Source Ecology of the Coral Reef System

Home , Acanthurus japonicus, Aesopia cornuta, Blast fishing, Bolinao, Canthigaster compressa, Creel (basket)

source Ecology of the Coral Reef System

Rodolfo B. Reyes, Jr. Kathleen N. Kesner

Association of Southeast Asian NationNnited States Coastal Resources Management Project International Center for Living Aquatic Resources Management

Fisheries Stock Assessment-CollaborativeResearch Support Program University of Rhode Island University of the Philippines Marine Science Institute Resource Ecology of the Bolinao Coral Reef System

JOHN W. Mc~~NUS CLETOL. NA~JOLA,JR. RODOLFOB, REYES, JR. KATHLEENN. KESNER

Published by the International Center for Living Aquatic Resources Management on behalf of the Association of Southeast Asian Nationmnited States Coastal Resources Management Project.

Printed in Manila, Philippines.

McManus, J.W., C.L. Naiiola, Jr., R.B. Reyes, Jr. and K.N. Kesner. 1992. Resource ecology of the Bolinao coral reef system. ICLARM Stud. Rev. 22,117 p.

Cover: (Front) Reef flat including Silaki Island. The spottiness results from patches of coral, sand and seagrass of various densities. (Back) A gillnet against the backdrop of a Bolinao sunset.

All photos by J.W. McManus.

ISSN 0115-4389 ISBN 971-8709-28-2

ICLARM Contribution No. 844 MSI Contribution No. 212 CONTENTS

List of Acronyms and Abbreviations

List of Tables vii List of Figures

Acknowledgments

Dedication

Foreword xvi

Abstract

Chapter 1. Introduction

Chapter 2. The Harvest of the Reef General Monitoring the fishery Slope fisheries Reef flat fishery Ove~allstudy results

Chapter 3. Reef Slope Fish Communities General Monitoring the reef slope Data calculations Fish abundances Species diversity Effects of harvest on fish species diversity

Chapter 4. Reef Flat Fish Communities General Monitoring the reef flat Fish abundances Species diversities Chapter 5. Reducing the Rate of Exploitation General Reducing fishing effort Types of overfishing How much harvest effort should there be?

Chapter 6. A Proposed Marine ReserveIPark System General The need for a reserve The reservelpark system Implications for fishery patterns Suggested implementation

Chapter 7. Recommendations for Management Action Overview Regulating tourism Developing alternative livelihoods Mariculture and agriculture Establishing marine reserves Blast and cyanide fishing Banning compressor diving Improving fish-handling facilities Reducing human population growth rates

References

Appendices. Fish Species Abundances 1. Combined list of all species sorted alphabetically. 2. Reef slope species by frequency. 3. Reef slope families by frequency. 4. Reef flat species by frequency. 5. Reef flat families by frequency. 6. Reef flat night trawl species by frequency. 7. Reef flat night trawl families by frequency. 8. Reef flat night trawl species by weight. 9. Reef flat night trawl families by weight. LIST OF ACRONYMS AND ABBREVIATIONS

Association of Southeast Asian NationdUnited States Coastal Resources Management Project CPUE catch per unit effort DA Department of Agriculture DAP Development Academy of the Philippines DAR Department of Agrarian Reform DENR Department of Environment and Natural Resources EIA environmental impact assessment FSA-CRSP Fisheries Stock Assessment-Collaborative Research Support Program GPS global positioning system hp horsepower MEY maximum economic yield MSY maximum sustainable yield NGO nongovernmental organization SEC Securities and Exchange Commission TURF territorial use rights in fisheries USAID United States Agency for International Development LIST OF TABLES

1.1 Seasonality of selected reef resources. Dates are approximate. Harvestore must often shift between target resources seasonally. 1.2 Prices of major marine and freshwater commodities set for the Bolinao fish market by the municipal government. Actual prices vary with avaiIability. 2.1 Differential catches of major reef flat gear. Numbers represent the percentage of the 1989-1990 catch that each taxon contributed to each gear (+ is 4%).Species shown are those which ranked in the top five for one or more gear. The table has been extracted from one sorted by reciprocal averaging, so that the species are grouped by similar gear affinities and the gear, by similar catches. 7.1 Potential products from mangrove forests (Saenger et al. 1983; Salm and Clark 1984). 7.2 Skilled labor available in the Bolinao municipality. A variety of human resources could be tapped for small-scale industry. Data are from a survey by DAR in 1991. LIST OF FIGURES

Map showing the extent of the Bolinao municipality and reef system. Map of reef areas and adjacent landmarks in Bolinao. Chart of the subsea topography around Santiago Island. Some fields of study which have direct relevance to CRM. Others which could have been added include public health, nutrition and food science. Education and occupation factors affecting development in Bolinao. Data from surveys by DA in 1990 and DAR in 1991. Human population growth in Bolinao. Based on a log-linear regression of historical levels. Data are from the National Census and Statistics Ofice. Mrljor full- and part-time occupations in Bolinao. Fishing is the least profitable full-time activity, but shellcraft is the most profitable part-time job. Data are from a survey by DA in 1990. Cumulative percentage of boats found at each distance from the reef crest. Monthly boat distances from the reef crest. Relationship between the distance from the reef crest and the area of the reef slope. The nearly straight relationship is a result of the trapezoid-like shape of the reef slope. Price of regular gasoline, based on purchases at a local filling station. The sharp rise in price in December 1990 occurred because of government pricing at the time of the Gulf War. Harvest and fishing effort for the Santiago Island reef flat and lagoon. Effort figures exclude traps and corrals. Catch rates (CPUE)of major gear. The most seasonal catch rates are those from spearfishing on the reef slope. The corral graph omits large catches obtained twice each year during the spawning migrations of Siganus fuscescens (rabbitfish, barangen). Distribution of fish lengths caught by handliners on the reef slope, indicating a decline in the numbers of fish longer than 30 cm. Harvest data on sea cucumbers and shells from Lucero on Santiago Island. The alternation of seasons leads to shifting occupations among some hawestors. Data are from a survey by de Guzman 1990. Primary gear use areas on the Santiago Island reef. A large area of the reef flat is rented to Ash corral owners by the municipality. During the spawning migrations of Siganus fuscescens (barangen), other harvest activi- vii ties in this area are prevented by the corral owners. Fishing gear use on the reef dope is more uniform than on the reef flat, but varies in intensity with distance from the reef crest. Comparison of seasonality in gillnet harvests in a nonreef soft-bottom area and on the eastern reef flat. Top species by weight in the shore landings of Bolinao. The irregular patterns are the result of the interplay of factors affecting the fish populations and hawest activities. Fish drawings are by Magnus Olsson-Ringby. Harvests from a nonreef longline fishery adjacent to Santiago Island. Profile of the Santiago Island reef, showing the flat and slope separated by a wave-breaking intertidal crest. The profile is based on a transect running northwards to the west of Silaki Island (3x vertical exaggeration). Seasonality in bottom salinity and temperature in 18 reef slope sites. Ver- tical bars are 95% confidence limits. Sampling sites around Santiago Island, Bolinao. S, F = slope and flat visual transect sites, T = trawl sites, L = fish landing sites. Permanent markings on the reef slope transect sites. Abundance and diversity by transect on the reef slope. Vertical bars are 95% confidence limits (left: all life stages; right: by life stage where black = recruits, dashed = subadults and dotted = adults). Nundances of fish recruits (top) and adult fish (bottom) on the reef slope. The decline of adults was interrupted only by a temporary pulse in May 1991 which quickly disappeared. Abundance and diversity of adult fish on the reef slope. Abundances of the most common species in the reef slope visual censusing (grey = combined sizes, black = recruits, dotted = adults). Fish drawings are by Magnus Olsson-Ringby. Diversity in combined transects on the reef slope. Vertical bars represent 95% confidence limits determined from variances obtained by jackknifing among sites (left: all life stages; right: by life stage where black = recruits, dashed = subadults and dotted = adults). Illustration of predator-mediated coexistence. The predator favors the more abundant species, preventing it from excluding the weaker competitor. Some possible effects of fishing on a community. Species are represented * by bars arranged initially in rank order by abundance. People can act as predators andlor as removers of predators to cause a variety of possible changes. Species abundance profiles of adult reef slope fish from year-end months. Many species appear to have "fallen off' as overall abundances declined. Bottom salinity and temperature measurements from six reef flat sites. Vertical bars are 95% confidence intervals. Abundances of selected commercially important invertebrates in 50 quadrats on the reef flat near Lucero. Vertical bars are 95% confidence limits. The graphs are based on unpublished data of A de Guzman 1990. Seasonal variation in seagrass cover on the reef flat. The greatest effect is seen as an annual loss of dense areas, indicating that thinning rather than contraction of seagrass areas explains the annual drop in cover. Reef flat fish abundances and diversities by transect from six transect sites. Vertical bars are 95% confidence intervals (left: all life stages; right: by life stage where black = recruits, dashed = subadults and dotted = adults). viii Reef flat fish diversities for combined transects. Vertical bars are 95% confidence limits determined by jackknifing among sites (lefe: all life stages; right: by life stages where black = recruits, dashed = subadults and dotted = adults). Reef flat fish abundances and diversities by transect from 7 roller trawl sites. Vertical bars are 95% confidence limits computed conventionally (left: all life stages; right: by life stage where black = recruits, dashed = subadults and dotted = adults). Reef flat fish diversities for combined trawl sites. Vertical bars are 95% confidence limits determined by jackknifing among sites (left: all life stages; right: by life stage where black = recruits, dashed = subadults and dotted = adults). Abundances of top species from the reef flat visual transects conducted during the day (grey = combined sizes, black = recruits, dotted = adults). Fish drawings are by Magnus Olsson-Ringby and J. McManus. Pictures next to unknown species are generalized for the taxa. Abundances of top species from the reef flat trawl collections made at night in the seagrass of the reef flat (grey = combined sizes, black = recruits, dotted = adults). Fish drawings are by Magnus Olsson-Ringby and J. McManus. Pictures next to unknown species are generalized for the taxa. Fixed price model for profit and cost in an open-access fishery. People tend to enter the fishery until profits are reduced to near the cost of fishing. If alternative livelihoods are available, the potential profit creates an additional "opportunity cost" to fishing, and the equilibrium point is pushed back to more desirable levels. 5.2 Symmetrical production curve. Bars represent possible ranges for effort reductions; arrows represent reductions from the indicated equilibrium effort levels. An effort reduction of 60% would be appropriate for the low- cost fishery (B) and conservative for the high-cost fishery (A). 5.3 Right-skewed production curve. Cost lines are the same as those in Fig. 5.2. Note that both points A and B fall more than 2.5 times the effort at MSY. A reduction of 70% or more may be optimal in such extreme cases. 5.4 LeR-skewed production curve. An effort reduction of 60% is appropriate for the high-cost fishery (A), but is conservative for the low-cost fishery (B). A reduction by 60% in the absence of information on the nature of the curve could serve to help establish the type of curve, permitting more optimal effort levels to be set later. Plot of coral reef fish catch vs. fishing intensity in western Indian Ocean sites (redrawn from Gulland 1979). Plot of coral reef fish catch per area vs. fishing intensity for 8 parishes around Jamaica (redrawn from Munro and Thompson 1983). Plot of coral reef fish catch vs. fishing intensity for 11 American Samoa villages (redrawn from Munro and Williams 1985). Note that the shape of the cuwe cannot be determined from the available data. Fixed price model for trawl fishing in Manila Bay. The optimal effort reduction was approximately 60%. Points represent yearly values prog- ressing from lea to right (redrawn from Silvestre et al. 1987). Fixed price model for the overall Philippine demersal fisheries (t x lo6). The optimal effort reduction was approximately 60%. Points represent groups of annual catches (left to right) from 1946 to 1984 (redrawn from Pauly and Chua 1988). 5.10 Fixed price model for Philippine pelagic fisheries. The suggested effort reduction was 70%. Data points progress generally from left to right, representing the years 1948 to 1985 (redrawn from Dalzell et al. 1987). 6.1 Proposed marine reserve and park system. The area has been chosen to encompass most of the range of habitat types, and is expected to substantially enhance yields of fish and invertebrates throughout the reef. The park would generate income and enhance the tourist trade to provide alternative livelihoods for reef harvesters. 6.2 Marine reservdpark system relative ta primary reef flat fishing areas. The eastern corner of the reserve would bisect the major migration path for Siganus fuscescens (barangen), ensuring a supply of adult fish for the gillnetters and spearfishers, and still permitting substantial harvests for the fish corral owners. 6.3 Yield-per-recruit analysis for Siganus fuscescens (barangen). Curves show that the fine-meshed fish corrals induce growth overfishing far more than gillnets and speafishing as locally utilized. Yields could be improved by favoring spearfishing over fish corrals. This would also improve income distribution because of the labor-intensive nature of spearfishing. Population parameters were based on monthly length histograms per gear weighted by annual catch and combined. Procedures followed those in del Norte and Pauly (1990), but involved an improved data set from mid-1988 to mid- 1989 (L,= 26.0, K = 0.84, M = 1.71). Natural mortality was estimated from the Pauly environmental formula, using T = 30°C. Individual catch curves by gear were used to estimate the length of the first capture (LC)per gear (Gayanilo et al. 1988). 6.4 Map of fishpond areas in Bolinao. Most of the ponds have been built on former mangrove forest areas, thereby reducing the potential resources available for small-scale income generation. 7.1 Tourist attractions in the Bolinno area. Tourism could be greatly enhanced with the provision of a scuba compressor, establishment of a marine park, and elimination of blast and cyanide fishing. 7.2 Selected products from a well-managed mangrove forest. 7.3 Many compressor (hookah) divers dive too deep for too long and acquire decompression sickness and other maladies. Paralysis and death are common. ACKNOWLEDGMENTS

This book was sponsored by the USAID Most of the fishery data were gathered by Fisheries Stock Assessment Collaborative research aides Jesse Cabansag, Elmer Dumaran Research Support Program (FSA-CRSP) Grant and Ferdinand Castrence, Jr. Our boatman, No. DAN-4146-G;-SS-5071-00, Biodiversity Pro- Eleuterio Dumaran, has served as a helpful gram Grant No. DAN-4024-G-SS-9113-00 and source of information. the ASEANAJS Coastal Resources Management Two Master of Science students, Miriam

Project ,(CRMP). Our thanks go to reviewers Balgos and Asuncion de Guzman, did excellent Daniel Pauly, John Munro and Gamy Russ for jobs of gathering data on the squid fishery and very helpful suggestions, many of which were reef flat invertebrate communities, respectively. incorporated into the final text. We have made considerable use of unpublished We would like to thank past members of data gathered by Mrs. de Guzman. Hayacin the University of the Philippines (UP) Marine Jaranilla provided estimates of diversity indices Science Institute CRSP team: and jackknifed variances. Fish drawings from the ICLARM FISHBASE computer program were drawn by Magnus Olsson-Ringby. Naniel Aragones Much of the information in the CRMP Clarissa Axida reports of the UP College of Social Work and Jerome Cabansag Community Development, especially Gladys Annabelle del Norte Campos Hingco's insightful reports, were useful. Wilfredo Campos Indispensable information on the town Vincent Hilomen came from the following: Marl Mammaril Joseph Pasamonte Quintin Caasi Chris Diolaso Programmers Jacinto Elefante Ruben Garcia Florante de Guzman Hayacin Jaranilla Mayor Merito Miguel Dondi Roa Gabriel Pamintuan Cecile Yalung We are grateful to the following for helpful Clerical staff technical advice, information and support: Jerry Llamado Mi ye1Miguel Porfirio AliAo Janet Poot Spiros Constantinides Lourdes Cruz Sony Caracas Elmer Ferrer Nida Padilla Mi ye1Fortes Perlita Soriano Edga~doGomez Lilian Ungria Gil Jacinto Alfredo Licuanan Binabalian Cesar Luna Quilita Angan-angan Liana McManus Jun Balmores .Donald McCreight Marcos Caasi Lambert Mefiez Terry Caasi Nemesio Montafio Entong Caracas John Rowntree Pipit Caracas Saul Saila Etong Castellano Michael Sissenwine Alyo de la Cnxz Ian R. Smith (t) Julio Paleb Minda Talaue Grace Tolentino Goyoden Gavino Trono Gideon Caasi Lamarr Trott Sabelo Caasi A1 Tyler Eduardo Cabana Ramon Valdestamon Latero Casta Cesar Villanoy Emang Corpa

We would like to thank Chua Thia-Eng for We are grateful to the following people for making the publication of this book possible. helpful information: The following editorial staff of the ASEANAJS CRMP, ICLARM, are also acknowledged: Marie Sol M. Sadorra, Cecille Legazpi, Pamela P. del Bolinao Public Library Rosario, Rachel D. Africa and Regina G. Odela Celso, Librarian Morales for editing work; Rachel C. Josue and Eloisa E. Ben Belaid for some typing work; and Bolinao Rural Health Center Rachel C. Atanacio for preparing the layout. Dr. Susan Pamintuan, Rural Health Our thanks are extended to the following 0ficer fish buyers who kindly gave us access to their Juanita Iloca, Public Health Nurse daily records: Department of Agrarian Reform Angela Cuesta, Statistician Dewey Juanita Rabaya, M,micipal Agrarian Fidel Castro Reform Officer Nonoy Castro Neneth del Rosario Arsing del Rosario Department of Agriculture Paquita Beltran, Agricultural Picocobuan Technologist Patria Albino Imelda Cacho, Aquaculturist I Elly Collado Moises Cacho, Municipal Agricultural Offker Lucero Recto Capua, Agricultural Iliang Caacbay Technologist/Municipal Sonia Padilla Planning Officer Olog Ritorio Justiniano Junio, Agricultural Pilar Technologist Paking Caaya Carolina Ramirez, Agricultural Nilie Callado Technologist Priscilla Reccion, Agricultural National Census and Statistics Ofice Technologist Tomas Africa, Administrator Domingo Tobias, Municipal Panelco Agricultural Officer Alberta Guiang, General Manager Napoleon Tugade, Municipal Dionisio Opolento, Member Services Agricultural OfIicer Director

Municipal Government Philippine Coast Guard Evangeline Arayi, Municipal Local Cmdr. Amadeo Ocuaman, Commander, Government Clerk 7th Coast Guard District Alejandro Caalim, Jr., Municipal Secretary Rivergold Shellcraft Samuel Calado, Revenue Collection Adoracion Caalim, Proprietor Clerk Ilalo Camangian, Secretary to the Violeta's Shellcraft Municipal Secretary Violeta Caasi, Proprietor Edwin Conte, Municipal Local Government Officer Honorable Merito Miguel, Municipal Mayor

xiii

DEDICATION

This book is dedicated to the people of Bolinuo. May they have the wisdom to see what must be done, so that their children benefit from the knowledge we haue gathered. FOREWORD

The coastal waters of Southeast Asian tats and reduced fish stocks. Indiscriminate cut- countries have some of the world's richest ting of mangroves for aquaculture, fuel wood, ecosystems characterized by extensive coral timber and the like has brought temporary reefs and dense mangrove forests. Blessed with gains in fish production, fuel wood and time warm tropical climate and high rainfall, these supply but losses in nursery areas of commer- waters are further enriched with nutrients from cially important fish and shrimp, coastal erosion the land which enable them to support a wide and land accretion, diversity of marine life. Because economic bene- The coastal zones of most nations in fits could be derived from them, the coastal MEAN are subjected to increasing population zones in these countries teem with human and economic pressures manifested by a variety settlements. Over 70% of the population in the of coastal activities, notably, fishing, coastal region lives in coastal areas where resources aquaculture, waste disposal, salt-making, tin have been heavily exploited. This situation mining, oil drilling, tanker traffic, construction became apparent between the 1960s and 1970s and industrialization. This situation is aggra- when socioeconomic pressures increased. Large- vated by the expanding economic activities sc-ale destruction of the region's valuable attempting to uplift the standard of living of resources has caused serious degradation of the coastal people, the majority of whom live below environment, thus affecting the economic life of the official poverty line. the coastal inhabitants. This lamentable situation is mainly the result of ineffective or poor Some MEAN nations have formulated management of the coastal resources, regulatory measures for their coastal resources Coastal resources are valuable assets that management (CRM) such as the issuance of permits for fishing, logging, mangrove should be utilized on a sustainable basis. Uni- sectoral overuse of some resources has caused harvesting, etc. However, most of these measures have not proven effective due partly to grave problems. Indiscriminate logging and enforcement failure and largely to lack of sup- mining in upland areas might have brought port for the communities concerned. large economic benefits to companies undertaking these activities and, to a certain extent, Experiences in CRM in developed nations increased government revenues, but could prove suggest the need for an integrated, interdisci- detrimental to lowland activities such as fish- plinary and multisectoral approach in develop- eries, aquaculture and coastal tourism-depen- ing management plans that will provide a dent industries. Similarly, unregulated fishing course of action usable for the daily manage- effort and the use of destructive fishing ment of the coastal areas. methods, such as mechanized push-nets and The ASEANIUS CRMP arose from the dynamiting, have seriously destroyed fish habi- existing CRM problems. Its goal is to increase existing capabilities within MEAN nations for mation gathered during a five-year study of a developing and implementing CRM strategies. heavily exploited fringing reef along the western The project, which is funded by USAID and coast of Luzon. The authors have examined the executed by ICLARM in cooperation with ecology of the fish communities, the dynamics of MEAN institutions, attempts to attain its goals the fisheries, and a variety of social and through these activities: economic factors in order to develop a set of spe- analyzing, documenting and dissemi- cific management recommendations for imple- nating information on trends in coastal mentation by the local municipality. Beyond resources development; this, however, the study has yielded increasing awareness of the importance unprecedented insights into the nature of of CRM policies and identifying, and overfishing under conditions of rapid population where possible, strengthening existing growth and growing poverty, a situation known management capabilities; as Malthusian overfishing. providing technical solutions to coastal Ecologists will find helpful presentations on resource-use conflicts; and diversity and abundance patterns of coral reef promoting institutional arrangements fishes over time. Sections on yield-effort that bring multisectoral planning to relationships will be of interest to fisheries coastal resources development. scientists and managers. The final two chapters In addition to implementing training and are concerned with the design and imple- information dissemination programs, CRMP mentation of marine reserves and other man- also attempts to develop site-specific CRM plans agement measures appropriate to small-scale, to formulate integrated strategies that could be open-access coastal fisheries. The book will thus implemented in the prevailing conditions in be particularly useful for those engaged in CRM each nation. studies in tropical developing countries. The present work, Resource Ecology of the Bolinao Coral Reef System, summarizes infor-

Chua Thia-Eng Project Coordinator ASW/US CRMP and Director, Coastal Area Management Program, ICLARM

xvii This book describes an intensive four-year program of monitoring the cornunity ecology and harvest patterns of a lalge fringing coral reef system in northeastern Philippines. Reef harvest methods included principally gathering, handlining, trapping, gillnetting, seining, corralling and spearfishing, both with and without air compressors. Blast and cyanide fishing have substantially diminished hard coral cover, as have coral-grabbing anchors. PmducLion on the reef flat was approximately 10 tAcrn21year, while that on the reef slope was roughly 3 tfkm21year.Catch rates on the reef flat were relatively constant, while those on the reef slope varied seasonally. It is shown that 60% effort reduction is a reasonable inltial management goal in cases such as this where a fishery subject to Malthusian overfishing produces minimal net profits, and the quantitative nature of the yield-effort relatiomhip is unknown. A simple conceptual framework is provided for analyzing the effects ofharvest on diversity. Visual censusing revealed that the number of adult fish on the reef slope declined substantially during the study period, m did the number of species with indwiduals reaching maturity. Recruitment on the reef slope occurrcd in a strong annual pulse around May. Visual and trawl sampling of the reef flat failed to show strong seasonal pulses or mterannual declines. Abundances were substantially lower than those reported in some reef arcas subject to less harvest pressure. Some dominant species may migrate between seagrass beds and corals seasonally or daily. 'Xbtal multispecies fish recruitment appeared to be more predictahlc between years than that of any single species on both the reef slope and reef flat. Invertebrate populations, including commercially important sea urchins (Zbipneustes gratilla), and gastropods important to the shellcraft industry, alternated in abundance seasonally. Seagrass beds underwent a seasonal thinning in densc areas. Management recommendations include a design for a proposed marine reservelpark and a program for establishing alternative livelihoods to employ at least 60% of the harvest force, including ventures in tounsm and mariculture. This book is designed for managers, researchers and students with minimal technical training.

xviii CHAPTER 1 INTRODUCTION

Coral reefs provide food, income and other dae, barangen) out of the seagrass beds to breed, benefits to millions of people worldwide. Most of Linkages are also reflected in the exploitation the people who depend on reefs survive on mar- system, as a fisher may shift from one ecosystem ginal incomes, and have few alternative means of to the other to catch fish or gather invertebrates. survival in the event of a decline in the viability of The Bolinao reef system (Figs. 1.2 and 1.3) the reefs. Yet, coral reefs are very vulnerable to provides for 35% of the employment in a munici- problems of excessive siltation, pollution and a pality of 50,000 people. The proportion of employ- myriad of abuses related to the ways in which ment in fisheries and gathering is expected to rise their resources are exploited. Villagers living sharply as the human population increases in the alongside reefs tend to have high population immediate future because opportunities in farm- growth rates, and reefs in many areas of the world ing and industry are limited. Thus, the trends we are being subjected to increasing levels of stress see today, such as excessive overharvesting, de- related to overharvesting. Because reef access is clining stocks and deteriorating environments, rarely effectively limited, reefs tend to accumulate may well accelerate in the next few years. increasingly larger dependant human populations The current study was initiated by the Fish- as other means of livelihood become less accessi- eries Stock Assessment - Collaborative Re- ble. Human populations are growing at accelerat- search Support F'rogram (FSA-CRSP) in order ing rates, thus we can expect the status of reefs in to facilitate the development of new ways to many countries to decline at accelerating rates as manage complex fisheries. Fieldwork was nec- well. essary to generate data for the program because The coral reef system, which is the subject of of a worldwide sparsity of long-term data on this book (Fig. 1.11, is typical of true fringing reefs heavily fished coral reefs. The study evolved in the Central Indo-Pacific, i.e., those with a sub- gradually, as both the ecosystems and the ex- stantial structure typified by a separation into ploihtion systems were extremely complex. reef flat and reef slope areas by an intertidal reef Considerable investigation and preliminary crest. True fringing reefs tend to be large, covering sampling were necessary at every stage. The tens or hundreds of square kilometers. Like many methodology included such approaches as reefs in the Philippines and eastern Indonesia, the satellite image analysis, surveys from an ultra- Bolinao reef system includes substantial beds of light aircraft, broad area assessments by towed seagrass. The fisheries tend to target seagrass fish divers, underwater fish counts, seagrass as well as coral-dwelling fish. The interdependen- trawling, mapping of fishing gear use, under- cies of the two systems are reflected in the daily water blast counts, weighing and measuring migrations of fish such as cardinalfish (Apogoni- harvested fish, copying notebooks from fish dae, bagsang) into the seagrass beds for foraging, buyers, distributing questionnaires and specific and the annual migrations of rabbitfish (Sigani- investigations as questions arose. Some vital Fig. 1 .I. Map showing the extent of the Bolinan municipality and reef system. data came from students whose Master's work involving blast fishing was obtained through a was sponsored by the program. In spite of the grant from the USAID Biodiversity Program. diversity of monitoring approaches, there were Other information which were considered in- still important considerations which could not cluded the 1990 census of the National Census and be covered by our small team. Statistics mice, and previous surveys by the De- ort tun at el^, the program coincided with a partment ofAgriculture (DA) and the Department complementary assessment project, for which of Agrarian Reform (DAR). Bolinao was a key element. The ASEAN/US The system of human and ecosystem interac- CRMP-Philippine component was directed specifi- tion at Bolinao is extremely complex. We have cally toward obtaining the information necessary summarized only the mGor points. For example, for a general management plan, which was to the many harvested and other ecologically impor- include the Bolinao area. This project included a tant species are recruited at different times of the heavier emphasis on sociological and economic year. This leads to substantial variations in the aspects than was possible, given the financial effort directed toward each species in any given limitations of the FSA-CRSZ! Much of this infor- month (Table 1.1). mation has been summarized in a book, The The market involves a broad range of species coastal enaironmental profile of Lingayen Gulf, with variable prices (Table 1.2). Murdy (1981) Philippines (McManus and Chua 1990), which studied the fish sold in the Bolinao market during should serve as a companion volume to the current monthly trips of a few days each for one year, and work. Agreat deal of information from the CRMP identified 286 species in 73 families. He classified was assessed and evaluated in preparing the man- 209 of these as reef or reef-associated species. The agement recommendations which provide the fo- most speciose families, with numbers of species in cus for this book. Information on problems parentheses, were: Labridae (44), Serranidae (171,

Table 1 .l.Seasonality of selected mef resources. Dates are approximate. Harvehrs mu& often shiR between target resources seasonally.

Remurce Source Events Seaweeds Jan. Feb. Mar. Apr.May Jun. Jul. Aug. Sep. Oct. Nov. Dec. Caulerpa spp. (arosep) 1 harvest maxima Hydrmlathrus clathratw 2 biomass maxima Hya?rmlathms termis 2 biomass maxima Sargassum spp. 3 biomass maxima Invertebratee Corals 4 mass spawning Shells 1 harvest maxima Strodus l*~8 5 spawning maxima Sttvmbus mew 6 population maxima Stmmbus labiatus 6 population maxima Cypraea dus 6 population maxima Cypraea mneta 6 population maxima midac~derasa (giant clam) 7 egg pmduction Sepwteuthis lessoniana (squid) 8 egg laying Sea cucumbers 1 harvest maxima Bipneustes gratiUa (sea urchin) 6 population maxima Fiuhee Reef slope fish (as a community) 9 major recruihqent Migratory rabbitfish (bamrgerz) Sigzwua fuacesnns 10 rnigratiori Siganus spinus 10 migration Sigmurs argentem 10 migration

sourcw: 1. F- et al. (1989). 6. de Guzman (1990). 2. G.L. lblentho, pers. annm. 7. S.S.M. hlingoa. HA Roa and D.k Bonga, pars. mmm. 3. Tmno and Ldsma (1990). 8. Balgos (1990). 4. P.M. Aliho and M.P. Atrigeuio, pers. mmm. 9. This study. 6. Lieuanan et al. (1991). 10. Aragones (1987).

Acanthuridae (12), Scaridae (ll), Gobiidae (11), see that at least 350 species are marketed, of Carangidae (ll), LuGanidae (10) and Mullidae which at least 270 probably come from the reef. (10). Thirty-four families had one species each in These estimates are undoubtedly conservative be- the market, As is apparent in Table 1.1, inverte- cause of the variety of seasonal or sporadically brates, seaweeds and sea turtles are also impor- encountered species which would have been tant components of the market. This does not missed in previous sampling efforts. include the mollusks harvested for the shellcraft We cannot possibly account for every factor of industry, the fish landed in other municipalities or interest in managing the reef resource system. sent directly to Manila and the even wider range Our study has been broad enough that we have of organisms eaten at home. In a 1.5-year study of adopted the term "resource ecology" in favor of the the reef flat of Santiago Island using repetitive more traditional "fisheriesecology," which seemed quadrat sampling, de Guzman (1990)encountered wholly inadequate to describe the range of details more than 160 species of rnacroinvertebrates, of necessary to reach even simple, practical conclu- which at least 35 were exploited commercially, sions about the system. The current approach The total may be extended to include some rough could well be a companion to counterpart studies estimates of marketed lobsters (S?), crabs (5?), in resource economics, resource sociology and oth- shrimps and prawns (7?), cephalopods (5?), sea- ers. Amore ideal relationship between these fields weeds (6?)and sea turtles (2), some of which are is shown in Fig. 1.4. not listed on the official market price board or are The data which have been gathered, are those found in areas not sampled by de Guzman. We can which were believed to be minimally essential to Table 1.2. Prices of major marine and fnxhwater commodities set for the Bolinao fish market by the municipal government. Actual prices vary with availability.

