Soliga people had access to the site (Table.4.1).

In Coonoor and Kotagiri regions the trees were in reserve forest areas that are under the control of the Forest Department (Table.4.1). The forests in Coonoor are part of the ancestral lands of the Kurumba community people and the terrain is steep and relatively inaccessible except to the Kurumbas. The trees in the forests of

Coonoor were found in the sacred groves of the Kurumbas. The Kotagiri forests were relatively easy to access as there were motorable mud roads that provide access for the many indigenous villages that are located in that region, even then no incident of resin harvest by a non-indigenous person was reported from the past or observed in

62 the study period. The forests of Nilambur region are under reserve forests and have relatively closed access because of the terrain and inaccessibility (Table.4.1).

Frequency and quantity of resin harvested varies highly across regions

(Chapter 2, Table.4.1). There are three main grades of resin in the markets and in indigenous people’s homes (Appendix B). First grade resin consists of large blocks which are highly priced and considered to be of best quality and can fetch a price of

INR 75 to 250. A block of resin that was nearly 12 inches long and 6 inches wide weighs approximately 750-1000gms (per obs). This first grade of resin is found mostly in Coonoor and Gudalur region of our study (Table.4.1). The second grade of resin (Appendix B), smaller blocks, is sold for INR 40 to 150 depending on location

(Table.4.1). The third grade of resin is a powder form which is collected by scraping the bark of the tree and is found available mostly in the Nilambur and Kotagiri regions (Table.4.1). This powder form fetches anywhere between INR40-70 depending on the demand. In the Kotagiri region we observed that this powder form of resin was offered to guests that were visiting the home of the resin harvester.

Through informal discussions with the harvesters we were able to assess an average quantity of resin that was harvested in a year by an individual harvester.

Results

Factors perceived by indigenous harvesters to affect resin quality

The resin harvesters of Chamrajnagar are used to collecting only one type of resin, the powder form with pieces of bark attached to it or grade three (Table.4.1,

4.2.a). In Chamrajnagar, factors relating to harvest method including tools and incision, rainfall and specifically the character of the tree (size and resin producing or not) are considered important in determining the resin quality (Table.4.2.). Indicator

63 species such as the insect ‘dhoopa nona’ (species unidentified; ‘dhoopa’ is the local name for resin) are important in determining timing of harvest (Table.4.2.).

Harvesters from Coonoor collect only big blocks of resin or grade one resin

(Table.4.1, 4.3.) called ‘gatidhupa’. In Coonoor region where the harvest is done without making incisions and is more opportunistic, time to harvest (waiting for resin to mature), and character of the tree (size, and presence of resin droplets on bark or at forks in the branches) are important factors that determine resin quality (Table.4.3.).

The harvesters also comment that, from experience, they know that only five out of a hundred trees yield resin. Harvesters also mentioned that leaving the resin not harvested if a bee colony was found on the tree, so as not to disturb the bees and since the resin harvesters were also honey hunters (Table.4.3.).

Kotagiri harvesters mentioned at least two grades of resin (Table.4.1, 4.4.) that is harvested from their forests. For the resin harvest in this region, factors relating to character of the tree (stance, size, resin producing or not), harvest method (incision height and depth, direction), season (harvest frequency, timing and incision making), and biology (red and broad leaves) were important in determining the quality of the resin (Table.4.4.). Harvesters insisted that making incisions was the only way to get resin (Table.4.4.).

Nilambur harvesters reported three distinct grades of resin and indicated that season (harvest and preparation of tree), harvest method (frequency and season) and character of the tree (sloping and size) were important factors in determining resin quality (Table.4.5.). No other indicators were mentioned by the harvesters of this region.

The resin harvesters of Chamrajnagar and Kotagiri reported that there were two kinds of resin trees, female (resin producing) and male (do not produce resin)

64 trees. Though the harvesters in Coonoor were clear that only a few trees yield resin, they did not classify them as male or female trees. The harvesters in Nilambur reiterated many times that all resin trees will yield resin.

Factors perceived by resin harvesters that allow for a high number of resin

trees

In our discussions about the resin trees the Soliga people of Chamrajnagar were not able to comment on what factors allow for a high number of resin trees because there were very few left in the forest according to them. The harvesters told us of how they walked long distances these days to find even one resin producing tree and resin harvesting was now more opportunistic. One elder in the group commented that the people were too busy working on local estates for wage labour and did not harvest resin as frequently as before. The harvesters informed us that when they see a tree in the forest they would stop by it and harvest the resin (if there was some resin left on the tree).

In the Nilambur when we asked about what factors allows for a high number of resin trees in the forest, Cholanayaka people reported that there “are too many trees in our forest and we need not worry about them”. The Kattunayaka people who live in another forest in the same region, talked about the lack of trees and remember how as young people they saw their parents carry large quantities of resin but today there are only very young trees left in the forest. When we asked them what could have happened, they clearly attributed the lack of trees to indiscriminate harvest methods related to high demand and good prices which caused a lot of trees to fall.

The resin harvesters of Coonoor and Kotagiri considered different physical and biological factors as influencing the high number of trees in their regions

(Table.4.6., 4.7.). In Coonoor region, the harvesters consider forest (evergreen and

65 large), water and associated species to be important factors. Many animals and birds that disperse seeds were mentioned and also the fact that these trees prefer to be in clusters was reported to be important (Table.4.6.). Harvesters of Kotagiri consider availability of water, forest (evergreen and mountainous) and climate to be important determinants to high numbers of resin trees (Table.4.7.). The role of seed dispersers was considered favourable for regeneration though there was some concern about predators like field rats (Table.4.7.) reducing the number of viable seeds. In these two regions, none of the harvester groups who contributed to this question mentioned that harvesting resin affects the number of resin trees in the forest.

Changes in resin harvest practises

The focus group discussions revealed that over the last decades and especially over the last 10 years many changes have occurred in the traditional harvesting practices (Table.4.8.). These changes appear to be influenced by laws of the state; influence of neighbours, changes in respect for ancestral lands, increase in number of harvesters, and market demand coupled with an overall decrease in the number of productive trees.

In the case of the Kattunayaka people (Nilambur region) who live near a coffee estate, they have stopped using fire to prepare their trees as was the tradition since there are now very strict laws about fire use and the region is now highly patrolled by the Forest Department. In the Coonoor region, harvest may be moving towards incisions due to communication and exchanges about practises among harvesters. For example, one harvester from a nearby village who is also a Kurumba has started to make incisions on the trees in one of our study site. The harvesters who accompanied us on field trips knew him and when we asked why this is happening, they told us that he had family in the Kotagiri area and was trying out the incision

66 method that he saw the harvesters of Kotagiri use, since he believed that this would increase the resin yield.

In Kotagiri and Nilambur regions the frequency of harvesting had increased and this has affected resin quality. In the Nilambur region the elder harvesters felt that the young were no longer respecting the traditional tenures and would harvest any tree that came in their way. Kotagiri harvesters felt the demand for resin has increased in the past years and this has lead to more frequent harvests. They also feel the number of harvesters has increased whereas trees have not. At the same time, the market may be driving changes in the opposite direction, as harvesters observe the high price offered for high quality resin and are willing to improve the quality of the resin (Table.4.3).

In the Chamrajnagar region the traditional methods are followed but harvesters do not wait for the resin to get bigger and harvest it as soon as they see it. They informed us that was also because they go less frequently to the forests these days and were not comfortable explaining to the Forest Department officials each time what they had gone for and what they had collected, so they maximise the harvest per trip

Discussion

Resin harvest is a prevalent practise amongst the indigenous people of the

NBR. When decisions about harvest of resin are taken factors that are perceived as important to the harvesters are phenology, season, regeneration, region and characteristics like size and stance. We compare these perceptions with the ecological findings of our study (Chapter 2, Table 6) in the following section. Our results also indicate that changes to resin harvest practises are effected by many social and ecological factors which we discuss in the light of implications to knowledge transmission and adaptation.

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TEK of resin harvest in relation to ecological conditions

Our results illustrate detailed knowledge of indigenous communities in the

NBR related to the quality of resin and the factors that allow for a high number of trees in the forest, although this varied across regions (Table.4.10.). In terms of resin quality, harvesters in all four regions recognized tree size as a determinant. Of the three regions where harvesters use incisions to tap resin, each recognized method as a determinant of resin quality but each mentioned local variations to their methods. This aspect, of specific methods, though reported to vary widely over space for a given species (Ticktin Johns 2002; Ghirmire et al. 2004), remains understudied in terms of the effects on the biology of the harvested species. In our ecological study (Chapter

2) we found no significant negative effects of harvest on vital rates of C. strictum and this did not vary across regions. Therefore even though the harvesters have different site specific methods this does not seem to affect the potential for sustainable harvests.

In regions that use incisions, season, especially with reference to harvest methods was important either in terms of preparation of the tree or in relation to the harvest time. In the Philippines resin of Canarium luzonicum is harvested commercially and although it can be collected throughout the year from the rain forest, it is preferably harvested in the wet season when there is a greater resin flow

(Langenheim, 2003). Resin trees are hypothesized to produce more resin when they are not growing, which in the tropics maybe less clear cut but excessive rain in the monsoon and less sunlight could mean a low growing stage (Langenheim,2003).

Further research on how the timing of harvest affect. C. strictum yields and vital rates would be valuable.

