MECHANISMS BY WHICH MARINE PROTECTED AREAS ENHANCE FISHERIES BENEFITS IN NEIGHBORING AREAS A DISSERTATION SUBMITTED TO THE GRADUATE DIVISION OF THE UNIVERSITY OF HAWAI‘I AT MĀNOA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY IN ZOOLOGY SPECIALIZING MARINE BIOLOGY December 2012 By Aileen Maypa Dissertation Committee: Charles Birkeland, Chairperson Kathleen Cole Kem Lowry Les Watling Alan White Keywords: marine protected areas, spillover, Caranx ignobilis, Caranx sexfasciatus 1 This dissertation is dedicated to the memory of my loving parents, Gloria M. Pascual Maypa who first taught me how to write and Leif F. Maypa who first taught me how to snorkel on a coral reef. 2 ACKNOWLEDGEMENTS This dissertation is my contribution to the agenda of food security, coral reef fisheries management and conservation in the Asia Pacific. I am grateful to the tireless support and expertise of my advisor, Charles Birkeland. I also thank my dissertation committee for helping me shape this work: Jeffrey Drazen, Katheleen Cole, Les Watling Kem Lowry and especially Alan T. White. This work could have not been completed without the tireless efforts and patience of my research team: Analie Candido-Regalado (Silliman University Institute of Environmental and Marine Sciences, SU-IEMS), Angela Fa’anunu (University of Hawaiʻi at Manoa, UHM; NSF – Igert Fellow and was partially funded by NSF- DGE – 054914 for this study; B.A. Wilcox, P.I.), Dean Apistar, Agnes Corine Sabonsolin, Danilo Delizo Jr., Roxie Diaz (Research and Monitoring team, REMOTE- of the Coastal Conservation and Education Foundation, CCE Foundation) Pablina Cadiz, Clarissa Reboton (SU-IEMS), R. Villarña, J.Villarña and V. Tampon and X. Suan, F. Candido and famiy, M. Aldeon and family (Apo Island community). Special thanks to the volunteer dive guides from Liberty’s Community Lodge and Paul’s Diving School (Dave, Sydney, Jed, Yhan, Dexel). The following institutions, non-government organizations, local government units, line agencies and individuals provided logistical support: CCE Foundation through R.L. Eisma-Osorio, J. Trangia, S. Tesch and R. Kirit, SU-IEMS through H.P. Calumpong and J. Luchavez, Silliman University –Angelo King Center for Research and Environmental Management (SUAKCREM) through R. Abesamis and J.L. Maypa, Negros Oriental DENR-PENRO through PASu V. Grefalde, Apo Island PAMB, Apo Island Barangay Council and community Hon. L Pascobello- Rhodes and Hon. M. Pascobello,, the Duin Municipality through R. Sanchez and Hon. R. Alanano, the local government units and MPA management bodies of Sumilon Island Fish Sanctuary, Oslob, Cebu; Balicasag Island Fish Sanctuary, Panglao Bohol; Saavedra Diveshop and A. Martinez. I also thank J. Claisse and R. Langston for their guidance in otolith preparation and ageing estimates, R. Humphreys for his insights and advice on Caranx otoliths and for his C. ignobilis samples and accompanying readings by D Lou of the Tropical Fish ageing at Townsville, Austraila, T. Carvalho and M. Dunlap of the Pacific Biosciences Research Center Biological Electron Microscope Facility at the 3 University of Hawai‘i at Manoa for the SEM training and use of facilities and the Friedlander lab (Zoology Department, University of Hawai‘i at Manoa) through A. Friedlander and M. Donovan for the use of their otlolith facilities. Special thanks to D. Ocampo for taking some of the C. sexfasciatus spawning videos, to A.D. Taylor for his statistics advice, to M. Djunaidy,V. Wong and N. Moravec for their support in Honolulu. Research funding was provided by Conservation, Food Health Foundation, PADI Foundation, Hawai‘i Cooperative Fisheries Research Center through C. Birkeland, GEF Coral Disease Component through LJR Raymundo, CCE Foundation - Saving Philippine Reefs/Earthwatch through E. White, The National Geographic Society through A.T. White. The Nature Conservancy Asia Pacific Region Marine Program and the Coral Triangle Support Partnership provided partial funds to review key coral reef and coastal pelagic fisheries movements relevant to effective MPA designs. The academic support of A.P. Maypa was provided by the Fulbright Philippine - Agriculture Scholarship Program, East -West Center (EWC) Degree Fellowship Program, Melga Torre and Buddy Gendrano Fellowship, EWC Association Alumni Scholars Award. I would finally thank my family and friends for their encouragement and support. 