English -em Local names Taxonomic group Pricw (p) (per kilo) 19S9 1991 Increase

Seaweeds A. Arorocep Caulerpa racemosa B. Culot Acanthophora spp., others C. Puk-puMo Codium edule StrawberryiMauritian canch Liswek Strombus luhuanus, S. decorus Spider conch Bariyawan Lumbis lambis Trapezium horse conch Nuga-nuga Fascidaria trapezium AU other kinds of edible shells Cuttle fish Kdanggotun Sepia latimatus Squid, white Laki Sepiotheutis lessionianu Squid, blown Bollpen Loligo spp. Octupw Kuritu Octopodidae Shrimps Orang, pasayan Metupenaeus spp., othem A. Large B. Medium C. Small Pmwn Sugpo, padaw Penaeus spp. Rock lobster A. Green, spotted white Orang kumpasan Panulirus ormtus B. Plain green Ot-ang kumpasan Panulirus uersicolor C. Red Orang kumpasan Panulirus longipes Crabs Ayama Scylla serrata Blue crabs Barisaway Portunus pelagicus Shark Pating, iyo Carcharhinus spp. Ray fish Pagui Dasyatis ~pp. Hawaiian ten-pounder Bayedbed Elops huwaiiensis Milkfish Bangus Chamchs A. Iarge B. Small Indian sardines Surdinella spp. A. Large B. Smd Shorbfinned gizard Nemutolosa japonicu A. Large B. Small Eel Igat Gymnothorax spp., others Sea eatfish It0 Plotosus spp. A. Large B. Smd Flying fi& Rayne Cypselurus spp. Halfbeak Bahot Hemimmphus spp. Gar ftsh Layday 2)losurus spp.. Strongylura spp. Gar fish Moulo !&losums spp., Strongylura spp. Ember fish Baya- baya Myripristis spp., Sargocentron spp. Grouper (lap-lapu) nth Epinepheluo spp. A. Large B. Small Red grouper (lapu-lapu) lbtokro Cephdophdis spp., VaridQ spp. Glass fish Damas, bagsangtaaw Apogon spp., Pempheris spp., &hew Largs cab alla !hlakitok .Carangidae, others

Continued Table 1.2 (Continued)

English names Local names Taxonomic group Price@ (B) (per kilo) 1989 1991 Increame

Scad Decapterus spp. A. Large B. Small Dolphin fish Durado Coryphna hippurus A. Whole B. Slice C. Head Slip mouth Leiognathus spp. A. Large B. Small Red snapper Mangngayat Luthnus argentimaculatus Large mouth snapper Mam-bituen Lutjanus riuulatus Snapper Rogso Lu&nus spp., Lethrinus spp. Spotted pomadasid Agu-ot Plectorhynchus spp. Fusilier Dnlagang bukid Caesio spp. Bream Besugo Nemipterus spp., Aphareus spp. Theadhn bmams Mcznarrat Nemipterus spp. Mojarras Batuan Germ abbreviatus Goat fish Gumian Parupemus spp. Rudder fish Ilek Kyphosus uaigiensis Mullet Burusi Liza spp. A. Large B. Small Barracuda Sphyraena barracuda A. Large B. Small Cichlid A. Large B. Smd Cigar wrasse Sangitan lawin Cheilio inermis Parrotfish Mulrnol tarektek Leptascanrs uaigiensis A. Large B. Small Parrotfish scarua spp. Black siganid Siganus guttatus, S. vermiculatus Yellow siganid B-en baka S. virgatus, S. punctutus Rabbitfish (sammaral) Bamngen dumnddan sig&us fuscescens A. Large 30.00 B. Small 20.00 Cutlass fish Pinka Dichiunrs lepturrrs 20.00 Yellow and black stripe Bdiwakwak Acanthurus spp., Ciknochaetus spp. 16.00 Surgeon fish Sungayan Naso literatus 15.00 Billfish Susay Istiophorus platypterus 20.00 lbna Bondying, orelerr Thunnus spp. 20.00 Yellow fin tuna Oreles Thunnus spp. 25.00 Spanish mackerel %nggui-gui Scomberomorus comrnerson 35.00 Kingfish Khaki Serida spp. 30.00 Spine fish Tbrtongan Diodon spp. 10.00 Sea turtle Pawikan Eretmmhelys imbrieata 20.00 Chelonia mydm Average increase: Average % increase: 7 ' The &or recommendations of the project are discussed in the last two chapters. They are summarized here: Establishment of a committee to plan and regulate the development of tourism to ensure that it is directed toward providing employment to fishers and maintaining local natural resources. Development of alternative livelihoods for at least 60% of the existing fishers and gatherers, and all future residents who would otherwise become occupied in harvesting marine resources. Development of nondestructive mariculture activities to provide food, income and livelihood, to alleviate some of the harvest pressures on the natural ecosystem, and to provide a strong incentive for the maintenance of a healthy marine environment. A complementary program of sustainable multicrop agriculture (permiculture) would provide for the optimal use of agri-

...... cultural lands to further reduce the har- Resource management vest pressures on marine resources. Establishment of reserve areas to provide undisturbed breeding grounds for reef Fig. 1.4. Some fields of study which have direct relevance to species and to augment stocks of fish and CRM. Others which could have been added include public invertebrates in surround,ing areas health, nutrition and food ecienoe. through larval dispersal and the emigra- tion of adults. Implementation of a program of public education and enforcement to completely understand ecological fundamentals important in eradicate blast and cyanide fishing from the management of the system. The data set is the area because of their destructive ef- relatively large, encompassing more than 7,000 fects on the organisms, their environ- pages. Future analyses of the data will undoubt- ments and the potential growth of diving edly turn up a smaller set of indicator variables tourism. which can be used by future researchers in moni- Banning of compressor diving (hookah) to toring other reefs. protect existing deepwater breeding popu- The current work will continue for as long as lations from overexploitation and to re- funds are available to support the monitoring. move the myriad of occupational hazards Only through long-term monitoring can we expect associated with this practice. to truly understand the dynamics of a system Improvement of fish-handling facilities so which is driven by annual pulses of juvenile re- as to reduce postharvest losses to spoilage, cruitment. Complex statistical analyses have been minimize health hazards from unsanitary avoided, so that the book will be useful to both conditions, increase local incomes by pro- researcher and resource manager alike. Some ma- moting more local processing, and in- terials of theoretical interest have been isolated in crease market value upon export by boxes within the chapters. Supplemental informa- meeting higher quality control standards. tion can be found in the more technical publica- Establishment of programs to reduce local tions stemming from the program (e.g., McManus human population growth rates so that as et al. 1988; del Norte et al. 1989; McManus 1989; total resource levels rise, so will the re- del Norte and Pauly 1990; Naiiola et al. 1990). turns of the individual harvesters. These recommendations could be critical steps with similar problems. Finally, we hope that the in avoiding a very distressing future scenario for methods and approaches we have used are evalu- the Bolinao municipality. However, it is hoped that ated appropriately and serve to guide those who they will also serve as a starting point for the intend to undertake related studies in the future. design of assessments on other coral reef systems

Recommended management actions:

1. Establish a tourism regulatory committee.

2. Develop alternative livelihoods.

3. Promote mariculture and improved agriculture.

4. Establish marine reserves.

5. Eradicate blast and cyanide fishing.

6. Ban compressor (hookah) diving.

7. Improve fish handling facilities.

8. Reduce the population growth rate. CHAPTER 2 THE HARVEST OF THE REEF

General shorelines where they are vulnerable to flooding and severe damage from storm winds. Sanitation is poor, and the implementation of proper sanitary Fishery-related occupations currently account facilities and training is difficult, given the crowd- for 31% of the employment in Bolinao (Fig. 2.1). ing and low-income levels. In many areas, includ- However, the population is rising rapidly (Fig. ing Silaki Island and parts of Santiago Island, 2.2). Educational achievement is low, with only 7% freshwater must be carried over in smalf boats of the population receiving training beyond high from the mainland. Most fishing families have no school, and 35% receiving no schooling at all (Fig. electricity. Remarkably, a few families in each 2.1). The farmlands, which currently support 49% village have television sets, often run on car bat- of the labor force are already virtually fully occu- teries which are periodically recharged in the pied. These fads make it very likely that most of main town. Lights are usually kerosene lamps, the incoming work force in the next few decades and cooking fires depend on the locally diminish- will attempt to enter the fishery. Thus, the propor- ing supply of small trees. tion of fishery-related occupations in Bolinao will probably rise sharply. This will accelerate the decline of the natural resource base, and may leave Monitoring the fishery tens of thousands of people living in deepening levels of poverty, Specific actions which can be Following an extensive program of prelimi- taken to avoid this situation are described in the nary investigation, a set of ten fish landing sites final two chapters of this book. were chosen and monitored from July 1988 to Fishing already provides the lowest average June 1991.The daily logbooks of major fish buyers monthly income of any major occupation locally were copied weekly. These books classified fish (Fig. 2.3). The mean monthly income of 81,830 is landed by weight into six broad categories of fish substantially below the estimated poverty level type. Supplemental data were obtained by sub- set by the Philippine government of B2,650/year. sampling each of five gear types at least three Families of fishers and gatherers generally live in times each month for catch composition by weight small, one-room nipa huts with floor areas of less and abundance at the species level. Inquiries were than 30 m2 and an average family size of 5 to 6 made routinely concerning the number of boats persons (McManus and Chua 1990). Because and fishers per gear and the number of hours and many of the fishing families are not native to days spentfishing. Much of this data was gathered Bolinao, having migrated from northern or central by research aides who were local fishers them- Philippines, very few own the land they live on. selves, and were therefore trusted by the local Houses are often densely packed against the villagers and buyers. Highest educational attainment

35% No schooling

Primary 32%

1% Postgraduate 1% Tochnlcal

26% Secondary

Principal occupation 49% Farming

2% Pensioners and others Fishery-related 31 3% Commerce 4% Trade and industry

Fig. 2.1. Education and occupation factors affecting development in Bolinao. Data from surveys by DA in 1990 and DAR in 1991.

Fig. 2.2. Human population growth in Bolinao based on a log-linear mgression of historical levels. Data am from the National Census and Statistics Office. Pensloner

Commerce

Farmlng

Trade and induetry

services

Flshlng

Shellcratt

Rope rnaldng

Charcoal rnaklng Part-time Salt rnaklng

Burl weavlng

Vegetable gardenlng

Fig. 2.3. Major full- and part-time occupations in Bolinao. Fishing is the least profitable full-time activity, but shellcraft is the mast profitable part-time job. Data are from a survey by DA in 1990.

Boats were mapped on the reef slope on one relative constancy between years has been random day each week. The mapping was done checked on a gear-by-gear basis with existing fiom a research boat using compass data, and appears to be a valid assumption. triangulation. Generally, the research boat lined up each fishing craft with a landmark and obtained a compass bearing to the landmark. Slope fisheries Then the research 'boat moved to a new location, lined up the same fishing craft with *other landmark, and obtained a second MAJOR TYPES OF GEAR bearing. These bearings were back-plotted in the laboratory to obtain precise positions for each craft. In the process, each craft was Hook and line identified as to the type of gear it supported. The estimations of catch rate (catch per People fishing on the reef slope must contend unit effort [CPUEI), effort and total yield were with the wave action of unprotected waters. Some based primarily on the records of the fish hook and line fishers use moderately sized (often buyers. In almost all cases, We buyers either 7 m) double outrigger boats (bangka)with small switched products or became inactive for some inboard engines (often 16 hp), usually requiring portion of the study period. At these times, fish low-octane gasoline. The majority of the boats are were marketed at unpredictable times and smaller and are paddled by hand or use sails. The places, often by the wives of the fishers, making fishing lines are held by hand without poles. The yield estimations difficult. The records for each gear consists of weighted nylon fishing lines of gear include some missing data, usually in various diameters, with one to three small, single groups of months. In order to preserve the point hooks usually baited with small shrimps or effects of seasonality, some records had to be pieces of squid. The bait is maintained near the filled in from one year to match months in bottom. The anchors from these and other boats another year. This could reduce apparent are constructed from iron-reinforcing rods and are interannual variability somewhat. However, the designed to catch corals, They cause substantial damage to the corals and thus reduce the long- a long tube to the diver, who uses the air without term viability of the fish resources. Some research a regulator. The divers frequently stay at depths should be initiated to find an alternative low-cost below 30 m for hours at a time, and are frequently anchoring system. crippled or killed by decompression sickness and Squid fishing involves trolling with hand lines. other diver-related maladies (see Chapter 7). pulling surface jigs resembling shrimp. This activity is highly seasonal (Balgos 1990). Octopus fishing (palaoy) involves using a small lure of rags Blast fishing shaped like an octopus, which is dragged along the bottom. In these and some other fisheries, series A broad variety of blasting devices are used of bamboo rafts are often towed over the reef slope locally to kill fish, ranging from handmade bombs in good weather by motorized bangka to provide to dynamite. However, the most common device is access for a wider range of fishers. The cephalo- a bottle filled with layers of sodium nitrate alter- pod-specific fishery catches have been omitted ing with layers of pebbles. The cord-type fuses are from the handline fishery calculations. usually commercially obtained. Sodium nitrate is sold legally to induce ripening in mangoes, and so is difficult to control. Each blast appears to kill Drive-in nets corals within a 2-3 m diameter. Fish kill distances are many times greater than this, especially for The principal form of drive-in net is theparis- fish with swim bladders. The blasts kill all sizes ris. This gear consists of a horizontal scare line of of fish, including juveniles. The fishing is very several hundred meters pulled by pairs of bangka wasteful because many dead fish living in or fall- in U-shape along the surface toward an area in ing down among the corals are difficult to see and which a floating net is subsequently laid. The net gather. More importantly, however, blasting re- forms a curved wall of a few meters depth and a duces coral cover and therefore has long-term few tens of meters length. The primary target fish effects on fish production. are needlefish (Belonidae, layalay) which frequent A common complaint is that the blast fishers the surface waters over the reef. come from municipalities outside of Bolinao. How- ever, our studies reveal that a major part of the blasting is by local fishers. Atisher can currently Spearfishing have returns of ten times or more on the investment in the blasting device, and substantially The local spearfishing gun is carved from better catches per hour than with traditional gear. wood, and is powered by large rubber strips re- However, the gain comes at a substantial loss to leased with a trigger. The spear is often a metal other fishers, particularly those of the next gen- rod sharpened at one end. The spearfishers use eration. It can take several decades for corals to small round goggles made of window glass, resettle and grow to the stat,es they were in before wooden frames for each eye and rubber strips. the blasting. These goggles can cause considerable eye damage Blasting rates were high at the start of the when used below a few meters depth because they study, such that our divers generally heard an cannot be equalized through the nose to compen- average of ten blasts per hour. Beginning in mid- sate for rising and falling external pressures. The 1989,blasting dropped by at least go%, apparently divers often use a single rigid wooden shield-like because of some extremely strict enforcement pro- paddle attached to one foot to assist them in swim- cedures. However, even the later rate of one blast ming. Divers traditionally use rocks to assist them per hour in a 2 to 3 km listening radius is too high in sinking to great depths (30-60 m) rapidly, often for ecological sustainability and the development resulting in considerable ear damage. of an active tourist trade. Most spearfishing on the reef slope involves The catch rates from blast fishing are difficult the use of air compressors, such as those used in to estimate, and so are omitted in our yield esti- vulcanizing shops and gasoline stations. The un- mations. However, they probably do not exceed filtered air passes a small reserve chamber which 15% of the total catch. The loss of corals undoubt- provides a final breath of air when motor trouble edly leads to the loss of fish yield, but this would stops the compressor. The air then passes through not be reflected in short-term estimations. Fish poisoning the values ranging as high as 26 t/km21year reported for some coralline areas in the Central A variety of fish poisons are used in Bolinao, Philippines (Alcala 1981), and the working value ranging from liquid detergents to natural plant of 15 t/km2/year summarized from a variety of derivatives. However, sodium cyanide is the over- studies on reefs worldwide (Munro and Williams whelmingly dominant poison. It is used on the reef 1985). However, it is within the general range of flat both for food and aquarium fish collecting, but 0.5-26 t/km2Iyear reported in the same summary. on the reef slope it is used more for the latter. It is The present value could be low because: applied by a skin or compressor diver to fish hiding 1. the reef does not support as much fish in corals by squirting as an emulsion from a plastic production as the average reef in previous bottle, or waving the tablet tied to the end of a studies because offactors such as low coral stick near the fish. The fish are stunned by the cover; poison and captured by hand. However, the fish 2. the reef has been fished for so long that tend to have a high mortality rate after shipping. gradual declines in production have oc- Thus, the practice does considerable harm to the curred; and international market for Philippine aquarium fish 3. the fishing effort is less than that in the (Albaladejo and Corpuz 1981; Rubec 1986; Hingco earlier studies. and Rivera 1991). It is also harmful to corals and It is unlikely that increasing fishing effort will other fish in the vicinity. As a gear which is harm- yield more fish in the long term. In fact, adult fish ful to the environment of the fish, sodium cyanide appear to be declining and may not be able to fishing should be prevented through management measures. As with blast fishing, annual yield rates are omitted in total yield estimations. However, they are probably insignificant in the overall mass of fish harvested. The important aspect of the gear that is used is its effect on the corals, and the threat it poses to future yields from the reef.

REEF SLOPE STUDY RESULTS

Fishing on the reef slope was generally uniform, with no particular gear dominating the fish- Distance from reef erest (km) ing effort in any given area. Fishing effort was concentrated near the reef crest, with an exponen- Fig. 2.4. Cumulative percentage of boats found at each tial decline proceeding outward (Figs. 2.4 and 2.5). distance frum thc reef crest. Throughout the study, 95% of the fishing tended to be within 2.7 km of the shore (Fig. 2.41, indicating that the mqjority of fishing was confined to approximately 42 km2 (Fig. 2.6). This limit is related to the cost of gasoline (Fig. 2.7)as well as considerations involving the spoilage of fish and safety from sudden inclement weather events. The monthly production mean of approximately 10 t translates to an annual production of 120 t, About 96% (114 t) of this mmes from 42 km2, for a yield of approximately 2.7 h2/year.We can check this figure by assuming that 50% of the catch comes from within 1km of the shore (Fig. 2.4), or 22 km2. Sixty tlyear would then come from 22 krn2, or 2.7 t/km21year as before. This contrasts sharply with Fig. 2.6. Monthly boat distances from the reef crest. than that on the reef slope, and for considerably less investment in gasoline and air compressors. The seasonality of the catch rates and the constancy of the effort lead to a seasonality in total catch over the year of a factor of two. Thus, there are times when the reef harvest translates to an annual equivalent of at least 4 #km21year. This may be a further indication that with less fishing, the catch rates could be improved by maintaining the interseasonal populations which are currently being fished to low levels.

Matance from reef crest (km) The adult fish populations have declined during the study period (Chapter 3), but the time Fig. 2.6. Relationship between the distance from thc rccf series on fish landings is not long enough to deter- crest and the area of the mef dope. The nearly straight mine for certain if the yield from-thereef slope has rclationship is a result of the trapezoid-like shape of the reef been declining as well (Fig. 2.8). The fishers have slope. not increased their range of operation to compensate for the sparsity of adult fish (Fig. 2.5) probably because of such factors as the effort needed to paddle the boats of the handliners, the effect of increasing gas prices on the motorized minority, and the increased risks involved in being caught far from shelter during a sudden storm. Instead, it appears that those few boats which once ranged more widely than the others have curtailed their long distance forays. A study of fish sizes caught by handlining (Fig. 2.10) indicates a possible decline in the number of large fish (30 cm) being caught. The long-term decline in fish sizes locally has been common knowledge to the elders in Boli- nao. Many people familiar with coral reef fish Fig. 2.7. Pricc of regular gasoline, based on purchases at a local have commented on the surprisingly small size of filling station. The sharp rise in price in Uecembcr 1990 oc the average fish in the markets (generally less curred bccauw of government pricing at the time or the Gulf than 20 cm). Similar comments are consistently War. made by experienced coral reef divers visiting the area, who are frequently shocked to see how scarce maintain former levels of juvenile recruitment to the fish are underwater, and how small the re- the slope (see Chapter 3). We must conclude that maining few appear to be. the low coral cover of the slope, and possibly the long-term effects of high fishing pressure (locally or regionally) combine to give unusually low fish- Reef flat fishery ing yields on this reef. The low coral cover could easily be related to the long history of intensive blast fishing in the area. MAJOR TYPES OF GEAR The total effort on the reef slope remained fairly constant during the study (Fig. 2.8). How- Hook and line ever, the catch rate varied radically between seasons. This indicates its dependence on the annual The handlines used on the reef flat are simil- recruitment pulse of fish in April and May. The ar to those described for the reef slope. However, uncertainty about catches may also help to limit the boats on the reef flat do not have to contend their entry into the slope fishery. It can be seen in with waves because of the protective intertidal Fig. 2.9 that there are seasons when the catch rate reef crest. The bangka here tend to be only a few from spearfishing inside the reef flat is higher meters long, and powered by paddle andlor sail. 1 Reef slope (based on boat counts)

Fig. 2.8. Hamest and fiahing effort for the Santiago Island reef flat and lagoon. Effort figures exclude trape and corrals.

Drive-in nets Blasting and poisoning

A reef flat counterpart to the parisris gear Blast and cyanide fishing are used widely on involves several fishers on raRs slapping the wa- the reef flat and do not differ substantially from ter and converging on a net. The target fish are what has been described for the reef slope. An hemiramphids. The catch is small relative to that exception to this is the fact that sodium cyanide is of other types of gear and will not be considered sometimes dispersed from a barrel on a boat in a further. radius of at least 10 m to capture fish for consumption. The poison is in the form of a slurry or mixed with fish and shrimp bits as "chum"on which the target fish feed. This undoubtedly poses a consid- Spearfishing erable health risk locally because the poison is very toxic to people. Another health risk involves The spear gun and its accessories are similar the practice of biting the tablet of sodium cyanide to those described for the reef slope. Additionally, to facilitate mixing it in plastic bottles for use in some fishers use metal rods with rubber strips the gathering of aquarium fish. More than 60% of attached instead of spear guns. Air compressors the lagoonal corals have been killed by blasting are unnecessary in the shallow waters of the la- and poisoning, greatly reducing the availability of goon and reef flat. Many fishers use kerosene coral reef fish to the fishery. lights mounted on boats or floats to help them spear at night in the seagrass. This is particularly Fish traps effective for the rabbitfish, Siganus fuscescens (barangen), which tends to turn sideways to the The local fish traps are approximately 30 cm light, presenting itself as an easy target. in length, and consist of a wicker box with an Spearfishing ... .. '3 F -. Reef slope :, :. ..: I (compressor) 1

7 Corrals 6 5 4 3 2

Gillnetting 1:: j*a A *, , , , , , , , , , <, , *, , , . , , , , , , <, , 0 0.5 F? - 0.04 Fish traps

0S " 0.3 1 0 9 : 1989 1990 1991 S 0.2 Handlining 0.1

Fig. 2.9. Catch rat- (CPUE)ofrnejorgear.The most wasonalcatches are those from spcarfishingon thc reef slope.Themrralgraph omits large catches obtained twice each year during the spawning migrations of Siganus fuscescens (rabbitfish, bamngen).

Fig. 2.10. Distribution of fish lengths caught by handliners on the reef slope, indicating a decline in the numbers of fish Ionfir than 30 cm. 17 entrance cone. These are used without bait in Fish corrals coralline and sandy areas. Fish enter out of curi- osity or to seek shelter. Their attempts to escape The fish corrals (baklad) of Bolinao are ar- attract other fish. The traps are generally left row-shaped fence structures whose angled sides, overnight and retrieved the following day. The and sometimes the stems, of the arrows extend for several hundred meters. The baklad depend on traps are very small compared to those of 2 m or mobile and migrating fish, and are often placed more found on some other Philippine reefs. The along migration pathways in the seagrass beds. traps in Bolinao have become substantially The favored sites are those which intercept the smaller in the last 12 years (J.McManus, personal Siganus observations). Because they are made of natural migrating adult rabbitfish, fuscescens (barangen), as they leave the reef flat to breed materials, they tend to be torn open by predatory fish if abandoned, and so do not pose serious twice each year. The Bolinao municipality leases the area on which the baklad are constructed. An threats to the fish community. Fish traps are gen- investor pays for the lease, and further leases out erally size selective and are otherwise favorable the rights to establish the baklad. The baklad at from a management standpoint, except when fish- times have virtually closed off large sections of the ers break corals to cover them. However, rocks are reef flat to eastward rabbitfish migrations along used more commonly. Fish traps are not commonly the reef flat north of Dewey. The baklad are usu- used on the reef slope. The small size and re- ally established by individuals or small consortia stricted use of the traps contrast markedly with with investment capital, and they compete for the situation in the Caribbean, where traps are seagrass fish with smaller-scale users of spears generally larger (122-229 cm) and dominate many and gillnets. coral reef fisheries (Munro and Thompson 1983). The difference in usage and the wide divergence in target species between this and other gear ~~k~dseining (Thble 2.1) call into question the utility of using standard traps to assess coral reef fishery poten- The rabbitfish return as juveniles to the reef tials, as is often proposed. flat twice each year, and are caught for use as fish

Tahle 2.1. Catches of major reef flat gear. Numbers represen1 the percentage of thc 1989-1YW catch that each taxon contributed to cach gcar (+ is c 1%).Species shown are those which ranked in the top five for one or more gear. The table has been extracted Crom one sorted by reciprocal averaging, so that gear arc grouped by similar catches, and species by similar tendencies to be caught hy each gcar.

Family Species Local name Gear: Traps Corrals Gillnet Spear Village: Goyoden Goyoden All Binabalian

Apogonidue Apogon sp. Ilngsarig Plotosi dae Plotos~slineatus It0 I~thrinidae Lethrinus hrak Rogso Scaridae Scarus ghnbban Molmol Scaridae Scarus rhorluropterus Molmol Scaridae Leptoscarus oaigiensis Molmol tarektek Labridae Choerdon anchorago Molmd rnangipcn Scaridae Calolornusjaponicus Molmol Siganidae Siganus fiscescens Barangen Loligonidae Sepioteuthis lessoniana Pi~sil Portunidae Portunus pelagicus Barisauray Plotosidae Plotosus canius It0 Octopodidae Octopus? spp. Coritn Genidae Gerres oyena L~rniolanong paste (bcycmng). One gear designed to capture women and children. Tools occasionally include these juveniles is the karokod seine. This is essen- push rakes to remove gastropods from the sea- tially a large plankton seine with a bag end pulled grass and bamboo raRs used in deeper waters, between two sailing bamboo rafts. This gear is especially for sea urchin gathering. The believed to be deleterious to the successful recruit- gathering of commercially valuable Tripneustes ment of the rabbitfish, and so has been banned gratilla sea urchins for roe was so intense that with increasing effectiveness during the final two by the end of the study period, some gatherers years of the current study period. had started using air compressors to provide access to a few deepwater seagrass beds. The principal gathered species are invertebrates, and their production is omitted in the Gillnetting yield estimations which follow. However, some information is available for the village of Lucero Local gillnets (tabar) usually have stretched on Santiago Island (de Guzman 1990), which mesh sizes that range from 4.5 to 5.4 cm. The nets indicates a strong seasonality in the harvests of are found in a variety of sizes and shapes. The sea cucumbers and shells (Fig. 2.11). usual net is weighted to rest on the bottom, and is approximately 100 m or more in length. The height is usually only approximately 1m. Gillnet- ting is a major fishery on the reef flat, but very little occurs on the reef slope. Gillnets are among the most desirable fishing gear from a management standpoint because each ' mesh size generally catches only one particu'lar \ Shells size of each fish species. In many cases, it is possible to regulate the mesh size to target a \ primary species (e.g., rabbitfish) at a size reached sometime after the age of first reproduction. This gives each fish an opportunity to contribute to the next generation of fish before being harvested. More precise analyses are possible to allow the harvest to be truly optimized through the control of mesh size. The mesh sizes in Bolinao reflect the small sizes of fish which remain on the reef flat under intensive fishing pressure.

Sea cucumbers '\ Gathering I \--

Gathering invertebrates and seaweeds by Sma cucumbors hand is probably the most important "fishing" method on the reef flat. Gathered products can match or exceed the total production of reef fish in some places (Savina and White 1986; McManus 1989a). The harvesting usually takes place at low )t Shells &. tide. Principal products include sea urchins, sea / \ cucumbers, octopus, some small species of fish, / \-@- a--a-.s Caulerpa seaweed and shells of many kinds. The Apr Jul Oct Jan Apr shells form the basis of the local shellcraft indus- 1989 1888 1988 I989 IQSa try, which ranks as the most successful of the local (Fig. Kg. 2.1 1. Harvcst data on sca cucumbels and shells from part-time industries 2.3). This gathering has Lucero on Santiago Island. The alternation of seasons leads to enticed the entry of many men into what was shifting occupations among some harvestom. Data are from a formerly a sustenance fishery dominated by survey by de Gueman (1990). Reef slolre ,.....

Fig. 2.12. Primary gear use areas on the Santiago Island ruef. A large area of the rccf flat is rcntud to fish corral owners by the municipality. During the spawning migrations of Siganus fuscesce~~s(Dararrgen), other harvest activities in this area are provented by lhe corral owneru. Fishing gear u$c on Ibc led slope is more uniform than on the reef flat, but varies in intensity with distance hmthe rccf crest.