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Interestingly the Soliga of Chamrajnagar knows that it is the right time to harvest the resin when the insect which they called the ‘dhoopa nona’ which literally translates to ‘resin insect’ has left the tree. We were not able to identify this insect.

These observations of the timing of plant or animal activities as indicators of management are an important part of resource management practices elsewhere

(Moller 2004; Grant et al 2007).

Harvesters in the regions of Nilambur and Kotagiri discussed the conditions associated with at least two grades of resin that was harvested and traded, as opposed to only one grade which was discussed by harvesters in Charmrajnagar and Coonoor.

This may be a result of the fact that harvest takes place at more intense levels in these regions - for Nilambur an estimated 100kgs is harvested per person annually whereas

Kotagiri harvesters only collect upto 20kgs(Table.4.1.). In addition to this the Kotagiri harvesters mentioned that the number of harvesters has gone up and the number of trees has decreased, suggesting that harvesters take whatever grade of resin they find on the tree to out compete others. When this was mentioned one of the harvester in the group disagreed, commenting that even though the resin with bark pieces or grade two resin fetched a lower price in the market, according to him it was an indicator that the harvester had deepened the incisions to ensure resin yield for the next visitor - e.g. as a courtesy. This example highlights the importance of understanding when social and ecological indicators meet.

The Irula harvesters of Kotagiri reported sex of the tree as a determinant of resin quality, commenting that there are male and female trees but only female trees yield resin. C. strictum is reported to have polygamous flowers (Shashidharan, 2006) and our observations in the field suggest that some trees have many more male flowers than female flowers. However, it is unknown whether the Irula categories for

69 male and female correspond to largely male or female trees in the botanical sense, and would be an interesting avenue for future research. The Kotagiri harvesters’ perception that trees with red flush produce better quality resin also merits further research. Red flush is often a defense against herbivores but does not require additional resources (Dominy et al 2013) and it is unclear how it may relate to the quality of resin produced.

In Kotagiri, (where incisions are used for tapping resin) and Coonoor (where only naturally formed resin is collected) harvesters discussed both abiotic and biotic factors as determinants to the presence of a high number of trees in the forest and the factors were similar between regions (Table.45.). Harvesters in both regions discussed factors associated with regeneration, including seed dispersal and seed predation.

Their observations of these processes point to the in-depth knowledge on this topic, and contrasts with other studies that have shown that observations by forest produce gatherers of the seedling regeneration process tends to be much less detailed than other kinds of ecological knowledge (Setty et al. 2008).

Complementarity in science and traditional knowledge has been discussed in human ecology writings, with many examples from fishers and temperate regions

(Berkes and Folke 1998; Turner 2000; Moller et al 2004; Grant & Berkes 2007). TEK has also been documented with a view to integrate it with sustainable management of natural resources (Moller, 2004; Sutherland, 2014). In more recent times especially with the rising level of awareness on climate change and pollinator declines, globally the need to integrate TEK is being recognised (Alexander et al 2009; Sutherland,

2014). Our results illustrate both how these two systems are complementary in the case of C. strictum resin harvest (Table.4.11.). Many of the indicators reported by the harvesters overlapped with the indicators that were tested in our ecological study

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(Chapter 2, Table.4.11.) and still others like habitat quality, size, water availability and plant-animal interactions which the harvesters reported could have only been a product of much longer term observations. We also find that even though the aspects of observation were common between TEK and ecological studies other factors that were taken into consideration were different (Table.4.11.). In the case of phenology and season, observations in TEK related to opportunities to improve or increase harvest yield and less to the effects on the tree. Animal sightings were also indicators to harvest timing. Various other studies have shown how TEK and ecological studies compare in terms of qualitative and quantitative assessments among other factors

(Rist et al 2010; Sundaram et al 2012). In our context we also observed that harvesters of one region were not necessarily aware of ecological context and harvest practises of nearby regions and therefore lacked some of the insights that can come from comparative research.

Changes and adaptation in TEK

There is a need to understand the conditions that support the generation of

TEK especially in the face of social-ecological changes that natural resource using communities are facing today (Gómez-Baggethun et al, 2013). The capacity of communities to respond and observe changes to the environment is an important part of TEK (Berkes et al. 2011) and one that is key to the new basis of community based conservation– one that does not rely so heavily on use but also brings in elements of observing the pollinators or the fruigvores or dispersers that may not be directly of use but are important to understanding the ecological processes that are undergoing shifts in the face of large environmental changes.

Knowledge production is a learning process where the application of knowledge to different scenarios produces outcomes that need to be evaluated and

71 adjusted as necessary (Grant and Berkes, 2007). Our results illustrate that harvesting practises in the NBR are changing in different ways depending on the context. These changes appear to be influenced by laws of the state, neighbours, changes in respect for traditional tenure, pressure on the resource base, and market demand (Table.4.8.).

The effects of these factors on resource management practices have been widely reported in the context of forest produce (Shackleton et al 2011). In some cases, different forces appear to be pushing practices in different directions. For example, in

Kotagiri and Nilambur region, an increase in the number of harvesters and market demand appears to be leading to new practises that are leading to lower quality resin and are likely unsustainable. At the same time, a new market in Kotagiri that pays better for high quality resin may be starting to cause shifts in management to obtain higher quality of resin (Table4.4.).

One incident I observed during field work is illustrative of how traditional practices are being adapted to changing climate conditions – although in this case related to honey gathering. In 2013 one group of young people from Kotagiri from the

Irula community, who also gather resin, had gone out for gathering wild honey and came back disappointed, since there was very little honey in the combs and the honey was very high in moisture content ( honey with lower moisture content is an indicator of mature honey and has longer shelf life and is preferred for sale fetching a higher price (Keystone, 2007)). An elder from the same village then went out to the forest to check what was happening and came back to report that that there was indeed something wrong with the way the Naeri (Syzygium sp.) trees were flowering - some of the branches had no flowers, while some had buds and still others had flowers. The

Naeri flower gregariously and this signals availability of bitter honey, a highly priced commodity in the local market. There was much discussion about this in the village

72 and the elders came to the conclusion that the youth had not behaved in an irresponsible manner by harvesting the honey before it was mature - but rather that things were changing in the forests. In this case, the young people are keeping alive the tradition of honey harvest because of the economic returns. And as they learn about the changes to the environment the elders do step in to validate their observations and dialogue with the young.

For TEK to be adaptive, it must be continually regenerated in response to changing conditions, and it must continue to be transmitted (Moller, 2004). In our discussions with the resin harvesters we observed that most were 30 years and above and the oldest harvester was over 70 years. Across all the four regions harvesters mentioned that “nobody taught us, we learnt by going to the forest”. Especially the harvesters in Coonoor and Kotagiri mentioned that it was not from their parents that they learnt but from uncles and friends with whom they went into the forest. When we asked if they would teach these to their children the comments ranged from were- ‘our children go to schools, where is the time?’; ‘there are too many elephants in the forest these days’; ‘we don’t want out children to look to the forest for their livelihoods’.

However in one of our focus group discussions in Kotagiri, with the Irula and

Kurumba people, a group of young people assembled around us and they asked their elders at the end of the meeting –“why don’t you teach us these things anymore”. The elders just laughed it away. The young people then met me later and asked if we could make charts with this information so that they could start to make written documents about the methods that were practised in the past. When asked if they went to the forest, they responded that they would go at least once a week even though they were employed on estates for wage work. This was important they felt to not only gather firewood but to keep their knowledge. The young men also said that they take their

73 children with them too. They also mentioned that often local medicinal plant traders would ask them to get herbs from the forest and they would supply them for the additional income. When asked how they recognised the plants, they confidently said they know most of the plants and when they were in doubt they would consult with an elder. Similar interest among youth in forest related activities has also been reported in Nepal with wild food gathering practises (Uprety et al 2012) and in the case of Jenu kurumba honey hunters of the Western Ghats, India (Demps 2012).

Our results illustrate some of the detailed knowledge that resin harvesters in the NBR have and the indicators they use to assess resin quality and the availability of resin trees in the forest. Community based monitoring programs that build on these kinds of knowledge and observations can help ensure strategies that allow for both use and conservation for the long-term while enabling communities to retain the capacity to regenerate and transmit the TEK that is part of their identity.

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Table.4.1. Details of harvester villages, communities, access, quality of resin and market price of resin across four regions in the NBR

Region State Number of Indigenous Access type Quality of ~Annual Market Price villages community resin rating harvested INR interviewed (also harvesters kgs/perso of resin) n Chamrajnagar Karnataka 2 Soliga Closed -protected area; Powder 5 40 indigenous territory Coonoor Tamil 2 Kurumba Closed-terrain restrictive; Big blocks 10 225-250 Nadu indigenous territory; reserved forests Kotagiri Tamil 2 Irula Partial-densely populated Small blocks 20 100-150 Nadu Kurumba villages in the area; trees are in reserved forests Nilambur Kerala 3 Cholanayaka Closed-terrain restrictive; Powder and 100 40-100 Kattunayaka 1* indigenous territory; small blocks Kattunayaka 2** reserved forests *Live close to forests, collect less resin these days since number of trees in their forest has reduced drastically **Live near a coffee plantation where there are resin trees. The resin in this area was of big blocks and access was of the open type since the trees were located in the estate.