4 ABSTRACT Marine protected areas (MPAs) are tools used for conservation, and have repeatedly shown effectiveness in restoring and protecting fishery stocks inside the MPA. However, MPA benefits to biodiversity and fisheries beyond its boundaries are less clear. Achieving objectives of increasing both fisheries and biodiversity is controversial, particularly for small community-based no-take MPAs (nt-MPAs). This dissertation investigates mechanisms contributing to fisheries benefits from small nt-MPAs in three areas: (1) density-dependent spillover through indirect evidence from density and biomass gradients of fish species and groups (2) density independent – spillover as exhibited by Caranx sexfasciatus life history patterns and (3) the association of C. ignobilis with the nt-MPA as determined by distribution, catch rates and reproduction patterns. Three general gradient patterns resulted: (1) an abrupt decline close to the nt- MPA boundary, (2) an extended and gradual decline from the nt-MPA boundary to fished areas and (4) a decline from the fished area and falling within the nt-MPA. It is estimated that fishery benefits from density-dependent spillover of small no-take MPAs in the Philippines generally occurs at a small scale (10s - 100s m). Larger scale benefits appear to come from density-independent spillover as shown in the life history patterns of C. sexfasciatus. Four size classes were identified in different habitat types. Patterns suggest that emigration from the MPA to fished areas is density-independent. Factors driving the habitat shifts include habitat complexity, predator densities, current velocity and the accompanying diet and reproductive shifts. Protection of the existing MPA is limited to C. sexfasciatus intermediate sizes (15 – 37 cm, S ) and excludes the recruits, younger L juveniles and spawning sizes. For C. ignobilis, distribution, densities and fish catch rates patterns suggest a home range center located on the northern traditional fishing grounds beyond the existing Apo Island nt-MPA. The location, size, and mixed use of this MPA make C. ignobilis more vulnerable to exploitation and a less sustainable fishery within the MPA than originally intended. Results suggest that more effective designs of MPAs for fisheries management must take into consideration fish assemblage movements and life history patterns of key coral reef and commercially important fishery species. 5 TABLE OF CONTENTS ACKNOWLEDGEMENTS.................................................................................................3 ABSTRACT.........................................................................................................................5 LIST OF TABLES………………………...........................................................................7 LIST OF FIGURES ............................................................................................................8 CHAPTER 1. General Introduction ..................................................................................9 CHAPTER 2. Trends of density and biomass gradients across three small no-take marine protected area boundaries in the central Philippines Abstract..............................................................................................................................12 Introduction........................................................................................................................13 Methods..............................................................................................................................15 Results................................................................................................................................20 Discussion..........................................................................................................................36 CHAPTER 3. Habitat, life history patterns and ecology of the bigeye trevally, Caranx sexfasciatus at Apo Island, central Philippines Abstract..............................................................................................................................42 Introduction........................................................................................................................43 Methods..............................................................................................................................45 Results................................................................................................................................59 Discussion..........................................................................................................................84 CHAPTER 4. Distribution, growth, reproduction and catch rates of the giant trevally Caranx ignobilis from Apo Island, central Philippines: implication to MPA spillover Abstract..............................................................................................................................93 