REEF FLAT STUDY RESULTS klad fish corrals overlaps somewhat with the spearfishing and handlining operating areas. Most gear are used in particular portions of However, the baklad owners do not permit other the reef flat. Gillnets and fish traps overlap in forms of fishing in their areas during the migra- some areas (Fig. 2.12). However, they tend to tions of the rabbitfish (Siganusfuscescens) twice target different species. The catch of the gillnets each year. Overlaps in species targeted by various tends to be strongly dominated by rabbitfish (Si- gear are illustrated in able 2.1. ganidae), while that of traps is dominated by small Gathering takes place throughout the non- wrasses (Labridae) and parrotfish (Scaridae). sandy parts of the reef flat, but different species Spearfishing areas include some seagrass regions are harvested in different areas. For example, and a strip of seasonal grounds along the reef west Caulerpa seaweeds are found primarily north of (Fig. 2.12). The handlining areas are generally Dewey on the eastern margin of the reef flat, while restricted to small areas in the northeastern reef Zlipneustes sea urchins are found mainly around flat and lagoon. However, occasional handlining Silaki Island and Lucero on the western portions can occur in other areas. The rental area for ba- of the flat. Most of the partitioning of the reef flat is years indicates that seagrass fish might be ex- because of the extreme heterogeneity and natural pected to recover in a strong pulse to higher popu- partitioning of the reef flat by the target organ- lation levels sometime during the few years isms. Villages tend to specialize strongly in which following the implementation of a marine reserve. sets of gear are used, based on such factors as Recovery of coral-dwelling fish may be slower be- distances to favored fishing grounds and exploita- cause of the longer periods of time necessary to tion patterns of other villages. However, none of reestablish coral habitats damaged by blast fish- the gear are exclusive to any village. Instead, each ing, cyanide fishing and coral-grabbing anchors. village tends to have a preponderance of one or two types of gear, and a minority of one or two others. This makes sampling difficult because all villages must be surveyed to obtain a reasonable picture Overall study results of the whole fishery. The average of appro xi mat el^ 26 timonth The overall harvest reflects the seasonality of translates to approximately 1tkm /month, or 12 the slope fisheries, which is buffered by the con- Vkm21year. This is similar to values previously stancy of the reef flat's yields (Fig. 2.8). Analyses calculated for the present reef flat (del Norte et al. of the individual catches illustrate the high degree 1989), and is close to the working figure recom- of uncertainty in the fishery at the species level mended by Munro and Williams (1985) of 14 (Fig. 2.14). The vagaries of irregular recruitment t/km2/year. The reef flat production may be kept success combine with the multitude of factors af- high by the fact that it is not cost-effective to use fecting the harvest procedures, such as weather blasting devices to capture fish dispersed through and market value, to produce very chaotic-looking the seagrass beds, and the fact that seagrasses patterns. However, the regularity of total recruit- tend to recover from various abuses (such as rak- ment is matched by a regularity in total harvest ing for shells) more rapidly than corals do from the which is remarkably predictable. stresses they must endure. A cursory look at the nonreef longline fishery There were no obvious long-term trends in indicates that harvests may have declined (Fig. either effort, catch rate or total catch (Fig.2.8). An 2.15). The time series of data is too short to be exception to this is that gillnets tended to have a certain of the long-term trend. However, there is peak in activity during August 1989 (Fig. 2.9). little hope for finding compensatory harvests in This peak was not found near Dewey on the east- other local ecosystems. ern reef flat, and is offset from the November-De- The irregularity ofharvests for particular spe- cember peaks found in nonreef soft-bottom areas cies has some implications for the development of (Fig. 2.13). This type of sporadic variation between the market system in Bolinao. The buying public

-I Gillnets I, 1.o 'I .; I

Jul Jan Jul Jan Jut 1988 1 98s 1909 1990 1990

Fig. 2.13. Compariaonofsoasonality in gillnet harvests in anonreef sofbbottom area and on the eastcrn reef flat. SIganus opinus

Eplnephelus merra

Fig. 2.14. Top species by weight in the shorn landings orBolinao. The imgular patterns are tho result of the interplay of factom afleeting the fmh populations and harvcst activilica. Fish drawing6 am by Magnus Olsson-Ringby. 1.2 Chellinus trllobatus F 1.o b 0.8 rm 0.6 E 1 0.4

0.2

Choerodon anchorago 1.5 LE 1.0 E 2 0.5 ?

10 ~asoun~cornis

Calotomus japonicus

Scarus ghobban

Fig. 2.14 (Continued) 1.0 - Soft bottom 8 0.8 - f 0.6 -

C 0.4 -

Fig. 2.15. Harvests from a nonreef longline fishery adjacent to Santiago Island. must be very flexible in order to benefit from the management measures are instituted, such as the fishery. Export to markets such as Manila may be establishment of a marine reserve. The vagaries limited somewhat by the fact that advance orders of the fishery could be avoided far more if a strong cannot be filled predictably. This situation may shift was made from a dependence on capturing change for some species, such as rabbitfish (ba- organisms to a reliance on raising them through rangen) and groupers (lapu-lapu),if appropriate mariculture techniques. CHAPTER 3 REEF SLOPE FISH COMMUNITIES

General sity coral cover (100%) exist in some places. The extent of these dense areas has decreased notice- The reef slope is the oceanward extension of ably in the last ten years because of the destruc- the reef which is separated from the reef flat by tive effects of blast fishing, cyanide fishing and an intertidal reef crest (Fig. 3.1). The reef slope of anchor damage. Other organisms covering the Bolinao is very large, extending northeastward slope include sponges, bryozoans, tunicates, hy- into a subsurface barrier for at least 15 km (Fig. drozoans, forams and algae, such that very little 1.1). In general, the slope is gradual down to the hard substrate is exposed at any time. There are edge of a drop-off, which ranges in depth from 10 large areas of sand and rubble in the pits and riRs to 20 m. The bottom of the wall below the drop-off of the slope, as is the natural case. ranges from 20 to 30 m in most areas. Beyond this The alternating monsoon seasons result lo- wall is a gentle talus slope of sand and coral cally in a period of dry weather from January to rubble, which extends for several kilometers to the May and rainy weather (with numerous typhoons) edge of the Luzon shelf. The talus is dotted with from June to December. Slightly out of phase with large outcrops of limestone substrate covered with this is an alternating pattern of temperature corals and other benthic life. which peaks in June and July, and drops to its The reef slope is formed from limestone ac- lowest in January and February (Fig. 3.2). Ty- creted over a base of ancient reef material. The phoons rarely hit Bolinao directly, but are turned ancient reef had been exposed to the air during the northward or southward by mountain ranges as previous ice age from approximately 45,000 to they approach Luzon from the east. However, the 6,000 years ago, so the reef we see today is no more peripheral winds, rains and wave action do affect than about 6,000 years old. The slope is creased the reef substantially, and typhoons have been with rifts or channels with depths that increase known to swing back toward Bolinao after aniv- outwardly from the crest. Alternating with these ing in the South China Sea. The storms account are broad ridges, such that the general morphol- for the presence of large boulders of dead corals, ogy resembles the toes of a person's foot. The wall sometimes exceeding 2 rn in diameter, and shifting structure is found only on the ridges, with the rifts sand bars or dunes found on many reef flats. opening directly into the talus slope. The ridges, Our studies of the fish community have re- rifts and numerous pits of various shapes and vealed that the species are distributed to some sizes on the slope show the combined effects of the degree by depth and by the amount of surface weathered ancient limestone and differential roughness, particularly in the 10-cm range (i.e., modern reef growth. small holes and cracks in which the fish and their The coral cover of the reef slope and wall is food organisms can hide). Despite these distribu- generally 15-30%, although patches of high den- tional tendencies, there is very little stratification 111 seagrass +- coral % Seaweed

Pig. 3.1. Pmfile of the Santiago Island reef, showing the flat and slope separated by a wave-breaking intertidal cmst. The profile is based on a transect ~unningnorthwards to the west of Silaki Island (3x vertical exaggeration). of the fish community into distinct subcommuni- those usually studied in ecological recruitment ties. Species abundance peaks are broad and over- studies (e.g., Doherty 1988). Our "recruits" are lap substantially. The assemblages also change smaller than those generally studied in fisheries significantly over time (Nafiola et al. 1990). studies, in which recruitment is defined in terms of the catchability of a gear (Sparre et al. 1989). A study of recruits focusing on the earliest settling Monitoring the reef slope stages would require supplemental sampling from very narrow transects (e.g., 1 m), which was be- The reef slope is monitored on alternating yond the scope of the current program. Analyses months by censusing fish along underwater tran- of the abundances of the smaller juveniles have sects. The divers usually swim in pairs along a been omitted from this book because of inadequate transect line, identifying and counting all fish data. within 5 m to each side and above the line. Fish From August 1987 until June 1990, each site are classed into life stages corresponding roughly was surveyed based on two transects laid at the to young juvenile, large recruited juvenile, time of the dive, each using a 100-m nylon meas- subadult and adult, based on relative sizes and uring tape on a reel. The depth at each site varied coloration patterns for each species. It is impor- somewhat because of slight inaccuracies in locat- tant to note that the group labeled "recruits" in ing the areas between samplings. By July 1990, this book refers to juveniles which are larger than all 18 sites on the slope had been marked with

Fig. 3.2. Seasonality in hothm salinity and hmpcralure in 111 reef sbpe sites. Vertical barn arc 95%confidence limits. Fig. 3.3. Sampling sites around Santiago Island, Bolinao. S, F = elope and flat visual transect sites, T=trawl sites, L = fish landing sites. permanent concrete markers anchoring bamboo some fishing operations. This was intended to buoys. Transects were permanently constructed reduce damage inflicted by irate fishers. However, from heavy nylon fishing line anchored with small some vandalism still existed, resulting in occa- concrete blocks (Figs. 3.3 and 3.4). The total tran- sionally missed samplings of certain sites at cer- sect length per site was reduced from 200 m to 100 tain times. Aglobal positioning system (GPS) was m because there was less variance between sam- used to document the coordinates of each site, but plings to be accounted for at each site. relocation often entailed waiting for the proper The bamboo buoys were designed to tilt over configuration of satellites to appear within hori- when struck by the horizontal scare line used in zon limits. Still, samplings are believed to have

(30~30x5em)

Fig. 3.4. Permanent markings on the reef slope transect sites. been adequate to represent changes over time in variance estimation (lbkey 1977; Pauly 1984). species and abundances on the reef slope. The method differed from that described by Zahl Temperatures were measured using labora- (1977) in that the variances were estimated by the tory-type liquid thermometers, and water samples successive omission of transects rather than spe- were taken for salinity analysis using a refracto- cies. As with the analyses of mean transect vari- meter. Depths were measured using capillary or ances, this was done in order to properly account Bourdon-tube depth gauges. Surface roughness for the variance among sites, which is conceptu- (heterogeneity) was measured at 1-cm, 10-cm and ally more relevant to our study than a variance 1-m scales with the use of a chain with 1-cm links, based on the way individuals are distributed a pair of 10-m sticks linked with a string, and a among species. The estimation of species number weighted meter stick, respectively. In each case, variances in this manner is mathematically the number of smaller sticks (links) was counted equivalent to the technique of Heltshe and For- which, when laid end to end across the substrate, rester (1983). In all cases, erratic results attribut- covered a linear distance ten times larger than the able to the sensitivities of the jackknife method to stick. The number was divided by ten to give a various data characteristics Wainer and Thissen measure of roughness at that scale. For example, 1975)prevented the use of the jackknifed diversity a roughness index of 1.4 at the 1-m scale meant index estimators. Thus, the graphs consist of di- that a meter stick was laid 14 times along a 10-m versities calculated normally, flanked by 95% con- linear distance, measured with a tape measure fidence limits based on jackknifed variances. held tangentially to the surface. The more rough the surface, the more short sticks or links are required to span the straight distance, and the higher the index. 7b reduce ambiguity, objects Fish abundances causing a tilt only within the first 20% of the short stick were ignored, and the 10-m distance was Graphs a and b of Fig. 3.5 show the variations measured tangentially to the substrate where it of fish abundances on the reef slope. Every year in was convex or as a chord where the surface was April and May, large numbers of juvenile fish are concave, such that each end was an equal distance recruited to the slope. A natural decline occurred vertically above the substrate. in the next few months in each case, probably because of the combined effects of losses to predation, harvesting and rapid growth to the subadult stage (Fig. 3.6). Data calculations The peaks in subadult abundances follow in July and August, reflecting the rapid growth of Certain diversity indices are very sensitive to most of the fish. The differences between the sample size and cannot be scaled up or down peaks for juvenile recruits and those for subadults without further field sampling (Pielou 1975,1977; represent primarily losses due to predation be- Magurran 1988). This fact contributed to our de- cause the fishing gear on the reef slope generally cision to include for all abundance and diversity target subadult and adult fish. The data series is analyses only transects (15 of 18) which had been too short to be certain of any trends in the heights sampled without omission throughout the study of the peaks of recruitment from year to year. dates included. Error bars on graphs based on The adult fish showed only minor seasonality mean transect abundances and diversities were (Fig. 3.7). This might have been expected natu- calculated based on the usual variance estimation rally because there is always a limit on how many procedures. Note that the variance used was that fish reach adulthood, at which they achieve a low among sites, not based on numbers of individuals rate of natural mortality. One reason for this is among species as is often used for the Shannon- that the adult fish tend to have well-established Wiener diversity index (Pielou 1975). The empha- and well-defended territories and hiding places. sis is therefore on the variability among transects, However, there was a decline in the abundances not on determining the uncertainty associated of adults over time, interrupted only briefly by a with applying the index to a sample unit. pulse in June 1991. Fishing pressure gradually The error bars on the diversity measures for reduced the populations of adult fish by approxi- combined transects were determined by jackknife mately 80%. By the end of the study, adult fish I \ ; ,.' '', 600 - I., ,---\ I* I

400 -

... 200 -

Fig. 3.5. Abundance and diversity by transect on the reef dope. Vertical bars am 95% confidence limits (Icft: all lifc staps; right: by life stage where black = recruits, dashed = suhadults and dottod = adults). were so scarce on the reef slope that they were becoming dificult for divers to find. The probability that an adult fish would encounter a hook or spearfisher declines rapidly with low abundances, particularly because they are scattered in essentially two-dimensional space. The one-dimensional search path that a spearfisher would have to take to encounter an adult fish would have to increase exponentially to account for a linearly declining two-dimensional abundance. This may be why the abundance remained fairly constant in the final year of the study. A major question arises as to whether the recruiting juvenile fish come primarily from the reef itself, from other fringing reefs or from elsewhere, as from the thousands of subsurface reefs of Philippine waters (McManus 1988). Recent studies indicate that most reef fish recruit on a scale of hundreds to thousands of kilometers (e.g., Doherty 1988). Therefore, it is unlikely that the reef is entirely "self-seeding". However, if the timing of reproduction were to be regulated to take Fig. 3.6. Abundances of fish mite(top) and adult fish (bot- advantage of offshore entrainment features, as tom) on the reef slope. The decline of adults was interrupted only by a temporary pulse in May 1991 which quickly disappeared.

Fig. 3.7. Abundance and diversity of adult fish on the reef slope. appears to be the case in Hawaii (Lobel and Rob- limitation is a controlling factor, and that the inson 1983), then recruitment success may be dominance of these resources is not guaranteed dependent on adult populations across reefs over from year to year by any particuIar species. In any a few hundred kilometers of coastline. The reefs case, it is surprising that the recruitment was so of Bolinao may be extensive enough relative to strongly seasonal. It is likely that there was some others within such a distance for the local adult driving factor, such as favorable current patterns populations to directly influence local recruit- or food availability which made this period par- ment. Of more importance and greater likelihood, ticularly successful for new recruits. however, is the limitation in recruitment expected to ensue from the broad-scale overfishing of reefs along much of the southwestern coast of Luzon. Species diversity May is in the midst of what is considered by local fishers to be the calm period of the year, and falls The overall mean number of species per tran- between monsoon seasons. This could influence sect on the reef slope appears to have dropped the timing of fish reproduction, such that larvae temporarily and then recovered. The total number are not broadly dispersed (Pauly and Navaluna of species in the combined transects known to 1983; Sinclair 1988). Currents affecting the area reach adulthood fell at least 33% (Fig. 3.7). The during the May recruitment period tend to pro- lack of a similar pattern in the number of species ceed northwards from the Central Philippines per 1,000 individuals (R) indicates that this drop (Wy-rtki 1961). The current structure of Lingayen is related to the general loss of individuals, and Gulf includes incoming currents from both the not necessarily a more complex ecological change north and south, which converge and generally driven by predation and competition. expel to the northwest, away from the Bolinao reef The Shannon-Wiener diversity is an indica- (de las Alas 1986). However, the possibility of tion of how likely an individual fish will encounter recruits arriving from the north on intermittent a high diversity of other species, and accounts for countercurrents remains. both the number of individuals per species and the The data series is too short to determine if the evenness with which they are distributed among recruitment is clearly decreasing with the decline species. This diversity measure showed little in local and regional stocks of adult fish. If further change over time. However, an analysis of the studies indicate continued high levels of recruit- evenness component of that index shows that for ment despite the increasing levels of coastal ex- a limited period, an increasing evenness balanced ploitation, then recruitment from offshore out the effect of the overall loss of species. This subsurface reefs may be indicated. There is a need increase in evenness is to be expected in any for longer-term transect data, investigations into situation in which predators, including people, the genetic structure of the local stocks of coral tend to harvest the most abundant species and to reef fish, and studies designed to pinpoint the switch from one to the other as each becomes sources of local recruitment. scarce (see technical box). The fact that so many An analysis of the top ten species by counts species are economically valuable locally tends to (excluding prerecruit juveniles and larvae) shows favor this process. that no one species accounted for a major part of The overall annual rise in the number of spe- the seasonal recruitment pattern (Fig. 3.8). Only cies in April and May coincided with the annual the goatfish, Parupeneus trifasciatys, and the pa- peaks of recruitment (Figs. 3.5 and 3.9). This macentrid, Pomachromis richardsoni, came close confirms that the recruitment tended to involve a to the appropriate pattern. The recruitment ap- multitude of species, approximately 10 to 20 out peared to consist of an annual "lottery for living of roughly 210, or 5'to 10%of the total slope species space," with success among individual species at the start of the monitoring. varying greatly between years (see also Sale Usually, the total number of species encoun- 1978). The total recruitment was fairly predict- tered was higher than that found per transect, a able, considering the potential effects of variabil- result of the restricted ranges of these species. ity in larval survival (Beyer 1989). However, the This heterogeneity in composition across the slope predictability as to which species dominated re- would result in an increase in the difficulty that a cruitment each year was low. This could be inter- spearfisher might have in finding a useful target. preted as indicating that some form of resource However, it also indicates that individuals of a Slganus splnus 40

Fig. 38. Abundances of the most common species in the reef dope visual censusing (grey = combined sizes, black = recruits, dotted = adults), Fi& drawings are by Magnus Olsaon-Ringby. Pomachromls rlchardsonl 20 E. m, a 6 15 a x 4 = - lo 5

Fig. 3.8 (Continued) L

I I 'I 100 - ...... 50 - ...... :....:. . - . .

0 -I.-... 0 :---'. '

100 - 500 : .a ...... * . 400 : "...... ,...... , 300 ...... : 'I .+ ... :...... : ., .. ' :.' ., ,* .* : 200 . *

0 -'"".'" ' ' LY...... I > c- 0 '.

I I . . 4.0 - ..+

3.5 -

3.0 - L......

...... '.. 0.9 - .' . '..". ,,.." ;+ : ...... ,\ . . : .' ".-y-

0.7 -

0.6 - 5 ' .'.....'.... '. 0.5 ' . ' - I988 lee9 1990 1991 1992 1900 1989 1990 1991 1992

Fig. 3.9. Diversity in combined transects on the reef elope. Vertical bars rapresent 95% mnfidence limits determined f~om variances obtained by jackknifing among sites (lelt: all life stages; right: by Iib stage whore black = recruits, dashed = subadults and dotted = adults). species may have difficulties finding mates with least occasional recruitment from other reef areas which to breed. It is possible that some species replenishes the supply. In areas where all reefs have reached or could reach population levels within a wide radius are heavily fished, this could below which reproduction is no longer successful. be a problem. The presence of unfished offshore If this were to occur on a scale large enough to reefs in the Bolinao area makes this unlikely to affect entire stocks of the fish, generally hundreds occur, except possibly for species dependent on to thousands of kilometers (Sinclair 1988), then shallow-water habitats for survival which are not this could result in local extinctions, unless at present on subsurface reefs. Effects of Harvest b. some of the abundant species normally act as switching predators on Fish Species Diversity which prevents competing species from competitive exclusion, i.e., one John W. McManus decimating the other in competition for food or space. This could be hap- pening here (see tilt-off). Add-on. Humans remove predators that normally had a fall-off effect. For example, In isolating the possible effects of fishing on removing most sharks from a reef (essen- the diversity of the fish, it is useful to define a set tially true in our case) might cause a gen- 3f simple effects which might be seen singly or in eral rise in successful recruitment zoncert. Many of the effects of fishing on a fish including that of species normally totally :ommunity known from the literature have been incompatible with the predators. This summarized by Russ (1991). I shall present here would cause a rise in abundances, species s. classification of some of the possible effects on richness and diversity, and have an un- diversity (Figs. 3.10 and 3.11), and then compare predictable effect on evenness. these possibilities with actual data regarding Tilt-on, Humans become switching changes in the composition of adult fish on the reef predators, causing an increase in even- slope. ness and freeing niche-space for other species. If the species pool is large, this could conceivably lead to an increase in species richness, simple diversity, Shannon-Wie- ner diversity, and of course, evenness. Otherwise, only the latter one or two of these would rise and the rest remain un- changed. Tilt-off. Humans remove existing switching predators, causing some species to become dominant relative to other competitors. If the switching predators are responsible for maintaining some of Fig. 3.10. Illustration of predator-mediatedcoexis- the species richness, then their removal tence. The predator (P) feeds more on the abundant might result in losses of richness, diver- species (A), preventing it from excluding the sity and evenness. This effect was widely weaker competitor (B) by dominating a resource. predicted based on studies of simple systems in which the removal of a predator appeared to have enhanced interspecies competition, as in barnacle communities with predatory snails (Connell1961), and HYPOTHETICAL EFFECTS similar rocky shore assemblages (Paine 1966; Menge and Sutherland 1976). How- 1. Fall-off. The abundance is reduced at ever, the loss of diversity predicted by many levels including both abundant and some to occur with the removal of top rare species, and some of the rare species predators from a reef fish community has are reduced to zero abundance. This yet to be clearly demonstrated empirically would only be expected if: (Bohnsack 1981; Russ 1991). It must be a. fishing was uniform regardless of the noted that a pulse of successful recruit- abundance of a species (false in our ment of a species in the midst of a fall-off case), and if many species are in- decline process could result in a tilt-off volved in the fishery (true). pattern. Fig. 3.11. Some possible effects of fishing on a co1ntnunity. Species are represented by bars arranged initially in rank order by abundance. People can act as predatom andfor as lemovers of predators to cause a variety of possible changes.

5. Terminate. Humans overexploit selected 6. Scramble. The dominance order of spe- species to local extinction. This might be cies is merely rearranged, with no sub- true especially when certain species are stantial net changes in abundance or very valuable, as with certain aquarium diversity. This could be the case ifrecruit- species, or if the docile seahorses of the ment was not strongly limiting and set- reef flat seagrass beds were to be collected tling space or other resources were. systematically for sale as folk medicine (a realistic danger), The activity would have to occur on a wide enough scale (hundreds to thousands of kilometers of coastline) to EMPIRICAL PATTERN impair recruitment processes. We would expect minor drops in richness and diver- A comparative analysis of diversity profiles sity, and conceivably a drop in evenness. from the inter-recnritment months of January- In all cases, this would hardly be notice- February (Fig. 3.12) highlights the dramatic drop able unless the original number of species in species encountered from 1988-1991.The num- was low or the number of selected species ber of species per 1,000 individuals increased until was high. 1990, a result of the fad that the number of January 198B

February IS90

January 1991

Fig. 3.12. Species abundance profiles of adult reef slope fish from year-end months. Many species appear to have "fallenoff' as overall abundancas declined. individuals encountered dropped faster (as per- changes in the degree of dominance. Considerable cent change per year) than the species number. "scrambling" took place among dominance ranks, This pattern reversed dightly in 1991, as the few as is apparent from the individual species graphs remaining species reaching adulthood reflected a (Fig. 3.8). However, this could easily be attribut- 10%increase in abundance. The Shannon-Wiener able to variability in recruitment success among diversity dropped from 4.0 to 3.2 (natural log the years. A possible example of a tilt-off transi- base). The evenness component of the Shannon- tion from 1988 to 1989, signaled by concurrent Wiener index dropped somewhat during the 1989 drops in diversity, richness and evenness, could be transition period, but returned to near its starting an artifact of recruitment variability in the midst value. This return indicates that the drop in the of a general fall-off process. The simplest explana- Shannon-Wiener diversity was more a result of tion for the overall loss of species is that they "fell the decrease in the number of species than net off' as abundances declined. CHAPTER 4 REEF FLAT FISH COMMUNITIES

General the ends and sedimentation throughout. The bottom of the lagoon is sand covered with microscopic algae, interrupted in places by patches of coral a The reef flat and lagoon are protected from few meters across. In 1978, most of these corals outside waves by the intertidal reef flat. Season- were alive and filled with dense schools of coral ality is as described for the reef slope (Chapter 3, reef fish. Our survey in 1986 showed that 60% of Fig. 4.1). The substrate throughout is mostly cal- the coral (in terms of cover) had been killed, pri- careous sand. Encircling the shore and encom- marily by blasting and cyanide fishing. These passing nearly all the fish ponds is a black, muddy activities have continued, and coral cover was substrate indicative of a time when mangroves believed to be far less by 1991. Unlike the case on were abundant. These are now virtually absent, the reef slope, there were very few newly settled with the exception of some seedlings recently corals to be found in the lagoon and reef flat. The planted by the Department of Environment and reasons for this are unknown, but possibilities Natural Resources (DENR). include organic pollution and siltation from The lagoon consists of what appears to be an coastal villages which may be harmful to plank- ancient riverbed modified by recent reef growth at tonic coral larvae or inhibitory to settling.

Jul Jan Jul Jan Jul 1988 IS89 1989 1990 1990

Fig,4.1. Bottom ~alinityand temperature measurements from six reef flat dCs. Vertical bars are 96% confidence intervals. The reef flat proper is dominated by seagrass. shift from pelagic to epibenthic within three more The beds are found in a variety of densities at weeks (Hasse et al. 1977). various depths, making satellite mapping of the Another group of fish of considerable impor- reef flat difficult (McManus 1989). Occasional tance is the cardinalfish, Apogonidae. These fish patches of living coral dot the seagrass beds, but generally are hidden during the day and disperse far more patches of dead coral abound. Very few at night for feeding (Thresher 1984). Although fish inhabit the dead hard coral, even when large their hiding places are generally in coral, they are abundances of algae and soft coral are present. An found in large abundances in the seagrass beds at exception to this are the dunes of coral rubble near night. This indicates that both types of habitat are the reef crests, especially north of Dewey, which essential to the populations. Thus, removal of the are inhabited by many herbivorous territorial coral from the reef flat could adversely affect the damselfish (Pomacentridae). fishery potential of the seagrass beds. The large sand areas exposed at low tide have Many species of fish in the seagrass beds and little seagrass or algal growth. Seagrass is also remaining coral patches form mixed-species absent from the backreef areas behind the reef schools which forage widely during the day. The crest. In some areas, the backreef consists of large herbivorous feeding activity appears to stir up solid coral colonies with "bald spots" on top where zooplankton in the substrate which are fed upon coral has been killed by exposure to air and fresh- by nonherbivores and herbivores alike. These water at low tide. These "microatolls" become schools of wrasses (Labridae), gaatfish (Mullidae), denser as one approaches the crest, finally coalesc- small rabbitfish (Siganidae), small parrotfish ing to form the raised crest itself. Other backreef (Scaridae) and others also frequent channels areas contain beds of Sargassum, a brown algae where other species lay benthic eggs. Many of which is not exploited locally but has market po- these are consumed. tential as a source of a variety of products. Pro- Another common schooling species is the ceeding across the crest in these areas, one passes striped catfish, Plotosus lineatus. This species from the wide beds of Sargassum into a thin band forms schools with others of similar species and of club-like llirbinaria, another brown algae. Fur- size, which comb through the seagrass and coral ther progress brings one to the intertidal crest beds in dense masses stirring up demersal itself, which is barren except for grayish-white zooplankton. These zooplankton, which live in the slippery algal coatings. Beyond the crest, the pat- substrate and migrate daily to and from the water tern often reverses, with another shallow-water column, are a major source of food in the reef flat, band of llirbinaria algae followed by a wider bed and probably on the reef slope as well. Many fish of Sargassum leading to the coralline reef slope. species are planktivorous throughout their lives In eastern areas of the reef, the crest consists of (e.g., small sea bass, Pseudanthias spp.; fusiliers, raised piles of dead coral rubble, oRen housing the Caesionidae). Others are more planktivorous as commercially important Caulerpa green algae juveniles and switch to eating seagrass as adults, (arosep). This algae consists of rhizomes with including some species of rabbitfish (Tsuda and berry-like projections. The algae is gathered for Bryan 1973; Bryan 1975). Studies have demon- use in salads, usually eaten with vinegar. strated that live coral tends to support more den- The primary fish species of the seapass beds sity of demersal plankton than either coral rubble is Sigctnus fuscescens, known as "rabbitfish or or sand (Porter and Porter 1977).Thus, damage to "spinefoot" in English and barangen (large) or the coral beds has a number of deleterious indirect padas (juvenile stages) in Bolinao. This rabbitfish effects on the total fish community of the reef flat, migrates out of the reef flat eastwardly, north of beyond the simple fact that living coral supports Dewey, on 2-4 nights after a new moon twice each greater fish densities than either dead coral or year in August-September (major spawning peak) seagrass. and March-May (minor spawning peak) (Arag- The invertebrate community of the reef flat is ones 1987; del Norte et al. 1989; del Norte and divided into species favoring seagrass, sandy, Pauly 1990). The fish are assumed to breed on the muddy and rocky (coral rubble) areas (de Guzman reef slope, but they have rarely been encountered 1990). The seagrass community is dominated by in the slope monitoring program. The juveniles herbivores, which vary in abundances seasonally. return to the reef flat within a few weeks, and The important commercial sea urchin, Dipneustes gratilla (kuden-kuden), maintains a low abundance throughout most of the year, but peaks in abundance in September and October (Fig. 4.2). This peak occurs just before an annual thinning of the seagrass beds in dense areas (Fig. 4.3), and may be one of the causative factors. The Zhp- neustes peak coincides with a peak in the abundance of Strombus labiatus. These are followed by a November peak in the abundance of another gastropod important in the shellcraft industry, Strombus urceus. The cowries used in shellcraft are found in rocky areas. Cypraea annulus, the ring cowrie, and Cypruea moneta, the money cowrie, both have broad peaks, the former being espe- Strombus urceus cially abundant in January (Fig. 4.2, data from de .llT Guzman 1990).