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Table.4.2. Factors affecting resin quality as perceived by resin harvesters from Chamrajnagar region, arranged in fuzzy logic sets

Chamrajnagar If tree is a resin producing tree (only some trees produce good resin) If it is big (above 30cm dbh) And incision can be made with axe And incision has been left open for 3 months Then resin can be harvested And the resin will have bark pieces along with it If the rain has come Then the ‘dhoopa nona’ insect will be driven away Then it is time for us to collect the resin Conditions: Incisions should be made only on one side not around the tree.

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Table.4.3. Factors affecting resin quality as perceived by resin harvesters from Chamrajnagar region, arranged in fuzzy logic sets

Coonoor If there are signs of resin forming on the tree at forks or cavities And there are no honey combs on the tree (we would not harvest the resin so as not to disturb the bees-anyway the bees leave in 3 months-we can always harvest the resin later) Then the resin from this tree can be harvested If the tree is bigger than 30cm dbh If the resin has been allowed to mature for 3-6 months Then we get big blocks of resin (gatidhupa) Conditions: Only a good resin tree will yield well - 5 out of 100 trees produce good resin.

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Table.4.4. Factors affecting resin quality as perceived by resin harvesters from Kotagiri region, arranged in fuzzy logic sets

Kotagiri If the tree is above 30cms (dbh) And is a female tree (resin yielding trees -tested by making trial incisions on the tree- are called female trees. Then incisions can be made 3 ft above the root and upto 10ft If incision is made on the East facing side of the tree with the Sun’s rays reaching till the base of the tree And the tree is sloping or has a tilted stance Then incision should be made on the side facing the forest floor And the incision is only as deep as the bark and is left undisturbed for 3 weeks And the harvest is done in pre monsoon months And the tree has produced many new red leaves Then good quality of resin can be collected If the incision is made on the West facing side of the tree And is harvested in less than 2 weeks Then resin of lesser quality is obtained Condition: Incisions can be deepened and prepared in the monsoons Only if harvest is done with incisions will the tree produce more resin Tree which have reddish and broad leaves will produce more resin

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Table.4.5. Factors affecting resin quality as perceived by resin harvesters from Nilambur region, arranged in fuzzy logic sets

Nilambur If it is monsoon And tree is large And tree is sloping toward one side And fire is set on the sloping side And the fire has been set in the summer And it is the first resin flow And resin is allowed to flow un harvested for 2 weeks (in summer the resin will be sticky and semi solid but in the monsoon it will be harder with layers of resin) Then we can get big blocks of resin (kaal pandham) If the big blocks of resin have already been collected And incisions have been be made on tree (upto the point that the flames have reached from the fire that was set in summer) Then we get smaller blocks of resin (katta pandham) If it is the subsequent summer after the fire was set And new incisions are made on the tree And resin has not been allowed to flow for 2 weeks or more Then we get resin powder and scraped from the bark (podi pandham or thari podi)

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Table.4.6. Factors affecting number of resin trees as perceived by resin harvesters from Coonoor region, arranged in fuzzy logic sets

Coonoor If the forests are cool and evergreen And the forests are large And there are Suruli (Messua ferrea) trees close by And running streams Then there will be a good population of resin trees Conditions: Fruit that has been ingested and seeds dispersed by flying squirrel seem to germinate better. The trees prefer to be in closed forests like groves, they are found close to each other. Civets, giant squirrels, field rats, hornbills feed on the fruit/seed.

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Table.4.7. Factors affecting number of resin trees as perceived by resin harvesters from Kotagiri region, arranged in fuzzy logic sets

Kotagiri If there is water close by And the forest is evergreen And forest is in the mountains Then there will be more number of trees If there are storms these trees fall easily And neighboring trees fall on them Then there will be lesser standing trees Other observations: Regeneration is not a problem for this tree, fruits are eaten and dispersed by giant squirrel, flying squirrel, civets. Some rodents (Chundelli) seem to eating the seeds on the forest floor and this may be affecting germination

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Table.4.8. Recent changes in resin harvest methods across the four regions in the NBR

Area Harvester group Traditional harvest methods and tools Changes from traditional harvest methods Chamrajnagar Soliga Incisions are made on the bark, 30cms above the The traditional practise is followed and resin is not ground and resin is harvested using small axe or harvested for sale. knife.

Coonoor Kurumba Trees are allowed to grow big and resin formed at In one cluster of trees one of the young harvesters fissures on the bark or at branch forks are harvested has started to make incisions on the tree. But when matured and in big blocks. Blunt end of the majority of the harvesters follow the traditional knife maybe used to detach the resin from the bark. practise. Kotagiri Irula + Kurumba Incisions are made on the bark above 50cms from Old method of harvest is followed but period of ground level. Incisions are made in circular patterns waiting for the resin to mature has reduced from 3 and moved around so that old incisions heal. Knife months to two weeks. In one incident a group of is used and a small axe in some cases. harvesters (Irula from Kotagiri) spent one month to specially collect the resin in blocks from old trees and brought 73kgs to the local market since they had seen that the blocks of resin fetch double the price they would normally get for their product. Nilambur Cholanayaka Setting low grade fires to the base of the tree, Traditional methods are followed but frequency of incising the bark with axe. Harvesting resin after it collection has increased leading to poorer quality of has matured and forms big blocks resin.

Kattunayaka Setting low grade fires to the base of the tree, Fire is no longer used, only incisions are made to (residing near incising the bark with axe. Harvesting resin after it the tree estate) has matured and forms big blocks. This was the practise 20 years ago.

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Table.4.9. Summary of factors perceived by resin harvesters important to resin quality

Factors Chamrajnagar Coonoor Kotagiri Nilambur Size of tree     Stance of the tree   Harvesting tool  Resin quality   Rain / Season    Animal indicator  Male and female tree  Flush colour 

Table.4.10. Summary of factors perceived by resin harvesters important to number of trees.

Factors Chamrajnagar Coonoor Kotagiri Nilambur Forest type   Forest size  Associated species  Water   Seed dispersers   Climate  Seed predators 

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Table.4.11. Comparison of perceptions in Traditional Ecological Knowledge and findings in Ecological studies in relation to aspects of resin harvest and resin trees

Aspects of resin harvest Traditional ecological knowledge Ecological Studies and resin trees Animal interactions Insects indicate time to harvest. Tree is a nesting site to Tree as a nesting site and young leaf and fruit as food for animals some animals. Young leaf and fruit are a source of food to was observed some animals Fruit production Fruits were eaten by birds and rodents Harvested trees produced more fruit Growth of tree Not described No significant factors affecting growth Phenology Red flush indicated good quality resin tree and time to Harvested trees show two periods of flowering. Trees that harvest produce green flush were more likely to be harvested than those with red flush but only if they were bigger in size. Regeneration Seed dispersers were important for seed germination. Seed Seeds from harvested trees showed higher germination rates. predation has a negative effect on regeneration. Region Size of the forest; forest type; springs and streams Trees of some regions were harvested more than others important; Presence of water; mountainous landscape Size of tree Larger trees were harvested Bigger trees yielded more resin; Bigger trees produced more fruit Trees with green colour flush tended to be larger than those with red colour flush; Seeds from bigger trees showed less germination rates Season Rain season yielded better quality of resin and trees were Flowering occurs before the rain and resin harvest was spread prepared for harvest before the rains across the months. There was a peak in the post rain period Tree stance Very important to make incisions; resin of high quality Not considered produced by sloping trees

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CHAPTER 5. CONCLUSIONS

My dissertation examines the consequences of harvesting practises on the ecology and phenology of a resin harvested species, Canarium strictum and explores the traditional ecological knowledge (TEK) that harvesters have in relation to resin harvesting. In this chapter I synthesise my findings from the previous chapters, discuss the implications for managing harvests from the wild, and explore the possibilities of integrating traditional ecological knowledge in community based ecological monitoring programs that will enhance community based conservation. I also describe the contributions of this research to the fields of sustainable use and management of wild resources integrating them with TEK. Finally, I discuss some of the limitations of this study and suggest further research that could improve our understanding of, and capacity to, include more participation of communities in conservation processes.

Main findings

I used data from an observational study on 89 C. strictum adult trees located in three regions with different harvesting practises in the Nilgiri Biosphere Reserve

(NBR). I used mixed effects modelling to identify significant predictors of resin production, resin quantity, growth, reproduction, survival and seed germination. I compared the phenology of harvested and not harvested resin trees to understand how harvest impacts the biology. I conducted focus group discussions and informal interviews with harvesters across my study sites to understand the TEK that exists in relation to resin harvest and ecology of the resin trees. I used a fuzzy logic approach to better understand the decision making processes that harvesters make in relation to harvests from the wild.

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Impacts of harvesting and ecology of a resin harvested species

In the forests of my study sites in the NBR, I observed that not all trees were harvested for resin and I was interested in exploring the characteristics of harvested trees. Overall C. strictum trees appear to produce more resin if tapping or making incisions was practised. I found that ‘region’ which is also a proxy for harvest method is a significant predictor of whether a tree gets harvested or not (Chapter 2). In the

Coonoor region where harvest is more opportunistic (no incisions were made on the tree), the chance that a tree gets harvested is significantly less than in Kotagiri and

Gudalur where incisions are made to tap for resin. I also found that trees that produced green colour flush as opposed to those that produce red colour flush

(differences in the colour of flush are reported in Chapter 3) are likely to be more frequently harvested. I also found that quantity of resin produced by individual trees is highly variable and found that size of the tree and number of times new incisions were made on the tree for harvest are significant predictors of the quantity of resin harvested (Chapter 2).

