Masking environmental feedback:

Misfits between institutions and in Belize and Thailand

Miriam Huitric

Doctoral Thesis in Management

Department of Systems Ecology Stockholm University SE-106 91 Stockholm, SWEDEN Stockholm 2004

Doctoral dissertation 2004 Miriam Huitric Department of Systems Ecology Stockholm University SE-106 91 Stockholm Sweden

© 2004 Miriam Huitric ISBN 91-7265-881-9 pp. 1-35 Printed in Sweden by Intellecta DocuSys AB, Sollentuna

ABSTRACT

The thesis analyses relationships between ecological and social systems in the context of coastal ecosystems. It examines impacts from the extraction of resources from these ecosystems, through and , and addresses management and governance behind resource exploitation. The main premises of the thesis are that a lack of ecological knowledge will to poor management and that the dichotomy between social and natural systems is an artificial one. The thesis illustrates the importance of basing resource management on the ecological conditions of the resource and its ecosystem. It also demonstrates the necessity of accounting for the human dimension in ecosystem management and the challenges of organising human actions for sustainable use of ecosystem services in the face of global drivers and economic incentives that push resource users towards short term extraction. Many reefs have been undergoing a shift from coral to macroalgal domination. An experiment in the of Glovers Atoll in Belize manually cleared patch reefs in a no-take zone and a fished zone (Papers I and II). The study hypothesised that has reduced herbivorous populations that control macroalgae growth. Overall, management had no significant effect on fish abundance and the algal reduction had a short-lived impact. The resilience of the desired coral-dominated state of Glovers Reef Atoll seems to be low. This study illustrated that the benefits of setting aside marine reserves in already impacted environments should not be taken for granted. Research suggests that this shift is in part a result of historical overfishing of herbivores, from large mammals to small fish. By studying the development of the and conch fisheries in Belize (Paper III), and the farming industry in Thailand (Paper IV), it was shown that feedback from the environment can be relieved and even masked to give the impression of resource abundance. In Belize was maintained through sequential exploitation of fishing grounds, of stocks, and of target , made possible by technology. The shrimp farming industry in Thailand responded to the spread of disease between farms by sequentially exploiting . In both cases inadequate property rights and artificially high economic returns, as these industries do not pay for their externalities, contributed to unsustainable resource use. The behaviour of both the and the shrimp farming industries show features of the pathology of natural resource management described for other industries. The final paper (V) compares the responses to changes in the resource by the lobster fisheries in Belize and Maine in terms of institutions, organisations and their role in management. The establishment of fishermen's co-operatives in Belize was found to have been a driver in sequential exploitation, demonstrating that social resilience of organisations does not ensure resilience of the social-ecological system (Paper III). In contrast to the Maine system, the Belize system seems to lack social mechanisms for responding in an effective fashion to environmental feedback. The results (Paper V) illustrate the importance of a diversity of organisations and institutions that incorporate ecological knowledge of the resource, respond to ecosystem feedback and provide a social context for learning and adaptation to deal with change in order to manage ecological and human processes at relevant scales.

TABLE OF CONTENTS

List of Papers 1

Introduction 2

Background 4

Ecosystems & Resilience 4

Patterns of Human Use & Measures to Control Use 6

Methods 9

Papers I and II 9

Paper III 10

Paper IV 12

Paper V 13

Summary and Conclusions of the Papers 13

Discussion of Major Findings 14

Ecosystem Dynamics & Management on Glovers Reef Atoll 14

Sequential Exploitation & the Pathology of Resource Use 16

Institutions & Organisations 17

The Problem of Fit Between Ecosystems & Institutions 19

Responding to Environmental Feedback 20

Measures to Prevent the Pathology of Resource Use 21

Concluding Remarks 22

Appendix I – Errata Papers I and II 25

References 28

Acknowledgements 35

LIST OF PAPERS

These papers are referred to by their roman numerals.

I McClanahan, T., K Bergman, M Huitric, M McField, T Elfwing, M Nyström, and I Nordemar, 1999: Response of to algal reductions at Glovers Reef, Belize. Marine Ecology Progress Series 206: 273-282.

II McClanahan TR, McField M, Huitric M, Bergman K, Elfwing T, Nyström M, and Nordemar I, 2001: Responses of , and fish to the reduction of macroalgae in fished and unfished patch reefs of Glovers Reef Atoll, Belize. Coral Reefs 19: 367 – 379.

III Huitric M, 2003: Lobster and conch fisheries of Belize – a history of sequential exploitation. Conservation Ecology (Ecology & Society) in revision.

IV Huitric M, Folke C and Kautsky N, 2002: Development, governmental policies and impacts of the shrimp farming industry on ecosystems of Thailand. 40: 441-455.

V Huitric M, 2004: Comparative study of the lobster fisheries in Maine and Belize. Identifying factors that contribute to social-ecological resilience. Manuscript.

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INTRODUCTION

Nature’s ability to supply ecosystem goods and services1 (Daily 1997) sets the preconditions for economic development (Berkes and Folke 1992). These goods and services are products of functional ecosystems and are finite if used faster than ecosystems’ regenerative capacity (Costanza and Daly 1992). The resources that extract are components of ecosystems, that is, they are embedded in ecosystem dynamics and the production of ecosystem services and therefore should not be managed as independent commodities (Berkes and Folke 1992). The main premises of the thesis are that a lack of ecological knowledge will lead to poor ecosystem management and that the dichotomy between social and natural systems is an artificial one. The five articles of the thesis illustrate the relationship between natural and social systems in the context of coastal ecosystems, examining in particular human impacts from the extraction of resources from these ecosystems, through fisheries and aquaculture, and the management of these ecosystems and their users, through institutions and their organisations. The overall theoretical framework is that of social-ecological systems (SESs) (Berkes and Folke 1998b). Analyses of SESs stress the importance of understanding ecosystem dynamics and resilience and the role that humans play in this context. It emphasises the need for capacity to respond to and manage feedbacks between ecosystems and society for sustainability. It emphasises the governance system behind feedback management and how it relates to the broader ecological and social environment, including drivers of change (figure 1). The thesis addresses these issues. The first two papers are about understanding ecosystem dynamics in a human dominated environment and are based on work on Glovers Reef Atoll in Belize. Papers III and IV describe the development of the lobster and conch fisheries in Belize and the shrimp farming industry in Thailand respectively and how these industries deal with feedback from the ecological and social systems and the governance systems behind management and drivers of change. The final paper (V) compares governance in coastal fisheries in Belize and Maine. Figure 1 presents the focus on feedback and management practice of the thesis and how this is embedded in the broader social-ecological system highlighting interactions across

1 Ecosystem services are the conditions and processes through which natural ecosystems, and the species that make them up, sustain and fulfil human life. They maintain and the production of ecosystem goods (, timber). Ecosystem services are also life-support functions (cleansing, re- cycling) (Daily 1997).

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scales. The thesis addresses certain aspects of these interactions and their match and mismatch.

Figure 1: The ecological and social systems as two interacting systems (modified from Berkes et al. 2003a).