Monitoring the reef flat

A set of six transect sites was monitored by visual censusing from August 1988 until July 1991 on alternate months. The techniques and data obtained match those described for the reef slope. Sites were permanently marked as of October 1989; however, the transects were not. There was one transect per site, extending for 100 m, serving as a guideline for a 10-m wide censusing swath. Familiarity with the area gave a high consistency to the process of locating the sites by triangulation and visual cues, so that depth variation was minimal between samplings. Some within-site substrate variability was caused by minor shifts in the transect positioning leading to mqjor shifis in the amount of coral intersected. However, the six sites combined give a fairly representative view of Fig. 4.2. Abundancee of aelectadmmrnenially important inver- changes over time in the daylight fish community tebrates in 50 quadrats on the reef flat near Lucem. Vertical excluding the dense seagrass beds. bars am 95%confidence limits. The graphs are basedonunpub- The difficulties with visually censusing fish in lished data of A. de Guzman. dense seagrass led to the initiation of a trawl sampling program from August 1988 to July 1991. The trawl had a width of 2 m, a rigid, rectangular opening height of 1 m, and a roller below the mouth to minimize scraping the seagrass and stalling as corals are encountered. Early trials Fish abundances indicated that the escapement rate was unrealis- tically high during the day, so trawling was sched- uled for nights during which the fish cannot see Contrary to the case on the reef slope, the the net until it is upon them. The trawling encom- abundances and diversities of reef flat fish show passed 7 sites of 7 minutes trawling time each very little consistent seasonality (Figs. 4.4 to 4.7). (approximately 175 m), which are sampled on There is also no particular trend over time. The alternate months. All fish caught were counted, reef flat has been fished far more intensely than weighed and measured. the reef slope for a longer time, and this may be a AP~ Jul Oct Jan 1988 1988 1988 19 %9

Fig. 4.3. Seasonal variation in seagrass cover on the reef flat. The greatest effect is seen as an annual loss of dense areas, indicating that thinnin~- rather than contraction of seagrass areas explains the annual drop in cover.

factor in the fact that no downward abundance amount of effort which local exploiters put into trend is visible. Alternatively, the dominance of harvesting every possible individual (see Chapter the flat by seagrass means that broadly dispersed 2). fish become especially difficult to eradicate below certain levels. Blasting does little damage to seagrass fish populations because they tend to be soli- tary or form schools which are small. However, the Species diversities fact that 60% of the coral cover had already been destroyed before the start of the study indicates No obvious trends occurred in species rich- that the community is far less productive than it ness, diversity or evenness during the study pe- should be. In areas of the Philippines where fish- riod (Figs. 4.4 to 4.7). Apparently, the reef flat fish ing is minimal, densities of coral reef fish gener- community has already long since been reduced to ally exceed 10,000ha (Aliiio, pers. comm.). The a level of diversity and abundance which has been reef flat abundances here, as on the reef slope, maintained over the three years of the study. It is rarely exceed 500ka. difficult to predict how long the current situation None of the 10 most abundant fish species in can be maintained ecologically with the rapidly the visual transects and trawl samplings shows growing human population and the systematic very regular seasonality of abundance (Figs. 4.8 destruction of coral by blasting and cyanide fish- and 4.9). Even the regularly migrating Siganus ing. We may expect some further changes in the fuscescens apparently has difficulty maintaining future as, for example, the amount of coral cover a regular pattern of successful recruitment (Fig. drops below the critical levels necessary to main- 4.9). This is not surprising, considering the tain the cardidfish populations during the day. Fig. 4.4. Reef flat fish abundances and diversities by transect fmm six transect sites. Vertical bars are 95% confidence intervals (leR: all life stages; right: by life stage where black = recruits, dashed = subadults and dotted = adults). Fig. 4.5. Reef flat fiah diversities for combinedtransecte. Vertical bars am 95%confidence limit^ determined by jadcknifmg among &es (left: all life ~tages;right: by life stages whem black = recruits, dmed = subadults and dotted = adults). Fig. 4.6. Reef flat fish abundances and diversities by transect from 7 mllcr trawl sites. Vertical bars are 95% confidence limits computed conventionally (lefk all life stages; right: hy life stage where black = recruits, dashed = sr~hadultsand dotted = adults). 250 04 .2 .-2 a 200 1 g !, o. 6 150 1 h -L

50 : 1 0 '-1.!.1.1.1.1.1 ...... I......

Fig. 4.7. Reef flat fish diversities for combined trawl sites. Vertical bars are 95% mnfrdence lirnit~determined by jackknifing among sites (left: all life stages; right: by life stages where black = recruits, dashed = subadults and dotted = adults). 0 Eupomacentrus nigricans 100

A 5.r eo I? e 9 2 60 Pg - 40 20

0 Dascyllus aruanus

Fig. 4.8. Abundances of top species from the reef flat visual transects conducted during the day (grey = combined sizes, black = recruits, dotted = adults). Fish drawings are by Magnus Olsson-Ringby and J. McManus. Pictures next to unknown species are generalized for the taxa. 1 @ Amblyglyphldodon curacao

Sardlnelle sp.

Fig. 4.8 (Continued) @ Apogon ranglcnsls 20

Fig. 4.9. Abundances of top speciea from the reef flat trawl wllectione made at night in the seagram of the reef flat (grey = combined ~izee,black = recruits, dotted = adults). Fieh drawidgs are by Magnus Olsson-Ringby and J. McManus. Pictures next to unknown species are generalized for the taxa. Fig. 4.9 (Continued) CHAPTER 5 REDUCING THE RATE OF EXPLOITATION

General of fishing, then the most desirable fishing level for most situations is that at which net yields (i.e., In broad terms, the reason for the poverty profit minus cost) are maximized. That point level among the harvesters of Bolinao is the open- (maximum economic yield -- occurs access nature of the fishery combined with a lack the leR Of the the gross profit curve of alternative employment (see Smith 1979). his (maximum sustainable yield -- MSW. However, in can be summarized simp]istical]y here. In a new an o~en-accessfisher~wherevirtually anyone can fishery, increasing levels of fishing yield join in, the number of fishers increases until the increasing incomes to a point, beyond which fur- average net returns are comparable to those that ther amounts of fishing result in diminishing total people could get from other of employment* gross returns (Fig. 5.1). If we assume that a con- In the Philippines, there is very little choice of stantly fising cost is associated with a rising level occupations for those with limited training and investment capital. Unemployment tends to be high, and there is no compensation for the average

~ost~profitequlllbrlum point unemployed laborer. A marginal income is better after provldln tor than no income. There is thus a tendency for alernat~velivel%ode increasing numbers of people to enter a fishery ilibrium until the average person in the fishery is making U' no more than a marginal income. This is particularly the case in fringing reef systems where a person can harvest with little or no initial invest-

This type of "bionomic equilibrium point" fishery (Smith 1979; Stevenson et al. 1982; Clark 1989) is not ecologically sound because harvests tend to be far beyond those which are sustainable in the long term. Furthermore, the best short-term competitive strategy for the individual fisher is Fishlng Intensity (etlon) often to find ways to cut costs at the expense of the Fig. 6.1. Fixed price model for pmfit and cost in an open-access community of fishers. For example, the fisher fishery. People tend to enter the fishcry until profits are might begin using blasting devices to hawest reduced to near the cost of fishing. If alternative livelihoob are more fish cheaply. This may improve individual available, the potential profit creates an additional bpportu- nity cost"to fishing,and the equilibriumpoint is pushed back to profits until the practice becomes widespread. more desirable levels. Then the resources will once again be exploited to the point of minimal returns -- at a new equilib- and family planning to change the scenario de- rium point lower on the gross profit curve than scribed above for the immediate future of Bolinao. before. This practice does considerable hamto the The alternative livelihoods would offer profits resource itself in the long run. which the fisher must "pass up" in order to fish. An alternative way of cutting costs is to give This cost of opportunity lost, "opportunity cost", low-interest loans to the fishermen to "improve must be considered by the fisher in deciding their gear". This again is a short-term solution whether or not to continue harvesting. The oppor- (usually with short-term political benefits to those tunity cost is effectively added on to the cost of who arranged for the loans). The fishermen gen- fishing. The "absolute cost" of fishing does not erally increase effort again until marginal in- change, but the total cost of fishing rises, forcing comes are the norm. Giving loans to fishers who the equilibrium point back to more desirable lev- are in an overfished situation usually makes the els (Fig. 5.1). Ideally, harvesters would then leave situation worse. The net result will actually be less the fishery until the available net profit to be made catch in the long run, despite increased effective by each remaining fisher meets or exceeds that effort. Additionally, the ecosystem may be pushed which could be made from the alternative liveli- into a state of less resilience to stresses and per- hood, However, other factors, such as job desirabil- turbations, natural- and human-induced, to which ity or the need for training must be accounted for. it is periodically subjected. Furthermore, the alternative livelihoods should be profitable enough that the fisher family could allow children to attend school rather than work. Reducing fishing effort Improvements in local school facilities would also encourage greater attendance, and ultimately im- There is good reason to believe that fishing prove occupational mobility. effort should be reduced by at least 60% from the current level and maintained that way in the future, i.e., at least 60% of the fishers and gather- Types of overfishing ers must leave the fishery (see technical box). In cases such as this, the general solutions to At least four types of overfishing have been the problem include: identified internationally: growth, recruitment, 1. offering unemployment compensation to ecosystem and Malthusian overfishing (Pauly et potential fishermen, which is not usually al. 1989). Growth overfishing involves harvesting economically feasible in the Philippines; in such a way that the mean size of the fish 2. taxing the fishery to raise the cost of fish- captured is suboptimal for providing effective ing, thereby protecting the ecosystem and yields from a fishery -- ie., the yield per recruit is stabilizing the resource supply -- this not optimal (Beverton and Holt 1957). Recruit- would leave many fishermen jobless, and ment overfishing occurs when the fishing effort is is not a realistic solution for most coastal so intense that the process by which the fishery is fisheries; restocked through reproduction and resettlement 3. forcing people out of fishing, which would is impaired (Ricker 1954, 1975; Schaefer 1954, be difficult to achieve, considering that 1957). Note that this would be most likely to occur most people locally view fishing as an in- when overfishing occurs on such a wide scale alienable human right -- this would also (hundreds to thousands of kilometers of coastline) lead to an unacceptably high level of un- that the "stock" or subpopulation providing the employment; and recruits is broadly affected (Sinclair 1988). Eco- 4. providing viable alternative forms of em- system overfishing causes a shift in community ployment and slowing down population structure from a fishery dominated by valuable growth. species to one dominated by species of less eco- The last solution is the most reasonable. nomic value or utility (Pauly 1979). Starting a series of local industries alone would Malthusian overfishing (Pauly et al. 1989; only be a short-term solution. It is unlikely that Pauly 1990) was named after the Rev. I.R. such industries could keep up with the currently Malthus (1766-1834), who clearly demonstrated rising population growth rate for long. Efforts that the exponential rise of human populations must be put into both alternative job development was a cause for concern. The definition of the overfishing condition is as follows (Pauly et al. yields on the reef slope, and fish populations 1989): throughout are far below what would be expected in a natural reef or one in a region fished optimally Malthusian overfishing occurs when poor fishermen, from a recruitment standpoint. The large schools faced with declining catches and lacking any other of milkfish (bangus),mullet (Mugilidae) and other alternative, initiate wholesale resource destruction in their effort to maintain their incomes. This may valuable species which historically had congre- involve in order of seriousness, and generally in gated in the area have nearly disappeared temporal sequence: (1) use of gears and mesh sizes (Quintin Caasi and others, pers. comm.). Finally, not sanctioned by the government; (2) use of gears the environmentally and self-destructive fishing not sanctioned within the fisherfolk communities andlor catching gears that destroy the resource baso; methods which abound are clearly symptoms of and (4) use of "gears"such as dynamite or sodium Malthusian overfishing. The strong causal rela- cyanide that do all of the above and even endanger tionship between poverty and this form of over- the fisherfolks themselves. fishing indicates that the most suitable corrective approach is an economically based one. This rein- All forms of overfishing are apparent in the forces the conclusion that the most appropriate Bolinao fishery, The fish in the markets are gen- means for reducing fishing pressure would be an erally small subadults. Adult fish are scarce on the effective program of alternative livelihood devel- reef slope, The fishery produces relatively low opment,

How Much Harvest would reduce the ratio of fishing mortality over Effort Should There Be? total mortality to less than 0.5, or a more precisely estimated value based on yield-per-recruit analysis (Gulland 1983;Pauly 1984; Sparre et al. 1989). John W. McManus One could then reassess the situation two or three years later, and readjust effort accordingly This is feasible because the monthly data on the lengths GENERAL of 30-50 fish can be gathered by a single worker as part of other duties, such as managing a marine In some studies, it is possible to produce quan- reserve or collecting fishery statistics. No matter titative curves for determining the relationships what course of management action is taken, it among yield, cost and effort. Doing this requires would always be wise to provide some minimal that a broad range of information on the relation- follow-up assessment and to assume from the ships is available. This may be obtained by moni- start that regulations will need readjustments toring a fishery from inception to an advanced every few years. However, it would be helpful to state. Alternatively, if data on a series of similar determine a "rule of thumb" for making an initial reefs are available, including those subject to a assessment of necessary effort adjustments. broad variety of effort levels, then the curves can be constructed quantitatively (Munro and Thomp- MELD/EFFORT CURVES son 1983). In either case, it is possible to assess the current status of yield, cost and effort, and to There are three fundamental shapes for a estimate the appropriate level of harvest effort yieldleffort curve (Pella and l'bmlinson 1969; (e.g., number of boats per day) necessary to maxi- Cushing 1981) which I shall refer to as symmetri- mize profits and ensure the longevity of the re- cal (Fig. 5.2), right-skewed (piled to the left, Fig. source. 5.3), and left-skewed (piled to the right, Fig. 5.4). In cases where this information is lacking, a The left-skewed curve implies that in the initial more indirect route may be necessary. One ap- fishery, small increases in effort lead to small proach would be to use some methods such as increases in catch until an optimum is reached, length-frequency analysis on some key species to beyond which yield falls off more abruptly. This determine if a system is overfished (Munro 1986). does not seem to be true of some coral reef fisher- One could then reduce effort arbitrarily to a cer- ies, where initial efforts produce rapidly acceler- tain level or to an estimate of the effort which- atikg yields until a maximum, beyond which yield MEY- MSY

L4 4B, 50% Effort reductlon ~-440, 60% Effort reduction -...... A c h4AB, 70% Effort reduction B Flshlng lntenslty (effort) Fishing intenslty (effort) Fig. 5.2. Symmetrical production curve. Barn represent pow Fig. 5.4. Left-skewed production curve. An effort reduction of sible rangea for effort reductions; amws represent wductions 60% is appropriate forthe high-mst fishery (A), but id mnsorva- from the indi~kdequilibriumeffortlevels. An effort reduction tive for the low-mst fishery (B). A reduction by 60% in the of 60% would be appropriate for the low-cost fishery (B) and absence of information on the nature ofthe curve could Bewe to conservative for the high-cost fiahery (A). help establish the type of curve, permitting more optimal effort levels to bz set later. 2. An excess, unemployed labor force is willing to enter the fishery as an occupation. 3. There is no formal or informal unemployment compensation which would keep people from wanting to work hard for marginal returns. 4. No noneconomic social force limits entry into the fishery. 5. There is a large demand for fish. 6. The system has been operating under the above factors for a few years. From this we can conclude that such is an "equilibrium point" fishery operating near the 60% Effort reduction point at which costs are almost equal to yields. ~~44~~ 70% Effort reductlon This is confirmed in our case by the fact that stocks Flshlng lntenslty (effort) of fish are declining and incomes are marginal among the harvesters. Fig. 5.3. Right-~kewedproduction cum. Cost lines are the Knowing that the equilibrium point generally same as those in Fig. 5.2. Note that both points A and B fall falls to the right of the top of the curve (MSY), and more than 2.5 times the effort at MSY. A reduction of 70% or that the most desirable point for a fishery is some- more may be optimal in such extreme cases. where to the left of MSY,we can observe the effect of arbitrarily choosing a reduction of 60% on a appears to taper off slowly, delayed by the fact that variety of curves (Figs. 5.2-5.4,5.8-5.10). As can be species tend to replace each other as they decline seen, a 60% reduction in effort from an equilib- in abundance (Figs. 5.5-5.7). This scenario would rium point never exceeds MSY unless the yield fit a symmetrical to right-skewed curve. curve is so strongly skewed to the right and the We now make six assumptions which are valid cost of fishing so low that the initial effort level (at in our case: the equilibrium point) is 2.6 times greater than 1. The fishery is open-access. the effort at MSY. Seychelles (East Coast)

Fig. 5.5. Plot of coral reef fish catch va. fishingintensity 0 1,000 2,000 3,000 in western Indian Owan sites (redrawn from Gulland 1979). Fishlng lntenslty (trap sets krn* year1)

Fig. 6.6. Plot of coral reef fish catch per area vs. fishing intensity for 8 parishes around Jamaica (redrawn fmm Mum and Thompson 1983). Fishing Intensity (canoes kmq)

Fig.intensity 6.7. Plotfor of11 coral American reef fish catchSamoa vs. villagesfishing 0 5LL-dL--0 0 10 20 30 40 50 60 70 (redrawn from Munm and Williarna 1985). Note that the shape of the curve cannot be determined from the available data. Fishlng intenslty (persons ha-l of reef) Fig. 5.8.Fixed price model for trawl fishing in Manila Bay. 0 0.5 1.5 2.0 2.5 The optimal effort reduction was approximately 60%. Points represent yearly values pmpssing fmrn left to L A, - -- right (redrawn fmm Silvestre et al. 1987). Fishing mortality (F = YII)

Fig. 5.9. Fixed price model for the overall Philippine demeiaal fieheties (tx 10'). The optimal effort reduction wan approximately 60%. Pointa repressnt pupa of 70% 60% 50% annual catchee (left to right) from 1946 to 1984 (18- drawn fmm Pauly and Chua 1988). Flshlng mortality (year1)

Fig. 6.10. Fixed price model for Philippine pelagic fisheries. The suggested effort reduction was 70%. Data points progress gcncrnlly from left to right, 70% 60% 50% representing the years 1948 to 1985 (redrawn Fleet horsepower (x loa) fmm Dalzell et al. 1987). If one chooses a 60% reduction in effort and off between favoring high net profits and being the unknown curve is: conservative enough to be certain of being to the symmetrical --the estimate will be conser- left of MSY in case the curve is strongly skewed. vative for a very low-cost fishery (Bso in Being conservative is a useful property be- Fig. 5.21, and close to the effort leading to cause: the MEY (EMEY)point otherwise (Aso in 1. schemes to reduce the effort may not work Fig 5.2). to 100% effectiveness; right-skewed (piled to the left) -- the 2. the total yield may not be precisely opti- estimate will be close to EMEYfor a low- mal, but the catch rate (CPUE) per fisher cost fishery as long as the equilibrium will be markedly better and this will have effort is less than ~.~*EMsY.In extreme a beneficial effect locally; cases where the effort at equilibrium is 3. populations tend to rise, and with them, greater than 2.5*E~sy,a reduction of 70% the pressure to find work for more fishers or more may be optimal (Fig. 5.3). will increase; and left-skewed (piled to the right) -- the 4. having now established an experimental estimate will be conservative, i.e., to the point to the leR of the leR of MSY,a better left of EMEYand EMSYfor a low-cost fish- estimate of the shape of the curve and ery (Bso in Fig. 5.4) and close to EMEYfor appropriate adjustments can be made in a high-cost fishery (Am in Fig. 5.4). subsequent years. Note that the lower the cost of the fishery, the closer MEY approaches MSY. Thus, we must trade (1) Rake net used to collect shells and emall fish fmm wagram bda. (2) Seagrass and branching coral (Acmpora)on the reef flat.

(3) Sea urchina (I).ipneustes gratilla) from the reef flat. The gonada are sold as Bod. A divefa foot paddle is shown on the right. (4) Small fish captured by gillnet on the reef flat. The average size iu 10-15 cm. Gillneta am very size-elective and can be designed to catch larger fieh if they am abundant.

(5) Fish trap camouflaged with mrals. Fish enter through the funnel and have difficulty finding the exit once inside. The wicker constmction lowers cost and limits long-term fishing once lost.

Creel opened to revcal scveral cap4 [all aim 36 am typical of local catchee. (7) Anchor designed to catch on corala. Anchor damage can be avoided in a marine resewelpark by establishing permanent moorings.

(8)Research aides Elmer C. Dumaran and Fernando I. Castrence, Jr. measure fish at a landing site. Incally hired personnel are esmntial to maintaining open lines of mmmunication with villagers.

(8) Portion of ref flat (foreground) recommended for a marine reserve. A reserve would greatly improve local harvests of fish and invertebrates. Sea turtles occasionally migrate through Malilnap Channel on the left to Piaalayan Point on the right. .. . (10) Spear guns are carved fmm wood, and are powered by rubber strips. The spear is often a bicycle spoke. (11) Gillnettar elapu the water to frighten fieh into the net.

(12) Close-up of a fish Fish 1 trapped in successi~ dler shaped chambers. CHAPTER 6 A PROPOSED MARUVE RESERVEPARK SYSTEM

General Oslob, Cebu. The reserve constituted approximately 25% of the coralline areas around the small The most reliable means available for enhanc- island. In 1984, a change in local government led ing resource production and sustainability on the to a breakdown of protective management. Fish- reef would be to set aside a substantial portion of ing was reintroduced to the reserve areas, and the the reef system as a nonfishing area (Russ 1985; range of gear was extended to include habitat-de- Alcala 1988). This site would serve as a protected structive techniques such as blast fishing and breeding ground, migration route and nursery muro-ami. In the latter method, corals are broken which would allow fish, invertebrates and sea- by large rocks on lines as fish are driven into nets. weeds to maintain natural population levels un- Both the total fish production of the island and the perturbed by human activities. The area would daily catch per fisherman dropped by more than permit recruited fish to reach larger sizes before 50%. This drop occurred despite the fact that the being caught. The migrations and other move- fishing area increased in size once the reserve was ments of adult fish out of the reserve area as abolished. This clearly shows that a resewe can populations grow should enhance catches by low- be an effective way of enhancing fishing yields and investment, size-specific gear such as gillnets, individual profits. thereby reducing the problem of growth overfish- Prior experience has shown that the chances ing. Additionally, many species of harvested inver- for success in the establishment of a marine re- tebrates and corals with short plankton stages are serve are greatly enhanced if there is a substantial likely to be highly dependent on local adult popu- involvement at the village level in the planning lations for their recruitment, and the young of and implementation stages (Casteiieda and Mi- these species would continually enhance the popu- clat 1981; White 1986; McManus et al. 1988; Mc- lation levels throughout the reef system. Finally, Manus 1988). There are now more than eight a system of such resemes along the southwestern municipal marine reserves and parks in the Phil- coast of Luzon would probably enhance the re- ippines (Alcala 1988;Wells 1988). Several ofthese cruitment of reef fish and invertebrates with long have been successful enough to foster the reestab- planktonic residence times. lishment of dense populations of large reef fish, The fact that a marine reserve can substan- which have not only increased fishing yield, but tially enhance fishery yields in adjacent areas has also generated substantial municipal income by been well demonstrated in a series of studies con- serving as tourist attractions. ducted in the Central Philippines (Alcala 1988; Presented here is a rough outline for a poten- Russ and Alcala 1989; Alcala and Russ 1990). A tial marine reserve and park system which is marine reserve had been established in 1974 for designed to enhance local harvests and incomes Sumilon Island by the nearby municipality of from the Bolinao Reef Complex (Fig. 6.1). Many Fig. 6.1. Proposed marine reserve and park system. The area has been chosen to encompass most of the range of habitat types, andis expected to substantially enhance yields of fish and invertebrates throughout the reef. The park would generate ihcome and-enhance the tourist trado to provide alternative livelihoods for reef harvesters.

details of management and implementation will 6. using future ecological and fishery data to depend on future action of the local municipality. evaluate the effectiveness of the plan as it The role of the ecologist can include: is implemented, and suggest modifica- 1. clearly establishing the fact that the reef tions and refinements as it proceeds. resources are being seriously depleted and that timely, effective action is necessary to ensure future sustainability; 2. determining an optimal location, shape The need for a reserve and size for the reserve to ensure that all critical habitats and migration routes are included which are necessary in the life The major factors which establish the need for cycles of commercially important species, a reserve can be summarized as follows: and the associated organisms on which 1. The densities of fish on the reef are more they depend; than one order of magnitude below those 3. presenting a general scenario for how found in reefs subject to low fishing pres- such a management scheme could be op- sures. erated, as a basis for further refinement; 2. Numbers of adult fish on the reef slope 4. . presenting the recommendations to the have declined sharply in three years, al- local government for appropriate consid- though fishing pressure was fairly con- eration and action; stant. The number of species reaching 5. serving as consultants during the plan- adulthood has declined by nearly 33%. ning and implementation stages to pro- Fishers are maintaining harvest rates by vide ecological analyses of details and turning to progressively smaller fish. This modifications to the plan as it develops; practice cannot be expected to be sus- and tained for long. 3. Coral cover in the lagoon is less than 40% Funds collected could further support the ranger of expected levels. An area of protection team. could be expected to support much higher densities of coral-dependent fish and in- SHORE HABITATS vertebrates than are currently found. The migratory nature of many species and the Mangrove forests were once an integral part dispersion of young fish could ensure a of the reef ecosystem. Asmall, uninhabited section constant supply of both recruiting young of coastline beginning along the eastern side of fish and harvestable adult fish in other Pisalayan Point has been set aside for replanting areas. as a mangrove forest. What is illustrated is a very 4. The large fish corrals tend to limit the minimal area for this purpose, but it would at least numbers of migratory fishes which are assure a supply of common mangrove-dependent able to reach sizes suitable for gillnetting species for other mangrove forests currently being and spearfishing This results in a system planted in less intensely managed areas nearby. which favors the harvest of small fish by Pisalayan Point (from the word "egg" in the sectors capable of higher than average Bolinao dialect) was until recently a viable breed- investments. This occurs at the expense of ing ground for sea turtles. The reserve protects economically restricted smaller-scale fish- what is believed to be the major route ofthe turtles ers whose harvest of adult fish would be from the ocean to the beach on the eastern tip of more ecologically favorable. A properly lo- the point. Aprogram of turtle rehabilitation along cated reserve area would ensure a depend- that beach would be a major asset to the plan. able supply of adult fish which would The eastern side of Pisalayan Point consists of favor the economically disadvantaged a rough, rocky outcrop (ancient reef limestone) fishers. covered with dense brush. Large monitor lizards (Varanus saluator) and a variety of birds inhabit the outcrop and a few small rocky islands nearby. The reservelpark system The preservation of these habitats would considerably enhance the diversity of protected organ- GENERAL isms.

The recommended reserve and park system is SEAGRASS HABITATS mapped in Fig. 6.1. The reserve consists of a four-sided section covering both reefflat and slope The areas immediately to the north and east areas. The size of the park is limited to that which of Pisalayan Point are dominated by seagrass. can be monitored visually on clear days from a Seagrass fish tend to be more widely dispersed small, central tower. A picturesque area near the than coral reef fish, thus maintaining minimal center of the reserve has been set aside as a populations may entail setting aside proportion- marine park, where tourist diving may take place. ally larger reserve areas. The included areas of The rental of permanent mooring sites (and a ban reef flat are by no means uniform. The seagrasses on anchors) would generate income to support a vary widely and abruptly in density and species rotating staff of rangers. These would be situated composition. Large and small patches of sand, in a small station/information center on stilts at rock, coral and algae in various combinations are the park center along the exit channel connecting interspersed throughout the reef flat. Each par- the reef flat and the reef slope. Boats would be ticular combination of these bottom types sup- permitted to pass through the reserve and park ports a unique assemblage of fish and areas along marked channels. However, no an- invertebrates. The fishery as a whole, and the choring or mobile harvest activities would be al- shellcraft industry in particular, are highly de- lowed anywhere in the reserve or park, except for pendent on the availability of a diverse range of scientific purposes (by permit) or emergencies. A species, which must be supported by a correspond- system of fines and other legal penalties could ingly broad range of habitat types (de Guzman ensure compliance. Fine collection could be as- 1990). The site outlined for the reserve includes sured by empowering the rangers to confiscate representative areas of most of the habitats of the and hold boats or equipment until compliance. reef flat. 60 SEAWEED HABITATS MIGRATION AND REPRODUCTION SITES

Significant stretches of the reef crest in the The placement of the eastern corner of the eastern portions of the reserve are dominated by reserve is especially critical. The species Siganus Sargassurn brown algae. This algae forms large fuscescens constitutes as much as 40% of the fish- beds which are scattered throughout the reef com- ery, and is important to the spearfishing, gillnet- plex and support unique biotas. The reserve is ting and corral industries. The entire northeast designed to permit the enhancement of this type sector of the reef flat is currently leased by the of biota. Sargassum is of commercial value, thus town to fish corral operators, and preferential stocks of the seaweed may require preservation in sites are those which intersect the migration the future. routes of this species. Analysis of corral catches The most important seaweed currently in the indicates that the bulk of the outward migration markets is Caulerpa (C. racernosa, arosep, and C. of adult fish to offshore spawning grounds is lentillifera, butones ). This algae consists of berry- through a narrow area along the reef crest. The like structures. on rhizomes, which form patches eastern corner of the reserve has been placed at in areas of moderate wave and current action. approximately the center of this area. Thus, the Significant patches of this seaweed are included outgoing stocks will be divided into large portions within the reserve area, principally along the for both potential capture (e.g., by fish corrals) and north-facing reef crest in the eastern portions. preservation. Several studies have indicated that many coral reef fish prefer reef channels and high points of reef structure for reproductive activities (Sale 1980; Johannes 1981; Thresher 1984). The crest CORAL HABITATS regions in the reserve north of the center are cut by several channels in a variety of sizes, and The area in the center of the reserve includes exhibit considerable structural relief. The major a lagoon which s~pportedhigh densities of coral channel (Malilnap Channel) in the center of the growth until about 1979 when blasting and cya- reserve was known to serve as a major route for nide diminished the stocks to a small living frac- the entry and egress of large schools of transient tion. The sides of the lagoon still support a broad reef species prior to 1980, and continues to be a variety of lagoonal hard and soft corals and asso- favored blast fishing site. ciated invertebrates. The large volume of tidal flushing in the lagoon, the abundance of hard substrate and the presence of seed populations of many coral species immediately beyond the lagoon Implications for fishery patterns on the reef slope result in a reasonable probability that rehabilitation will occur. The lagoon opens into a very heterogeneous reef slope with considerable topographic relief. GENERAL The walls and channels are covered in places with a high diversity of corals. Damage from blast Important considerations in the construction fishing is particularly noticeable, but small re- of the reservdpark system are the existing system cruiting coral colonies are found in abundance. of territorial use rights in fisheries (TURFS)(Fer- This area would be a, very effective attraction for rer 1991), and existing traditional knowledge of tourist divers, particularly if feeding stations were fish distributions and behaviors (Lopez 1985). established to maintain large, tame fish. Both of these factors tend to be reflected in the Our research confirms that depth is an impor- current-day fishing patterns. The proposed re- tant variable in the distribution of fish species serve park system was developed to account for (NaAola et al. 1990). The northern corner of the information on fish abundances and migrations, reserve has been extended into depths of over 30 as indicated by the way particular gear are de- m. The enclosed slope area therefore includes a ployed, as well as to minimize the disruption to be broad range of depths and a correspondingly wide caused by the sudden restriction of the fishing variety of fish species. grounds. Further investigations into traditional knowledge and TURFS will be helpful in refining ducing alternative livelihood and population con- aspects of the design of the reservdpark system, trol must be implemented before the full effect of as well as in guiding the approaches toward im- improved resource availability will be apparent to plementation. the individual gatherer. A major advantage of the reserve is that no single m@orvillage dominates fishing in the area. Rather, fishers from several villages use the site REEF FLAT HANDLINING to supplement fisheries in other parts of the reef. The fact that the area is a desirable fishing ground Approximately one-half of the handlining on for a broad variety of gear is indicative of the the reef flat will be curtailed by the creation of the heterogeneity and productive nature of the area. reserve (Fig. 6.2). This reflects the fact that the It is necessary that a reserve support organisms area includes some of the few remaining habitats which are desirable to the fishers, otherwise it amenable to supporting adult fish under exploita- would not be easily justifiable. Therefore, some tion pressure. We expect that the large fish mi- conflicts with existing use are inevitable. The im- grating out of the reserve area once it is portant point is that the area should not, and in operational will favor expansion of the handlining this case does not, completely monopolize any of grounds in the future. the fisheries which it is intended to help sustain. REEF FLAT SPEARFISHING REEF SLOPE The case with spearfishing will be quite similar to that of handlining (Fig. 6.2). The reserve will Fishing along the reef slope is very homogene- ous, with no particular gear predominating in any cut back primarily on the seasonal spearfishing grounds. The seasonality of the target fish in those area. MGor methods and gear include spearfish- areas is indicative of critical life-history events, ing, drive-in nets (parisn's), handlining, and blast fishing especially reproduction, which causes them to fishing. Approximately 95% of the occurs amass there. The fish are particularly susceptible within 4 km of the reef crest (Fig. 2.4); the inten- to overharvesting at that time. Therefore, these sity drops off sharply beyond that distance. The areas are particularly desirable for inclusion in resewe will cut off only a small portion of that the reserve. A major target of the year-round fishery. spearfishery is Siganus fuscescens (bamngen), which is expected to flourish with the establishment of the reserve and to migrate outward in REEF FLAT GATHERING mature stages. Therefore, the spearfishing industry can be expected to gain far more than it loses The reserve includes several areas which are with the establishment of the reserve (Fig. 6.3). subject to harvest by gatherers, particularly in regions east of Silaki Island. The gatherers are currently enticed to travel considerable distances REEF FLAT GILLNETTING to reach these areas because sites closer to their villages are often too heavily harvested and are Sl;ganusfiscescens is also amajor target of the depauperate in desirable invertebrates. The pat- gillnetting industry, along with a variety of other terns of tidal currents on the reef flat indicate that migratory seagrass species. Gillnetting is very the reserve will bolster the stocks throughout the size selective, thus the fishery will adapt to target- reef flat as planktonic larvae are dispersed. This ing larger individuals as they become abundant. will provide increased harvests closer to home for Similar effects will be seen with spearfishing, the gatherers. However, the present rate of human trapping and, to a lesser degree, handlining. All population growth and the lack of alternative live- four of these gear share the characteristics of lihood will lead to overexploitation of the gathered being low-investment, widely dispersed fisheries resources per gatherer no matter what level the with a tendency to target large fish when avail- stocks achieve. The reserve will add to the total able, and with sharply declining effectiveness harvest, and prevent the complete depletion of when stocks are reduced. These are all desirable most species. A complementary program of intro- fisheries from a management point of view, which Fig. 6.2. Marine resewelpark system relative to primary reef flat Eshing areas. The eastern corner of the reserve would bisect the major migration path for Siganus fuseescens (bamngen), ensuring a supply of adult fish for the gillnettem and spearfishem, and still permitting substantial harvests for the fish corral owners.