I found that harvesting of resin has no negative effect on the growth rate of the

C. strictum trees. Higher rates of growth are observed in trees located in closed and partial light as against open light conditions. Trees that produced high number of fruits and green colour flush also showed higher growth rates (Chapter 2). The number of fruits produced by the C. strictum tree is dependent on the size of the tree

(large trees produced more fruit than smaller ones) and interestingly, trees which were harvested for resin showed higher levels of fruit production and also higher seed germination (Chapter 2). I found no significant effects of harvest on survival, however of the six adult trees that dried up and fell to storms, during the study period, all were harvested for resin.

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My observational studies on the phenology of C. strictum shows that, trees begin to produce young leaves or flush towards the end of December, going into

January sometimes continuing to March. I observed that the flowering period is relatively short in C. strictum trees, lasting not more than 10 days, with the maximum percentage of trees in flower in February, March and April across all sites, this occurs before the onset of the rain period. Fruiting begins new in the months of April and no new fruits were observed on the canopy after July (Chapter 3). I found that harvested trees of C. strictum appear to flower more frequently than not harvested trees although in terms of flush and fruit timings, there were no differences between harvested and not harvested trees.

Across all study sites I observed that individual trees produced two distinct flush colours, green and red. The probability for a tree to flush green was significantly greater for bigger trees than small trees. Openness of canopy, fruiting intensity

(whether a tree fruited or not) and region were not significant predictors of flush colour (Chapter 3). My observations on other animals that were dependent on the C. strictum trees either for forage or shelter were similar to observations by local harvesters of resin. The Langurs and Indian Giant Flying Squirrel (Petaurista philippensis) forage on the young leaves; the Golden backed Woodpecker or

Flameback (Dinopium benghalense) has built nests in the trunk of the tree; Great Pied

Hornbill (Buceros bicornis ) feeds on the C. strictum fruits. One of the most common sights during the fruiting season was the Malabar Giant Squirrel (Ratufa indica) feasting on the fruits in the canopy (Chapter 3).

TEK of resin harvest

I found that resin harvesting is a prevalent practise amongst the indigenous communities of the Nilgiri Biosphere Reserve (NBR). In some locations like

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Nilambur in Kerala, for the Cholanayaka people resin is an important part of their livelihood and collection is all through the year (Chapter 4). In other regions like

Chamrajnagar because of wage options and issues with access to the forests, resin collection is still practised but not with the same intensity as in other regions of my study (Chapter 4). Harvest methods are followed by members of the communities and harvesters correlate these methods with the quality of the resin (Chapter 4). They also made linkages between the ecology of the species and the presence of high number of trees in the forest (Chapter 4). The harvesters of all the study sites discussed that harvesting practises affect the quality of the product and some discussed attributes of the ecology of the species that determined the population status of the trees (Chapter

4). I found that harvesters attributed the same set of factors to influencing production of resin as I had independently assessed in my ecological studies on the species. I also found that harvesters attributed factors such as habitat quality, size of the forest and availability of water as key indicators of the status of resin tree populations (Chapter

4).

Further studies are warranted overall to assess the long term and short term factors that affect resin production and conservation status of lesser studied species like C. strictum. Resin harvest is an old practise and has been practised for more than

100 years in the experience of the harvesters of the NBR. Many of the older accounts of the people of the region mention harvest of resin as a communal activity. One of the further steps would be to document what they remember from the past. This would help in creating a historical timeline of the changes and understanding what the indigenous people perceive as the causes of those changes.

88

Contributions to scientific literature

My dissertation focuses on the ecology of a lesser studied species, C. strictum, and examines the relationship between harvest of an exudate on the ecology and phenology of the species. I examine the effects of the harvest on individual trees, across regions located in different management types. I also bring in the perspective of harvesters, who have interacted for much longer periods with the trees and the forests that they are located in, on the ecology and management of the tree.

Trees produce resin for a number of reasons, primarily to avoid auto toxicity, defense from parasites, herbivores and predators and resins also play a role in attracting pollinators and seed dispersers (Langenheim, 2003). The high cost of production of this primary metabolite is offset by the benefits of pollination, dispersal and defense from pathogens for the tree. Harvesting resin has been shown to impact reduction in number of fruits and viable seeds (Rjikers, 2006) and kept populations in a vegetative state (Cunningham et al 1993, Cocks et al 2004). In contrast some studies show that removal of resin does not affect flower or fruit production (Bitariho et al

2006; Schumann et.al 2010; Bladauf et al 2014). My study shows that harvesting of resin from C. strictum has no negative effect on the overall growth, but that resin harvested trees have high fruit production and germination rates of seeds than in not harvested trees. This suggests that removal of resin has no negative impact on the growth, reproduction and survival of the C. strictum trees. My study also shows that harvested trees flower more than not harvested trees and also produced more fruit.

Disentangling anthropogenic factors responsible for changes in the timing of phenological events needs to be further investigated. Nearly one third of plant species in tropical forests delay the greening of their leaves until full expansion (Coley and

Kursar, 1996). This delayed greening strategy is thought to provide young leaves with

89 protection from herbivores; this is particularly the case with shade tolerant species

(Kursar and Coley 1992). This protection is derived from keeping the young leaves devoid of nutrition and not by investment in expensive physicochemical defences as previously thought (Dominy et al 2013). In the forests of the NBR especially in the months of January to March many of the C. strictum trees are distinct because of their bright red young leaves. My results show that not all C. strictum trees produce red flush. This feature of the trees has not been reported elsewhere. The occurrence of leaf colour forms within a single plant population is of both ecological and evolutionary interest. Intra species difference in colour of young leaves has been reported by

Karageorgou and Manetas (2006) in Quercus coccifera. Smith (1986) studied the distinct leaf colour morphs in Byttneria aculeta and concluded that the green colour morph was more heavily attacked by herbivores and is related to the frequency of occurrence of that morph in a particular habitat. Interestingly in the case of the

Quercus sp. photosynthetic efficiency was higher in the red coloured young leaves and the young green leaves were more prone to herbivore attacks (Karageorgou and

Manetas, 2006). A recent study from the Himalayas showed that individuals from the

Canarium genus showed signs of folivory as early as the Miocene period (Khan, et al,

2014). This indicates that insect plant interactions have been ancient with this genus and resulted in defence mechanisms. This is of particular interest to the study of evolutionary ecology. My results show that probability of a tree to have green flush was significantly higher if it was larger. I also observed green flush trees flower gregariously and produce more resin and less fruit (Chapter 3).

My study shows that intensive harvesting leads to higher production of resin and has no negative effects on the biology of this species. This is consistent with earlier studies which hypothesized that collection of plant exudates like resin if

90 harvested in prescribed ways (either by traditional knowledge or science) has the least impact on the species and its populations as compared to harvest of other vegetative or reproductive parts (Peters, 2001). Harvesting methods and their effects on resin production are under studied but the observations of harvesters in relation to the ecology of the species and the changes to the environment are even more understudied. In many communities decision-making is guided by their TEK which deals with complexity through rules of thumb and indicators which are used to simplify questions about natural resource use, consistent with fuzzy logic thinking

(Berkes 2009). My study is the first of its kind to use a fuzzy logic approach to document the TEK of resin harvest.

Canarium strictum trees are found in natural habitats which are under pressure from changing weather patterns, as observed in the drying up of seasonal streams and springs. This necessitates investing in the long term monitoring of the biology of the species in order to understand the linkages with the environment. In the case of C. strictum previous studies have recorded the importance of fruits from as a food source to rare birds and primates (Kannan, 1989; Kitamura 2011). In my study sites I report additional findings of young leaves being foraged by primates and nesting spaces provided by the trees. My results clearly illustrate the presence of fruit and flush through the year, and as such make this species a good candidate for a “keystone plant resource”. Although my study was carried out over 24 months, longer term monitoring would shed more light on inter annual variation and on the relationships between phonological patterns and local climate. This would contribute greatly to the understanding and conservation of this economically and ecologically important species.

91

Linking TEK and sustainable use and management

Indigenous communities are portrayed as custodians of conservation and increasingly community based conservation are gaining ground, especially through community conserved areas (Shahabuddin and Rao, 2010). Participation of forest- dwelling indigenous communities in the conservation process is undergoing a rapid change in the Indian context due to the recently implemented Scheduled Tribes and

Other Traditional Forest Dwellers Recognition of Forest Rights Act, 2006 also referred to as the Forest Rights Act of 2006. More efforts based on strengthening local knowledge based institutions and practises will empower local communities to better manage biological diversity and put forward many suggestions on how this may be achieved in the light of the new legislation (Bawa et al 2011). One of the important suggestions I make is to align local knowledge and conservation science within the citizen science work. There is a lack of space that is lacking for the participation of some sections of society in the growing citizen science movement and this needs to be addressed (DeVictor, 2010). My results illustrate some of the detailed knowledge that resin harvesters in the NBR have and the indicators they use to assess resin quality and the availability of resin trees in the forest. Community based monitoring programs that build on these kinds of knowledge and observations can help ensure strategies that allow for both use and conservation for the long term.