Human interaction with nature inevitably changes it. Change is an inherent part of ecological systems (Holling 1973, Connell 1978). However, the rate and type of anthropogenic change is of concern (Likens 1994). Certain effects of human activities are more obvious, or direct, such as converting mangroves into shrimp . Others are indirect. Ecological , when a species is so reduced that it no longer fulfils its function , have resulted in trophic cascades in fisheries (Pace et al. 1999). Trophic cascades are predator-prey effects that cause inverse patterns in abundance or across more than one trophic link in a web (Paine 1966). Global fisheries have moved down trophic levels from large piscivore fish to smaller invertebrates and planktivorous fish (Pauly et al. 1998). In other words, there are cross-scale effects where change on one scale can affect other scales. Human interaction with the environment can change entire landscapes (van der Leeuw and Team 2000, Gordon et al. 2003) and seascapes, for example, the Caribbean (Hughes 1994, Jackson et al. 2001, Pandolfi 2002), which complicates implementing measures to restore local ecosystems such as coral reefs (papers I and II). We now talk about human domination of global ecosystems (Botsford et al. 1997, Vitousek et al. 1997, Steffen et al. 2004). Resource management needs ecological knowledge in order to sustain flows of goods while maintaining functional ecosystems. This requires the ability to recognise when use, whether exploitation or emission, exceeds the ecosystem’s regeneration capacity. Yet

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humans are not very good at managing the ecosystem capacity that produces the extracted goods (Hanna and Jentoft 1996). This is often due to our focus on the flow of goods such as shrimp. Attempts to control resource use are often based on limited ecological information such as stock sizes, forest area, and species diversity. Such information only captures parts of ecosystem structure, but does not reveal the state of the ecosystem. This requires understanding and appreciation of ecosystem processes and dynamics (Christensen et al. 2003) as well as of the history of the system (Jackson et al. 2001). The thesis illustrates the importance of basing resource management on the ecological conditions of the resource and its ecosystem. The thesis confirms, through analyses of dynamics and the development of coastal fisheries and aquaculture, unsustainable human behaviour observed by others (Beveridge et al. 1994, Flaherty and Karnjanakesorn 1995, Roberts 1995, Aiken et al. 1999). It also demonstrates the necessity of accounting for the human dimension in ecosystem management and the large difficulties of organising human actions for sustainable use of ecosystem services in the face of those global drivers and economic incentives that push resource users towards short term extraction (López and Scoseria 1996, Reed 1996, Geist and Lambin 2001, Lambin et al. 2001). The thesis starts with a background section on ecosystem resilience, resource exploitation and governance. This is followed by a description of the methods used and a brief summary of the major conclusions of each paper of the thesis. In the next section I will discuss ecosystem dynamics with an emphasis on the coral reef experiments at Glovers Reef Atoll, Belize, the human dimension of ecosystem management in relation to the findings of the work on fisheries and aquaculture in Belize and Thailand, as well as the comparative study with Maine lobster fisheries. New findings and hypotheses that the thesis has generated are presented in the concluding section.

BACKGROUND

Ecosystems & Resilience Ecosystems are complex assemblages of biodiversity interacting with the surrounding physical and chemical environment (Golley 1993). These interactions cover different temporal and spatial scales and often complicate defining ecosystem boundaries as no ecosystem is closed or isolated, so the term is used loosely here to encompass the natural

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system used by a resource unless otherwise specified. For example, both the , Homarus americanus, and the Caribbean , Panuliris argus, undergo several ontogenetic shifts during their life-cycles as well as migrate seasonally and even diurnally between ecosystems (Paper V). Disturbance edits community structures by affecting recruitment, predation, competition, and, physical and chemical factors (Nyström et al. 2000). A disturbance is defined here as “any relatively discrete event in time that disrupts ecosystem community or population structure and changes resources, substrate availability, or the physical environment” (White and Pickett 1985). Intermediate levels and frequencies of disturbances increase diversity in ecosystems in space and time (Connell 1978). Thus, disturbance contributes to the dynamic character of ecosystems as well as enhances the maintenance of ecosystem functioning in a resilient system (Gunderson and Pritchard 2002). Resilience is defined here as the amount of disturbance a system can absorb whilst still remaining within the same stability domain, the ability of the system to self-organise (versus lack of organisation or organisation forced by external factors) and the degree to which the system can develop and increase the capacity for learning and adaptation (Carpenter et al. 2001). The loss of local diversity, and changes in species composition, can affect ecosystem processes at both local and global scales (Chapin III et al. 1997), reducing their resilience. If a system’s resilience has been reduced, a disturbance normally absorbed may instead cause a shift in stability domain (Scheffer et al. 2001, Carpenter 2003). Throughout the Caribbean, coral reefs are shifting from coral to algal domination (Hughes 1994, Bak and Nieuwland 1995, Shulman and Robertson 1996, McClanahan et al. 1999), with little evidence of recovery despite management interventions (Porter and Meier 1992, Connell 1997). A similar shift has occurred over the last 25 years on Glovers Reef Atoll, Belize (McClanahan and Muthiga 1998). The reef assemblage in Belize and the Caribbean was dominated by Acropora sp. for several millennia until the 1980s (Aronson and Precht 2001, Pandolfi 2002). It has been suggested that historical fisheries exploitation and emissions of waste have pushed coastal ecosystems towards “weedy systems” (Jackson 1997, Knowlton 2001) subject to larger environmental variability and with a reduced capacity to sustain society with ecosystem services (Moberg and Folke 1999, figure 2). Events that can have affected this shift on Glovers Reef Atoll include: reduced herbivory by overfishing (Hughes 1994) and the Diadema antillarum die-off during the 1980s (Lessios et al. 1984), coral disease (Koltes et al. 1998), enrichment (Mumby 1998) and others such as coral disease carried in dust from (Shinn 2000) and rises in

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(Hoegh-Guldberg 1999). This atoll may have already been altered two hundred years ago by the arrival of Europeans who systematically eliminated large mammals, turtles and more recently large grazing fish, all important herbivores in ecosystem processes (Jackson 1997). The systematic exploitation from larger to smaller herbivorous organisms has reduced the resilience of the reefs (Elmqvist et al. 2003) and seems to be a critical factor explaining the shift from coral to algal dominated reefs in several areas of the Caribbean (figure 2).

Figure 2. Schematic representation of changes on Caribbean reefs.

As systems are embedded, changes at one scale can affect others. For example, the loss of mangroves affects the functioning of remaining mangroves and of surrounding ecosystems (Rönnbäck 1999). Indeed, the global cover of mangroves has decreased alarmingly (Alongi 2002). In the case of shrimp farming, the shrimp ponds produce externalities that further affect their surroundings (Flaherty and Karnjanakesorn 1995, Paper IV). Environmental feedback as a result of these changes, such as reduced yields and increased frequency and scale of disease, should warn users and managers about change. How have industries managed to avoid developing effective responses to the signals of resource and environmental deterioration?