Fig. 6.3. Yield-pelcrecruit analysis for Sigcmus fuacesce~(barangen). Curves &ow that the fine-meshed fish corrals induce growth overfishing far mom than gillnets and spearfishing as locally utilized. Yielda could be improved by favoring speafi&ing over fish corrals. This would also improve income distributionkause of the labolcintensive nature of spearfishing Population parametem were based on monthly length histograms per gear weighted by annual catch and combined. Procedures followed those in del No& and Pauly (1990),but involved an improved data mt from mid-1988 to mid-1989 (L-= 2f3.0, K = 0.84, M = 1.71). Natural mortality was estimated fmm the Pauly environmental formula, using T = M'C. Individual catch curvea by gear were wed to estimate the length of firat capture (L) per gear (Gayaniloet al. 1988). will benefit directly from the reserve. The benefits cluded and replanted as a mangrove forest. As to gillnetters, in particular, will far outweigh the with the rest of the reserve, it would be important loss of some fishing areas (Figs. 6.2 and 6.3). to prevent the harvesting of the forest, so that natural populations could be maintained as a way of reinforcing neighboring mangrove forest popu- REEF FLAT TRAP FISHING lations. There is currently an effective program of replanting mangrove forests throughout the Boli- Trap fishing tends target coral-dependent to nao area, since many of these areas will be open fish. There has been a very noticeable drop in the to the exploitation of the fish, invertebrates and sizes of traps used over the last 12 years as the plant products they support. The mangrove areas sizes of the target species have dropped. The rein the reserve should be set aside and protected serve actually covers very little of the existing trap from exploitation in order that they may "seed"the fishing grounds (Fig. 6.2). However, the coralline biotas of the exploited areas. areas protected by the reserve should provide an abundance of large fish which will migrate out to revitalize the trap fishery. Suggested implementation

REEF FLAT FISH COW GENERAL The reserve will cut back on the area desirable for fish corrals (baklad) by about one-half (Fig. The mior distinction between a "paperwork" 6.2). Fish corrals tend to target fish in the process reserve and an effective one lies in supervision. The proposed reserve has been designed such that of migration and to be located along important it can be surveyed conveniently from a small tower migratory routes. It is inevitable that an effective erected on the reef flat near Malilnap Channel at reserve would be aimed at protecting those same the center of the reserve. From this point, small routes. The southeastern side of the reserve has boats can be dispatched to investigate possible been located so as to bisect the major migratory - violations of anchoring or harvesting regulations. route of Siganus fuscescens. This will permit the continued harvest by corral owners of some of the The majority of powered boats passing through the reserve will be following the Malilnap Chan- stock, while protecting the rest and channeling it nel, which includes the only useful eastern exit toward exploitation by smaller-scale gillnetters route to the ocean at most tide levels. Therefore, and speafishers. The town currently derives an a ranger station located at this point will be very income from the corral area leasing arrange- effective in controlling activities by fishers. ments. This income may be reduced to some de- A small visitors' center could be included in the gree initially. However, some expansion of the building complex to provide information on the available stocks due to protection may later in- reserve and on the need for conservation and crease the desirability of other corral fishing resource sustainability The suitability of the site grounds in the future, and the income may then for supporting a building on stilts is demonstrated be recovered. Initially, however, the reserve is by the fact that a small shack on stilts at the site expected to assist the smaller-scale fishermen by (used as a trading station for fishers) has survived redirecting some of the stock away from the cor- for more than two years through several typhoons rals, which tend to benefit a higher economic because of the protection of the reef crest and strata because of their area rental, implementa- shallow waters. tion and maintenance costs.

FISHPONDS MARTNE PARK

The reserve is anchored along the eastern A small area extending from the ranger sta- edge of Binabalian and borders on a significant tion to the reef slope near the mouth of the Malil- fishpond area (Fig. 6.4). This area was formerly a nap Channel could be set aside as a marine park. very complex mangrove forest. It would be very As with the reserve, the park area would be pro- beneficial to the reserve if the pond could be in- tected from all forms ofharvest. However, the park f-'

Fig. 6.4. Map of fishpond areas in Bolinao. Most of the ponds have been built on former mangmve forest aroas, thereby reducing the potcntial resources available Ibr small-scale income generation.

would be available for nondestructive recreational management. Bolinao has an order of magnitude diving. Aset of permanent mooring buoys could be more suitable diving area and site diversity than established, and the use of anchors prohibited so the entire Batangas-Puerto Galera area com- as to protect the corals. Afee could be charged for bined. Divers would be easily attracted to make the use of the buoys, based on tickets dispensed at the 5-hour trip from Manila to Bolinao if three the ranger station. This income could help main- conditions were met: tain the station and support the staff. 1. availability of a diving compressor, prefer- A major benefit from establishing a marine ably operated by a knowledgeable diving park would be the generation of alternative liveli- expert or instructor; hood along the Bolinao coastline. Bolinao has a 2. effective curtailment of blast fishing; and very high potential for development as a diving 3. abundance of large fish. tourist area supported by visitors from Manila. The latter two conditions would hold true in The diving population of Manila includes thou- the park. If all forms of exploitation were effec- sands of business employees making repetitive tively eliminated, especially spearfishing which trips to low-cost resort areas. A major diving makes fish avoid divers, then large fish would ground is the Anilao, Batangas area south of Ma- accumulate within 3-5 years of operation. The nila. This area requires approximately 2 to 3 hours process could be greatly enhanced with the estab- of travel by road, and has flourished because of the lishment of regular feeding stations. Asimple rou-- industry. However, diving sites in the area are tine of feeding by divers on a regular basis at fixed limited. The divers often travel a total of 3 to 5 points can rapidly establish a dense population of hours by road and ferry from Manila to Puerto large fish which can be easily approached and Galera, and risk becoming stranded when sea photographed. With this and other enhancements conditions become hazardous. Here again, diving such as underwater trail markers, the park can sites are limited and degrading because of poor become a major attraction for tourists in Bolinao. SHORESIDE PROTECTION There is strong scientific evidence that establishing a marine reserve will provide better har- The establishment of mangrove forests along vests from the Bolinao reef system. This the shore will restrict visibility and make information can be simplified and disseminated shoresidz protection difficult. A substantial secu- widely through media such as pamphlets and rity fence would be necessary along the shorevvttrd comic books, school presentations, meetings, pub- limit of the reserve. The reserve should extend lic hearings and so forth. However, the material several hundred meters onto land in order to pro- must be presented in appropriate ways. Each per- tect turtle-nesting beaches and shoreside bird, son involved will have to be in a position to use the plant and other biota. Therefore, the fence would evidence to convince herself or himself that pre- be mostly on dry ground, which will facilitate viously held concepts are wrong; e.g., that larger maintenance. fishing grounds and more fishing effort yield more fish. The decision to set aside some areas to in- MARINE MARKERS crease yields in others will be a difficult one, involving cognitive dissonance (Festinger 19721, The boundaries of the reserve and park could i.e., a challenge to existing value systems. Most be marked on the reef flat by permanent struc- people tend to avoid cognitive dissonance, particu- tures with warning signs every 100 m or so. In larly when the avoidance is reinforced by short- deeper waters, permanent buoys may be neces- term rewards such as the immediate benefits of sary, requiring more frequent maintenance. It is daily harvest activities in the proposed reserve essential that everyone entering the reserve know area. Changes in value systems can often be ef- clearly that he or she has done so, and that no fected through societal interactions which direct anchoring or harvesting be allowed, except for peer pressure toward convincing individuals to scientific purposes as authorized by carefully con- realize the need to change them (Asch 1972).Care- trolled permits. fully guided discussion groups can be effective in this manner (Fever 1989; Ortigas 1991). Follow- up information campaigns and public activities PUBLIC AWARENESS can be equally important, as it is necessary to reinforce changes in value systems in order to Management tends to be most effective when stabilize them (Cabanban and White 1981). The violations of regulations are not only illegal, but resewelpark system must incorporate a continual socially unacceptable as well (McManus et al. information dissemination effort to remind the 1988). Social unacceptability of an action arises public of the need to maintain the system. It must most easily when it is clear to every member of a inform them of the benefits attributable to the society that the action is detrimental to the mem- reservelpark as data become available on such bership as a whole. This clarity is often achieved matters as increased harvests or job opportuni- when the membership itself shares in the respon- ties. It is important to avoid ningas kogon (liter- sibility for imposing and arranging for the enforce- ally, grass fire), or the tendency to act with ment of a regulation. This procedure bypasses the enthusiasm in the short term, but lose interest common tendency of local groups to increasingly over time. In order to be effective, the plan for the mistrust the motives of progressively higher reservelpark system must be thoroughly inte- authoritative bodies over which they exert dimin- grated into the long-term planning and govern- ishing levels of control. ance of the municipality. CHAPTER 7 RECOMMENDATIONS FOR MANAGEMENT ACTION

Overview pressor can represent a liability to a resort opera- tor lacking expertise in diving. However, there are The management of the coral reef resources of several diving instructors in Manila who earn Bolinao can be improved with the following spe- salaries on an unpredictable basis, who might be cific actions: attracted to more stable job opportunities associ- 1. Establish a tourism regulatory commit- ated with resort operations. An interim solution to tee. the problem of eliminating blast fishing and at- 2. Develop alternative livelihoods. tracting fish would be the creation of a marine 3. Promote mariculture and improved agri- reserve and park (see Chapter 6). This would culture activities. provide a safe area for divers, and could serve as 4. Establish marine reserves. a focal point for attracting tourists to Bolinao. 5. Eradicate blast and cyanide fishing. Once Bolinao gains a reputation as a safe, 6. Ban compressor (hookah) diving. attractive diving area, one can expect a rapid 7. Improve fish-handling facilities. period of increased tourism, as was seen in the early 1980s in Anilao, Batangas and Puerto Galera, Mindoro. However, both of these areas Regulating tourism suffered from a lack of regulation in the development of the industry, particularly with respect to The Bolinao area has a very high potential for the prasemation of the marine and shoreside en- tourism development. The area includes nearly vironments. For example, the many isolated, re- 200 km2 of coral reef, 17 km of sandy beaches, mote beaches in Puerto Galera adjacent to small large sheltered harbor areas, several underwater patches of coral were rapidly crowded with dense, shipwrecks, two scenic lighthouses, an early 17th unsanitary living and eating facilities. The corals century church, and numerous caves and water- were substantially damaged through associated falls (Fig. 7.1). The town is approximately 5 hours siltation, gathering and breakage from boat an- of driving time from Manila, roughly the same chors. This type of difficulty arises because of the amount of time necessary to reach popular tourist tendency of many investors to favor quick profits sites to the south such as Puerto Galera. from short-tern investments in the face ofunregu- Some major factors which currently limit div- lated competition. ing activities include the lack of an air compressor, A much different problem has arisen with the abundance of blast fishing, and the scarcity of some tourist developments in Bohol, Cebu and fish. These constraints can be eliminated through elsewhere. Large areas which were previously a proper investment and management. An air com- source of livelihood to economically disadvantaged Fig. 7.1 Tourist attractions in the Bolinao area. Tourism could be greatly enhanced with the provision of a scuba compressor, establishment of a marine park, and elimination of blast and cyanids fishing. people have been purchased for expensive resort which would ensure the success of such an invest- hotel operations, sometimes with the aid of politi- ment include access to sanitary, fresh food; the cal pressure. In many cases, profits are tightly certainty that the quarters are clean, screened restricted to the outside investors, who provide and vermin-free; and the provision of running monopolized transportation to the resort, and all water and clean toilet facilities. These conditions the boats, food and services needed by the tourists. are rarely met in a village nipa hut in the Bolinao The resorts tend to hire well-trained staff mem- area. However, collaborative investment among bers from Manila and elsewhere, and employment neighbors or relatives could produce indigenously of local labor is very minimal. In these cases, there designed cottages incorporating the necessary lev- is very little benefit to the local populace. For this els of convenience and sanitation. Electricity may reason, it may be best to discourage development be helpful, but is by no means essential provided aimed at attracting highincome overseas tourists, that an adequate variety of fresh food is available and to concentrate instead on carefully controlled and kerosene or gas lamps and stoves are avail- developments aimed at attracting visitors from able. Some tourists prefer to "rough it", but are Manila. rarely willing to forfeit accustomed levels of sani- There is also a suitable target group of inter- tation to do so. The important point is that the national tourists, such as many of those already accommodations are not misrepresented when ad- visiting Bolinao regularly, who prefer economical vertised. A few cases of misrepresentation and tourist facilities. There are areas in the Caribbean poor quality control in the Bolinao area would do and elsewhere where small-scale tourist facilities considerable damage to the industry. It would be ("ecotourism") are highly successful (Boo 1990). desirable to form cooperatives for the purposes of For example, a coastal dweller who owns one quality control and advertising both nationally house for his or her family may build a second and internationally. These cooperatives could be house nearby for rental to tourists. Some factors regulated and assisted by the town government. In order to ensure the longevity of the local nearly any environmentally sound industry which resources and optimal benefit to the town and the provides higher salaries and more stable incomes majority of its people, it will be necessary to regu- than fishing is likely to have a positive effect on late development along the coast. This can be the resource ecology of the reefs. achieved through methods such as zoning, selec- The major natural resource of the municipal- tive licensing, provision of economic incentives, ity is its 200-km2 coral reef. This could be used and strict requirements for and evaluation of en- effectively to build a viable tourism industry, as vironmental impact assessments (EIAs) for all discussed above, especially if a marine park were proposed construction. This type of planning and to be implemented (Chapter 6). control could be vested in a small committee ern- Aside from tourism based on living reefs, Boli- powered appropriately by the local government. It nao has a potential for limestone production, would be important that the committee ade- based on fossil reefs. At the time of this writing, quately represent the views of both local mer- there is an ongoing survey which may lead to the chants and economically disadvantaged fishers development of an open pit mine immediately who must profit from the development in order to south of the Bolinao Marine Station. The proposed justify it. One or more scientists from the Bolinao mine, which will be financed principally by inves- Marine Laboratory could be involved to help en- tors from Zgiwan, would cover an area of several sure that natural resources are enhanced rather tens of square kilometers. A thorough study would than degraded by proposed activities, be necessary to ensure that any silt which leaks An appropriately directed tourism program from the operation areas will not remain in sus- would allow the present fishers to use their boats pension until it is carried over the reef slopes. to support recreational diving rather than for fish- Siltation can block the light needed by the algae ing. In Anilao, Batangas, small boat owners were living in coral tissues, thereby hindering the able to earn a gross income of P800lday when growth of the corals (Johannes 1975; Yap and diving tourism began to grow in the early 1980s. Gomez 1985). Silt which settles out of the water This was at a time when the peso was worth more column too quickly to be removed by the mucous than twice its current value in spending power. and polyp actions of the corals can kill the coral Boat owners in Bolinao today rarely earn that colonies (Alifio 1983). Losses in coral cover can much for rentals. The prices in Anilao fluctuated lead directly to loss in harvestable fishes and under the competing forces of local inflation, invertebrates. which was discouraging many potential tourists, Of equal concern is the effect of the mine on and price war declines, which threatened incomes. the land biota. The Bolinao area supports a rich The prices were eventually stabilized by a local plant and animal biota. The area under considera- boat owners association. tion supports a broad variety of birds and populations of the endangered monkey (Macaca). Many of the plants are valuable sources of natural me- Developing alternative livelihoods dicinal drugs. An adequate environmental impact study should involve surveys by knowledgeable Even if blast and cyanide fishing were to be botanists and zoologists before the project is ap- completely curtailed, the reef environment would proved. Additionally, the possibility that the town continue to be degraded because of anchor dam- subterranean water supplies might be adversely age, and the fish and invertebrate stocks would affected should be investigated by a competent continue to decline because of overexploitation. hydrologist. There are too many fishers in Bolinao. On the positive side, the mine would provide The families dependent on harvesting reef a few years of employment to many hundreds of organisms tend to live on marginal and unpre- workers. Should the mine be implemented, it dictable incomes. Fishing ranks the lowest in an- would be important that steps be taken to ensure nual incomes of the major occupations in Bolinao that local labor is employed wherever possible. (Fig. 2.3). The willingness of many to shift occupa- Otherwise, the mine will serve to draw immi- tions has been well illustrated by the fact that grants from other areas into the Bolinao munici- often more than 50 men abandon fishing to seek pality. This would exacerbate the current resource work whenever a new phase of construction is problems, especially after the mine closes down initiated at the Bolinao Marine Laboratory, Thus, again and ceases to be a source of employment. It is important to emphasize that the purpose and a demand might be generated at progressively of an EIAis not to hinder development, but rather larger scales to national or international levels. to enhance long-term, rational development. The Once established, the industry might move on- assessment provides access to several sides of the ward to mariculture activities and the production total development picture, so that optimal deci- of more refined products such as medicinal chemi- sions may be made. Without environmental as- cals. Asimilar industrial potential may exist in the sessments, the interests of a minority, usually an form of jellyfish, which abound locally and can be economically advantaged group, are facilitated at processed initially for sale to Chinese and Japa- the expense of the environment, which inevitably nese communities for food. Jellyfish such as adversely affects the economically disadvantaged. seawasps produce biochemicals associated with This would be particularly true in Bolinao, where stinging cells which may eventually prove to be of fishers and farmers, who depend directly on the considerable medicinal value (Walker 1988). maintenance of a healthy environment, constitute The current program of planting mangroves 80% of the human population (Fig. 2.1). throughout the Bolinao coastline could lead to a Another potential source of employment broad variety of cottage industries. The range of would be to expand various cottage industries. products available from mangroves is broad (able Currently, the most profitable part-time cottage 7.1) (Saenger et al. 1983; Salm and Clark 1984 ). industry is shellcraft (Fig. 2.3). This industry is However, an effective industry based on gathered probably operating near the limit of the available mangrove products would require organizational resource supply, and could not be extended further efforts, such that the products of individual collec- until total shell production is enhanced by such tors are amassed and delivered to appropriate means as the establishment of a marine reserve. processing facilities and markets. This would be The shellcraft industry has the desirable charac- especially true for wood products such as ship- teristic of maintaining local workers in producing building materials. A high demand for firewood a refined end product, In this way, the town bene- for the salt making industry in adjacent munici- fits optimally from a limited resource. If the prod- palities brings an immediate danger of overexploi- uct was to be exported in its raw state, much of the tation of the planted mangroves. This must be profit to be made would be lost to the town. The acted upon immediately through controls such as fact that end products are completed locally also restrictive regulation and licensing. Additionally, makes this industry complementary to the devel- the provision of a marine reserve incorporating opment of tourism. mangrove areas would provide for renewed popu- Another industry of high potential involves lations of mangrove species on a continual basis: seaweed gathering and processing. More than 15 Educational achievement is limited in the mu- km of reef slope in the southwestern portions of nicipality, where more than one-third receive no the municipality are highly dominated by Sargas- education (Fig,2.1). However, a substantial num- sum seaweed (aragan).This algae can be used for ber of people maintain skills useful in the devel- a broad variety of purposes ranging from feeding opment of small-scale industries (Table 7.2). Some cows to the production of medicines. The addition of these skills are passed on locally, while others of the seaweed to chicken feed can replace the are acquired during periods of employment in expensive beta carotene often added to enhance Manila or overseas, including training in the Phil- yolk production. However, it would be desirable to ippine or US. military, and work experience on initiate an industry requiring local processing to Saudi Arabian oil fields. produce a widely saleable product. One such in- Occupational mobility could be enhanced con- dustry would be liquid fertilizer. The seaweed can siderably by improving local schools to encourage be cooked and filtered to produce a concentrate. attendance. Most of the schools are greatly in need When mixed with water and sprayed on plants of repairs, new desks and chairs, books and sani- twice monthly, it can reduce dehydration and in- tary plumbing, Funds for such improvements sect damage, induce budding and fruiting, and could be solicited from various sources, including reduce the need for commercial fertilizers, espe- international sources targeting nongovernmental cially when cooked with a source of calcium such organizations (NGOs) and various civic groups in as ash from burned coconut fronds ( Dr. Nemesio developed countries. In some circumstances, a lo- Montaiio, pers. wmm.) This type of backyard in- cal parent-teacher organization might qualify to dustry would involve little capital investment, acquire the necessary funds. In others, it may be Table 7.1. Potential products from mangrove forests (Saenger et al. 1983; Salm and Clark 1984). Mangrove plant producb

Food, drugs and beverages Consltruction materials Fimhing equipment Sugar Timber, scaffolds Poles for fish traps Alcohol Heavy construction timber E'ishing floats Cooking oil Railroad ties Fuel for smoking fish Vinegar Mining pit props Tannins for net and line preservation Tea substitute Boatbuilding materials Wood for fish drying or smoking racks Fermented drinks Dock pilings Dessert topping Beams and poles for buildings Household itmms Condiments from bark Flooring mture Sweetmeats from propagules Paneling, clapboard Glue Vegetables from pmpagules, Thatch or matting Hairdressing oil fruits or leave^ Fence posts, water pipes, lbol handles Cigar substitute chipboards, glues Mortars and pestles 'Ibys Textiles and leather Fuel Matchstick8 Synthetic fibers (e.g., rayon) Firewood for cooking, heating Jncense Dye for cloth Cha~oal Tannins for leather preservation Alcohol Other products Packing boxes Agriculture Paper products Wood for smoking sheet rubber Fodder, green manure Paper of various kinds Wood for firing bricks Medicine fmm bark, leaves and fruits

Mangrove wildlife products

Fish Oysters Insects Birds Crabs . Mussels Honey Mammals Shrimp Shells Wax Reptiles and reptile skins

possible to create an appropriate NGO by compli- Many potential industries, such as clothes or shoe ance with the regulations of the Philippine Secu- manufacture, would be better supported if an or- rities and Exchange Commission (SEC). Afurther ganized effort was put into the development of step toward improving school enrollment would be shipping arrangements for raw materials and end to ensure that alternative livelihoods provide har- products. The excellent harbor behind Santiago vesters with adequate incomes to make it unnec- Island could facilitate this. A major renovation of essary for a family to employ its children to the portside facilities has recently been completed. acquire food or income. Currently, children repre- Some additional modifications may be necessary, sent a major workforce in the communities, and however, because the current dock is located in this works against the long-term improvement of water too shallow for any reasonable ocean-going local life-styles. vessel. It is likely that an effective alternative liveli- Amajor portion of the fishing population lives hood program would require the active develop- on Santiago Island. This island has electricity only ment of markets for existing or proposed products in the southeastern corner, and this is very'spo- (DAP 1978;Kotler and Armstrong 1989).Products radic because of the exposure of the lines to such as shellcraft creations sell widely not because weather as they cross wide channels to and from they are outstandingly useful, but because they Siapar Island, There are many areas which have have public appeal. Public opinion is often no fresh water during the dry months from Febru- strongly influenced by advertising. The success or ary to May. Many people bring in water from the failure of a cottage industry may depend less on mainland in small containers by boat. There is no the need for the product than on the effectiveness bridge connecting the island to the mainland. with which local producers are able to interact Thus, it would greatly improve the chances of with marketing agencies and companies with ad- success in a program of alternative livelihood de- vertising and outlet distribution capabilities. velopment if a bridge could be constructed to the Table 7.2. Skilled labor available in the Bolinao municipality. A variety of human resources could be tapped for small-scale industry. Data are from a survey by DAR in 1991.

Type of skill Skilled Barangays individuals involved

Factory/industrial Shellcraft Construction Ropemaking Charcoalmaking Matweaving Copra Bamboocraft Drivers Fig. 7.2. Selected pmducta from a wellmanaged mangrove forest.