Future directions

As a two year case study on the impacts of harvest on a tree that has been harvested for centuries perhaps, my dissertation necessarily has limitations and my findings suggest further avenues for study.

A better understanding of several aspects of C. strictum ecology would allow me to make more precise and improved management recommendations. Over the

92 duration of the study there was a high degree of variation in the rainfall patterns of the three regions where the trees were located. The year 2012 was a particularly dry year and the monsoons failed to arrive at the usual time. Given the substantial effect of climate on phenological events (Parmesan et al, 2010) more long term studies need to be undertaken spanning several years before management suggestions can be made.

The variation in management conditions across my study sites provided useful comparison but confounded management and region effects. A larger number of study sites and more replicates in terms of management practises would have been helpful for better understanding the effects of harvest methods. I have taken several steps to ensure that the results of my study can contribute to management decisions. This research was designed and carried out in collaboration with Keystone Foundation, with whom I have been affiliated for more than 10 years now. Keystone is a voluntary agency that works on issues related to conservation, enterprise and livelihoods in the

NBR and across India. I have shared my results broadly with the NGO and presented these results especially focussing on the need to include observations of local communities in ecological assessments to NGO groups in the state of Orissa, North

East India and also to South East Asian network of forestry managers. I met with harvesters of the regions in my study site to explore the possibility of a project on community based ecological monitoring in the region. I received funding for this project in 2013 from the Criticial Ecosystems Partnership Fund to support a group of

‘barefoot ecologists’ who are currently monitoring the quality of their forests and recording changes to the same. It is my intention that the results of my dissertation in addition to informing ecological theory will facilitate decision making about the management of this important tree species. This will contribute to the community

93 based ecological monitoring program that I have initiated in the NBR. I hope it will have long term implications for community based conservation in a biosphere reserve.

94

APPENDIX A. MAPS OF STUDY REGIONS WITH APPROXIMATE LOCATION OF SITES

Western Ghats, India Nilgiri Biosphere Reserve, Western Ghats, India

Chamrajnagar

Gudalur Kotagiri

Coonoor

Nilambur

S

tat

e 95

bor

der

s APPENDIX B. PHOTOS OF STUDY SITE, RESIN, RESIN TREE, AND HARVEST METHODS

B.1 Vegetation characteristics of study sites 96

First grade of resin

Third grade of resin

Second grade of resin

B.2. Grades of resin found in the region

97

Adult tree in the forest

Red flush Green flush

Seedlings in the forest

Buds and flowers

Fruits

B.3. Characteristics of resin tree, Canarium strictum

98

B.4. Resin harvesting tools and practises

99

APPENDIX C. DETAILS OF METHODS AND RESULTS FROM THE LINEAR MIXED EFFECTS MODELS ON PREDICTORS

OF RESIN HARVEST AND EFFECT OF RESIN HARVEST ON GROWTH, FRUIT PRODUCTION AND SEED GERMINATION

ON THE WILD DAMMAR TREE (CANARIUM STRICTUM)

Table C.1. Specifications of full linear and generalised linear mixed effects models (LMM and GLMM) used to predict resin harvest and effect of resin harvest on growth fruit production and seed germination in C. strictum

Model Form Response variable Random Main effect Interactions effect Harvest Status Binomial Resin harvested or Site Logdbh+Flush colour+Region+Intensity Logdbh*Flush colour GLMM not (1/0) of fruiting Region*Logdbh Region*Flush colour Resin quantity Negative Total quantity of resin Site Logdbh+ Region+ Fruiting intensity+ Logdbh*Flush colour Binomial harvested per tree Number of incisions made+ Flush colour GLMM over two years Growth LMM Growth (difference Site Logdbh(year1)+Region+Resin quantity Logdbh*Flush colour dbh year 1 and year 2) harvested+Canopy status+Flush colour+ Region*Logdbh Intensity of fruiting Region*Flush colour Flush colour*Fruiting intensity Fruit Negative Fruit production(total Site Logdbh1+ Region +Harvest status + Logdbh*Flush colour Binomial number of fruits Flush colour + Canopy openness Region*Logdbh GLMM produced in 2012) Germination Binomial Seed germinated or Tree id / Logdbh+Region+Flush colour +Harvest None GLMM not (1/0) Site status

100

Table C.2. Estimates and standard errors for main effects and interactions that were non-significant (p>0.05) by likelihood ratio test during model reduction in Chapter 2. All terms were significant in Growth model.

Model Main effect Estimate S.E Harvest Status Normal fruiting intensity 0.8970 0.8035 Low fruiting intensity -0.4029 0.7699

Resin quantity Flush colour red 0.029170 0.076623 Low fruiting intensity 0.098546 0.085573 Region Kotagiri 0.080061 0.124429

Fruit Region Gudalur -0.7657 0.5028 Canopy open 0.4390 0.6316 Canopy partial -0.2110 0.6726 Germination Logdbh 1.4724 0.9252 Flush colour red 0.2268 0.5200

101

APPENDIX D. UNIVERSITY OF HAWAIʻI COMMITTEE ON HUMAN SUBJECTS RESEARCH EXEMPTION

102

BIBLIOGRAPHY

Alexander, Clarence, Nora Bynum, Elizabeth Johnson, Ursula King, Tero

Mustonen, Peter Neofotis, Noel Oettlé, et al. 2011. “Linking Indigenous and

Scientific Knowledge of Climate Change.” BioScience 61 (6): 477–84. doi:10.1525/bio.2011.61.6.10.

Allendorf, Fred W, and Jeffrey J Hard. 2009. “Human-Induced Evolution

Caused by Unnatural Selection through Harvest of Wild Animals.” Proceedings of the

National Academy of Sciences of the United States of America 106 Suppl 9987–94. doi:10.1073/pnas.0901069106.

Aravind, N A, K N Ganeshaiah, and R Uma Shaanker. 2013. “Indian

Monsoons Shape Dispersal Phenology of Plants Indian Monsoons Shape Dispersal

Phenology of Plants.” Biology Letters 9:20130675 (October).

Ashton, Peter, and Chris J. Kettle. 2012. “Dipterocarp Biology as a Window to the Understanding of Tropical Forest Structure: Where Are We Looking Now?”

Biotropica 44: 575–76. doi:10.1111/j.1744-7429.2012.00913.x.

Augustine, J., and P. G. Krishnan. 2006. “Status of the black dammar tree

(Canarium strictum Roxb) in Periyar Tiger Reserve, Kerala and the uses of black dammar”. Indian Forester 132(10):1329–1335

Baldauf, Cristina,Alexsandra Salvador da Silva Julia C. Sfair, Rosij ania

Ferreira, and Flavio Antonio Ma€ es dos Santos. 2014. “Harvesting Increases

Reproductive Activity in Himatanthus Drasticus (Mart.) Plumel (Apocynaceae), a

Non‐Timber Forest Product of the Brazilian Savanna.” Biotropica 46 (3): 341–49. http://onlinelibrary.wiley.com/doi/10.1111/btp.12109/full.

Barrett, C. B., Travis, A. J., & Dasgupta, P. (2011). Biodiversity Conservation and Poverty Traps Special Feature: From the Cover: On biodiversity conservation and

103 poverty traps. Proceedings of the National Academy of Sciences, 108(34), 13907–

13912. doi:10.1073/pnas.1011521108

Bawa, K S, W J Kress, N M Nadkarni, and S Lele. 2004. “Beyond Paradise -

Meeting the Challenges in Tropical Biology in the 21st Century.” Biotropica 36: 437–

46. doi:10.1646/Q1609.

Bawa, Kamaljit S, Nitin D Rai, and Navjot S Sodhi. 2011. “Rights,

Governance, and Conservation of Biological Diversity.” Conservation Biology : The

Journal of the Society for Conservation Biology 25 (3): 639–41. doi:10.1111/j.1523-

1739.2010.01640.x.

Bawa, Kamaljit S, W John Kress, Nalini M Nadkarni, Sharachchandra Lele,

Peter H Raven, Daniel H Janzen, Ariel E Lugo, Peter S Ashton, and Thomas E

Lovejoy. 2004. “Tropical Ecosystems into the 21st Century.” Science 306: 227–28. doi:10.1126/science.306.5694.227b.

Belcher, B., & Schreckenberg, K. (2007). Commercialisation of Non-timber

Forest Products: A Reality Check. Development Policy Review, 25(3), 355–377. doi:10.1111/j.1467-7679.2007.00374.x

Berkes, F, and M Berkes. 2009. “Ecological Complexity, Fuzzy Logic, and

Holism in Indigenous Knowledge.” Futures 41 (1): 6–12. doi:10.1016/j.futures.2008.07.003.

Berkes, F, J Colding, and C Folke. 2000. “Rediscovery of Traditional

Ecological Knowledge as Adaptive Management.” Ecological Applications 10: 1251–

62. doi:10.1890/1051-0761(2000)010[1251:ROTEKA]2.0.CO;2.

Berkes, F. 2004. “Rethinking Community-Based Conservation”. Conservation

Biology, 18(3), 621–630. doi:10.1111/j.1523-1739.2004.00077.x

104

Berkes, F. 2008. “Sacred ecology: traditional ecological knowledge and resource management”. Taylor & Francis.

Berkes, Fikret, and Carl Folke. 1998. “Linking Social and Ecological Systems for Resilience and Sustainability.” Linking Social and Ecological Systems:

Management Practices and Social Mechanisms for Building Resilience, 1–25.