Patterns of Human Use & Measures to Control Use Economically successful resources with high yields and returns will encourage production and attract new users (Papers III – V). Under certain conditions this influx of new users and increased effort has negative effects on the resource and its environment and therefore on the users themselves (Coswig Kalikoski et al. 2002, Papers III - V). The

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industries studied in Papers III-V are dependent on fossil fuel and technology to maintain yields. As capitalisation by an industry increases, the economy becomes more vulnerable to a collapse of the natural system (Gunderson et al. 1995, Hanna and Jentoft 1996) and the managing body is under greater pressure to protect the industry and, therefore, to decrease the impacts of nature’s variability (Folke and Kåberger 1992). This pattern has been termed the ‘pathology of regional development’ (Holling and Bocking 1990, Holling and Meffe 1996, Holling and Sanderson 1996) and can be summarised as: a lack of inhibition on investments in good periods, but strong pressure not to dis-invest in bad years (Ludwig et al. 1993). Incomplete valuation of ecosystems, lack of incorporation of environmental feedback and the assumption that resources were renewable, or would recover when exploitation could be reduced, were found to support this behaviour. The work in papers III and IV identified conditions that “masked” environmental feedback (Berkes and Folke 1998a). One response to local declines in production is to move to new areas. This pattern of resource use is termed sequential exploitation (Grima and Berkes 1989) and is described in Papers III and IV. Sequential exploitation in these studies was facilitated by technology (Paper III) and unclear property rights (Paper IV) as well as an emphasis on short-term profit maximisation versus long-term sustainability. New technology relocates users to new areas (Papers III and IV) or intensifies extraction by providing access to previously inaccessible resources (Paper III). In this way, technology, and even capital markets (Paper IV), can mask users’ dependence on natural capital and can even be perceived to replace natural capital, for example, the perception that aquaculture can replace (Folke 2003). This may work in the short-term, but it increases the scale of problems that will catch up in the long-term (Holling and Meffe 1996). By not adjusting resource use to feedback, (unexpected) crashes in the resource can occur, as illustrated in forestry (Regier and Baskerville 1986), fisheries (Ludwig et al. 1993, Finlayson and McCay 1998) and several other sectors and ecosystem-linked activities (Gunderson et al. 1995). This was described in Thailand for the regional crashes in the shrimp farming industry (Paper IV). Such crashes are in part due to positive feedback loops caused by responding to the wrong feedback. Shrimp farms in Thailand stock their ponds with fry that are hatched from wild caught gravid females. The conversion of mangroves to ponds reduces the area of nurseries for fry that will in turn reduce the abundance of adult and gravid female shrimp in the wild. Thus shrimp farming in mangroves can reduce its own supply of fry. Whilst visiting a shrimp farm and hatchery area in Thailand in 1997, I was informed that a boat was on 24-hour call to collect gravid

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females that might be caught by trawlers at . These environmental changes may be difficult to revert (Hutchings 2000, Paper II). Property rights are important institutions in regulating natural resource use (Bromley 1992). Institutions are here defined as “humanly devised constraints that shape human interaction” (North 1990) and are therefore important driving forces of natural resource use (Reed 1996). Property rights determine who has access to a resource and control their use of the resource (Hanna and Jentoft 1996). Resources without rights are said to be open access and are predicted to become over-exploited because individuals will maximise their individual benefits at the cost of the resource base, or “free-ride” (Hardin 1968). There are few resources for which there is true open access, but if rules controlling access and use are not enforced, or do not match the resource, it can become de facto open access (Papers III - V). There are three main types of property rights: private, state and common property. Private property restricts access to a single user who can deny others access to the property (Paper IV). Private property is not appropriate when exclusion from the resource is difficult and when there is subtractability, that is, what a user removes cannot be used by others. Resources exploited in fisheries are typical examples, as these resources tend to be mobile and patchy in distribution (Brown et al. 2002). For these resources, state governance or common property are appropriate (Papers III – V). There are many examples of groups that have self-organised to control resource use as a group, often as common property systems (Feeney et al. 1990, Bromley 1992, Berkes and Folke 1998b, Castilla and Fernandez 1998, Paper V). Resource users have isolated “resources” from their ecosystems for too long (Papers I and II). The support of ecosystems for resource production has often been taken for granted. should incorporate knowledge of ecosystem dynamics and environmental feedback (Paper III and V) as emphasized in the ecosystem approach promoted in several recent governmental reports (National Research Council Committee on Ecosystem Management for Sustainable Marine Fisheries 1999, SOU 2003). Management must recognise that humans are part of these ecosystems by identifying the factors that drive users (Papers III and IV) like institutional and economic incentives (Lambin et al. 2003) and by finding ways of organising governance to redirect unsustainable patterns of exploitation to improve management of resources and ecosystems (Paper V). Successful management is defined here as exploitation that sustains an industry without compromising ecosystem function and resilience.

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METHODS

The methods used in the thesis varied from ecological fieldwork producing quantitative data in Papers I and II to literature searches, interview methods from the social sciences producing qualitative accounts of the history as well as semi-quantitative information on the lobster and conch fisheries in Belize (Papers III, IV and V). Papers III and IV use a historical approach and Paper V uses a comparative approach.

Papers I and II This study took place on patch reefs located in the no-take zone, “Conservation Zone”, of the and in the “General Use Zone” on the Glovers Reef Atoll, Belize (figure 3). Within each zone, a subset of these patch reefs was manually cleared of fleshy macroalgae (see Paper I) and functional groups were monitored over the following year (Paper II). The sample unit for analysis in both papers was the patch reef, except for analyses of fish behaviour that compared two locations on the patch reef (Paper I). While it is not possible to run field experiments over corals’ lifespans (>100yrs), the spatial scale of the experiment aimed to compensate for this (Done 1992, Porter and Meier 1992). The spatial scale studied ranged from benthic cover (centimetres) along transects (10m), to patch reef (~0.001 km2), to management zone (~5 km2), to the total area studied (~10 km2). While this led to variation in the results, it was a more relevant scale for the management aspect of the study. On a temporal scale, we studied the immediate effects (2 weeks later) of the algal reduction and monitored all patch reefs on three subsequent occasions over a year (total of 5 survey periods). By monitoring for a year, our time scale incorporated seasonal cycles in order to distinguish these from experiment-induced changes (Hughes and Connell 1999).