There are many mariculture techniques which cause minimal disruption of natural ecosys- island, bringing water pipes, electrical lines, and tems. Many of these involve very little investment, ready transportation to and from the mainland. and are suitable for implementation by villagers. A meaningful effort in developing alternative Examples include maintaining pens for crabs, lob- livelihoods must involve a strong effort in market sters, conchs, sea urchins, sea cucumbers and analysis in Manila and overseas. It will also be giant clams; stick culture of oysters and mussels; necessary to invest in efforts to advertise existing rack culture of seaweeds; and cage culture of products and to attract investors for others. A fishes. It is conceivable that most ofthe gastropods number of bilateral aid agencies could be tapped and bivalves involved in the shellcraft industry for funds to assist in these areas, particularly with could be maintained in some form of controlled the current emphasis on supporting privatization enclosures and fed for optimal ~owth. (e.g., Australia) and NGOs (e.g., United States). Potentially, at least 30 km of reef flat and a few square kilometers of protected harbor waters could be used in one way or another for maricul- Mariculture and agriculture ture. Potential problems from mariculture include disruption of local currents and localized pollution, especially where feeding is necessary (e.g., Marine and brackishwater aquaculture which grouper culture). However, these problems could involves the destruction of productive marine be monitored as the industry grows and develop- habitats, such as mangrove forests and estuaries, ment altered as necessary. A major benefit from are referred to as "destructive mariculture activi- having efforts directed toward mariculture by a ties". For example, nearly all of the formerly ex- large segment of the population is that strong tensive mangrove forests in the Bolinao area have incentives will be developed to maintain a healthy been displaced by ponds for growing milkfish environment and to prevent disruptions from (bangus) and prawns. This has severely reduced blast and cyanide fishing. A further benefit of the availability of a myriad of plant and animal considerable consequence is that more food will products which would otherwise be available for become available, leading to reduced health prob- harvest by local villagers (able 7.2, Fig. 7.2). lems and absorbing some of the demand for reef Instead, the profits from the enclosed areas now fish. go directly to large-scale pond owners with very Amajor research thrust of the Bolinao Marine little diversion to laborers such as guards and Laboratory of the Marine Science Institute of the occasional maintenance people, University of the Philippines is in the area of small-scale mariculture. Some potential maricul- food crops for consumption by the family, and then ture organisms which have been investigated in- to sell the excess production. Intensive multispe- clude giant clams (n-idacna spp., Hippopus spp., cies gardens can be designed in such a way that taklobo), sea urchins (5"Fipneustesgratilla,kuckn- they require decreasing levels of maintenance kuden), sea scallops (Amusium pleuronectes, over time -- a major goal of the internationally kapis), abalone (Haliotu8 asina), lobsters (Pa- growing practice of permaculture (Mollison and nulirus spp., kising-kising), rabbitfish (Siganus Slay 1991). Fertilizer costs can be eliminated fuscescens, bamngen) and seaweed (especially through the use of mulch and seaweed products. Eucheuma aluemsi, tomsao). Currently, programs Appropriate technologies and crop choices can are underway to encourage local small-scale eliminate the need for expensive fertilizers. Acon- mariculture of giant clams and Eucheuma centration on perennial rather than annual crops through training programs and the provision of can lead to reduced maintenance efforts and a spat or propagules. Similar programs for other constant supply of a variety of food products. The species are expected to follow, which could help street market system of Bolinao consists of small considerably in efforts to promote local maricul- stalls selling overlapping varieties of crops. This ture. system lends itself well to the sale of small quan- As with other forms of alternative livelihood, tities of various fruits and vegetables produced in success in mariculture depends on the provision of small family plots. an adequate market and product transportation. Complementary agricultural approaches in- The latter is particularly complex in some cases, clude the small-scale crop-livestock-aquaculture such as the supply of live groupers to restaurants techniques developed at ICLARM and elsewhere in Manila. In other cases, an emphasis on local for use in tropical areas (Edwards et al. 1988). processing, such as canning, could help to reduce Possibilities include integrated rice-fish, live- postharvest losses and internalize economic re- stocklpoultry-fish, vegetable-fish, and all combi- turns within the municipality. The development of nations of these (Pullin 1989). As with markets and advertising campaigns would re- permaculture, the general goal is to minimize quire funding and could be facilitated through the investments and waste by producing groups of formation of cooperatives and assistance and ac- complementary products. These systems could tion by the local government. Additionally, educa- provide for better nutrition and incomes from ag- tional programs would be necessary to encourage ricultural lands, and reduce the pressure to ex- fishers to become mariculturists and to broaden ploit the marine environment. the diets of the local populace to absorb the new products and promote better health. A further step toward alleviating harvest pressure from the marine environment would be Establishing marine reserves an intensive program to improve the use of local agricultural lands. For example, many hectares of Marine reserves could potentially improve the land are currently devoted to growing maguey, a local fisheries and provide for a continually high plant used to provide fibers for constructing inex- diversity of hawestable species. A sample plan for pensive ropes. The market for this fiber has been a marine reserve centered on Malilnap Channel poor recently, but local farmers have been slow to has been outlined in Chapter 6. Once this system refocus on more profitable crops. has demonstrated its merits, it may be useful to In economic settings such as Bolinao, it is a establish others. One excellent site which has questionable practice for a low-income family to been proposed elsewhere (McManus 1989b) would devote available lands to producing single crops, be Cangaluyan Island, an area which would sup- such as rice, maguey or coconuts. A substantial port reefs in both the Bolinao and Anda munici- proportion of the money gained from such produc- palities. The island falls under the latter's tion goes toward buying other foods necessary to jurisdiction. Other potential sites include a4 area sustain the family. The sporadic and risky nature along the reef crest northwest of Lucero, an off- of the incomes leads to periods when purchases of shore reef several kilometers east of Silaki, and fruits and vegetables are minimized, leading to selected areas both east and west of the Balin- malnutrition. In many cases, it would be better for gasay River. However, as considerable educational the family to concentrate on growing a variety of and political effort is required for each reserve area, it may be desirable to finish establishing the The eradication of these destructive fishing primary reserve at the Malilnap Channel prior to methods must involve both public education and undertaking new programs. This area is ecologi- publicly acceptable forms of enforcement. Areduc- cally and oceanographically the most suitable of tion in blasting by 90% during the study period is the potential sites in the Bolinao municipality. largely attributable to fears generated by the ru- mor that five people involved in blast fishing or transporting blasted fish were summarily executed by unknown parties. This form of extreme Blast and cyanide fishing enforcement is not likely to endear the people to promoters of resource management programs. A much prefered approach would be one of commu- Blast and cyanide fishing are both nonselec- nity organization and public information involv- tive, environmentally damaging fishing methods. ing publications (in the Bolinao and Ilocano The explosions and poisons kill all life history dialects), village meetings and school assemblies stages of the target species and most other organ- (Cabanban and White 1981; Ferrer 1989, 1991). isms nearby. The corals, which form the basis for Blasting is not commonly viewed with the level of the ecological habitats of the species, are also seriousness necessary to prevent its open use in destroyed (Thlbot and Goldman 1972; Carpenter the villages (Galvez 19891, and efforts must be et al. 1981; McManus et al. 1981; Naiiola et al. directed toward making it not only illegal, but 1990). Corals are very slow at recolonization and socially unacceptable as well (McManus et al. growth, and complete recovery may take several 1988). decades (Johannes 1975; Yap and Gomez 1985). The living coral cover in the reef flat and lagoonal areas has been reduced by 60% because of these Banning compressor diving fishing methods. The methods compete directly with the use of more desirable gear such as gillnets, traps, hook and line, and spearfishing. A growing number of fishers use air compres- Ultimately, however, blast and cyanide fishing sors with long hoses to facilitate underwater har- should be completely eradicated because of their vesting. The most prominent uses are for effects on tourism. The tourist industry holds the spearfishing, lobster gathering, aquarium fish greatest promise for providing alternative em- catching, and recently, for sea urchin gathering. ployment and removing harvesters from the reef. The air compressors are of the type commonly Theoretically, a frequency of about one blast per used for filling tires at gasoline stations. week might have very little direct ecological effect International scuba diving norms dictate that on the reef system as a whole. However, tourism a compressor should involve an air intake extend- is built on reputation and expectation. If a diver ing several meters upwind of the compressor and from Manila hears a single blast during his lim- a series of filters to remove particles from the air. ited stay in Bolinao, the chances are great that These precautions are necessary because concen- news of the event will spread throughout the div- trations of gases such as carbon monoxide and ing clubs of Manila within a few weeks. A similar carbon dioxide which have negligible effects at sea effect would arise from a tourist diver encounter- level, can become fatal if inhaled under pressure ing a fisher squirting cyanide underwater to catch during a dive. Both gases are produced by the aquarium fish. 'Iburists are not usually concerned compressor itself, as well as by boat engines and with the statistical adequacy of the sampling of an tobacco smoke. They cannot be filtered out of the event. Rather, they tend to react to signs that air under most conditions, and must be carefully previously held beliefs are valid. It has become avoided. Oils which enter the compressor can common knowledge that some divers in the last cause lipo-pneumonia as they accumulate in the few years have been seriously injured and killed lungs. by blast fishing in the Philippines. Fear of the International standards dictate that diver as- danger of being injured or killed by blast fishing cents and descents must be carefully regulated to or by ingesting poisoned water is prevalent. Thus, avoid ear and sinus damage, and similar injuries. a few unfavorable anecdotes could seriously dam- Nitrogen narcosis often leads to diving accidents, age the diving tourist industry in Bolinao. particularly in waters below 30 m, because the euphoric feelings it induces cause judgement to be considered for implementation at a national scale altered. The use of medical drugs, alcohol or to- as well. bacco smoke in the 12 hours prior to a dive can lead to difficulties during the dive. Medical conditions must be checked frequently to avoid heart attacks and other heavy work or pressure-related Improving fish-handlingfacilities injuries. Underwater times, depths and rates of ascents must be strictly limited to prevent decom- A substantial portion of the catch in Bolinao is pression sickness, which often leads to paralysis lost to spoilage. Conditions in the fish landing or death. Frequency of diving is limited to avoid sites are unsanitary, and a significant public degradation of the bone marrow. Training is par- health risk exists. This is a common situation ticularly concentrated on reducing the likelihood throughout the Philippines (Santos 1988). of a diver holding his or her breath during emer- The fishers on the reef generally fish approxi- gency ascents, which often leads to lung bursts, mately 6 hours each day. Those involved in spear- producing emphysema and air embolisms, the lat- fishing and hook-and-line fishing often carry ter of which is a frequent cause of death. Under- boxes of ice, especially during the day. However, water asthma attacks are yet another cause of air upon arrival at the landing site, the fish are often embolisms and death. laid on top of the ice rather than properly buried The compressor divers of Bolinao often dive and interspersed with the ice. Ice is ground or in depths of 30 to 60 rn for a few hours at a time. chopped under unsanitary conditions. Most fish- They are susceptible to all of the above-mentioned ers who use other gear do not carry ice. diving hazards. The most commonly known prob- In the main fish market in Bolinao, many fish lem is death or paralysis from decompression sick- are spread on table tops with no ice, and are ness, which is locally called kuriente. The local exposed to flies. Those in boxes or washtubs of ice name refers to the fact that the divers associate it with electrocution, and often believe that it is caused directly by temperature changes in the Time underwater (hours) water. Actually, it is common because the divers 0 1 2 3 routinely exceed the so-called "no decompression" limits used by knowledgeable divers (Fig. 7.3). The air from the compressors is laden with oils and dangerous gases. The lack of a regulator at the diver's end of the air hose invites problems ofpanic and associated air embolism. There is no ~racticalway for Bolinao fishers to be properly trained and equipped for commercial diving. The only feasible means of avoiding the overwhelming number of safety and health hazards associated with compressor diving is to ban it entirely. Banning compressor diving would have a beneficial effect on fisheries ecology in the Bolinao area. There is currently a rapid decline in the number and diversity of fishes reaching adult sizes on the reef slope. This decrease includes a 50% drop in species richness and an 80% drop in Compressor divers abundance in three years. A ban on compressor diving would serve to curtail this decline and help safeguard the breeding populations which help to supply the coastal reefs along Western Luzon with annual recruits. Thus, a ban on compressor diving Fig. 7.3. Mnny compressor (hookah) divem dive tuo deep for would be beneficial for both resource management too long und ilcquirc dccomprcssion sickness and other mala- and humanitarian reasons. Such a ban should be dies. IJarillysis and death am common. are not usually interspersed. The floors are invari- forms a natural limit on the number of people who ably slippery with material dripping from the can be employed in farming. With the current tables. Fish are handled without protection and no scarcity of alternative employment, most of the sanitary facilities are available. There is a faucet incoming labor is expected to seek employment in with running water, but no hose is available for fishing. There are already roughly twice as many washing the floors. Fish which are purchased for fishers on the reef as the system can sustain in the Manila are usually shipped in ice trucks. Local long term. Doubling this again will cause a very fish processing is limited and generally involves rapid decline of the major resource of the munici- open air drying andlor salting. pality. Those already dependent on the reef will be The fish market should be reconstructed to left with diminishing catches and incomes as more include raised water taps and sinks, and properly competitors join the fishery work force. A great drained floors. Tables should be designed to facili- deal of conflict and difficulty is expected to result tate holding trays of ice to presellre fish on display. in the next few decades. A single full-time employee could periodically Conceivably, the Philippines could enter into clean the floors, provide soap for the sinks, and a period of rapid economic growth and jobs could generally maintain sanitary conditions. Inexpen- hecome available in cities which will draw people sive ice should be made available and its use out of Bolinao. However, the growth rate in Boli- required in the market. It may also be possible to nao is matched by an equally high rate throughout provide disposable plastic gloves and bags with the country. A disproportionately large amount of heat sealers to minimize spoilage and health haz- the incoming national labor force is expected to ards upon purchase. These changes might entail migrate to cities. Therefore, it is unlikely that raising the current fee for market usage by a small evem a very high rate of economic growth nation- amount to finance the maintenance of the facili- ally will result in enoughjob opportunities in cities ties. However, funding for the initial construction to alleviate the population problem in Bolinao. could be sought from bilateral aid agencies. In addition to creating alternative sources of Ice may become more readily and inexpen- employment locally and restricting the entry of sively available in the future because of the recent laborers to indigenous people, steps can be taken construction of a new ice plant. An information to encourage birth control. The national program campaign about the use of ice and the need to of family planning has had little impact locally. maintain sanitary conditions in the market could Occasional attempts at developing educational be implemented, including posters and school programs and distributing birth control devices presentations. Training in sanitary fish-handling have been short-lived and on too small a scale to methods could be requested from the Bureau of substantially change traditional social values. Fisheries and Aquatic Resources, the Home Eco- The average resident still depends on having a nomics Department of the University of the Phil- large number of grateful children as a way of ippines, or the University of the Philippines in the ensuring a source of income in retirement. It is felt Visayas College of Fisheries, all of which maintain that it is far more fruitful to invest in progeny than appropriate specialty staff members. A local in- in savings accounts and other economic invest- vestment in fish processing, such as canning, ments. A strong educational campaign would be might help internalize economic returns from the necessary to convince young couples that invest- resource within Bolinao. ing more in fewer children and in personal economic growth is a rational strategy for success in later years. Other programs aimed at avoiding Reducing human population teenage pregnancies would be helpful as well. It is widely believed that the birth rate will growth rates decline as the local economy grows. This could very well be the case. Unfortunately, the popula- The human population of Bolinao is rising at tion growth rate is being matched by a rapid an accelerating rate (Fig. 2.2). The current popu- decline in available resources. An active program lation is approximately 50,000, of whom 31% are of alternative livelihood generation and the estab- involved in fishery-related employment, and 49% lishment of marine reserves and parks could con- in farming. The population is expected to reach ceivably slow the decline in resources. However, it 100,000 in 30 years. Farm land is limited, and is not likely that such programs would compensate 76 for the accelerating population growth rate. Even increase the likelihood that average personal in- with a general strengthening of the local economy, comes would rise, and thus that population it is unlikely that the average life-style will change growth rates might diminish more passively in the significantly under current circumstances. An ac- future. tive program of encouraging birth control would REFERENCES

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Jackknifing an index of diversity. Ecology 58907- Smith, I.R. 1979. A research framework for traditional fisheries. 91 3. ICLARM Stud. Rev. 2.45 p. APPENDICES: FISH SPECIES ABUNDANCES Appendix 1. Combined list of all species sorted alphabetically. Abundances are in indJ1,OOO ma. Weights are in g/1,000 mx. Most identifications were based on Allen 1975;Rau andRau 1980; Schroeder 1980;Masuda et al. 1984; Randall et al. 1990; and Myers 1991. (* denotes uncertain identification).

By frequency By weight

Rank IndJtraneect No. Species Family Slope Flat Trawl Slope Flat Trawl Rank Trawl

1 Abudefduf coelestinus Pomacentridae 2 *~bu&fdufleucotonus Pomacentridae 3 Abudefduf smtilis Pomacentridae 4 ilbudefduf septemfasciatus Pomacentridae K Acanthurid Acanthuridae 6 Acanthurid sp.1 Acanthuridae 7 Acanthurid sp.6 Acanthuridae 8 Acanthurid sp.6 Acanthuridae 9 Acanthurid sp.9 Acanthuridae 10 Acanthurus bariene Acanthuridae 11 Acanthurur dussumieri Acanthuridae 12 Acanthurus gahhm Acanthuridae 13 *Acanthurus glaucopareius Acanthuridae 14 *Acanthurusjaponicus Acanthuridae 15 Acanthurus lineatus Acanthuridae 16 Acanthurus rnata Acanthuridae 17 Acanthurus nigrofiscus Acanthuridae 18 Acanthurus olioaceus Acanthuridae 19 Acanthurus pyrofems Acanthuridae 20 Acanthurus sp.1 Acanthuridae 21 Acanthurus triostegus Acanthuridae 22 *Acanthurus xanthopterus Acanthuridae 23 Acreichthys tomentosus Monacanthidae 24 Aediscus strigatus Centriscidae 25 Aesopia cornuta Soleidae 26 Aluteres scriptus Monacanthidae 27 Amblyapistus taenianotus Congiopodidae 28 *Arnblyeleotrisfirsciatu Gobiidae 29 *Amblyeleotrisjaponica Gobiidae Amblyglyphidodon uureus Pomacentridae Amblyglyphidodon curacao Pomacentridae Amblyglyphidodon leucogaster Pomacentridae Amblygobius albirnaculatus Gobiidae Amblygobius phalaena Gobiidae Amblygobius sp. Gobiidae Amphiprion clarkii Pomacentridae Amphiprion frenatus Pomacentridae Arnphiprion ocelluris Pomacentridae Amphiprion perideraion Pomacentridae Arnphiprion sandarocinos Pomacentridae Ammpses coeru~eopunctatus Labridae Anampses geographicus Labridae Anampses meleagrides Labridae Anampses twistii Labridae Antennarius moluccemis Antennariidae Antennarius nummifer Antennariidae Antennnrius sp.1 Antennariidae Anthias sp. Serranidae Apogon amboinensis Apogonidae Apogon bandanensis Apogonidae Apogon coccineus Apogonidae Apogon compresstls Apogonidae Apogon cyamsomu Apogonidae

Continued 83 Appendix 1 (Continued)

By frequency By weight

Rank IndJtransect No. Species Family Slope Flat Trawl Slope Flat Trawl Rank Trawl

54 Apogon nwemfasciatus Apogonidae 191 3"8 35 0.05 2.38 0.94 55 Apogon sangiensis Apogonidae 3 - - 25.89 56 &won sp. Apogonidae 29 35 123 1.12 2.83 0.03 57 Apo@n sp.1 (Schroeder 1980) Apogonidae - 214 - 0.03 - 58 Apogon sp.5 (Schroeder 1980) Apogonidae 30 18 37 1.04 8.01 0.91 59 Apogon sp.8 (Schroeder 1980) Apogonidae - 121 - 0.03 60 Apogonid Apogonidae 15 79 151 1.97 0.67 0.01 61 Apogonid sp.10 Apogonidae - 127 - 0.03 62 Apogonid q.11 Apogonidae - 106 - 0.04 63 Apogonid q.2 Apogonidm - 170 63 - 0.06 0.33 64 Apogonid sp.3 Apogonidae - 221 - 0.02 - 65 Apogonid sp.4 Apogonidae - 198 - 0.03 - 66 Apogonid sp.5 Apogonidae 232 21 12 0.03 7.74 4.69 67 Apogonid 5p.6 Apogonidae - 238 * 0.02 * 68 Apggonid sp.7 Apogonidae 89 - 0.07 69 Apogonid sp.8 Apogonidae - 235 * 0.02 - 70 Archamia lineolata Apogonidae 68 - 0.17 7 1 Ariosoma anagoides Colocongridae 56 - 0.30 72 Arothron hispidus Tetraodontidae 373 283 44 cO.01 0.01 0.67 73 Arothron immaculatus Tetraodontidae - 147 16 - 0.09 2.93 74 Arothron mappa Tetraodontidae - 161 - 0.01 75 Arothron nigropunctatus Tetraodontidae 123 140 125 0.17 0.10 0.03 76 Arothron sp. Tetraodontidae - 287 * 0.01 - 77 Arothron sp.2 Tetraodontidae * 163 - 0.01 78 Arothron stellatua Tetraodontidae 224 197 133 0.03 0.03 0.03 79 Aspidontus taeniatus Blenniidae 302 249 - 0.01 0.01 - 80 *Astermpteryx semipunctatus Gobiidae 313 - 54 0.01 - 0.33 81 Atherinid Atherinidae 68 5 - 0.42 16.98 - 82 Atule mate Carangidae 356 - - c0.01 - 83 Aulostornus chinensis Aulostomidae 249 179 108 0.02 0.05 0.04 84 Balistapus undulatus Balistidae 103 - - 0121 - 85 Balistid Balistidae 246 - 141 0.02 - 0.02 86 Balistid sp.1 Balistidae 190 - - 0.05 - 87 Balistid sp.4 Balistidae 362 - - <0.01 - 88 Balistid sp.6 Balistidae 276 - - 0.01 - 89 Blenny Blenniidae 146 236 - 0.11 0.02 - 90 Blenny sp.2 Blenniidae 362 - - <0.01 - 91 Blenny sp.7 Blenniidae 309 - - 0.01 - 92 Bodianus axillaris Labridae 287 - - 0.01 - 93 Bodianus bilunulatus Labridae 264 - - 0.02 - 94 Bodianus hirsutus Labridae 239 - - 0.02 - 95 Bodianus mesothorax Labridae 139 144 - 0.14 0.10 - 96 Bodianus sp. Labridae 289 - - 0.01 - 97 Bolbomelopon bicolor Scaridae 189 189 - 0.05 0.04 - 98 Bothus pantherinus Bothidae - 160 - 0.01 99 Caesio caerulaurea Lutjanidae 64 - - 0.45 - 100 Caesio etythrogaster Lutjanidae 82 201 177 0.33 0.03 0.01 101 Cuesio sp. Lutjanidae 297 - - 0.01 - 102 Caesio tile Lutjanidae 129 84 - 0.15 0.49 - 103 Calloplesiops dtivelis Plesiopidae 345 - - 0.01 - 104 Motomus carolinus Scaridae 254 - - 0.02 - 105 Calotomus japnicus Scaridae 32 27 42 1.02 5.14 0.59 106 Cdotornus sp. Scaridae 179 - - 0.05 - 107 Cantherhines dumerilii Monacanthidae 359 - * ~0.01 - 108 Cantherhines pardnlis Monacanthidae 157 233 - 0.09 0.02 - Continued Appendix 1(Continued)

By frequency By weight

Rank IndJtransect No. Species Family Slope Flat Trawl Slope Flat Trawl Rank Trawl

109 Canthigmter bennetti Tetraodontidae 110 Canthigaster compressa Tetraodontidae 111 Canthigaster coronata Tetraodontidae 112 Canthigaster janthinopters Tetraodontidae 113 * Canthigaster solundri Tetraodontidae Canthigaster valentini Tetraodontidae Carangid Carangidae Carangoides fulooguttattrs Carangidae Caranx rnelampygus Carangidae Centrogenys oaigiensis Percichthyidae Centropyge bicolor Pomacanthidae Centropyge bispimsus Pomacanthidae Centropyge herddi Pomacanthidae Centropyge tibicen Pomacanthidae Centropyge urolicki Pomacanthidae Cephalophlis argus Serranidae 1% * ~e~halo~holi8boenrrck Serranidae 126 Cephalopholis mi~iuta Serranidae 127 Cephdophdis puchycentron Serranidae 128 Cephalopholis sp. Serranidae 129 Cephalopholis urodela Serranidae 130 * Chaetodon adiergastos Chaetodontidae 131 Chaetodon auriga Chaetodontidae 132 Chaetodon baronessa Chaetodontidae 133 Chaelodon bennetti Chaetodontidae 134 Chaetodon citrinellus Chaetodontidae 135 Chaetodon ephippium Chaetodontidae 136 Chaetodon kleinii Chaetodontidae 137 Chaelodon lineolatus Chaetodontidae 138 Chaelodon lunula Chaetodontidae 139 Chaetodon melannotus Chaetodontidae 140 Chaetodon mertensii Chaetodontidae 141 Chaetodon octofasciatus Chaetodontidae 142 Chaetodon ornatissimus Chaetodontidae 143 Choetodon punctatofasciatus Chaetodontidae 144 Chaelodon raflesi Chaetodontidae 145 Chaelodon sp. Chaetodontidae 146 Chaetodon trifascialis Chaetodontidae 147 Chwlodon trifasciatus Chaetodontidae 148 Chaetodon ulietensis Chaetodontidae 149 Chaetodon unirnaculatus Chaetodontidae 150 Chaetodon uagabundus Chaetodontidae 151 Chaetodon xanthurus Chaetodontidae 152 Cheilinus bimaculatus Labridae 153 * Cheilinus celebicus Labridae 154 Cheilinus diagrammus Labridae 155 Cheilinus fusciatus Labridae 156 Cheilinus rhodochrous Labridae 157 Cheilinus sp. Labridae 1BS Cheilinus trilobatus Labridae 169 Cheilinus undulatus Labridae 160 Cheilio imrmis Labridae 161 Cheilodipterus macrodon Apogonidae 162 Cheilodipterus quinquelineatus 'Apogonidae 163 Chelonodon pa&a Tetraodontidae 0.13 33 6.91 Continued Appendix 1 (Continued)

.- By hquency BY weight

Rank IndJtransect No. Species Family Slope Flat Trawl Slope Flat Trawl Rank Trawl

164 Choerodon anchorago Labridae 125 45 27 0.16 1.77 1.34 23 165 * Chowodon shoenlei&i Labridae 166 Chmmis caerulea Pomacentridae 167 Chromis lepidolepis Pomacentridae 168 Chromis rnargaritifer Pomacentridae 169 Chmmis sp. Pomacentridae 170 Chromis weberi Pomacentridae 171 Chmrnis xuulthra Pomacentridae 172 Chrysiptera leucopma Pomacentridae 173 Cirrhilabrus cyanopleum Labndae 174 * Cirrhilabrus polyzona Labridae 175 Cirrhitichthys aprinus Cirrhitidae 176 Cirrhitichthys falco Cirrhitidae 177 Cirrhitichthys serratus Cirrhitidae 178 Cirrhitops hubbwdi Cirrhitidae 179 Cirripectes polyzona Blenniidae 180 * Cirripectes varidosus Blenniidae 181 Clupeid Clupeidae 182 Conger cinereus Congridae 183 Conger sp. Congridae 184 Coris aygula Labridae 185 Coris dorsumacula Labridae 186 Coris gaimadi Labridae 187 Coris uariegatu Labridae 188 Corythoichthys haematopterus Syngnathidae 189 * Corythoichthys schultzi Syngnathidae 190 Ctenochmtus binotatus Acanthuridae 191 Ctenochaett~sstriatus Acanthuridae 192 Dampieria eyelophthdma P~eudochromidae 193 Dampieria sp. Pseudochromidae 194 hcyllus aruanus Pomacentridae 195 hcyllus mdanurus Pomacentridae 196 DascyUus reticulatus Fomacentridae 197 Dascyllus trimaculatus Pomacentridae 198 Decaptems sp. Carangidae 199 &ndmchirus zebra Scorpaenidae 200 Diodon hystrix Diodontidae 201 Diploprion biwciatus Grammistidae 202 Dischislodus chrysopoecilus Pomacentridae 203 Dischistcdus notopdhalrnus Pomacentridae 204 Dischistodug perspicillatus Pomacentridae 205 Dischist& prosopotmnia Pomacentridae 206 * Dischist& pseudoclrrysopoeciltrs Pomacentridae 207 Drepane longimam Ephippidae 208 Dumker~c~tnpusdactylbphorus Syngnathidae 209 Echidna nebulma Muraenidae 210 * Eleotris fusw Gobiidae 2 11 Encheiliophis uerrnicuturis Carapidae 212 Engradid Engraulididae 215 Epibulus insidiator Labridae 214 Epinephelus fmiatus Serranidae Serranidae Serranidae Serranidae Serranidas Continued Appendix 1 (Continued)

By frequency By weight

Rank Indltransect No. Species Family Slope Flat Trawl Slope Flat Trawl Rank Trawl

219 Epinephelus megachir Serranidae 305 220 Eaineahelua merra Serranidae Serranidae 222 Epinephelus sexfasciatus Serranidae 223 Epinephelus sp. Serranidae 224 Epinephelus tauvina Serranidae 225 Escudosa thmacata Clupeidae 226 Eupornacentrus lividus Pomacentridae 227 Eupornacentrus nigricam Pomacentridae 228 Exallias breuis Blenniidae 229 Exyrias bellissirnus Gobiidae 230 Exyrias puntang Gobiidae 231 Fistularia petirnba Fistulariidae 232 Forcipiger fluvissimus Chaetodontidae 233 Fowleria variegata Apogonidae 234 * Gerres ocinaces Gemidae 235 Gerres oyena Gerreidae 236 Glossogobius olivaceous Gobiidae 237 Glyphidodontops biocellatus Pomacentridae 238 Glyphidodontops cyamus Pomacanthidae 239 Glyphidodontops hemicyaneus Pomacentridae 240 Glyphidodontops leuwpomus Pomacanthidae 241 Glyphidodontops rollandi Pomacanthidae 242 * Glyphidodontops starcki Pomacentridae Gnathodenter uureo2ineatus Lethrinidae Goby Gobiidae Goby sp. Gobiidae Goby sp.11 Gobiidae Goby sp.12 Gobiidae Goby sp.4 Gobiidae Goby sp.5 Gobiidae Goby sp.6 Gobiidae Goby sp.7 Gobiidae Goby sp.8 Gobiidae Goby sp.9 Gobiidae Gomphosus uarius Labridae Grammistes sexlineatus Grammistidae Gymnomumena zebra Muraenidae Gymnothorax fimbriatus Muraenidae Gymnothorax meleagris Muraeni dae Gymnothorax pictus Muraenidae Halicamphus dunckeri Syngnathidae Holichoeres biocellatus Labridae Hdichoeres hortulanus Labridae Hdichmres margaritaceus- Labridae Hdichoeres marginatus Labridae Hdichoeres melanochir Labridae Halichceres melanurus Labridae Hdichoeres nebdosw Labridae Hdichmrer poecilopterus Labridae Halichoeres prosopsion Labridae Halichoeres scapularis Labridae Hdiehwres sp. Labridae Halichoeres sp.2 (Schroeder 1980) Labridae Halichoeres si.3 Labridae -. -- Continued 87 Appendix 1 (Continued)

By frequency By weight

Rank IndJtransect No. Species Family Slope Flat Trawl Slope Flat Trawl Rank Trawl

Halichoeres trimaculatus Labridae 158 26 - 0.08 6.23 - Hemiglyphidodon plagiometopon Pomacentridae Hemigymnus fasciatus Labridae Hemigymnus melapterus Labridae Hernipteropotus taeniurus ' Labridae Neniochus chrvsostomus Chaetodontidae Heniochus oarius Chaetodontidae Hippichthys spicifer Syngnathidae Hippocampus histrix Syngnathidae Hippocampus ku& Syngnathidae Hippocampus sp. Syngnathidae Histrio histrio Antennariidae Hologymmsus annulatus Labridae Hologymrwsus doliatus Labridae Hologymrwsus sp. Labridao Hypoatherina bleekeri Atherinidae Hypodytes rubripinnis Congiopodidae Istigobius ornatus Gobiidae Labrichthys unilineatus Labridae Labrid Labridae Labrid sp.17 Labridae Labroides bicolor Labridae Lubroides dirnidiatus Labridae Lubropsis rnanabei Labridae Lactoria cornuta Ostraciidae Leptoscarus uaigiensis Scaridae LethPinid Lethrinidae 301 * Lethrinus hmatopterus hthrinidae 302 Lethrinus hard Lethrinidae 303 Lethrinus lenljan Lethrinidae 304 Lethrinus mahsena Lethrinidae 306 * Lethrinus nebulosus Lethrinidae 306 * Lethrinus nernatucanthus Lothrinidae 307 * Lathrinus obsdetus Lethrinidae 308 Lethrinus ornatus Lethrinidae 309 * Lethrinus reticulatus Lothrinidae 310 Lethrinus sp. Lethrinidae 311 * Lethrinus uariegatus Lethrinidaa 312 Lutjanid Lutjanidae 313 Lutjanus biguttatus Lutjanidao 314 Lutjanus bohar Lutjanidae 315 Lutjanus &cussatus Lutjanidae 316 Luljanus fuluiflamma Lutjanidae 317 Lutjanus fuluus Lutjanidae 318 Lutjanus gibbus Lutjanidae 319 Lu2janus kasmira Lutjanidae 320 Lutjanus lineolatus Lutjanidao 321 Lutjanus htjanus Lutjanidae 322 Lutjanus mrwstigma Lutjanidae 323 Lutjanus russellii Lutjanidae ,224 Lu$janus sp. Lutjanidae 326 Lutjanus uitta Lutjanidae 326 Macolor niger Lutjanidae Labridae Labridae Continued Appendix 1(Continued)

By frequency By weight

Rank IndJtransect No. Species Family Slope Flat Trawl Slope Flat Trawl Rank Trawl

329 Malacanthus breuirostris Malacanthidm Meiacanthus grammistes Blenniidae Melichthys uidua Balistidae Monacanthid sp.1 Monacanthidae Monotaxis grandaculis hthrinidae Mulloidichthys flavolineatus Mullidae Myrichthys aki Ophichthidae Myripristis berndti Holocentridae Myripristis murdjan Holocentridae Myripristis sp-1 Holocentridae Nmbreuirostris Acanthuridae Nam lituratus Acanthuridae Naso sp. Acanthuridae Naso unicornis Acanthuridae Nemateleotris magnifica Gobiidae Neoniphon sammara Holocentridae Nouaculichthys macrolepidotus Labridae Nouaculichthys hniurus Labridae Oostethus brachyurus Syngnathidae Ophichthus sp. Ophichthidae Ophichthus urolophus Ophichthidae Ostracion cubicus Ostraciidae Ostracion meleagtis Ostraciidae Paracirrhites arcatus Cirrhitidae Paracirrhites forsteri Cirrhitidae Paraglyphihrlon behni Pomacentridae * Pamglyphidodon carlsoni Pomacentridae Pamglyphidodon rnelas Pomacentridae Pomacentridae 358 ~aragl~~hidodonpolyacanthus Pomacentridae 359 * Paraplyphidodon- -- thoracotaeniatus Pomacentridae Parapercis cephalopunctata Mugdoididae Parapercis clathrata Mugiloididae Parapercis cylindrica Mugdoididae Parapercis polyophthdma Mugiloididae Pampercis sp. Mugiloididae Parapercis tetracantha Mugiloididae Pardachirus pauoninus Soleidae Parupeneus barberimides Mullidae Panrpeneus barberinus Mullidae Parupenem bifosciatus Mullidae Parupeneus cyclostomus Mullidae Parupeneus heptacanthus Mullidae Parupeneus indicus Mullidae Parupeneus pleurostigma Mullidae Pampeneus lrifmciatus Mullidae Pelatus quadrilineatus Teraponidae Pempheris oudenais Pempherididae Pentapodus macrums Nemipteridae Peruagor aspricaudus Monacanthidae Peruagor janthimsoma Monacanthidae Petroscirtes breviceps Blenniidae Petroscirtes SD. Blenniidae ~1a~iotremu;rhimrh~mhs Blenniidas 385 Plagiotrernus tapeinasoma Blenniidae Continued 89 Appendix 1 (Continued)

By frequency By weight

Rank Ind Itransect NO. Species Family Slope Flat Trawl Slope Flat Trawl Rank Trawl

384 Plataz orbicularis Ephippidae Plahpinnatus Ephippidae Platycephalus inolicus Platycephalidae Plectorhynchus chaetodontoides Haemulidae Plectorhynchus diagramus Haemulidae Plectorhynchm goldmanni Haemulidae Plectorhynchm lineatus Haemulidae Plectorhynclurs sp. Haemulidae Plectroglyphiohhn dickii Pomacentridae Plectroglyphidodon lacrymatus Pomacentridae * Plectroglyphidodon leucozona Pomacentridae Plectropornus leoparh Serranidae Plotosus canius Plotosidae Plotosus lineatus Plotosidae Pornacanthus imperator Pomacanthidae Pornacanthus semicirculatlrs Pomacanthidae Pomacentrid Pomacentridae Pomacentrid sp.1 Pomacentridae Pomacentrid sp.10 Pomacentridae Pomacentrid sp. 11 Pomacentridae Pomacentrid sp.12 Pomacentridae Pomacentrid sp.2 Pomacentridae Pomacentrid sp.4 Pomacentridae Pomacentms amboinensis Pomacentridae Pomacentms bankunensis Pomacentridae Pomacentrus coelestis Pomacentridae Pomacentms flauicauda Pomacentridae Pomacentms gramrnorhynchus Pomacentridae Pomacentrus kbiutus Pomacentridae Pornacentrus lepiabgenys Pomacentridae Pomacentrus melanopterus Pomacentridae Pomacentms moluccensis Pomacentridae Pornacentrus nagasrrkiensis Pomacentridae Pomacentms philippinus Pomacentridae Pomacentms smithi Pomacentridae Pomacentms sp. Pomacentridae Pornacentrus tueniomeboponm Pomacentridae Pomwentms trirnaculatus Pomacentridae Pomacentrus tripurntatus Pomacentridae Pomacentrus vaiuli Pomacentridae Pomachrornis richardioni Pomacentridae Priacanthus macracanthus Priacanthidae Pseudanthias squamipinnis Serranidae Pseudobalistes flwimarginatus Balistidae Pseudohalistes fucus Balistidae Pseudocheilintcs evanidus Labridae Pseudocheilinus hexataenia hbridae Pseudocheilim octotaenia Labridae Pseudochromid sp.2 Pseudochrumidae Pseudochromis sp. Pseudochrornidae Pseudojuloides cerasiw Labridae Pseudomonucanthus macrurus Monacanthidae Ptereleotris evides Gobiidae Pterocaesio chrvsozona Lutianidae Ptemmslo Continued Appendix 1 (Continued)