Bitariho, Robert, Alastair McNeilage, Dennis Babaasa, and Robert Barigyira.

2006. “Plant Harvest Impacts and Sustainability in Bwindi Impenetrable National

Park, S.W. Uganda.” African Journal of Ecology 44: 14–21. doi:10.1111/j.1365-

2028.2006.00597.x.

Brummitt, N, and S Bachman. 2010. “Plants under Pressure a Global

Assessment.” … Red List Index for Plants. Natural History Museum, UK

Brundtland, G. 1987. Report of the World Commision on Environement and

Development: Our Common Future. Oxford paperbacks (Vol. Report of, p. 400). doi:10.2307/2621529

Calinger, Kellen M, Simon Queenborough, and Peter S Curtis. 2013.

“Herbarium Specimens Reveal the Footprint of Climate Change on Flowering Trends across North-Central North America.” Ecology Letters 16 (8): 1037–44. doi:10.1111/ele.12135.

Champion, H G and Seth, S K.1968. “ A Revised Survey of the Forest Types of India”. Forest Research Institute, New Delhi, 1968.

Chao, Sophie. 2012. “Forest Peoples: Numbers across the World.” Forest

Peoples Programme, 27.

Chapman, Colin a., Lauren J. Chapman, Thomas T. Struhsaker, Amy E.

Zanne, Connie J. Clark, and John R. Poulsen. 2005. “A Long-Term Evaluation of

105

Fruiting Phenology: Importance of Climate Change.” Journal of Tropical Ecology 21

(1): 31–45. doi:10.1017/S0266467404001993.

Cincotta, R P, J Wisnewski, and R Engelman. 2000. “Human Population in the

Biodiversity Hotspots.” Nature 404 (6781): 990–92. doi:10.1038/35010105.

Cocks, M., and T. Dold. 2004. “The informal trade of Cassipourea flanaganii”.

In T. Sunderland, and O. Ndoye (Eds.). Forest products, livelihoods and conservation:

Case studies of non-timber forest products systems, pp. 73–90. CIFOR, Bogor,

Indonesia.

Coley, P. D., & Kursar, T. A. 1996. “Causes and consequences of epiphyll colonization” in Tropical forest plant ecophysiology (pp. 337-362). Springer US.

Cunningham, A B, and F T Mbenkum. 1993. Sustainability of Harvesting

Prunus Africana Bark in Cameroon : A Medicinal Plant in International Trade. People and Plants Working Paper 2.

Cunningham, A.B. 2001. “Applied Ethnobotany: People, wild plant use and conservation”. Earthscan Publications Ltd., London and Sterling, VA.

Cunningham, Anthony B. "Non-timber Products and Markets: lessons for export-oriented enterprise development from Africa." In Non-Timber Forest Products in the Global Context, pp. 83-106. Springer Berlin Heidelberg, 2011.

Daniels, R. J. R., Madhav Gadgil, and N. V. Joshi. 1995. “Impact of Human

Extraction on Tropical Humid Forests in the Western Ghats Uttara Kannada, South

India.” Journal of Applied Ecology 32: 866–74. doi:10.2307/2404826.

Danielsen, F., Jensen, A. E., Alviola, P. a., Balete, D. S., Mendoza, M.,

Tagtag, A., … Enghoff, M. 2005. “Does Monitoring Matter? A Quantitative

Assessment of Management Decisions from Locally-based Monitoring of Protected

106

Areas”. Biodiversity and Conservation, 14(11), 2633–2652. doi:10.1007/s10531-005-

8392-z

Danielsen, F., Burgess, N. D., Balmford, A., Donald, P. F., Funder, M., Jones,

J. P. G., … Yonten, D. 2009. “Local participation in natural resource monitoring: a characterization of approaches”. Conservation Biology : The Journal of the Society for Conservation Biology, 23(1), 31–42. doi:10.1111/j.1523-1739.2008.01063.x

Darimont, Chris T, Stephanie M Carlson, Michael T Kinnison, Paul C Paquet,

Thomas E Reimchen, and Christopher C Wilmers. 2009. “Human Predators Outpace

Other Agents of Trait Change in the Wild.” Proceedings of the National Academy of

Sciences of the United States of America 106: 952–54. doi:10.1073/pnas.0809235106.

Das, A., Krishnaswamy, J., Bawa, K. S., Kiran, M. C., Srinivas, V., Kumar, N.

S., & Karanth, K. U. (2006). Prioritisation of conservation areas in the Western Ghats,

India. Biological Conservation, 133(1), 16–31. doi:10.1016/j.biocon.2006.05.023

De Beer, J. and M.J. McDermott. 1996. “The economic value of non-timber forest products in Southeast Asia”. Amsterdam: Netherlands Committee for IUCN

Demps, K., Zorondo-Rodriguez, F., García, C., & Reyes-García, V. 2012. The

Selective Persistence of Local Ecological Knowledge: Honey Collecting with the Jenu

Kuruba in South India. Human Ecology, 40(3), 427–434. doi:10.1007/s10745-012-

9489-0

Demps, Kathryn, Francisco Zorondo-Rodríguez, Claude García, and Victoria

Reyes-García. 2012. “Social Learning across the Life Cycle: Cultural Knowledge

Acquisition for Honey Collection among the Jenu Kuruba, India.” Evolution and

Human Behavior, April. Elsevier Inc. doi:10.1016/j.evolhumbehav.2011.12.008.

107

Dominy, Nathaniel J, Peter W Lucas, Lawrence W Ramsden, Pablo Riba- hernandez, Kathryn E Stoner, Ian M Turner, and Costa Rica Kathryn. 2013. “FORUM

FORUM.” Oikos 98: 163–76.

Drew, Joshua A. 2005. “Use of Traditional Ecological Knowledge in Marine

Conservation.” Conservation Biology 19: 1286–93. doi:10.1111/j.1523-

1739.2005.00158.x.

Duraiappah, A. K. (1998). Poverty and Environmental Degradation : A

Review and Analysis of the Nexus. World Development, 26(12), 2169–2179.

Ella, A.B. and A.L.Tongacan 1992. “Techniques in tapping almaciga (Agathis philippinensis Warb.) for sustained productivity of the tree: the Philippine experience”. Forest Products Research and Development Institute Journal 21: 73-79

Encontro, V I, Alice Dantas, and Brites Procam. 2012. “The Ecological

Effects of Harvesting Non-Timber Forest Products from Natural Forests : A Review of the Evidence.”

Eshete, Abeje, Frank J. Sterck, and Frans Bongers. 2012. “Frankincense

Production Is Determined by Tree Size and Tapping Frequency and Intensity.” Forest

Ecology and Management 274 (June). Elsevier B.V. 136–42. doi:10.1016/j.foreco.2012.02.024.

Freese, Curtis H., ed. 1997. “Harvesting wild species: implications for biodiversity conservation”. Baltimore, Maryland: Johns Hopkins University Press.

Gadgil, M. 2014. Western Ghats Ecology Expert Panel. Economic & Political

Weekly. Retrieved from https://xa.yimg.com/kq/groups/13367407/1128840523/name/Western_Ghats_Ecology

_Expert_ Panel.pdf

108

Gadgil, Madhav. 1992. “Conserving Biodiversity as If People Matter: A Case

Study from India.” Ambio 21 (3): 266–70.

Ganesh, T, and Priya Davidar. 2001. “Dispersal Modes of Tree Species in the

Wet Forests of Southern Western Ghats.” Current 80: 394–99. http://www.ias.ac.in/currsci/feb102001/394.pdf.

Gaoue, Orou G., and Tamara Ticktin. 2007. “Patterns of Harvesting Foliage and Bark from the Multipurpose Tree Khaya Senegalensis in Benin: Variation across

Ecological Regions and Its Impacts on Population Structure.” Biological Conservation

137: 424–36. doi:10.1016/j.biocon.2007.02.020.

Gaoue, Orou G., and Tamara Ticktin. 2008. “Impacts of Bark and Foliage

Harvest on Khaya Senegalensis (Meliaceae) Reproductive Performance in Benin.”

Journal of Applied Ecology 45: 34–40. doi:10.1111/j.1365-2664.2007.01381.x.

Ghimire, Suresh Kumar, Doyle McKey, and Yildiz Aumeeruddy-Thomas.

2005. “Heterogeneity in Ethnoecological Knowledge and Management of Medicinal

Plants in the Himalayas of Nepal : Implications for Conservation.” Ecology And

Society 9: 6.

Gómez-baggethun, Erik, and Victoria Reyes-garcía. 2013. “Reinterpreting

Change in Traditional Ecological Knowledge.” doi:10.1007/s10745-013-9577-9.

Grant, S, and F Berkes. 2007. “Fisher Knowledge as Expert System: A Case from the Longline Fishery of Grenada, the Eastern Caribbean.” Fisheries Research 84

(2): 162–70. doi:10.1016/j.fishres.2006.10.012.

Groenendijk, Peter, Abeje Eshete, Frank J. Sterck, Pieter a. Zuidema, and

Frans Bongers. 2011. “Limitations to Sustainable Frankincense Production: Blocked

Regeneration, High Adult Mortality and Declining Populations.” Journal of Applied

Ecology, November, no–no. doi:10.1111/j.1365-2664.2011.02078.x.