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Figure 3. Glovers Reef Atoll and the general areas of the experiment (CZ: Conservation Zone – no-take area; GUZ: General Use Zone)

Paper III Fisheries Statistics: Catch and export statistics were used to compile a time series for production of lobster and conch in Belize in Papers IV and V. Production is under-estimated throughout this time series. Until the 1950s, the colonial government only collected data on exports. The production data, which are available from 1977, exclude national sales that bypass the co-operatives. Until the 1980s, the catch bypassing the co-operatives is thought to have mainly consisted of illegal catch (undersized and berried females) that was taken for consumption at home, as the co-operatives offered the best prices. The growth of the tourist industry since the 1980s has increased the national market for both lobster and conch and a growing number of fishers are reported to be selling directly to restaurants. Interviews with co-operative board members revealed that restaurants are paying competitive prices with the co-operatives’ first payment (fishers are paid in two instalments), this means fishers are being paid less but as their landings are smaller, the fuel costs of landing the catch at the co-

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operative may not make this worthwhile. It was not possible to estimate the size of this market, although all of the co-operative board members interviewed reported that this was a growing problem. The number of fishers are estimates by the colonial government and in the literature and statistics from the Fisheries Department (1946-55: (Colonial Government 1947, 1954, 1955, 1957); 1957, 1960s, mid-70s (Gordon 1986), 1973: (Baird 1973); 1985: (Daly Price 1986); 1989: (Pinto and Vasquez 1989); 1993-1998: Fisheries Department Statistics in (CSO 1999); 1999: Fisheries Department Statistics in: (CZMAI 2000). The estimates are thought to underestimate the total number of fishers as most studies reported in the literature have focused on fisheries in the north of Belize. These figures are likely to be a better approximation of the number of fishers targeting lobster until the mid-1960s, when the co- operatives were established and increased the number of fishers throughout the country.

Interviews: Due to insufficient data to calculate changes in and the reliability of existing data, scientific studies and interviews were used to provide qualitative data to complement, contrast or verify existing data. Also, as there has been a bias in the literature on the fisheries in the north, a goal of these interviews was to collect data on the development of the in the south. Interviews were carried out during five field visits between 1998-2002. Between 1998 and 2000, various interview methods were tested (including surveys, questionnaires). Qualitative and semi-structured interviews (Kvale 1996) were chosen as they produced detailed accounts of change, which was particularly relevant for building up a history of the fishery and key events in its past. Stakeholders interviewed were fishers, co-operative board members as well as Fisheries Department officials and NGO representatives. These informants provided a wealth of background information on past and recent events in the fishery at local and national scales. Initially, co-operative board members were targeted. Co- operative board members interviewed in Belize City, and later in San Pedro, Placencia and Punta Gorda, were initially randomly selected as it depended on who was available at the time. All of these informants were re-visited during subsequent field visits and became key informants. Fishers were interviewed in Belize City at the landing sites at the two co- operatives and on the local market. These fishers were randomly chosen, depending on who was present. Fishers were also interviewed on Caye Caulker. Due to limited time and details of fishers on the , fishers were selected using snowball sampling (Bernard 1994). A researcher on the island recommended the first fisher. Subsequently, each interviewed fisher

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was requested to recommend three others. Usually only one of these was interviewed, three were requested in case people were unavailable or at sea. In 2002, sixteen key informants were interviewed and of these 11 were interviewed in depth. Key informants are here defined as: observant, reflective and articulate and having the ability to intellectualise their own culture and to facilitate access to other important informants (Bernard 1994). Due to time constraints in the field, individuals able to place the fishing in a broader context were targeted. A long history in the fishery or involvement with other stakeholder groups was criteria that were desired. All informants had at least ten years experience of the either as fishers and/ or as board members of the co- operatives. At least one board member from each co-operative was included. Co-operative board members were rich sources of information. Most had long personal experiences as fishers and were knowledgeable on their membership and trends in catches at local and national scales. These board members also had working experience of the economic driving forces, such as export markets and the economic effects of hurricanes and changes in catch on the fishers. The co-operative in Punta Gorda was recently established at the time of this fieldwork, and the chairman of the board had not been a member of the former co- operative. A local NGO recommended a former fisher, now tour guide. This informant had over fifty years of experience of fishing in the area, had been a productive member of the former co-operative in the 1960s and had knowledge of local history that has affected fishing in the region. Due to the interest in developing a more complete history of the fishery’s development in the south, informants from Fisheries Department, Co-operative Department and local NGOs working in the region with the fisheries were interviewed and provided information on recent events and issues in the fishery. Interviewees were questioned on the history of the fishing industry and key events in the past, the organisation of and their roles in the industry. These interviews produced a wealth of information that was then separated into different categories that allowed comparing the data between interviews as well as with the literature. There were no important differences in data between the interviews or between different types of stakeholders, with the exception of views on the efficiency of marine reserves that differed between those in the fisheries and NGO representatives.

Paper IV The analysis is based on existing scientific literature, other written sources and official statistics of Thailand as well as meetings and interviews with stakeholders dealing

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with the industry in Thailand. Information gathered from written sources were verified and complemented during a visit to Thailand in November 1997. This was done through a seminar at the Department of Agriculture and Resource Economics, Kasetsart University (Bangkok), and through individual meetings with representatives from NGOs, universities, and government officials and organisations. Visits to the Thai Development and Research Institute (TDRI) and the Network of Aquaculture Centres in -Pacific (NACA) libraries allowed the collection of further information.

Paper V The information for the analysis of causes of changes in lobster yields in Belize was collected from the scientific literature (Espeut 1994, Jacobs 1999, Heyman and Graham 2000, Huitric et al. 2003), statistics on catch and number of fishers and interviews with stakeholders in Belize (see above for details). This results were compared with past interview studies in Belize (see Appendix 1 in Paper V). The information for the Maine fishery was collected from the scientific literature. The main sources used were: (Acheson 1988, 2003, Steneck and Wilson 2001).

SUMMARY AND CONCLUSIONS OF THE PAPERS

The main hypotheses of Papers I and II were: 1. Overfishing in the lagoon of Glovers Reef Atoll has reduced herbivorous fish populations and greater fish abundance was expected in the marine reserve. 2. Fish abundance, of herbivores in particular, was expected to increase on experimental patch reefs. These herbivores would control subsequent macroalgae growth. 3. Manual reduction of algae is a viable management option at local scales in areas with increased . Overall, management had no significant effect on fish abundance and the algal reduction had short-lived impact on these patch reefs. The effect of large-scale disturbances on the results could not be determined, however, the resilience of the desired coral-dominated state of Glovers Reef Atoll, seems to be low. Papers I and II illustrate that the benefits of setting aside marine reserves in already impacted environments should not be taken for granted. Setting aside reserves will not be sufficient to conserve the reef heritage (Allison et al. 1998,

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Castilla 2003). Instead reserves should be viewed as one tool amongst others to improve and enhance ecosystem services in the overall seascape, recognising that human actions are part of and not excluded from ecosystems. Papers III and IV expected a mismatch between the control and the effect of resource use. It was also hypothesised that environmental feedback would be missed or neglected by the institutional structure controlling users. Both studies concluded that the introduction of new technology under inadequate property rights masks environmental feedback and contributes to pathological resource use. A major conclusion of Paper III was that social resilience of organisations does not ensure ecological resilience if management of resource use is not based on the ecological system of the resource. Paper V was based on the findings from Papers III and IV and used a comparative approach to identify factors that may contribute to successful resource management. A result of Paper V was the difference in incorporation of ecological knowledge of processes affecting lobster abundance in the regulations governing the two fisheries. A conclusion of this study was that institutional structures should incorporate existing knowledge as well as enhance the generation of new knowledge and learning. In all three case studies of Papers III-V, a single centralised governing body failed to implement existing regulations. A result of Paper V was the importance of a diversity of organisations and institutions to manage ecological and human processes at relevant scales.

