Ecology and Implications of Recovering Previously
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ECOLOGY OF RECOVERING DEGRADED REEF COMMUNITIES WITHIN NO-TAKE MARINE RESERVES by JONATHAN ALBURO ANTICAMARA B.Sc., University of the Philippines in the Visayas, 1994 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY in THE FACULTY OF GRADUATE STUDIES (Resource Management and Environmental Studies) THE UNIVERSITY OF BRITISH COLUMBIA (Vancouver) May 2009 © Jonathan Alburo Anticamara, 2009 Abstract No-take marine reserves are a highly advocated tool to recover degraded marine ecosystems, but we have limited evidence as to how marine reserves facilitate recovery of marine communities. To address this limitation, we conducted monthly underwater visual censuses over three years on 423 reef fish species in eight sites where fishing had been excluded for different lengths of time. We then used our data to assess four impacts of protection within no-take marine reserves in the central Philippines: (1) magnitudes and rates of reef fish community recovery; (2) changes in reef fish diversity; (3) patterns of reef fish community succession; and (4) shifts in community interactions, based on distributions of pairwise correlations among reef fish species biomass. We found that total fish biomass increased with the duration of protection, but total fish abundance and species richness or diversity were both more influenced by site location than by reserve age. In addition, large-bodied herbivores drove the biomass recovery in older marine reserves, while small-bodied zoobenthivores and zooplanktivores influenced the higher abundance in offshore sites. Moreover, our results showed that ubiquitous large-bodied herbivore species (e.g. Chlorurus bleekeri) increased in biomass dominance in older reserves, whereas ubiquitous medium-bodied species (e.g. Thalassoma lunare) lost biomass dominance. Our non-metric multidimensional scaling (MDS) representation of reef fish community trajectories with duration of protection showed convergent trends in sites within similar locations relative to the mainland. Finally, the frequency distribution of pairwise correlation values among species biomass time-series within each site showed positive mean values regardless of protection-duration, as is typical of disturbed or high diversity systems. Indeed, less than ten percent of common species (those present in ii ≥50% of the 33 monthly surveys) within each site showed significant decline over time, while about 40 percent showed significant increase. In summary, our research provides comprehensive evidence on how marine reserves recover depleted reef fish communities. However, it also emphasizes that understanding of reef ecological processes could improve marine reserve site selection and design in order to meet specific conservation goals of marine reserve establishment. iii Table of Contents Abstract..........................................................................................................................ii Table of Contents ........................................................................................................iv List of Tables ..............................................................................................................vii List of Figures........................................................................................................... viii Acknowledgements .....................................................................................................ix Co-authorship Statement ........................................................................................ xiii 1. Introduction..............................................................................................................1 Theoretical framework...................................................................................................2 Disturbance and recovery of ecological systems ......................................................2 No-take marine reserves: issues, theory, empirical evidence ...................................4 Thesis context ................................................................................................................8 Thesis development .....................................................................................................12 References....................................................................................................................14 2. General methods .....................................................................................................29 Study sites ....................................................................................................................31 Benthic habitat survey..................................................................................................32 Fish Underwater Visual Census...................................................................................34 Conversion of fish length into biomass .......................................................................35 Categorization of fish species into body size groups and trophic groups....................36 Standardization of surveyor’s skills.............................................................................36 General data analyses...................................................................................................38 Tables...........................................................................................................................42 Figures..........................................................................................................................48 References....................................................................................................................53 3. How much and how quickly can reef fish communities recover within no-take marine reserves? .............................................................................................................58 Introduction..................................................................................................................59 Methods........................................................................................................................61 Data analyses ..........................................................................................................62 Results..........................................................................................................................63 Whole assemblage ..................................................................................................63 Body size classes.....................................................................................................64 Trophic groups........................................................................................................67 Discussion....................................................................................................................69 The influence of protection-duration on fish communities.....................................69 The influence of location on fish communities.......................................................74 Interactions between the effects of protection-duration and site location ..............76 Implications for conservation and management .....................................................76 Figures..........................................................................................................................80 References....................................................................................................................86 iv 4. Quantity versus quality: spatio-temporal variation in reef fish diversity within no-take marine reserves .................................................................................................92 Introduction..................................................................................................................93 Methods........................................................................................................................96 Species accumulation curves ..................................................................................96 Species diversity indices.........................................................................................97 Abundance-Biomass Comparison (ABC) curves ...................................................98 Results..........................................................................................................................99 Species accumulation curves ..................................................................................99 Species diversity indices.......................................................................................100 Abundance-Biomass Comparison (ABC) curves .................................................100 Discussion..................................................................................................................101 Figures........................................................................................................................106 References..................................................................................................................111 5. Patterns of reef fish succession within no-take marine reserves ......................120 Introduction................................................................................................................121 Methods......................................................................................................................125