Introduction........................................................................................................................94 Methods..............................................................................................................................96 Results..............................................................................................................................103 Discussion........................................................................................................................119 CHAPTER 5. Summary of management applications and future research....................128 REFERENCES ................................................................................................................132 6 LIST OF TABLES 2.1 Description and general characteristics of 3 island nr-MPAs………...….…………..17 2.2 Mean density and biomas of target species/species groups…………………….…....23 2.3 Results of Two-Way Repeated Measures ANOVA.....................................................25 2.4 Results of significant linear regressions………….……………...…..….....…….......27 2.5 Details of the best-fit logistic decay models................................................................33 2.6 Results of significant mutiple linear regression tests...................................................35 3.1 Description and general characteristics of C. sexfasciatus habitats………..…...........60 3.2 Summary of the stomach contents analysis of Caranx sexfasciatus............................61 3.3 Results of ANOSIM ……………………………………………................................68 3.4 Mean current speeds at different Caranx sexfasciatus habitats……...........................70 3.5 Percentage of C. sexfasciatus at different maturing stages..........................................78 3.6 Widths and statistics tests results of C. sexfasciatus growth increments……...…......83 4.1 Fish densities, benthic characteristics…………………………………………........107 4.2 Apo Island annual catch composition and yield…………………………….….......108 4.3 Statistical test results fish yields, fishing effort and CPUE comparisons……..........112 4.4 Variation in C. ignobilis sex ratio according to maturity stages................................114 7 LIST OF FIGURES 2.1 Map of Apo, Balicasag and Sumilon Islands...............................................................16 2.2.1 Density and biomass gradients (abrupt decline close to MPA boundary) ...............30 2.2.2 Density and biomass gradients (extended and gradual decline to fished areas)…...31 2.2.3 Density gradients (from the fished area to the nt-MPA)….......................................32 3.1 Map of Apo Island and the location of study sites…...................................................47 3.2 Densities of the different size classes of Caranx sexfasciatus ....................................65 3.3 Gut content (% weight of prey) per C. sexfasciatus size class…................................67 3.4 C. sexfasciatus spawning pair......................................................................................72 3.5 C. sexfasciatus spawning timing……………………..................................................73 3.6 Length at first maturity (L ) for C. sexfasciatus.........................................................74 50 3.7 Monthly (mean ± SD) gonadosomatic index (I ) for C. sexfasciatus…….................75 G 3.8 Caranx sexfasciatus total fecundity.............................................................................78 3.9 Scanning electron micrograph of a C. sexfasciatus otolith section…………….........81 3.10 Growth and the von Bertalanffy growth curve derived for C. sexfasciatus...............82 4.1 Map of Apo Island showing locations and densities of Caranx ignobilis…...…........97 4.2 Monthly densities of C. ignobilis according to three size classes..............................104 4.3 Dendogram and MDS plots of C.ignobilis sites………………………………..…..105 4.4 C. ignobilis yield, fishing effort and catch per unit effort ………..………..............110 4.5 Size frequency of C, ignobilis....................................................................................113 4.6 Length at first maturity of C. ignobolis …………………….……….......……........115 4.7 C. Daily I for Caranx ignobilis ……………………………………………................117 G 4.8 Caranx ignobilis total fecundity ……………………………………………................118 4.9 Growth and the von Bertalanffy growth curve derived for C. ignobilis………........120 4.10 Scanning electron micrograph of C. ignobilis otolith….…………………….........121 5.1 Fish movement scales…….………………………………..