DISCUSSION OF MAJOR FINDINGS

Ecosystem Dynamics & Management on Glovers Reef Atoll The immediate effects of the study on Glovers Reef Atoll were short-lived and over the course of the year the sites returned to their pre-reduction conditions (Paper II). The general conclusions of this study were that herbivory had no impact on the experimental shift and that manual reduction is not a viable management option on these patch reefs at present levels of herbivory. We could not test the management effects on overfishing as we only had one management replicate and therefore differences between management zones could simply be due to different environmental conditions in the lagoon (this was the case for coral cover see: Paper II). Also the reserve on the atoll was established in 1993 (McField et al. 1996), which is little time to restore the herbivore functional group. Three possible explanations for the fast return of macroalgae cover were the interference of a hurricane and bleaching event with the experiment,

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that past and/or present large-scale processes of and overfishing beyond the scale of the experiment have caused this shift or a combination of these factors. The experiment was affected by a massive bleaching event in September 1998 that bleached most corals in the lagoon area (Huitric and McField 2001). Bleaching is defined as “massive” when >25% of coral colonies are bleached over geographic areas, with some individuals more than 90% bleached (Hoegh-Guldberg 1999, McField 1999). A month after the algal reduction, Hurricane Mitch’s storm surge caused structural damage to the atoll’s forereefs, although it appeared to have caused comparatively little damage in the lagoon (personal observation, April 1999). The hurricane also led to heavy rainfall on land that caused a freshwater plume that extended from the mainland and covered half of the atoll for several days (Andrefouet et al. 2002). From December onwards, grazing and the abundance of Acanthuridae (herbivores), coral and turf cover decreased (Paper II). As these events were unplanned, our experimental design could not assess these disturbances’ effects on our results. The hurricane’s storm surge may have caused , which allowed macroalgae to recover. Although the lack of visible structural damage on patch reefs in the lagoon, attributed to dissipation of wave activity by the shallow eastern reef crest and Middle Caye (figure 3), indicated less wave activity in the lagoon, it might have been enough to affect fish. The freshwater plume following the hurricane is expected to have been an important nutrient pulse to the atoll and may in part explain the monitored in December 1998 (Paper II). Together with coral mortality and reduced resilience due to both the bleaching event and the hurricane, this may have allowed the rapid return of macroalgae into the areas that were algae had been removed. These disturbances demonstrate the atoll’s interaction with systems at different scales. The low herbivore abundance and the return to pre-reduction levels suggest that the algal stability domain is resilient to change. If this shift is a result of past ecological extinctions, in this case the herbivore functional group, then the scale of this experiment could neither detect this nor control the shift. This study confirms a shift in the baseline environment from coral- to macroalgal- dominance that is difficult to shift back. It may be irreversible. These conclusions are relevant to the systems studied in Papers III and IV, shrimp farming and lobster fisheries, as both are linked to the problem of fishing down the food web. These results, and uncertainties, illustrate the limitations of marine reserves as management tools in disturbed environments. Due to the scale of the shift, spatially in the lagoon and Caribbean and possibly temporally (Jackson et al. 2001), it will not be enough to remove

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fishing pressure, as the small-scale is overwhelmed by the conditions at larger scales. Restoration will require larger interventions than a local marine reserve.

Sequential Exploitation & the Pathology of Resource Use A result in papers III and IV was the description of spatial sequential exploitation (see figures 4 in Paper III, IV). In both cases this movement was driven by local declines in and resulted in dependence on fossil fuels to maintain production. Paper III developed the sequential exploitation concept by finding additional dimensions to spatial sequential exploitation. These were the sequential exploitation of technology from large to small targets and finally changing targets. These dimensions have been described in resource exploitation literature (Regier and Baskerville 1986, Ludwig et al. 1993, Holling and Meffe 1996), although not necessarily in the context of sequential exploitation. These additional dimensions helped to indicate changes in the resource as well as possible ecosystem effects of exploitation, for example, the replacement of bully nets with skin diving for targeting lobster on the reef in Belize (Paper III), which is useful when data are limited (Berkes et al. 2001). A second result was identifying sequential exploitation as a key factor leading to the pathology of resource use (Papers III, IV). In Thailand, collapses on farms due to and disease were solved with sequential exploitation, as it was cheaper to move and restart than to invest in water treatment (Paper IV). These pollution problems were related to the loss of mangroves, which act as a water filter and barrier (Robertson and Phillips 1995, Rönnbäck et al. 1999). This is an example of how externalities can accumulate to become disturbances at larger scales and may create a positive feedback loop if sequential exploitation continues. While these environmental problems had reduced the average lifetime of a farm to five years, the return on investments required a single harvest and made the risk worthwhile. The potential profits and price of land drove the shrimp farming industry and illustrates the role of economic incentives and property rights as driving forces to pathological resource use. As this industry grew relative to its ecosystems, by means of capitalisation, dependence on constant or increasing yields rose (Costanza and Folke 1996). This was also observed in Belize where the co-operatives responded to stabilised catches by increasing their capitalisation and supporting members’ debts (Paper III). In other words, the industries responded to feedback from the social system rather than from the environment (Paper IV). This pattern of resource use works in the short-term, but it increases the scale of

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problems as seen in examples of fishing down food chains (Koslow et al. 1988) and the effects of mangrove loss (Primavera 1997, Mumby et al. 2004). The effects of these activities are often gradual (Jackson et al. 2001), which can result in shifting baselines, as users do not perceive the change (see p. 23 Paper III). Lack of knowledge and understanding of ecosystem function can also delay response to change as well as lead to the use of incorrect baselines for management (Pauly 1995).

Institutions & Organisations None of these case studies were true open access. However, one conclusion was that the lobster fishery in Belize and mangrove use by the shrimp farming industry in Thailand had de facto open access to these resources. This was due to mismatches between the rules controlling resource use, property rights, the structure of the implementing organisations and the industry’s and natural systems’ respective states (Papers III-V). Also, the ability to prevent free riding does not ensure the sustainable use of a resource (Paper V). Property rights and rules developed for sustainability are useless without ecological knowledge. The first regulations controlling shrimp externalities in Thailand were introduced in 1991, late in the industry’s development (Paper IV). Of these rules, some were inappropriate (Paper IV). Furthermore, the waste treatment rules only applied to farms larger than 8 hectares, <5% of the number of farms, and therefore failed to match the scale of the industry (Paper IV). Comparison of the conservation rules used in the lobster fisheries of Maine and Belize showed a lack of ecological knowledge in Belize’s rules (Paper V). The effect of these conservation rules in Belize could not be judged due to lack of enforcement. Three governance systems were represented in the thesis, with rule-making and enforcement carried out by: 1. A central government (all cases) 2. The users themselves at a local level (Maine, northern Belize) 3. Both the local level and the central government (Maine) In all of these examples, the legal enforcing body was a centralised organisation. The success of state-owned and state-run property assumes that its control is effective in management and that enforcement will be complete (Adger et al. 1997). Central governments often lack funds and personnel to ensure full monitoring of both the resource and users, leading to de facto open access (Shanmugaratnam 1996, Papers III-V). As the resource system increases in size, environmental and social heterogeneity increases, which in turn raises enforcement costs.