By frequency By weight

Rank Ind./transect No. Species Family Slope Flat Trawl Slope Flat Trawl Rank Trawl

Pterois uolituns Scorpaenidae Rhinecurithus aculeutus Balistidae Rhinecanthus rectungululus Ualistidae Rhinecanthus sp. Balistidae Rhinecanthus oerrucosus Balistidae Salarias fasciatus Blenniidae Salarias sp. Blenniidae Surdinella sp. Clupeidae Sargocentron caudirnuculaturn Holocentridae Sargocentron diadenra Holocentridac Sargocentron ittodui Holoccntridae Sargocentron rubrurn Holocentridae Surgocentron sp. Holocentridnc Sargocentron sp.3 Holoccnt~idae Saurida gracilis Synodontidae Saurida sp. Synodontidae Scarid Scaridae Scarid sp.10 Scaridae Scarid sp.15 Scaridae Scarid sp.18 Scaridae Scarid sp.2 Scaridae Scarid sp.7 Scaridac Scarns bowrrsi Scaridae Scarus chlorodon Scaridae Scarus dimidiotus Scaridac Scurus fusciatus Scaridae Scurus forsteni Scaridae S~urusghobbu~i Scaridae Scarus gibbus Scaridae Scarus globiceps Scaridae Scurus harid Scaridae Scarus lepidus Scaridae Scarus longiceps Scaridae Scarus ooifrom Scaridac Scarus prasiognathus Scaridao Scarus psittacus Scaridac Scarus quoyi Scaridae Scurus rhduropterus Scaridae Scarus rubrouiolaceus Scaridae Scrrrus schlegeli Scaridao Scarus sordidus Scaridao Scarus sp. Scaridae Scarus sp.2 Scaridae Scarus sp.3 Scaridae 483 * Scarus tricolor Scaridae 484 Scolopsis bilineatus Nemipteridae 485 Scolopsis cancellatus Nemipteridae 486 Scolopsis ciliutus Nemipteridae 487 Scolopsis sp. Nemipteridae 488 Scolopsis sp.2 Nemipteridao 489 Scolopsis sp.3 Nemipteridae 490 Scorpaena sp. Scorpaenidae 491 scorpaena sp.1 Scorpaenidae 492 Scorpaenid Scorpaenidao 493 * Scorpaenopsis cirrhosa Scorpaenidao . . Continued Appendix 1 (Continued)

By frequency By weight - Rank IndJtransect No. Species Family Slope Flat Trawl Slope Flat Trawl Rank Trawl

494 Scorpaerwpsis sp. Scorpaenidae Selar crumenoahthcrlmus Carangidae Serranid Serranidae Serranid sp.4 Serranidae Serranid sp.5 Serranidae Siganid Siganidae Siganus argenteus Siganidae Siganus fuscescens Siganidae Siganus guttatus Siganidae Siganus puellus Siganidae Siganus punctatus Siganidae Siganus spinus Siganidae Siganus virgatus Siganidac Siganus vulpinus Siganidae Solenostomus paradarus Solenornostidae Sphaeramia nernatoptera Apogonidae Sphaeramia orbicularis Apogonidae Sphyraena barracuda Sphyraenidae Sphyraena jell0 Sphyraenidae Stenogobius sp. Gobiidae Stephanolepis tomentosus Monacanthidae Stethojulis bandunensis Labridaa Slethojulis sp. Labridae Stethojulis sp.5 Labridae Stethojulis strigiuenter Labridae Stethojulis trilineata Labridae Stolephorus indicus Engraulididae Sumamen bursa Balistidao Suflamen chrysopterus Balistidae Suflamen fraenatus Balistidao Synaptura mrginata Soleidae Syngnathoides biaculeatus Syngnathidae Synodus oariegatus Synodontidae Takifugu rubripes Tetraodontidae Tetraodontid sp.2 Tetraodontidae Thdassoma amblycephalum Labridae Thalassoma hardwickii Lahridae Thalassoma janseni Lahridae Thalassoma hare Labridae Thalassoma lutescens Labridae 634 * Thalassoma purpureum Lahridae 535 Thalassoma quinqueuittatum Labridae 536 Thalassoma sp. Labridae 537 Tylosurus acus melanotus Belonidse 538 Upeneus tragula Mullidae 539 * Vdenciennw longivpinnis Gobiidae 640 Vdenciennea strigata Gobiidae 541 Vderwiennea wadi Gobiidae 542 Yongeichthys criniger Gobiidac 643 Zanclus cornu'tus Zanclidne 544 Zebrasoma scopas Acanthuridae 545 Zebrasoma veliferun~, Acant huridac

Totals 132.30 467.89 314.34 1,247.48 Total no. of species 373 336 186 186 Total no. of families 41 46 48 48 Appendix 2. Reef slope fish recorded from visual census from October 1989 to June 1991 and sorted by fmquency of occurrence (ind./1,000 ma). (* denotes uncertain identification).

Slope Overlap

Upper Mid Lower Flat Trawl Rank Species Family Cum% % Total 1-5 m 5-16 m 16-26 m Total Total

Halichoeres nebulosus Labridae Ctenochuetus binatatus Acanthuridae Parupeneus trifasciatus Mullidae Plotosus linealus Plotosidae Thalassoma hardwickii Labridae Scarus sordidus Scaridae Pomacentrus vaiuli Pomacentridae Pomacentrus coelestis Pomacentridae Pomachrornis richrdsoni Pomacentridae Scarus rhodrrropterus Scal-idae Ctenochaetus striatus Acanthuridae Cheilinus trilobatus Labridae Scarus harid Scaridae Pomacentrus bankanensis Pornacentridac Apogonid Apogonidae Siganus spinus Siganidae IIalichoeres hoeoeni Labridao Sufflan~enchrysopterz~s Balistidno Ch.eilodipterus quinquelineatus Apogonidae Chuetodon kleinii Chactodontidae Thalassorna amblycephalurn Labridae Chaetadon mertensii Chaetodontidae Cirrhilabrus cyanopleura Labridae Puracirrhites arcatus Cirrhitidae Macropharyngodon rneleagris Labridae Thalassoma quingueuittatum Labridae Labroides dimidiatus Labridae Pseudochsilinus hexataenia Labridae Apogon sp. Apogonidae Apogon sp.6 (Schroeder 1980) Apogonidae Halichwres melanochir Labridae Calotomus japonicus Scaridae Stethojulis trilineata Labridae Darnpieria cyclophthaln~a Pseudocllromidae Clupeid Clupeidae Pornacentrus fluvicuudu Pomacentridac Scarid Scaridae Chromis weberi Pomacentridae Parapercis clathrata Mugiloididae Amphiprion clarkii Pomacentridae Nemateleotris rnagnificu Gohiidac Epinephelus merra Serranidae Gomphosus varius Labridae Paraglyphillodon melag Pomacentridae Cephalopholis urodela Serranidae Thalusmma lunare Labridae Chmmis margaritifer Pomacentridae Zanclus cornutus Zanclidae Canthigmter ualentini Tetraodontidae Coris oariegata Labridae Stethojulis bandanensis Labridae Plectroglyphidodon lacymatus Pomacentridae Epinephlus fasciatus Serranidae Thalassoma ianseni Labridae 77.37 0.39 1.56 0.10 1.36 0.10 Continued Appendix 2 (Continued)

Slope Overlap

Upper Mid Lower Flat Trawl -Rank Species Family Cum% % Total 1-5 m 5-16 m 16-26 m Total Total

55 Meimanthus grmmistes Blenniidae 56 Zebrasoma scopas Acanthuridae 57 Cheilinus bimaeulatus Labridae 68 Cirrhitichth s fdco Cirrhitidae 59 Smlopsis biimeutus Nemipteridae 60 Salarias fasciatus Blenniidae 61 Acanthurus gahhm Acanthuridae 62 Chaetodon punctatofasciatus Chaetodontidae 63 * Cheilinus cetebicus Labridae Caesio caerulaurea Lutjanidae Gnathodentex aureolineatus Lethrinidae Dascyllus trimaculatus Pomacentridae Halichoeres marginatus Labridae Atherinid - Atherinidae Halichoeres hortulanus Labridae Pomacentms smithi Pomacentridae Ammpses geographicus Labridae Scarus longiceps Scaridae Cheilinus diagrammus Labridae Pomacentrus lepidogenys Pomacentridae Hemigymnus melapterus Labridae Halichoeres poecilopterus Labridae Pomacentrus moluccensis Pomacentridac Ptereleotris eoides Gobiidae Parapercis cephalopunckta Mugiloididae Valenciennes strigata Gobiidae Pomacentrus philippinus Pomacentridae Caesio erythrogaster Lutjanidae Parapercis polyopht~alma Mugiloididae Coris gaimardi Labridae Siganus argenteus Siganidae Labrid Labridae Stethojulis slrigioenler Labridae Dascyllus reticulatus Pomacentridae Pomacentrid sp.4 Pomacentridae Labrichthys unilineatus Labridae Scarus sp.2 Scaridae Anampses caeruleopunclatus Labridae Cheilio inermis Labridae Centropyge heraldi Pomacanthidae 95 * Acanthurus japonicus Acanthuridae 96 Pervagor janthirwsorna Monacanthidae 97 Chaetodon uagabundus Chaetodontidae 98 Plectroglyphidoobn dickii Pomacentridao 99 Centropyge orolicki Pomacanthidae 100 Parupemus barberinus Mullidae 101 Halichoeres melanurus Labridae 102 Myripristis murdjan Holocentridae 103 Balistapus undulatus Balistidae 104 Acanthurus olioaceus Acanthuridae 105 Naso lituratus Acanthuridae 106 Paraglyphidodon behni Pomacentridae 107 Chmmis mnthura Pomacentridae 108 Plqgiotremus rhinorhynchos Blenniidae 109 Scarus bowersi Scaridae Continued Appendix 2 (Continued)

Slope Overlap

Upper Mid Lower Flat Trawl Rank Species Family Cum% % Total 1-5 rn 5-16 m 16-26 m Total Total

Scarus ghobban Scaridae Scolopsis ciliatus Nemipteridae Centropyge bispinosus Pomacanthidae Amblyglyphihhn curacao Pomacentridae Pomacentms laeniornetopon Pomacentridae Chaetodon octofasciatus Chaetodontidae Epibulus imidiator Lahridaa Scarus sp. Scaridae Amphiprion ocellaris Pomacentridae Hologymmsu~annulatus Labridae Parapercis cylindrica Mugiloididae Chaetodon citrinellus Chaetodontidae Ostracion meleagris Ostraciidae Amthron nigropunctutus Tetraodontidae Parupemus cyclostomus Mullidae Chmrodon anchorao Labridae Dischistodus chrysopoecilus Pomacentridae Canthigaster bennetti Tetraodontidae Chaetodon melannotus Chaetodontidae Caesio tile Lutjanidae Acanthurus pyroferus Acanthuridae Chaetodon trifasciatus Chaetodontidae Scarus schlegeli Scaridae Peroagor aspricaudus Monacanthidae Scarus ovifrons Scaridae Pterocaesio chrysozona Lutjanidae Acanthurus lineatus Acanthuridae Chaetodon xanthurus Chaetodontidae * Cirripectes oariolosus Blenniidae Bodianus mesolhorcrr Labridae Eupomacentrus lividus Pornacentridae Ostracion cubicus Ostraciidae Forcipiger flavissimus Chaetodontidae Paraglyphidodon nipris Pomacentridae Chrornis caerulea Pomacentridae Blenny Blenniidae Ammpses twistii Labridae Chuetodon auriga Chaetodontidae Plagiotremus tapeinosoma Blenniidae Scarus prasiognathus Scaridae Melichthys vidua Balistidae Pterocaesio pisang Lutjanidae Cheilodipterufi macrodon Apogonidae Plectorhynchus linwtus Haemulidae Apogon cyanosoma Apogonidae Pleclropomus leopardus Serranidae Syndus variegatus Synodontidae Cantherhines pardalis Monacanthidae Halichoeres trimaculatus Labridae Naso unicornis Acanthuridae 160 * Acanthurus glaucopareius Acanthuridae 161 Centropyge tibicen Pomacanthidae 162 Dendrmhirus zebra Scorpaenidae 163 Plectorhynchus diagrammus Haemulidae 164 Euoomacentrus nipricans Pomacentridae . Continued Appendix 2 (Continued)

Slope Overlap

Upper Mid Lawer Flat Trawl Rank Spcies Family Cum% % Total 1-6 m 6-16 m 16-26m Total Total

Blenniidae Parucihiies fozsteri Cimhitidae Abu1fiauf dills Pomacentridae lna.lasm?na lute80em Labridae Macropbyqpah mgmsemls Labridae Acdhurus nr';grofscus Acanthuridae Halichoeres prompewn Labridae Cheilinus fasclatus Labridae Abudefduf codestinus Pomacentnidae Zebrasoma ueliferum Acanthuridae Rhinecanthus muleatus Balistidae Parupeneus bifosciatus Mullidae Chaetodm unirnaclllatus Chaetodontidae Calotomus sp. Scaridae Rhinecanthus verrucosus Balistidae Goby Gobiidae Pomacentrid sp.1 Fbmacentridae Dischistodus prosopotaenia Pomacentridae Heniochus chrysostomus Chaetodontidae Saurida gracilis Synodontidae Diploprion bifasciatus Grammistidae Novaculichthys laeniurus Labridae Bolbometopon bicolor Scaridae Primanthus macmcanthus F'riacanthidae Apogon nooemfasciatus Apogonidae Balistid sp.1 Balistidae 192 * ~mbl~ele&isfascia& Gobiidae Chaelodon ornatisslmus Chaetodontidae Siganus fuscescens Siganidae Scarus quoyi Scoridae Cephalopholis pachycentron Serranidae Scarus lepidus Scaridae Parupeneus pleurostiha Mullidae Scarus fasciatus Scaridae Pomacentrid Pomacentridae Pamcrcentrus trirnacdatus Pomacentridae Pornacentms sp. Pomacentridae Centrogenys uaigiensis Percichthyidae Scarus firstmi Elcaridae ' Paraglyphi&n carlson Pomacentridae Luwus dectwsatus htjanidae Cepholopholis argus Sewanidae Sargocentmn rubrum Holocentridae Lu-us fulvifiamma Lutjaddae Mdoidichthys flavolineatus Mullidae Pornacentme amboimnsis Fbmacentridae Siganw virgCCru8 Siganidae Scarus sp.8 Scaridae Pseudmheilinus evanidus Labridae Aediscus strigatus Centrlscidae Thalasmma sp. Iabridae Halichoeres margaritaceus Ltabridae Eomacentridae 2 19 scaris-psittactrs Scaridae Continued Appendix 2 (Continued)

Slope Overlap

Upper Mid Lower Flat Trawl Rank Species Farnil y Cum% % Total 1-5 rn 5-16 m 16-26 m Total Total

Heniochus varius Chaetodontidae 98.63 0.02 0.08 - 0.01 0.07 Pseudojuloides cerasinus Labridae Gymnothorax pictus Muraenidae Pentupodus macrurus Nernipteridae Chaetohn trifascialis Chaetodontidae Cirrhitichthys aprinus Cirrhitidae Arothron stellatus Tetraodontidae Scarus dimidiatus Scaridae Naso sp. Acanthuridae Hemiglyphihdon plagiomelopon Pomacentridae Salarias sp. Blenniidae Apogonid sp.5 Apogonidae Pseudunthias squamipinnis Serranidae Chlysiptera leucopoma Pomacentridae Chromis sp. Pomacentridae Selar crurnenaphthdm.us Carangidae Monotaxis grandoculis Lethrinidae Centropyge bicolor Pomacanthidae Grammistes skxlineatus Grarnrnistidae Lutjanid Lutjanidae Istigohius ormtus Gohiidae Siganus vulpinufi Siganidae Fistularia petimba Fistulariidae Bodianus hirsutus Labridae Macolor niger Lutjanidae Pseudocheilinus octotaeniu Labridae Malacanthus breuirostris Malacanthidae Purupeneus barberinoides Mullidae Balistid Balistidae Auloslornus chinensis Aulostomidae 260 * Thulassoma purpureum Labridae 251 * Scarus tricolor Scaridac 252 Serranid Serranidae 253 Dascyllus aruanus Pomacontridae 254 Colotornus carolinus Scaridae 265 Pseudobalistes flauimarginntus Balistidae 256 Ammpses meleagrides Labridae 257 Acanthurus mata Acanthuridae 258 Hologymnosus daliatus Labridae 259 Acanthurus triostegus Acanthuridae 260 Amblygobius dbimaculatus Gobiidae 261 Acanthurid sp.6 Acanthuridae 262 Lethrinus hard Lethrinidae 263 Sargocentron diadema Holocentridae 264 Gymnomuraena zebra Muraenidae 265 Glyphidodontops leucopomus Pomacanthidae 266 Chaetodon lunula Chaetodontidae 267 Valemiennea wardi Gobiidae 268 Bodianus bilunulutus Labridae 269 Myripristis berndti Holocentridae 270 Suflamen bursa Balistidae 271 Stephanolepis tomentosus Monacanthidae 272 Cheilinus unrlulatus Labridae 273 Monacanthid se.1 Monacanthidae 214 Epinephelus s; Serranidae 99.48 0.01 0.04 0.03 - 0.02 Continued Appendix 2 (Continued)

Slope Overlap

Upper Mid Lawer Flat Trawl Rank Species Family Cum% % Total 1-5 m 5-16 m 16-26 m Total Total

Lutjanus gibbus Lutjanidae Balistid sp.6 Balistidae Apogon bandanensis Apogonidae Cork dorsurnacula Labridae kthrinus mahsenu hthrinidae Scarus rubroviolaceus Scaridae Hemigymnus fasciatus Labridae Epimphelus sexfasciatus Serranidae Pomacentrid sp.12 Pomacentridae Amphiprion perideraion Pomacentridae Bodianus uxillwis Labridae Parapercis tetracantha Mugiloididae Parapercis sp. Mugiloididae Bodianus sp. Labridae Chaetodon ulietensis Chaetodontidae Blenny sp.7 Blenniidae Diodon hystrix Diodontidae Lutianus uitta Lutjanidae Amphiprion sandaracims Pomacentridae Rhinecanthus rectangulux Balistidae 295 * Cirrhitichthys serratus Cirrhitidae 296 Pomacanthus semicircula;irs. Pomacanthidae 297 Parupemus heptacanthus Mullidae 298 Anthias sp. Serranidae 299 Acanthurid sp.5 Acanthuridae 300 Cephalopholis miniata Serranidae 301 Stethojulis sp. Labridae 302 * Arnblyeleotris japolu'ca Gobiidae 303 Chaelodon baronessa Chaetodontidae 304 Pomacentrus tiipunctatus Pomacentridae 305 Pomacentrid sp.2 Pomacentridae 306 Caesio sp. Lutjanidae 307 Chaetodon sp. Chaetodontidae 308 Canthigaster coronata Tetraodontidae 309 * Asterropteryx semipunctatus Gobiidae 310 Cirripectes polyzona Blenniidae 311 Cheilinus sp. Labridae 312 Goby sp.12 Gobiidae 313 * Cirrhilabrus polyzona Labridae Pomacentrzls nagasakiensis Pomacentridae Hologymnosus sp. Labridae Aspidontus taeniatus Blenniidae Nooaculichthys macrolepidotus Labridae Cirrhitops hubbardi Cirrhitidae Stethojulis sp.5 Labridae Chmmis lepidolepis Pomacentridae Upeneus tragula Mullidae Lutjanus BP. Lutjanidae Lutjanus lutjanus Lutjanidae Scolopsis sp. Nemipteridae Cephalopholis sp. Serranidae Myripristis sp.1 Holocentridae Lutjanus lineolatus Lutjanidae 328 * Acanthurus xunthopterus Acanthuridae 329 Canthigaster cornpressa Tetraodontidae. . Continued Appendix 2 (Continued)

Slope Overlap

Upper Mid Lnwer Flat Trawl Rank Species Family Cum% % Total 1-5rn 5-16m 16-26 m Total Total

330 &orpaenid Scorpaenidae 99.86 ~0.01 0.02 - - 0.02 0.05 351 Amphiprion frenatug Pomacentridae 99.85 <0.01 0.02 - - 0.02 332 Glyphidodontops.rollandi Pomacanthidae 99.86 <0.01 0.02 - - 0.02 0.31 -

353 * Cephalopholis knack Serranidae 99.86 4.01 0.02 - + 0.02 Calloplesiops dtivelis Plosiopidae Scarus gibbus Scaridae Nmniphon sammara Fblocentridae Parupemus indicus !Nullidae Serranid sp.5 Serranidae Pornacanthus imperator Pomacanthidae GyrnnoUlwvu fimbriotun Muraenidae Naso bmvimstris Acanthuridae Labmpsk mcmabei Labridae Caram melarnpygus Carangidae Lethrinid bthrinidae Pseudobalistes fupcw Balistidae

346 * Scorpaempsis cirrhosa Scorpaenidae 99.91 <0.01 0.01 + 0.01 - 0.08 0.18 hthrinus ornatus ~ethrinidae Pterois oditans Scorpaenidae Amthmn hispidus Tetraodontidae Plectorhynchus chaetodontoides Maemulidae Atule mate Carangidae Cantherhines dumerilii Monacanthidae Blenny sp.2 Blenniidae &capterns sp. Carangidae Slemgobius sp. Gobiidae Pomacentrid sp. l 1 Pomacentridae Halichms ~p. Labridae Glyphidothntops cyaneus Pomacanthidae Halichmes scapularis fiabridae Carangoides fuluoguttatus Carangidae Pomacentrus grammorhynchlur Pomacentridae Balistid sv.4 Balistidae Coris ayg& Labridae 364 * Canthigaster solandri Tetraodontidae 365 Sargocentmn sp. Holocentridae 366 Plectorhynchus sp. Haemulidae 367 Goby sp.7 Gobiidae 368 Cheilinus rhadochrotur Labridae 369 Plectorhynchus goldmanni Haemulidae 370 Carangid Carangidae 371 Gymnothorax meleagris Muraenidae 372 * Scarus globiceps Scaridae 375 SuffZamen fraenatus Balistidae

Others Totals Appendix 3. Reef slope fish families recorded from visual census from October 1989 to June 1991 and sorted by frequency of omrrence (ind/1,000 mB).

Upper Mid Lawer Rank Family Cum% % Total 1-5 m 5-16 m 16-26 m

1 Labridae 2 Pomacentridae 3 Acanthuridae 4 Scaridae 5 Apogonidae 6 Mullidae 7 Chaetodontidae 8 Plotosidae 9 Siganidae 10 Balistidae 11 Serranidae 12 Cirrhitidae 13 Blenniidae 14 Mugiloididae 15 Gohiidae 16 Lutjanidae 17 Tetraodontidae 18 Pseudochromidae 19 Pomacanthidae 20 Clupeidae 21 Nemipteridae 22 Zanclidae 23 Monacanthidae 24 Lethrinidae 25 Atherinidae 26 Holocentridae 27 Ostraciidae 28 Haemulidae 29 Synodontidae 30 Scorpaenidae 31 Gramrnistidae 32 Muraenidae 33 Carangidae 34 Priacanthidae 55 Percichthyidae 36 Centriscidae 57 Fistulariidae 38 Malacanthidae 39 Aulostomidae 40 Diodontidae 41 Pleriopidae 100.00 4.01

Totals 100.00 396.89 187.43 122.59 86.88 Appendix 4. Reef flat fish recorded from visual census from August 1988 to June 1991 and sorted by frequency of occurrence (ind./1,000 m2).Bottom cover, A = corals and sand, B = corals and seagrass, C = seagrass and D = Sargrrssum spp. (* denotes uncertain identification).

Flat Overlap

Slope Trawl Rank Species Family Cum% % Total A B C D Total Total

Eupomacentrus nimicans- Pomacentridae 7.59 7.69 142.10 ~n~graulid Engmulididae Sardinella sp. Clupeidae Dascyllus aruanus Pomacentridae Atherinid Atherinidae Chmmis caerulw Pomacentridae Halichwres hcleueni Labridae Scarus harid Scaridae Clupeid Clupeidae Hypmtherina bleekeri Atherinidae Pornacentrus f2auicaud.a Pornacentridae Plotosubi li~atus Plotosidae Eupomacentrus liuidus Pomacentridae Scarus rhodwopterur Scaridae Arnblyglyphidodon curacao Pomacentridae Scarid Scaridae Stolephorus indicus Engraulididae Apogon sp.5 (Schmeder 1980) Apogonidae Cheilodipterus quinquelineatus Apogonidae Dischistodus chrysopoecilus Pomacentridae Apogonid sp.5 Apogonidae Thrrlassoma hardu~ickii Labridae Parapercis cylindrica Mugiloididae Stethojulis strigiuenter Labridae Parupemus trifasciatus Mullidae Hdichaeres trirnaculatus Labridae Cdotomusjaponicus Scaridae Scarus sordidus Scaridae Epinephelus rnerra Serranidae Siganus spinus Siganidae Pornacentrus grammorhynchus Pomacentridae Halichoeres scapulwis Labridao Pomacentrus bankanensis Pomacentridae Apogon cyanosoma Apogonidae Apogon 8P. Apogonidae Stethojulis trilineata Labridae Labroides dirnidiatus Labridae Apogon novemfzsciatus Apogonidae Cheilinus trtlobatus Labridae Cork uariegata Labridae Pomacentms tripunctatus Pornacentridae Ctemchaetus bimtatus Acanthuridae Paraglyphidodon rnelas Pomacentridae Plectroglyphidodon lacrymatus Pomacentridae Choerodon anchorwo Labridae Onathodentex aureolineatus hthrinidae Saltarias fasciatus Blenniidae Scolopsis bilineatus Nemipteridae Pornacentrus taeniornetopon ~omacentridae Siganus fuscescens Siganidae Pterocaesio chrysozona Lutjanidae Stethoiulis bandunensis Labridae

Pornacentrus nwluccensis Pomacentridae ~ ~ Continued Appendix 4 (Continued)

Flat Overlap

Slope Trawl Rank Species Family Cum% % Total A B C D Total Total

Apogon bandanensis Apogonidae Scarus ouifrons Scaridae Ctenochadus striatus Acanthuridae Dischistodus prosopotaenia Pomacentridae Hemigymnus melapterus Labridae Abuhfduf codestinus Pomacentridae Chaetodon trifasciatus Chaetodontidae Scarus sp. Scaridae Hdichoeres hortulanus Labridae Hdichoeres nebulmus Labridae Zanclus cornutus Zanclidae Thalassoma hnare Labridae Abudefduf saxatilis Pomacentridae Hdichoeres marginatus Labridae ChaeWn rnelannotus Chaetodontidae Dischistodus notopthaltnus Pomacentridae Pomachromis richardsoni Pomacentridae Cheilio inermis Labridae Cheildpterus macrodon Apogonidae Scarus dimidiatus Scaridae Chaetodon auriga Chaetodontidae Neoniphon sarnmara Holocentlidae Canthigmter ualentihi Tetraodontidae Pseudocheilinus hexataenia Labridae Scarus schlegeli Scaridae Apogonid Apogonidae Pornacentrus coelestis Pomacentridae Gomphosus varius Labridae hcyllus trimacul~tus Pomacentridae Dampieria cyclophthalma Pseudochrornidae Caesio tile Lutjanidae Dascyllus reticulatus Pomacentridae Zebrasoma scopas Acanthuridae Parupemus barberinbides Mullidae Hdichoeres poecilopterus Labridae Pomacentrus philippinus Pomacentridae Chaetodon kleinii Chaeietodontidae Escualosa thoracata Clupeidae 92* Cirriwctes uariolosus Blenniidae ~hae&lon citrinellus Chaetodontidae Amphiprion ocellaris Pomacentridae, Parupeneus barberinus Mullidae Lethrinus ornatus Lethrinidae Scarus sp.2 Scaridae Aeanthurus gahhm Acanthuridae Chaetwlon vagabulzdus Chaetodontidae Glyphidodontops cyaneus Pomacanthidae Lethrinus hak hthrinidae Pomacentms trimaculatus Pomacentridae Pomacentms amboinemis Pomacentridae Chaetodon merbnsii Chaetodontidae Acreichthys tomentosus Monacanthidae Pornacentrus sp. Pomacentridae Meiacanthus gramrnistes Blenniidae Scarid sp.18 Scaridae Continued Appendix 4 (Continued)

Flat Overlap

Slope Trawl Rank Species Family Cum% % Total A B C D Total Total

Aeoliscw strigatus Centriscidae 97.96 0.06 0.89 0.47 0.36 0.06 - 0.09 3.81 Amphiprion clarkii Fbmqcentridae Paraglyphidodon nigmris Pomacentridae Pornacentms uaiuli Pomacentridae Epibulus insidiator Labridae Coris gaimardi Labridae Plectroglyphidodon leucozonu Pomacentridae Pampercis clathrata Mugilaididae Corythoichthys haematopterus Syngnathidae Myripristis niurdjan Holocentridae Scolopsis cancellatus Nemipteridae Epinephelus ongus Serranidae Plectorhynchus diagrummus Hae'mulidae Chaetodon xanthurua Chaetodontidae Glyphidodontops biocellatus Pomacentridae Atadmpses geographicus Labridae Halichoeres sp. Labridae Paraglyphidodon carlsoni Pomacentridae Monotaris grandoculis hthrinidae Pamglyphidodon behni Pomaceqtridae Lutjanus fuluus Lutjanidae Pomacentms labiatus Pomacentridae Hemiglyphidodon plagiometopon Pomacentridae Dascyllus rnelanurus Pomacentridae Pomacentms smithi Pomacentridae Goby Gobiidae Zebrasoma ueliferum Acanthuridae Grarnmistes sexlineatus Grammistidae Plectorhynchus lineatus Haemulidae Diechistodus pseudochrysopoecilus Poinacentridae Pomacentms k%?unopterus Pomacentridae Arothron nigropunctatus Tetraodontidae Bodianus mesothorax Labridae Hemigymnus fasciatus Labridae AburEefduf septamfmciatus Pomacentridae Amblygobius albirnaculatus Gobiidae Ostracion cubicus Ostraciidae Scarid sp.7 Scaridae Arothron imrnaculatus Tetraodontidae Tylosunrs acus melanotus Belonidae Macropharyngodon meleagris hbridae Parupemus bifasciatus Mullidae Rhinecunthus muleatus Balistidae Lethrinus mahsena Lathrinidae Sigunus atgenteus Siganidae Canthigaster bennetti Tetraodontidae Fistularia petimba Fistulariidae Gymnothorax pictus Muraenidae Synodus uariegatus Synodontidae Glyphidodontops rollandi Pomacanthidae ~tethojulissp. Labridae Scarus fasciatus Scaridae Halichoeres prosopeion Labridae