109

Gunarathne, R M U K, and G A D Perera. 2014. “Climatic Factors

Responsible for Triggering Phenological Events in Manilkara Hexandra ( Roxb .)

Dubard ., a Canopy Tree in Tropical Semi-Deciduous Forest of Sri Lanka” 55 (1): 63–

73.

Ibrahim, J., Abu Said, A., and A. Abdul Rashid. 1987. “Tapping of oleoresin from Dipterocarpus kerrii”. Malaysian Forester 50: 343-353

Jansen, Merel, Pieter A. Zuidema, Niels P R Anten, and Miguel Mart??nez-

Ramos. 2012. “Strong Persistent Growth Differences Govern Individual Performance and Population Dynamics in a Tropical Forest Understorey Palm.” Journal of Ecology

100: 1224–32. doi:10.1111/j.1365-2745.2012.02001.x.

Kabra, A. (2009). Conservation-induced displacement: A comparative study of two Indian protected areas. Conservation and Society, 7(4), 249. doi:10.4103/0972-

4923.65172

Kannan, R. 1992. “Burning out the black dammar, Canarium strictum Roxb.”.

Journal of Bombay Natural History Society 91(1):159.

Karageorgou, Panagiota, and Yiannis Manetas. 2006. “The Importance of

Being Red When Young: Anthocyanins and the Protection of Young Leaves of

Quercus Coccifera from Insect Herbivory and Excess Light.” Tree Physiology 26 (5):

613–21. http://www.ncbi.nlm.nih.gov/pubmed/16452075.

Keystone Foundation, 2006. “Forest Plants of the Nilgiri Biosphere Reserve-

Eastern-An Illustrated Field Guide”. Keystone Foundation.

Keystone Foundation, 2008 – “Forest Plants of the Nilgiri Biosphere Reserve-

Northern-A Pictorial Field Guide”. Keystone Foundation.

Keystone Foundation. 2007. “Honey Trails in the Blue Mountains”. Keystone

Foundation, Kotagiri

110

Khan, M. A., Spicer, R. a., Spicer, T. E. V., & Bera, S. 2014. “Fossil evidence of insect folivory in the eastern Himalayan Neogene Siwalik forests”.

Palaeogeography, Palaeoclimatology, Palaeoecology, 410, 264–277. doi:10.1016/j.palaeo.2014.05.043

Kitamura, Shumpei. 2011. “Frugivory and Seed Dispersal by Hornbills

(Bucerotidae) in Tropical Forests.” Acta Oecologica, February. Elsevier Masson SAS,

1–11. doi:10.1016/j.actao.2011.01.015.

Kursar, TA, and PD Coley. 1992. “Delayed Development of the

Photosynthetic Apparatus in Tropical Rain Forest Species.” Functional Ecology. http://www.jstor.org/stable/2389279.

Langenheim, J. 2003. “Plant resins: chemistry, evolution, ecology, ethnobotany”. Timber Press, Portland, Oregon, USA.

Langenheim, J. 2003. “Plant resins: chemistry, evolution, ecology, ethnobotany”. Timber Press, Portland, Oregon, USA.

Lantz, Trevor C.,and Nancy J. Turner.2003.”Traditional phenological knowledge of aboriginal peoples in British Columbia”. Journal of Ethnobiology

23:263-86

Lengerke, Hans J von and Blasco, F. 1989. “The Nilgiri Environment in: Blue

Mountains: the Ethnography and Biogeography of a South Indian Region”. (ed) Paul

Hockings, Oxford University Press, New Delhi.

Mackinson, Steven. 2001. “Integrating Local and Scientific Knowledge: An

Example in Fisheries Science.” Environmental Management 27: 533–45. doi:10.1007/s002670010168.

111

Madhusudan, M. D., & Shankar Raman, T. R. 2003. “Conservation as if

Biological Diversity Matters: Preservation versus Sustainable Use in India”.

Conservation and Society, 1, 49–59.

Mandle, Lisa, Tamara Ticktin, Snehlata Nath, Siddappa Setty, and Anita

Varghese. 2013. “A Framework for Considering Ecological Interactions for Common

Non-Timber Forest Product Species: A Case Study of Mountain Date Palm (Phoenix

Loureiroi Kunth) Leaf Harvest in South India.” Ecological Processes 2 (1). Ecological

Processes: 21. doi:10.1186/2192-1709-2-21.

Marquis, R. J. 2013. “Phenological Variation in the Neotropical Understory

Shrub Piper Arielanum : Causes and Consequences.” Ecological Society of America

69: 1552–65. doi:doi:10.2307/1941653.

Moller, Henrik, Fikret Berkes, Philip O Brian Lyver, and Mina Kislalioglu.

2004. “Combining Science and Traditional Ecological Knowledge : Monitoring

Populations for Co-Management.” Ecology And Society 9 (3).

Morsello, C., Ruiz-Mallén, I., Diaz, M. D. M., & Reyes-García, V. 2012. The effects of processing non-timber forest products and trade partnerships on people’s well-being and forest conservation in Amazonian societies. PloS One, 7(8), e43055. doi:10.1371/journal.pone.0043055

Murali, K. S., and R. Sukumar. 1993. “Leaf Flushing Phenology and

Herbivory in a Tropical Dry Deciduous Forest, Southern India.” Oecologia 94: 114–

19. doi:10.1007/BF00317311.

Nantel, P, D Gagnon, and A Nault. 1996. “Population Viability Analysis of

American Ginseng and Wild Leek Harvested in Stochastic Environments.”

Conservation Biology 10: 608–21. doi:10.1046/j.1523-1739.1996.10020608.x.

112

Nebeker, T. E., R. A. Tisdale, and R. F. Schmitz. 1995. “Comparison of

Oleoresin Flow in Relation to Wound Size, Growth Rates, and Disease Status of

Lodgepole Pine.” Canadian Journal of Botany. doi:10.1139/b95-038.

Newton, Peter, Andrew R. Watkinson, and Carlos a. Peres. 2011.

“Determinants of Yield in a Non-Timber Forest Product: Copaifera Oleoresin in

Amazonian Extractive Reserves.” Forest Ecology and Management 261 (2). Elsevier

B.V. 255–64. doi:10.1016/j.foreco.2010.10.014.

Pain, A.2009. “What is driving change in the Nilgiri Biosphere Reserve and what effects might such change have on the role of NTFP in the livelihoods of indigenous people?” In Proceedings of the Biodiversity and Livelihoods Conference.

Ed. Dutt, R., Seeley, J. and P.Roy. Coonoor, The Nilgiris, India

Parmesan, Camille, and Gary Yohe. 2003. “A Globally Coherent Fingerprint of Climate Change Impacts across Natural Systems.” Nature 421 (6918): 37–42. doi:10.1038/nature01286.

Peters, C.M. 2001. “Lessons from the plant kingdom for conservation of exploited species” in Conservation of Exploited Species. Ed. Reynolds, J.D. Mace

G.M., Redford, K.H. and Robinson, J.G. Conservation Biology 6. Cambridge

University Press, UK

Peters, Charles M., Alwyn H. Gentry, and Robert O. Mendelsohn. 1989.

“Valuation of an Amazonian Rainforest.” Nature. doi:10.1038/339655a0.

Pinheiro, Jose, Douglas Bates, Saikat DebRoy, Deepayan Sarkar, and R

Development Core Team. 2012. “Nlme: Linear and Nonlinear Mixed Effects

Models.” R Package Version 3.1-105.

Plowden, C., C. Uhl and F. de. A. Oliviera. 2002. “Breu resin harvest by

Tembe Indians and its dependence on a bark boring beetle”. In Ethnobiology and

113

Biocultural diversity pp 365-380, eds. J.R. Stepp, F.S. Wyndham and R.K. Zarger.

University of Georgia Press. Athens.

Prakbhakar, R. 1994. Resource Use, Culture and Ecological Change: A case study of the Nilgiri Hills of Southern India, CES, IISc Bangalore, March 1994,

Unpublished Ph.D. thesis

Primack, Daniel, Carolyn Imbres, Richard B Primack, Abraham J Miller-

Rushing, and Peter Del Tredici. 2004. “Herbarium Specimens Demonstrate Earlier

Flowering Times in Response to Warming in Boston.” American Journal of Botany

91 (8): 1260–64. doi:10.3732/ajb.91.8.1260.

Primack, RB. 1987. “Relationships among Flowers, Fruits, and Seeds.”

Annual Review of Ecology and Systematics 18 (1987): 409–30. http://www.jstor.org/stable/2097138.

R Development Core Team, R. 2011. “R: A Language and Environment for

Statistical Computing.” R Foundation for Statistical Computing. doi:10.1007/978-3-

540-74686-7.

Ravikumar, K. & D.K.Ved. 2000. “Illustrated Field Guide-100 Red Listed

Medicinal Plants of conservation concern in Southern India”. FRLHT, Bangalore,

India.

Rathcke, B., & Lacey, E. P. 1985. “Phenological Patterns of Terrestrial

Plants”. Annual Review of Ecology and Systematics. doi:10.1146/annurev.es.16.110185.001143

Rijkers, Toon, Woldeselassie Ogbazghi, Marius Wessel, and Frans Bongers.