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In addition, legal property rights did not limit entry. This allowed influxes of new users when production rose, whether naturally or due to new technology or prices, and in turn contributed to sequential exploitation (Papers III, IV). In comparison, sequential exploitation was slower in the Maine lobster industry despite the existence of technology to access more distant grounds and unlimited entry in the legal property rights (Acheson 2003). In reality, entry to the fishery and enforcement of state and local regulations were controlled at the harbour level (Acheson 1988). Paper V concludes that this local property rights structure slowed sequential exploitation as it increased the costs of expanding one’s grounds, fuel and gang wars, compared to in Belize and Thailand. Decentralisation reduces the group size managed by each authority (Ostrom 1993) (Paper V). Social sanctioning, as took place in Maine, lowers the costs of enforcement and monitoring (Ruttan 1998). This requires shared generalised rules of reciprocity and trust among users (Ostrom 1993, Trosper 2003). This kind of social capital, however, is often missing in large heterogeneous groups. Co-operation becomes more difficult when the size of the group increases and/or is not constant (Ostrom 1993), as actors do not have repeated contact with the same individuals (North 1990). For example in the fisheries in Belize, there is a large number of individual fishers working throughout the and in the Thai shrimp farming industry there are many individual shrimp farms and a high turnover of farms in local areas (Lebel et al. 2002). This seems to limit the ability for collective action (Paper IV). As the size of harbour gangs and their territories grew in Maine, the traditional property rights began to break down and the state of Maine introduced co-management of the lobster fishery (Acheson 2003). This property regime is a mid-way point between common property and state property, involving both stakeholders and officials in decision-making. In this case a new management level was introduced that bridged the central government agency with the local harbour gangs. This created several management areas that allow experimenting with policy, the ‘Learning-by-doing’ process of adaptive management (Gunderson et al. 1995, Walters 1997). With a single governing body, policymakers experiment simultaneously with all of the natural resources in their jurisdiction. If management is delegated, the risk of failure is spread, reducing the risk of large-scale disasters. An additional advantage of decentralising resource control to smaller-scale units is being able to adapt rules to local conditions (Paper V). In this sense co-management increases collaboration across scales without losing the ability to provide institutional frames for local action, which Folke and others (2003) refer to as ‘framed creativity’. Organisations involved in resource use and its governance may have conflicting goals to those of sustainable resource use. Paper IV identified measures taken by the Thai

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government that directly and indirectly subsidised the shrimp farming industry, but conflicted with measures to control the industry. Incomplete valuation of ecosystems, lack of environmental feedback and the assumption that resources are renewable, or would recover when exploitation could be reduced, were found to lead to the pathology of resource use (Papers III, IV). This is also related to an emphasis on short-term versus long-term profit maximisation at individual, organisation and state levels. In Belize, co-operatives provided members with subsidies and incentives to capture economic rent despite falling catches per fisher (Paper III). The co-operatives were established to increase income to members and not to manage resources in a sustainable manner. An important conclusion of Paper III was that socially resilient organisations do not necessarily result in ecological resilience of the ecosystems they exploit.

The Problem of Fit Between Ecosystems & Institutions is a result of a mismatch of scales, when management and natural resource appropriation do not correspond to the spatial, temporal or functional scales of natural phenomena (Lee 1993). A sustainable harvest in the context of the capacity of the ecosystem, rather than a desired yield from a system, should be management’s goal and is the basis of ecosystem management (Christensen et al. 2003). As seen in Papers I and II, no ecosystem is closed, or isolated, from its surroundings. Even heavily-managed ecosystems remain dependent on subsidies from other ecosystems. This is taken to an extreme in intensive shrimp farming that relies on fishmeal that can be imported from, for example, to Thailand (Naylor et al. 2000). Monocultures produce a high output of a specific product, but are less resilient than complex systems. Disease spreads rapidly within and between shrimp farms in Thailand due to the high stocking densities in ponds and high densities of farms with little water treatment, as well as being related to the loss of mangroves that act as filters (Dierberg and Kiattisimkul 1996). Similarly, high densities in the Caribbean could control macroalgal growth but also set ideal conditions for the spread of disease. A holistic approach to managing resource use is, therefore, needed in order to maintain the flow of goods and services into the future. Complexity and diversity maintain ecosystem function and resilience. Yet humans tend to focus on a small subset of ecosystems, set management boundaries that do not coincide with the ecological boundaries (Jansson 1991, Odum 1993) and apply a small set of rules to a wide range of resources and appropriators (Ostrom 1998). Management can then fail unexpectedly if a disturbance comes from beyond the boundaries it has set itself. The

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findings from Papers I and II suggest that both complexity and diversity have, in the past, been reduced on the patch reefs of Glovers Reef Atoll. Together with the effects of the disturbances, which showed that this system is embedded in larger scale systems, this study demonstrates the limitations of marine reserves to deal with the shift to algal domination. This finding is relevant at a regional scale, as this shift has been reported throughout the Caribbean.

Responding to Environmental Feedback Social-ecological systems respond to feedback from the resource and social systems. In all of the examples in the thesis, environmental feedback signalled negative effects of exploitation. They differed, however, in whether there was an adaptive response or not. Coping is getting along with change. Adaptation involves learning. In a sustainable social- ecological system there is learning about ecosystem dynamics and how to respond to environmental feedback. In each case, the users initially responded to changes in resource through some form of sequential exploitation. In Thailand and Belize, this was spatial while, in Maine, this was from large to small individuals in local grounds. In Belize, institutional response has been to establish marine reserves although, as seen in Papers I and II, this may have little effect due to the history and spatial scale of fishing pressure. The institutional response in Thailand came late and was not well suited to the situation. In both of these cases, this is effectively the same as no response (Berkes and Folke 2002). The analysis of the lobster industry in Belize showed that despite awareness of the change and its impacts on users, there was no appropriate institutional response based on their understanding of the cause of change. Governance systems should be capable of adapting to changes (Brown et al. 2002). Social resilience is built and invested in robust institutions (Janssen et al. 2003). In Maine, there have been two important examples of response in the history of the industry. The change in perception in the fishing community regarding conservation rules that set the path for the development of a “conservation ethic” (Acheson 2003) in the industry. Some local harbour gangs have established additional conservation rules (Acheson 2003). This response is based on learning (Berkes and Folke 2002) from existing regulations that have been further adapted or complemented. The territorial system broke down as the users within and at the boundaries of the territories increased beyond a certain threshold. One finding of these studies is the need to incorporate social mechanisms in the institutional setting that can respond to environmental feedback as well as to changes in the social system (Pretty and

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Ward 2001, Brown 2003). The finding is in line with earlier work (e.g. Berkes and Folke 1998b).