………………….........131 8 CHAPTER 1 GENERAL INTRODUCTION Global concern for marine conservation and food security has resulted in efforts to explore marine protected areas (MPAs) effectiveness and designs that can meet multiple objectives (e.g., Gaines et al. 2010, Halpern et al. 2010, Salomon et al. 2011, Fernandes et al. 2012). Marine protected areas (MPAs; also known as marine reserves, marine refuges, marine preserves) are areas in the marine environment wherein forms of human physical disturbance and use are banned or managed (Russ 2002, Gell and Roberts 2003, Hughes et al. 2006). MPAs can be used for conservation, fisheries management (Gell and Roberts 2003). Recently, MPAs are advocated for coral reef resilience brought about by climate change (Mumby 2007, Green and Bellwood 2009, Stockwell et. al. 2009), and as an alternative livelihood tool for those MPAs that are able to generate enough revenues from tourism and related activities (e.g., Cadiz and Calumpong 2000, White et al. 2000, CCE Foundation 2006, Gravestock et al. 2008). Conservation benefits from MPAs such as biodiversity increase, abundance, biomass and reproductive potential as animals increase in size, are well documented within its boundaries (Gell and Roberts 2003, Halpern et al. 2003, Lester et.al 2009, Babcock et al. 2010). However, benefits of MPAs to biodiversity and fisheries beyond their boundaries are less clear (Russ 2002, Sale et. al. 2005, Gaines et al. 2010). To be functional as fisheries and management tools, MPAs are expected to enhance adjacent fishing areas through net export of adult and juvenile organisms (spillover) and/or net export of eggs and larvae (recruitment subsidy) (Bohnsack 1993, Russ 2002, Sale et al. 2005). The empirical evidence for spillover and studies on fish movement patterns have greatly increased over the past two decades (e.g. reviewed by Palumbi 2004, Gruss et al. 2011). Fish movements from tagging and tracking are able to quantify home range and daily movements (e.g Aprion virescens: Meyer et al. 2007, Cranx ignobilis: Wetherbee et al. 2004, Meyer et al. 2007; C. melampygus: Holland et al. 2006; Naso spp: Meyer and Holland 2005, Marshell et al. 2011; Plectropomus leopadus : Samoilys 1997, Zeller; Sparisoma cretense: Alfonso et al. 2008; sharks: 9 Meyer et al. 2009, Heupel 2010 ) or habitat utilizations long-term dispersal patterns (e.g., Holland and et al. 1996, Meyer et al. 2000) (reviewed by Maypa et al. 2012). Despite these advances, only a few studies document that the adjacent fisheries are actually benefiting from spillover (e.g., Attwood and Bennett 1994, McClanahan and Mangi 2000, Roberts et al. 2002, Russ et al. 2003, Kaunda-Arara and Rose 2004, Russ and Alcala 2004, Abesamis et al. 2006a, Abesamis et al. 2006b, Harmelin-Vivien 2008, La Mesa et al. 2011). Studies on fish larval dispersal patterns are rare, yet these recent few studies suggests that a significant amount of self-recruitment occurs in marine populations (Shultz and Cowen 1994, Jones et al. 2005, Almany et al 2007, Pineda et al. 2007, Cowen and Sponaugle 2009, Christie et al. 2010). Further, Jones et al. (2009) concluded that coral and fish connectivity may be independent from pelagic larval duration (PLD) but geographical setting may strongly influence it. These results will greatly influence MPA network configuration. In the Philippines, no take MPAs (nt-MPAs) have been established to provide fisheries benefits, yet are small (90% are < 1 km2) (PhilReefs 2008, Weeks et al. 2009), thus, their effectiveness is being questioned. While science recommends large MPAs, the economic and socio-political realities in the Coral Triangle countries limit the establishment of large MPAs (Maypa et al. 2012a). As MPAs are being put forward as primary fisheries management tools in the Philippines (i.e., the Fisheries Code of 1998, RA 8550, requires that at least 15% of the municipal waters be set aside as MPAs) there is an urgent need to assess their design and effectiveness. To date only a few studies investigated the efficacy of nt-MPAs in the Philippines for fishery benefits (e.g., Russ and Alcala 1998,, Russ et al. 2003, 2004, Abesamis and Russ 2005, Abesamis et al. 2006a, 2006b, Aurellado et al. 2009) considering its importance in the fisheries management framework of the country. This dissertation attempts to fill in the gaps in knowledge by investigating the mechanisms that contribute to the fisheries benefits from small nt-MPAs in three areas: (1) density-dependent spillover through indirect evidence from density and biomass gradients of fish species and species groups, (2) density independent – spillover as exhibited by the life history patterns of Caranx sexfasciatus, and (3) the nt-MPA use of C. ignobilis, the patterns of distribution, growth, reproduction and catch rates. 10
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