2006. “The Effect of Tapping for Frankincense on Sexual Reproduction in Boswellia

Papyrifera.” Journal of Applied Ecology 43 (6): 1188–95. doi:10.1111/j.1365-

2664.2006.01215.x.

114

Rist, Lucy, R Uma Shaanker, and Jaboury Ghazoul. 2010. “The Use of

Traditional Ecological Knowledge in Forest Management : An Example from India.”

Ecology And Society 15 (1).

Root, Terry L, Jeff T Price, Kimberly R Hall, Stephen H Schneider, Cynthia

Rosenzweig, and J Alan Pounds. 2003. “Fingerprints of Global Warming on Wild

Animals and Plants.” Nature 421: 57–60. doi:10.1038/nature01333.

Roy, P., Leo, R., Thomas, S. G., Varghese, A., Sharma, K., Prasad, S.,

Davidar, P. (2011). Nesting requirements of the rock bee Apis dorsata in the Nilgiri

Biosphere Reserve , India. Tropical Ecology, 52(3), 285–291.

Sanyal, P., & Sinha, R. 2010. “Evolution of the Indian summer monsoon: synthesis of continental records”. Geological Society, London, Special Publications,

342(1), 153-183.

Sasidharan, N. 2006. “Illustrated manual on tree flora of Kerala supplemented with computer-aided identification”. Research Report 282. Kerala Forest Research

Institute, Peechi, p 698.

Schmidt, Isabel B., and Tamara Ticktin. 2012. “When Lessons from

Population Models and Local Ecological Knowledge Coincide - Effects of Flower

Stalk Harvesting in the Brazilian Savanna.” Biological Conservation 152: 187–95. doi:10.1016/j.biocon.2012.03.018.

Schmidt, Isabel B., Lisa Mandle, Tamara Ticktin, and Orou G. Gaoue. 2011.

“What Do Matrix Population Models Reveal about the Sustainability of Non-Timber

Forest Product Harvest?” Journal of Applied Ecology 48 (4): 815–26. doi:10.1111/j.1365-2664.2011.01999.x.

115

Schmidt, Isabel B. “Effects of Local Ecological Knowledge , Harvest and Fire on Golden-Grass ( Syngonanthus nitens , Eriocaulaceae ), a Non-Timber Forest

Product ( ntfp ) Species from the Brazilian Savanna.” Unpublished Phd Dissertation.

Schumann, Katharina, Rüdiger Wittig, Adjima Thiombiano, Ute Becker, and

Karen Hahn. 2010. “Impact of Land-Use Type and Bark- and Leaf-Harvesting on

Population Structure and Fruit Production of the Baobab Tree (Adansonia Digitata L.) in a Semi-Arid Savanna, West Africa.” Forest Ecology and Management 260 (11).

Elsevier B.V. 2035–44. doi:10.1016/j.foreco.2010.09.009.

Setty, R Siddappa, Kamal Bawa, Tamara Ticktin, and C Made Gowda. 2008.

“Evaluation of a Participatory Resource Monitoring System for Nontimber Forest

Products : The Case of Amla ( Phyllanthus Spp .) Fruit Harvest by Soligas in South

India.” Ecology And Society 13 (2).

Shackleton, Sheona, C.O. Delang, and A. Angelsen. 2011. “From Subsistence to Safety Nets and Cash Income: Exploring the Diverse Values of Non-Timber Forest

Products for Livelihoods and Poverty Alleviation.” In Non-Timber Forest Products in the Global Context, 55–81. doi:10.1007/978-3-642-17983-9_3.

Shahabuddin, Ghazala, and Madhu Rao. 2010. “Do Community-Conserved

Areas Effectively Conserve Biological Diversity? Global Insights and the Indian

Context.” Biological Conservation 143 (12). Elsevier Ltd: 2926–36. doi:10.1016/j.biocon.2010.04.040.

Sheil, Douglas, and Anna Lawrence. 2004. “Tropical Biologists, Local People and Conservation: New Opportunities for Collaboration.” Trends in Ecology and

Evolution. doi:10.1016/j.tree.2004.09.019

Siebert, Stephen F. 2000. “Abundance and Growth of Desmoncus

Orthacanthos Mart. (Palmae) in Response to Light and Ramet Harvesting in Five

116

Forest Sites in Belize.” Forest Ecology and Management 137: 83–90. doi:10.1016/S0378-1127(99)00316-3.

Smith, Alan P. 2014. “Oecologia Ecology of a Leaf Colour Polymorphism in a

Tropical Forest Species : Habitat Segregation and Herbivory” 69 (2): 283–87.

Subash Chandran, M. D. 1997.” On the ecological history of the Western

Ghats”. Current Science, 73, 146–155.

Sundaram, Bharath, Siddhartha Krishnan, Ankila J. Hiremath, and Gladwin

Joseph. 2012. “Ecology and Impacts of the Invasive Species, Lantana Camara, in a

Social-Ecological System in South India: Perspectives from Local Knowledge.”

Human Ecology, October, 931–42. doi:10.1007/s10745-012-9532-1

Sutherland, William J., Toby a. Gardner, L. Jamila Haider, and Lynn V.

Dicks. 2013. “How Can Local and Traditional Knowledge Be Effectively

Incorporated into International Assessments?” Oryx, November, 1–2. doi:10.1017/S0030605313001543.

Terborgh, J. 1986. “Community aspects of frugivory in tropical forests”. In

Frugivores and seed dispersal (pp. 371-384). Springer Netherlands.

Ticktin, T. 2004. “The Ecological Implications of Harvesting Non-Timber

Forest Products.” Journal of Applied Ecology 41 (1): 11–21. doi:10.1111/j.1365-

2664.2004.00859.x.

Ticktin, T., Ganesan, R., Paramesha, M., & Setty, S. 2012. “Disentangling the effects of multiple anthropogenic drivers on the decline of two tropical dry forest trees”, 1–11. doi:10.1111/j.1365-2664.2012.02156.x

Ticktin, Tamara, and Timothy Johns. 2002. “Chinanteco Management of

Aechmea Magdalenae: Implications for the Use of TEK and TRM in Management

117

Plans1.” Economic Botany. doi:10.1663/0013-

0001(2002)056[0177:CMOAMI]2.0.CO;2.

Ticktin, Tamara, and Charlie Shackleton. "Harvesting non-timber forest products sustainably: opportunities and challenges." In Non-timber forest products in the global context, pp. 149-169. Springer Berlin Heidelberg, 2011.

Turner, Nancy J., Marianne Boelscher Ignace, and Ronald Ignace. 2000.

“Traditional ecological knowledge and wisdom of aboriginal peoples in British

Columbia.” Ecological Applications. doi:10.1890/1051-

0761(2000)010[1275:TEKAWO]2.0.CO;2.

Uprety, Yadav, Ram C Poudel, Krishna K Shrestha, Sangeeta Rajbhandary,

Narandra N Tiwari, Uttam B Shrestha, and Hugo Asselin. 2012. “Diversity of Use and Local Knowledge of Wild Edible Plant Resources in Nepal.” Journal of

Ethnobiology and Ethnomedicine 8 (1): 16. doi:10.1186/1746-4269-8-16.

Uricchio, Vito F., Raffaele Giordano, and Nicola Lopez. 2004. “A Fuzzy

Knowledge-Based Decision Support System for Groundwater Pollution Risk

Evaluation.” Journal of Environmental Management 73: 189–97. doi:10.1016/j.jenvman.2004.06.011.

van Schaik, C. P., Terborgh, J. W., & Wright, S. J. 1993. “The phenology of tropical forests: adaptive significance and consequences for primary consumers”.

Annual Review of Ecology and Systematics, 24(1), 353-377.

Varadarajan, D. B. 2014. “REDD , Climate Change and the Rights of Tribal

Communities” in. Journal of Studies in Dynamics and Change, 1(1), 15–22.

Varghese, Anita, and Tamara Ticktin. 2008. “Regional Variation in Non-

Timber Forest Product Harvest Strategies , Trade , and Ecological Impacts : The Case

118 of Black Dammar ( Canarium Strictum Roxb .) Use and Conservation in the Nilgiri

Biosphere Reserve , India.” Ecology And Society 13 (2).

Vedeld, P., Angelsen, A., Sjaastad, E., & Berg, G. K. 2004. “Counting on the environment - forest incomes and the rural poor”. Environment Department Papers. doi:10.1177/1420326X04041346

Wunder, S. 2001. “Poverty Alleviation and Tropical Forests -What Scope for

Synergies ?” World Development, 29(11).

Wunder, S. 2014. “Forests , Livelihoods , and Conservation : Broadening the

Empirical Base”. WORLD DEVELOPMENT, xx. doi:10.1016/j.worlddev.2014.03.007

Zadeh, L. 2008. “Is There a Need for Fuzzy Logic?” Information Sciences 178

(13): 2751–79. doi:10.1016/j.ins.2008.02.012.

Zuidema, Pieter a, Roel J W Brienen, Heinjo J During, and Burak Güneralp.

2009. “Do Persistently Fast-Growing Juveniles Contribute Disproportionately to

Population Growth? A New Analysis Tool for Matrix Models and Its Application to

Rainforest Trees.” The American Naturalist 174 (5): 709–19. doi:10.1086/605981.

Zuur, A., Ieno, E. N., Walker, N., Saveliev, A. A., & Smith, G. M. 2009.

“Mixed effects models and extensions in ecology with R”. Springer.

119