Measures to Prevent the Pathology of Resource Use Institutions and property rights should manage feedback between ecological and human systems. Ecological knowledge is never complete and needs to be constantly updated. The institutional structure should be able to incorporate new knowledge of the ecological system or to adjust its structure as the industry changes (Paper V). Governance’s goals should be for the long-term as users’, and their organisations’, goals are rarely to conserve per se (Ruttan 1998, Papers III-V). Different resource conditions demand different property rights and rules. As seen in the development of the lobster fishery of Maine, there is not a standard institutional set-up for resource management, rather it must be able to adapt to changing conditions in the social and environmental systems (Berkes et al. 2003b). An institutional context for monitoring and responding to feedback is needed. Currently, there are proposals for adaptive management approaches combined with co-management. Such adaptive co-management systems can develop within decadal time scales (Olsson et al. 2004). A difficult question that policy makers will have to determine is whether management will make a difference or not. In the case of the patch reefs in the lagoon on Glovers Reef Atoll, can coral dominance be restored or is the macroalgae state too resilient? The algal reduction was an intensive effort, albeit at too small a scale, but the speed of return of macroalgae cover suggests restoring these patch reefs will be difficult. Will a large-scale intervention be enough and what should that be? Simply restoring herbivore populations through reduced fishing pressure is unlikely to suffice. What additional measures can be implemented at such large scales as the Caribbean? The recovery in lobster catch in Maine has been attributed in part to conservation measures by fishers. Additional factors have been the removal of predators in nursery areas and higher water temperatures. The latter is beyond management’s control. Would the conservation measures have had any effect without these environmental and ecological changes, and would fishers have developed the conservation ethic at all had recovery taken longer than the ten years it did? Additional questions have emerged during these studies that are either fuel for future study or factors that are known to have effected the observed results but lie beyond the scope of the author(s):

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• Is over-exploitation inevitable? Global trade is an example of a driver creating cross-scale interactions in social-ecological systems. It has also helped “masking” environmental change as users and consumers can easily switch between producers at a global scale, sometimes without even being aware of this switch. The feedback is exported out of sight in time and space (Holling et al. 1998). Tiger shrimp production is largely produced for the global market. Does this industry need to be managed at a global scale in order to internalise externalities (see Paper IV)? Similarly, does spiny lobster fishing need to be managed at a regional scale? Belize’s neighbours also have lobster fisheries and therefore markets. When their regulations do not match those of Belize it creates opportunities for fishers to fish illegally. Also due to the potential spatial scales of source and sink populations, the measures taken in one country may not benefit its own fisheries.

• Is crisis necessary for adaptive response? While crisis can spark institutional learning (Holling and Sanderson 1996), will crisis alone spark reformation or can resilience be developed to reduce the intensity/occurrence of a crisis (Gunderson 2003)? Can the fisheries in Belize learn from others’ mistakes and successes, or must it undergo a crisis itself to do so?

• Is a history of social capital a prerequisite for successful local/co-management (Pretty and Ward 2001)? As seen in the lobster fishery of Maine, when participants share generalised norms of reciprocity and trust, this social capital can be used as a foundation in resource management within the group. In the 1930s when fishers decided to comply with regulations, these could be incorporated into their existing institutional structure. This social capital is often lost or absent in rapidly growing industries, such as the shrimp farming industry in Thailand and the fisheries in Belize. In the case of Belize, fishers were able to mobilise into co-operatives despite a lack of support from the government. Is this social capital still present and how can it be mobilised and will fishers want to?

CONCLUDING REMARKS

The thesis has studied ecosystem dynamics and resource users’ and managers’ interactions with these dynamics. The different papers illustrate that and coastal ecosystem management is no longer only a matter of local decisions. The success of such efforts is a reflection of the state and condition of the larger seascape in which

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resources and ecosystem services are embedded. Furthermore, the thesis also illustrates that although knowledge and understanding of resource and ecosystem dynamics is a precondition for successful ecosystem management it is not a sufficient condition. The social dimension needs to be accounted for as well. However, analysing the social dimension alone does not provide sufficient answers and direction for sustainable resource use. For example, the organisation of local fishermen in Belize into co-operatives and investments into fossil fuel based technology may seem to be socially sustainable responses, but these exacerbated the exploitation of the resource. The direction of resource exploitation (like fishing down food webs) and land use change (like removing mangrove ecosystems) will impact the resilience of the seascape. Incentives, drivers and policies further influence ecosystem governance. The social-ecological perspective applied in the thesis highlights the interactions between ecosystem dynamics, ecological knowledge of these dynamics, and incentives, institutions and governance systems that are in tune with these dynamics. The necessity to address cross-scale issues becomes obvious. More specifically the conclusions of the thesis are as follows. (A) The failure of the experiment on Glovers Reef Atoll to locally shift from macro-algae to coral dominance was due to the fact that the small-scale was overwhelmed by the large-scale. This study demonstrated the limitations of marine reserves as a means of restoring damaged environments. The goals of marine reserves need to be based on the state of the seascape. It seems that in the case of the patch reefs a larger intervention than just adding herbivores is needed to revert to coral dominance. (B) The thesis combined different patterns of sequential exploitation (resource, spatial and technological) and linked these to the pathology of resource use, with particular focus on the role of technology to accelerate use and increase the disconnection between user and resource. The concept of pathology of resource use (Holling and Meffe 1996) allowed me to combine sequential exploitation (Grima and Berkes 1989) with shifting baselines (Pauly 1995), and the social dimension of resource use and ecosystem management with technology and institutions into a linked social-ecological system (Berkes and Folke 1998a). C) The thesis identifies conditions leading to the pathology: Insufficient incorporation of ecological knowledge in the rules controlling resource use, easy access to the resource, including the use of technology, without appropriate property rights, and the absence of enforcement of rules. A conclusion of the thesis is that under certain conditions a

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lack of ecological knowledge can lead to overexploitation and this is enhanced if feedback from the ecological system is not included in institutions and governance. (D) Ecological or social resilience is not the same as social-ecological resilience. An ecosystem may lose resilience and shift into a less desired state. But if the social system can reverse this shift back into the desired state the social-ecological system has resilience. A resilient social-ecological system has understanding of ecosystem dynamics that is incorporated in the institutional setting. Such an understanding was lacking in the working rules of the co-operatives or the territories in the north of Belize. This is an important result for social analysis of resilience and vulnerability. It is not sufficient to draw conclusions about sustainability based on studies of e.g. the ability of groups to mobilise into collective action. Their ability to respond to environmental feedback without degrading the ecosystem needs to be accounted for as well. (E) Paper V illustrates that there are no set property rights regimes for given resources. Rules governing use should be based on ecological knowledge of the resource and its ecosystem and the structure of the user and governance systems should match environmental conditions and social structures. Co-management arrangements and organisational diversity seem to improve the capacity to deal with changes in the natural and social systems.

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APPENDIX I – Errata to Papers I and II

PAPER I ERRATA Results P 276: Benthic cover: …Management did not affect benthic cover (ANOVA, p > 0.05) except that coral cover was higher in the general use zone (p < 0.05). Should read: …Initial analysis found no significant differences between management zones in September and October respectively (t-test results: September – vs. General Use: hard coral p < 0.05 (ns); fleshy algae p < 0.01 (ns); October – Wilderness vs. General Use: fleshy p < 0.05 (ns)). Management did not affect benthic cover (ANOVA, p > 0.05) except for coral cover which was higher in the general use zone (p < 0.005*).

PAPER II ERRATA

Data analysis (Paragraph 1, sentence 6)… Each benthic category was tested for normality and transformed as necessary prior to analysis of variance (Tables 1, 2 and 3) using Statistica 4.0 software package.

(Paragraph 1, sentence 8)… Multidimensional scaling plots were created from square-root- transformed benthic community data (percent cover) using PRIMER ecological statistics as described in Clarke and Ainsworth (1993). Patch reef data (coral species, fleshy algae genus and the remaining categories) were averaged to give percent cover per treatment per management zone per time period.

(Paragraph 2, sentence 6)… In this study, both fish and benthic data consisted of numerous variables, therefore, the sequential Bonferroni technique, which adjusts the p-value according to the number of variables included, was applied (Holm 1979). For the presentation of statistical results, we use the symbols * p < 0.05; ** p < 0.01

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and *** p < 0.001, in order to relate the adjusted p-values to their unadjusted counterparts.

Results Benthic cover and algal biomass (Paragraph 2, sentence 2)… Fishing exclusion had little significant effect on benthic cover, with the exception of calcareous algae, which had higher cover, and stony coral, which had lower cover in the Wilderness Zone. While several interactions were significant, none were greater (lower p-values) than the single factor effects.

(Paragraph 2, sentence 10)…There was a significant increase in branching coralline algae over time (p = 0.0000 ***; Tables 1 and 2) with significantly lower abundance in experimental patches (P=0.0008 **; Table 1 and 2).

(Paragraph 4, sentence 2)… These net changes (% cover September 1999 to September 1998) included: increases in Lobophora 5.8% (P< 0.003*), branching coralline 2.5% (P< 0.001**), encrusting coralline 1.4% (P< 0.006*), Dictyota 2.9% (P< 0.0001***), Padina 1.1% (P< 0.002*), fleshy algae (grouped taxa) 12.3% (P< 0.008**), and a decrease in stony coral 9.5% (P< 0.0001***).

(Paragraph 5, sentence 2)… Despite this, there were significant effects of time (P=0.0000***) and treatment (P=0.0001***) for the total algal biomass (Table 3) with lower algal biomass in the experimental versus control reefs and increased algal biomass over time after the reduction.

Fish population estimates (Paragraph 1, sentence 2)… Management significantly affected four of the 30 species presented, all of these were more abundant in the Wilderness Zone (Table 4).

(Paragraph 2)… Among the species pooled into families, the Acanthuridae were significantly more abundant in the Wilderness than General Use Zone (P = 0.0000005***, df = 4). Acanthuridae and Labridae both exhibited significant effects of treatment (P =< 0.00041* and p = 00007** respectively) as well as time (P < 0.001*** each). The large parrotfish

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S. viride (scraping and excavating turf algae) was affected by both treatment (P = 0.000003***) and time (P = 0< 0.001***) (Table 4).

(Paragraph 3, sentence 1) …The Acanthuridae and Labridae also exhibited a significant effect of time (P = 0 < 0.001 ***, df = 4).

Discussion (Paragraph 3, sentence 2)…Management effects could be confounded by differences in wave energy, water or some other geographically influenced environmental factor between the two zones, which were separated by approximately 7 km. There were no significant differences between the two management zones in September 1998.

(Paragraph 4, sentence 2-4)… The effect of protection was apparent for three species of the fish species examined, all of which were more abundant in the Wilderness Zone. These species were herbivores, and included the scraping , Scarus viride. Among the families, surgeonfish (Acanthuridae) were more common in the Wilderness Zone.

(Paragraph 6, sentence 1-2)…The three species and the Acanthuridae family as a whole, and the Labridae family, were affected by the treatment and exhibited a higher mean abundance on experimental reefs. Among them were the most common scraper/excavator Sparisoma viride and Thalassoma bifasciatum.

(Paragraph 9, sentence 5)… These findings suggest that management may have an effect on fish assemblages, but that it may require additional time before this effect cascades down to the .

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ACKNOWLEDGEMENTS

It’s almost all over. I can almost hear your sighs of relief. I am fully aware that this ordeal has been just as big for many of you as it has for me. And come what may in the next decisive weeks, it was worth it. I have lived a lifetime in these years, and met people I hope to share the next lifetime with. As it was all drawing to an end I wondered how I came to be here. This would make quite a good story in itself, but another time. I do need to thank fate, Lisa and Nils. These brought me to Sweden to do my masters. Nils, thank you! As I got started Calle came into my office and asked if I wanted to continue and do a PhD. I said “yes”. I am not sure I’ve ever told you, but my mind was asking “what the hell is she saying?!”. So you changed my path. Thank you. You let me do something I never imagined I would. Thank you also for your support in these last few months and your super-vision. Thanks to all of you at the department, special thanks to Barbara Pol for having made everything so easy – I hope you understand how much this has helped. Lisa, Maria and Line - you have been my critics, editors, sources of ideas and knowledge, and true friends. I look forward to carefree meetings and time together. Åsa and Lisa – if all else fails, I am counting on a music career with the Karamellerna – now there is another thing I never thought I would do! I would like to thank the Beijer Institute for having taken me in. More importantly, I want to thank each of you for making the Beijer such a great place to be. Anna and Christina thank you for all of your help and care. Jessica, it is almost over, the wine is chilling and we will recover slowly in the sun. Thank you. Melanie, you are a role model and I would love to have the chance to work with you again, or at least time to improve our tofu flambé recipe. I also want to thank you, Ian and Mr and Mrs McField for having made Belize a home. I owe special thanks to Nina. You have followed this thesis’ progression very closely. (Any grammar mistakes are mine and mine alone – it is my insatiable need to re-edit everything). I look forward to our next meeting. Finally my family. Lisa, Gustaf and Anna-Britta, goodness, what can I say? This would never have been possible without you. Never. Papa, this is why I have been out of touch – but I’ll soon be back at the end of the line! I am one of those few to have the best brother and sisters in the world – each of you have helped in your own way, as you always have. If I had not had you to influence me, I would be a very dull person today.

It is now time for me to go home and take care of you, Jorge, who has put up with this never-ending thesis. It’s our turn now. Até já querido X.

This work has been funded by a grant to the Beijer Institute from the John D. and Catherine T. MacArthur Foundation and the Swedish Research Council for Environment, Agricultural Sciences and Spatial Planning (Formas) and by the Pew Fellowship Award. I owe special thanks to: Dr. Paul Erftemeijer; Jasper Goss; Khun Tipparat Ponthanapich; Khun Buntoon Srethasirote; Khun Siri Tookwinas; Yad Fon Association; NACA for use of their library. In Belize I would like to thank the Fisheries Department and Coastal Zone Management Unit who have always taken the time to meet with me, special thanks to Janet Gibson, Rachel Graham, TIDE and Wil Heyman. To the interviewees in Belize, I would like to have thanked each of you in person. The wisdom that you hold is invaluable, and I hope that someone will be able to transform it into a library one day soon.

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