Nnm lituratus~-~-- Acanthuridae Cheilinus birnaculatus Labridae 99.23 0.01 0.26 0.01 0.11 0.06 0.08 1.48 0.22 Continued Appendix 4 (Continued)

Flat Overlap

Slope Trawl Rank Species Family Cum% % Total A I3 C D Total Total

Saurida macilis Synodontidae 99.24 0.01 0.26 ~canthuiuslineatus Acanthuridae Labrid Labridae Apogonid sp.2 Apogonidae Wdichmres matgaritaceus Labridae Apogon compressus Apogonidae Nouaculichthys tueniurus Labridae Plagiotremus rhinorhynchs Blenniidae Thalessom amblycephalurn Labridae Pterois oditans Scorpaenidae Lutjanus decussatus Lutjanidae Plectroglyphidodon dlckii Pomacentridae Heniochus vwius Chaetodontidae Aulostomlls chiuensis Aulostomidae Heniochus chrysostomus Chaetodontidae Acanthurus mata Acanthuridae Scarus prasiognathus Scaridae CIiaetodon adiergastos Chaetodontidae Scarid sp.2 Scaridae Chrornis mnthura Pomacentridae Echidna nebulosa Muraenidae Naso unicornis Acanthuridae Dampieria sp. Pseudochrornidae Labrichthys unilineatus Labridae Scarus ghobban Scaridae Gymnothorax firnbriatus Muraenidae Bolbornetopon bicdor Scaridae Pornmentms lepidogenys Pomacentridae Chaetodon lunula Chaetodontidae Sufflamen chrysopteru Balistidae Halichoeres rnelanurus Labridae Thalassorna quinqueuittaturn Labridae Plectorhynchus chaetodontoides Haemulidae Caesio erythrogwter Lutjanidae Glyphidodontops hemicyaneus Pomacentridae Apogonid sp.4 Apogonidae Arothron stellatus Tetraodontidae Chrornis weberi Pomacentridae Siganus virgatus Siganidae Epinephelus fasciatus Serranidae Parapercis polyophlhalrna Mugiloididae Mulloidichthys flauolineatus Mullidae Paraglyphidodon polyacanthus Pomacentridae Scarus longiceps Scaridae Sargocentron ittodai Holocentridae Apogvn sp.1 (Schroeder 1980) Apogonidae Lutjanus monostigma Lutjanidae Lutjanus biguttatus Lutjanidae Centropyge herddi Pomacanthidae Cheilinus fasciatus Labridae Sargocentron rubrum Holocentridae Halichoeres melamchir Labridae Dischistodus perspicillatus Pomacentridae Acanthurid sp.1 Acanthuridae Naso sp.1 Acanthuridae Continued 104

Appendix 4 (Continued)

Flat Overlap

Slope Trawl Rank Species Family Cum% % Total A B C D Total Total

Chaebdon ephippium Chaetodontidae 99.74 0.01 0.11 - 0.03 0.03 0.06 Apogon coccineus Apogonidae Chaeibdon ulietensis Chaetodontidae Apogonid sp.3 Apogonidae Sargocentron diadema Holocenttidae Halichceres sp.2 (Schmeder 1980) Labridae Exyrias puntang Gobiidae Lutjanus fulviflamma Lutjanidae Dendrochirus zebra Scorpaenidae Lutjanus gibbus Lutjanidae Chaebdon punctatofasciatus Chaetodontidae Parupeneus cyclostomus Mullidae Apogonid sp.8 Apogonidae Petroscirtes sp. Blenniidae Apogonid sp.6 Apogonidae Acanthurus dussumieri Acanthuridae Halichoeres sp.3 Labridae Blenny Blenniidae Canthrhines oardulis Monacanthidae 238* Scorpaenopsis cirrhosa Scorpaenidae 239 Cheilinus celebicus Labridae 240 Rhinecanthus oerrucosus Balistidae 24 1 Amblyglyphidodon leucogaster Pomacenttidae 242 Cheilinus diagrammus Labridae 243 Plectorhynchus goldmanni Haemulidae 244 Ostracion mleagris Ostraciidae 245 Diploprion bifwciatus Grammistidae 246* Glyphidodontops starcki Pomacentridae 247 Yongeichthys criniger Gobiidae 248* Lethrinus nematacanthus Lethrinidae 249 Centropyge tibicen Pomacanthidae 250 Acanthurus triostegus Acanthuridae 251* Lethrinus obsdetus hthrinidae 252 Myripristis berndti Holocentridae 253 Acanthurid sp.6 Acanthuridae 254 Leptoscarus uaigienais Searidae 255 Exallias breuis Blenniidae 256 Aspidontus taeniatus Blenniidae 257 Pomacentrid sp.10 Pomacentridae 258 Canthigwter janthinoplera Tetraodontidae 259 Chaetodan'limdatus Chaetodontidae 260* Vdenciennea longispinnis Gobiidae 261* Acanthurus japonicus Acanthuridae 262 Chaelodon raflesi Chaetodontidae 263 Sargocentmn caudimaculatum Holocentridae 264 Amblygobius phnllrentr Gobiidae 265 Glyphidudontops leucopomus Pomacanthidae 266 Valenciennes strigata Gohiidae 267 Hemipteromtus trreniurus Labridae 268 Scarid sp.16 Searidae 269 Lethrinus haernatoptem Lethrinidae 270 Gymnornumna zebra Muraenidae 271 Pseudochrornid sp.2 Pseudochrornidae 99.93 ~0.010.03 0.03 - 272 ~ara~l~~l~~hidodonthomcotaeniutus Pomacentridae 99.93 d.01 0.03 - 0.03 - 273 Vdenciennwr umrdi Gobiidae 99.93 ~0.01 0.03 - - 0.03 . . . Continued aoooopooooooooooooooopooooooooooooooooooooooooooooooooo ~oooooooooobooooooooooo~oooo~o~o~~~~o~oooooooooooooooo ~wr~~w~~~~c~~~r-~~wcLcLrw~w~~o~~~w~wcQww~~~wZw~~w~wwwwwwww Appendix 4 (Continued)

Flat Overlap

Slope Trawl Rank Species Family Cum% % Total A B C D Total Total

329 Serranid sp.4 Serranidae 330 &by ~p.4 Gobiidae 331 Scolopsi~sp.3 Nemipteridae 332 Scarid sp.10 Scaridae 333 Acanthurus bariene Acanthuridae 334 Lutjanus russellii Lutjanidae 335 Epi~phehsmaculatus Serranidae 336 hbroides bicolar Labridae

Others Totals Appendix 6. Reef flat fish families mcorded from visual census from August 1988 to June 1991 and sorted by frequency of occmmnce (indJ1,OOO m2).

Corals and

Rank Family Cum% 46 Total sand seagrass Seagrass Sargassum

Pomacentridae Labridae Clupeidae Scaridae Engraulididae Apogonidae Atherinidae Plotosidae Siganidae Mullidae Mugiloididae Serranidae Acanthuridae Chaetodontidae Lethrinidae Blenniidae Lutjanidae Nemipteridae Tetraodontidae Holocentridae Zanclidae Pseudochmmidae Pornacanthidae Gobiidae Haemulidae Monacanthidae Centriscidae Muraenidae Syngnathidae Balistldae Synodontidae Grarnmistidae Ostraciidae Scorpaenidae Belonidae Fistulariidae Aulostornidae Ophichthidae Gerreidae Percichthyidae Cirrhitidae Ephippidae Diodontidae Soleidac 45 Solenostomidae 100.00 cO.01

Totals 100.00 1,871.66 405.63 661.14 409.60 396.19 Appendix 6. Reef flat fish caught at night by a roller beam trawl in the seagrass beds of Santiago Island from October 1988 to June 1991 and sorted by frequency of occurrence (indJ1,OOO m2).(* denotes uncertain identification).

Trawl Overlap

Seagrass density and depth Dense Sparse Dense Sparse Slope Flat Rank Species Family Cum% % Total d.5 m d.5 m >1.5 rn 21.5 m Total Total

Sinanus fuscescens Siganidae- 28.34 28.34 356.32 161.10 78.29 107.15 9.78 0.13 5.63 ~o&leriauariegata Apogonidae Apogon sangiensis Apogonidae Plotosus lineatus Plotosidae Syngnathoides biaculeatus Syngnathidae Sphaerarnia orbicularis Apogonidae Acreichthys tomet~tosus Monacanthidae Apogon coccineus Apogonidac Cheilodipterus quinquelineatus Apogonidae Siganus uirgatus Siganidae Petroscirtes breoiceps Blenniidae Apogonid sp.6 Apogonidae Lethrinus hard Lethrinidae Parupeneus barberinus Mullidae Siganus spinus Siganidae Arothron immaculatus Tetraodontidae Centrogenys uaigiensis Percichthyidae Lethrinus ornatus Lethrinidae Scarus rhoduropterus Scaridae Sauridn madis S~nodontidac 21* hthrin& obsoletus Gthrinidau 22 Siganus argenteus Siganidae 23* Eethrinus reticulatus Lethrinidae 24 Scarusghobban Scaridae 25 Apogon bandanensis Apogonidae 26 Hyptherina bleekeri Atherinidac 27 Choerodon anchorago Labridae 28 Apogon amboimnsis Apogonidae 29 Scarus longiceps Scaridae 30 Parapercis cylindrica Mugiloididae 31 Epinephlus merm Serranidae 32 Gymnothorax pictus Muraenidae 33 Upeneus trugula Mullidae 34 Aediscus strigatus Centriscidae 36 Apognn nwemfasciatus Apogonidae 86 &O@n CyUMSOt?l5 Apogonidao 37 Apogran sp.6 (Schroeder 1980) Apogonidae 38 Epinephelus ongus Serranidae 39 Cheilinus trilobatus Labridae 40 Stethnjulis strigiventer Labridae 41 Parupeneus trifasciatus Mullidae 42 Calotomusjaponicus Scaridae 43 Pelatus quadrilineatus Teraponidae 44 Arothron hispidus Tetraodontidae 45 Siganus punctatus Siganidae 46 Corythoichthys haematophrus Syngnathidac 47 Pardachirus paooninus Soleidae 48 Parupemus barberimides Mullidac 49 Gerres oyena Gemidae 50 Lutjanus fuluiflamma Lutjanidae 51 Sphaeramia nemabptera Apogonidac 52 Lethrinus mahsena Lethrinidac 53 Apogonid sp.2 Apogonidae Continued Appendix 6 (Continued) Trawl Overlap

Seagrass density and depth Dense Sparse Dense Sparse Slopo Flat Rank Species Family Cum% % Total c1.5 m c1.6 m >1.5 m 3.1.5 m Total Total 54* Asterropteryx semipunctatus Gobiidae 97.59 0.10 1.31 - - 1.26 0.05 0.03 55 Lethinu lentmn Iathrinidae 97.69 0.10 56 Ariosoma amgoides Colocongridae 97.78 57 Leptoscarus oaigiemis Scaridae 97.86 58 Darnpieria cyclophthalma Pseudochromidae 97.93 59 Amblyapistus taenianotus Congiopodidae 98.00 60 Cheilodipterus mumodon Apogonidae 98.07 61 Surdinella sp. Clupeidae 98.13 62 Cheilio inermis Labridae 98.19 63 Amblygobius albimaculatus Gobiidae 98.25 64 Scolopris bilineatus Nemipteridae 98.31 65 Canthigaster bennetli Tetraodontidae 98.36 66 Rzyrias puntang Gobiidae 98.42 67 Goby sp.11 Gobiidae 98.47 68 Archamia limolala Apogonidae 98.52 69 Escualosa thracula Clupeidae 98.68 70 Dunekerocampus dmtyliophorus Syngnathidae 98.63 71 Parupeneus indicus Mullidae 98.67 72 Synaptura marginata Soleidae 73 Scarus prasiognathus Scaridae 74 Chelonadon patoca Tetraodontidae 75 Scolopsis ciliatus Nemipteridae 76 Scorpaena sp. Scorpaenidae 77 Plotosus canius Plotosidae 78 Chaetodon auriga Chaetodontidae 79 Sphymena jello Sphyraenidae 80 Conger cinereus Congridae 81 Aluteres scriptus Monacanthidae 82 Synodus uariegatus Synodontidae 88 Mulloidichthys flauolineatus Mullidae 84 Platycephalus indicus Platycephalidae 85 Halicamphus dunckeri Syngnathidae 86 Scolopsis cancellatus Nemipteridae 87 Goby Gobiidae 88 Canthigaster ualentini Tetraodontidae 89 Apogonid sp.7 Apogonidae 90 Siganus guttatus Siganidae 91 Lactoria cornuta Ostraciidae 92 Ostracion cubicus Ostraciidae 93 Hippocampus histrix Syngnathidae 94 Dischistodzu chrysopoecilus Pomacentridae 96 Cheilinus bimaculatus Labridae 96 Exyrias bellissirnus Gobiidae 97 Encheiliophis uerrnicularis Carapidae 98 Hypodytes rubripinnis Congiopodidae 99 Goby sp.9 Gobiidae 100 Yongeichthys criniger Gobiidae 101* Scorpaempsis cirrhosa Scorpaenidae 102 Naso sp. Acanthuridae 103 Plectorhynchus lineatus Haemulidae 104 Chaetodon melannotus Chaetodontidae 105 Antennarius nummifer Antennariidae 106 Pseudobalistes fuscus Balistidae 107 Hippocampus 8udu Syngnathidae 99.51 0.01 0.16 - 0.10 0.06 - Continued Appendix 6 (Continued)

Trawl Overlap

Seagrass density and depth Dense Sparse Dense Sparse Slope Flat Rank Species Family Cum% % Total c1.6 m d.5 m >1.5 rn >1.5 m Total Total

108 Apogonid sp.11 Apopnidae 109 Aulostomus chinensis Aulostomidae 110* Lethrinus nematacanthus Lethrinidae 111 Nasr, unicornis Acanthuridae 112 Pterois volituns Scorpaenidae 113 Fistulwia petimba Fistulariidae 114 Lutjanus &cussatus Lutjanidae 11b Lutjanus lineolatus Lutjanidae 116* Lethrinus variegatus Lethrinidae 117 Grammistes sexlineatus Grammistidae 118 Goby sp8 Gobiidae 119 Scarid Scaridae 120 Apogon sp.8 (Schroeder 1980) Apogonidae 121 Clupeid Clupeidae 122 Apogwn sp. Apogonidae 123 Sphyraena bamuda Sphyraenidae 124 Parupeneus heptmanthus Mullidae 126 Apogonid sp.10 Apogoni dae 126 Antennarius moluccensis Antennariidae 127 Histrio histrio Antennariid* 128 Arothron nigropunctatus Tetraodontidae 129* Corythoichthys schdtzi Syngnathidae 130 Bupomacentrus nigricans Pomacentridae 131 Platux orbicularis Ephippidae 132 Arothron stellatus Tetraodontidae 139 Plectorhynchus chaetodontoide#s Haemulidae 134 Oostdhus brachyurus Syngnathidae 135 Acanthurid Acanthuridae 136* Eleotris fusca Gobiidae 137 Saurida sp. Synodontidae 138 Siganid Siganidae 139 Labrid Labridae 140 Myrichthys aki Ophichthidae 141 Balistid Balistidae 142 Hippichthys spicifer Syngnathidae 143 Chaetodon lunula Chaetodontidae 144 Conger sp. Congridae 145 Chaelodon trifasciatus Chaetodontidae 146 Abudefduf samtilis Pomacentridae 147 Penlapodus macrurus Nemipteridae 148 Antennarius sp.1 Antennariidae 149 Bothus pantherinus Bothidae 150 Scorpaenid Scorpaenidae 151 Scarus ovifrons Scaridae 152 Plectorhynchus sp. Haemulidae 153 Pempheris oudensis Pempheridae 154 Dihnhystrix Diodontidae 156 Dischistodus notopthalmus Pomacentridae 156 Apogonid Apogonidae 157 Lutjanus gibbur Lutjanidae 158 Dendrochirus zebra Scorpaenidae 169 Epimphdus tauvim Serranidae 160 Oohichthus sp. Ophichthidae 161 Glossogobius olioa.ceous Gbiidae 99.91 c0.01 0.05 - 0.05 - Continued Appendix 6 (Continued)

Trawl Overlap

Seagrass density and depth Dense Sparse Dense Sparse Slope Flat Rank Species F~Y Cum% % ~otdc1.5 m c1.6 m >1.5 m 21.5 m ~otd ~otd 162 Arothran spL2 Tetraodontidae 99.92 4.01 0.05 - - 0.05 - 168 Pseudobalistes f2avimmginadus Balistidae 99.92 4.01 0.06 - - 0.06 - 0.06 0.01 164 Arothron rnappa Tetraodontidae 99.98 4-01 0.06 - - 0.05 - 165 Eupomacentrus lividus Pomacentridae 99.93 4.01 0.06 0.05 - - 0.36 46.94 166 Drepam longimana Ephippidae 99-99 4.01 0.05 0.OK - 167 Tokifugu rubripes Tetraodontidae 99.94 4.01 0.05 0.05 - 168 Aesopia comta Soleidae 99.94 4.01 0.05 0.06 - 169 Chilinus fasciatus Labridae 99.94 4.01 0.05 0.06 - - 0.18 0.11 170 Amblygobiua sp. Gobiidae 99.95 4.01 0.05 0.05 - 171 Tetraodoritid sp.2 Tetraodontidae 99.95 c0.01 0.05 0.05 - 172 Scorpaena sp.1 Scorpaenidae 99.96 4.01 0.05 0.05 - 173 Luwus hrnira Lutjanidae 99.w 4.01 0.04 0.04 - 174 Siganus puellus Siganidae 99.96 4.01 0.04 0.04 - 176 Naso lituratm Acanthuridae 99.97 4.01 0.04 0.04 - - 0.61 0.26 176 Acanthus gahhm Acanthuridae 99.97 <0.01 0.04 0.04 - - 1.39 1.25 177 Scarus schlegeli Scaridae 99.97 4-01 0.04 0.04 - - 0.43 2.29 178 Pmtdornonaeanthus mcrurus Monacanthidae 99.97 4.01 0.04 0.04 - 0.01 179 Amblygobius phalae~ Gobiidae 99.98 4.01 0.04 0.04 - 0.06 180 Caesw erythrogaster Lutjanidae 99.98 4.01 0.04 0.04 - - 0.98 0.14 181* Lethrinus nebulmus bthrinidee 99.98 ~0.01 0.04 0.04 - 182 Clenochaetus binoWus Acanthuridae 99.99 c0.01 0.04 0.04 - - 33.12 7.93 188 Pomacentrus tripunctatus Pomacentridae 99.99 c0.01 0.04 0.04 - - 0.03 8.86 184 Lethrinus sp. bthrirridae 99.99 ~0.01 0.04 0.04 - 185* Choerodon shoenleinii Labridae 99.99 <0.01 0.04 0.04 - 186 Epinephelus fuscoguttatus Serranidae 100.00 4.01 0.04 0.04 -

Others Totals 112

Appendix 7. Reef flat fish families caught at night by a roller beam trawl in the seagrass beds of Santiago Island and sorted by frequency of occurrence (ind/1,000 m2).

Seagrass density and depth Dense Sparse Denso Sparse Rank Family Cum% % Total ~1.5m d.5 m >1.6 m >1.5 m

1 Apoaonidae 36.81 462.81 121.25 26.57

~fitosidae hthrinidae Syngnathidac Monacanthidao Mullidae Scaridae Blenniidae Tetraodontidae Labridae Percichthyidae Synodontidae Serranidae Atherinidae Gobiidao Mugiloididae Muraenidae Centriscidae Teraponidao Soleidae Lutjanidae Clupeidae Gerreidao Nemipteridao Colocongridac Congiopodidae Scorpaenidae Pseudochromidae Chaetodontidae Sphyraenidae Acanthuridae Pomacentridae Ostraciidae Congridae Antennariidae Haemulidae Platycephalidac Balistidae Carapidae Aulostomidae Fistulariidac Ephippidae Grammistidae Ophichthidae Diodontidae Pempherididae 48 Bothidae 100.00 c0.01

Totals 100.00 1,257.38 482.29 154.29 491.15 128.66 Appendix 8. Fkef flat fish caught at night by a roller beam trawl in the seagrass beds of Santiago Island from October 1988 to June 1991 and sorted by weight (g/1,000 ma).(* denotes uncertain identification). Seagrass density and depth Dense Sparse Dense Sparse Rank Species Family Cum% % Total ~1.5m r1.5 m b1.5 m rl.6 m 1 Fowleria oarie~ata- Apogonidae 2 Arothron immaculatus Tetraodontidae 3 Apogon sangicnsis Apogonidae 4 Siganus fuscescens Siganidae 5 Sphramia orbicularis Apogonidae 6 Syngnathoides biaculeatus Syngnathidae 7 Acreichthys tomedosus Monacanthidae 8 Arothron hispidus Tetraodontidae 9 Centrogenys uaigiensts Percichthyidae 10 Lethrinus hard Lethrinidae 11 Gymnothorax pictus Muraenidae 12 Siganus uirgatus Siganidae 13 Saurida gracilis Synodontidae 14 Apgon coccineus Apogonidae 15 Siganus spinus Siganidae 16 Scarus ghobban Scaridae 17 Petrascirtes breviceps Blenniidae 18 Plotosus lineatus Plotosidae 19 Parupemus barberinus Mullidae 20 Epinephelus rnerra Serranidae 21 Scarus rhoduropterus Scaridae 22 Parapercis cylindrica Mugiloididae 23 Choerodon anchorago Labridae 24 Cheilodipterus quinquelineatus Apogonidae 25 Cdotomus japonicus Scaridae 26 Epinephelus ongus Serranidae 27 Pardachirus pauoninus Saleidae 28 Lethrinus reticulatus Lthrinidae 29 Apogonid sp.5 Apogonidae 30 Siganus pumtatus Siganidae 31 * Lethrinus obsoletus Let hrinidae 32 Conger cinereus Congridae 33 Chelonodon patoca Tetraodontidae 34 Apogon bandm~ensis Apogonidae 35 Lethrinus ornatus Isthrinidae 36 Diodon hystrix Diodontidae 37 Siganus guttatus Siganidae 38 Siganus argenteus Siganidae 39 Apogon arnboi~nsis Apogonidae 40 Scarus longiceps Scaridae 41 Cheilinus trilobatus Labridae 42 Platycephalus indicus Platycephalidae 43 Synaptura marginata Soleidae 44 Cheilio inermis Labridae 45 Hypoatherina bleekeri Atherinidae 46 Arothron stellatus Tetraodontidae 47 Lutjunus fuhiflmma Lutjanidae 48 Upenem hagula Mullidae 49 Ariosomu anqoides Colocongridae 50 Plotosus canius Plotosidae 61 Amblyapistus taenianotus Congiopodidae 52 Apogon sp.5 (Schroeder 1980) Apogonidae 63 Apogon nouemfascidus Apogonidae 64 Parupeneus trifosciatus Mullidae 65 Asterropteryx semipunctatus Gobiidae 66 Apogon cyanosoma Apgonidae 95.96 0.18 8.91 1.31 0.66 0.16 Continued Appendix 8 (Continued) Seagrass density and depth Dense Spame Denw Sparse Rank Species Family Cum% % Total ~1.6m c1.5 m r1.5 m >1.5 m Amthmn nigropunctatus Tetraodontidae Eqt-ias puntahg Gobiidae Lactoria cornuta Ostraciidae Lethrinus lentjan Lethrinidae Parupemus barberimides Mullidae Pelatus quadrilineatus Teraponidae Aluterea scriptus Monacanthidae Parupeneus indicun Mullidae Leptoscarus uaigiensis Scaridae Canthigaster bsnnetti Tetmdontidae Stethojulis strigiventer Labridae ~mbl$gobius~bimmulatus Gobiidae Corythoichthys haematopterus Syngnathidae Scorpaena sp. Scorpaenidas Gerres oyena Gerreidae Dampieria cyclophthalrna Pseudochromidae Synodus variegatus Synodontidae Scorpoempsis cirrhosa Scorpaenidae Lethrinus mahsena Isthrinidae Aeoliscus strigatus Centriscidae Hypodytes rubripinnis Congiopodidae Adostomus chinensis Aulostomidae Archamia lineolata Apogonidae Sphaeramia nernatoptera Apogonidae Dendrochirus zebra Scorpaenidae Plectorhynchus lineatus Haemulidae Grammistes ~xlineatus Grammistidae Scarus prasiognathus Scaridae Dunckerocampus dactyliophorus Syngnathidae Goby Gobiidae Epinephelus fusmguttatus Serranidae Exyrias bellissirnus Gobiidae Scolopsis ciliatus Nemipteridae Goby sp.8 Gobiidae Enchsiliophir verrnicularis Carapidae Yongeichthys criniger Gobiidae Ophichthue sp. Ophichthidae ~Ppocarnpuskuda Syngnathidae Apogonid sp.2 Apogonidae 96 + kthrinus variegatus- Lethrinidae Apogonid sp.7 Apogonidae Sphymena jello Sphyraenidae Hippocampus histrix Syngnathidae Arothron mappa Tetraodontidae Condhigaeter valentini Tetraodontidae Scorpaenid Scorpaenidaa Naso unicornis Acanthuridae Dischistodus chrysopoecilua Pomacentridae Tetraodontid sp.2 Tetraodontidae Goby sp.11 Gobiidae Preudobalistes fireclrs Balistidae Apogon sp.8 (Schroeder 1980) Apogonidae Takifugu rubrips Tetraodontidae Scolopsis bilineatus Nernipteridae Cheilodipterus macrodon Apogonidae Ostracion cubicus O~traciidae 113 Plectorhynchu cktodontoides Haemulidae Continued Appendix 8 (Continued) - Seagrass density and depth Dense Sparse Dense Sparse Rank Species Family Total c1.6 m 4.5 m >1.5 m >1.S m 114 Antennurius molllccemis Antennariidae 0.96 - 115 Chaetodon awiga Chaetodontidae 0.94 0.24 116 Chaetodon melannotus Chaetodontidae 0.88 - 117 Antennatius nurnrnifer Antennariidae 0.85 - 118 Histrio histrio Antennariidae 0.86 - 119 Pterois volitons Scorpaenidae 0.82 0.40 120 Glossogobius diuaceous Gobiidae 0.79 - 121 Lu*nus &uSsatlLa Lutjanidae 0.77 0.37 122 SardineUa sp. Clupeidae 0.76 - 123 Escualosa th.omata Clupeidae 0.66 0.04 124 Lutjanus kmirnira Lutjanidae 0.60 0.60 125 Cheilinus bimoculatus Labridae 0.59 0.14 126 * Lethrinus nentrrtacanthus hthrinidae 0.68 - 127 Halicamphus dunckeri Syngnathidae 0.67 - 128 Aempia cornuta Soleidae 0.57 0.57 129 Antennatius sp.1 Antennariidae 0.53 - 130 Apogonid sp.11 Apogonidae 0.53 - 131 Epinephelus tcuruina Serranidae 0.63 - 132 Bothw pantherinus Bothidae 0.63 - 133 Naso lituratus Acanthuridae 0.52 0.52 134 Goby sp.9 Gobiidae 0.51 - 135 Plaicrr orbicularis Ephippidae 0.49 0.33 136 Pen&podw mcrurus Nemipteridae 0.48 - 137 Pomacentrus tripurntatus Pomacentridae 0.48 0.48 138 Scorpaena sp.1 Scorpaenidae 0.48 0.48 IS9 Amblygobim sp. Gobiidae 0.48 0.48 140 Conger sp. Congridae 0.44 - 141 Amblygobius phalaena Gobiidae 0.44 0.44 142 Sphymena barracuda Sphyraenidae 0.41 - 143 Fistularia petirnba Fistulariidae 0.40 0.24 144 Eleotris fusw Gobiidae 0.40 - 145 Mulloidichthys pauolineatus Mullidae 0.38 osa 146 Pseudobalistes flauimwginatus Balistidae 0.37 - 147 Eupomacentrus lividus Pomacentridae 0.38 0.53 148 Myrichthys aki Ophichthidae 0.32 - 149 Naso sp. Acanthuridae 0.30 - 150 Parupemus heplrrcanfhus Mullidae 0.29 0.10 151 Lutjanus gibbus Lutjanidae 0.26 - 152 Dischistodus mtopthalmus Pomacentridae 0.26 - 153 Bupomacentrus nigricans Pomacentridae 0.26 0.20 154 Drepane longirnana Ephippidae 0.24 0.24 155 Cheilinus fmialus Labridae 0.24 0.24 153 Scolopsis cancellatus Nemipteridae 0.23 0.17 157 Amthron sp.2 Tetraodontidae 0.21 - 158 Corythoichthys schdtzi Syngnathidae 0.21 - 159 Cuesio etythrogmter Lutjanidae 0.20 0.20 160 Siganus puellus Siganidae 0.20 0.20 161 Acanthurus gahhn Acanthuridae 0.20 0.20 162 Pseudomo~canthusmacrurus Monacanthidae 0.20 0.20 163 Choerodon shoenleinii Labridae 0.20 0.20 164 Balistid Balistidae 0.19 - 165 Lutjanus lineolatus Lutjanidae 0.16 0.08 166 * Lethrinus nebulosus Lthrinidae 0.12 0.12 167 Ctenochaetus bimtatus Acanthuridae 0.12 0.12 168 Apogon sp. Apogonidae 0.12 - 169 Scams ovifmns Scaridae 0.11 - 170 Apogonid ~p.10 Apogonidae 0.11 - Continued Appendix 8 (Continued)

Seagrass density and depth Dense Sparse Dense Sparse Rank Species Family Cum% % Total 4.5 m d.5 m >1.5 rn z1.5 m

Pempheris oualemis Pempheridac Acanthurid Acanlhuridae Saurida sp. Synodontidae Oostethus brachyurus Syngnathidae Scarus schlegeli Scaridae Clupeid Clupeidae Scarid Scaridae Chaetodon lunula Chaetodontidae Hippichthys spicifer Syngnathidae Chaetodon trifasciatus Chaetoduntidae Apogonid Apogonidae Abudefduf srucatilis Pomacentridae Plectorhynchus sp. Haemulidae Siganid Siganidae Labrid Lahridac Lethrinus SD. Le thrinidae

Totals Appendix 9. Reef flat fish families caught at night by a roller beam trawl in the seagrass beds of Santiago Island and sorted by weight (g/1,000ma).

Seagrass density and depth Dense Sparse Dense Spame Rank Family Cum% % Total ~1.5m d.5 m >1.5 m 21.5 m

1 Apogonidae 40.08 40.08 2000.01 2 Tetraodontidae 51.96 3 Siganidae 4 Syngnathidae 5 Isthrinidae 6 Monacanthidae 7 mdae 8 Percichthyidae 9 Muraenidae 10 Synodontidae 11 Serranidae 12 Mullidae 13 Labridae 14 Plotosidaa 15 Blenniidae 16 Soleidae 17 Mugiloididae 18 Gobiidae 19 Congridae 20 Diodontidae 21 Platycephalidae 22 Lhtjanidae 23 Congiopodidae 24 Scorpaenidae 25 Atherinidae 26 Colocongridae 27 Ostraciidae 28 Teraponidae 29 Gerreidae 30 Pseudochromidae 3 1 Nemiptmidae 82 Haemulidae 33 Antennariidae 34 Centriscidae 36 Aulostomidae 36 Pomacentridae

Grammistidae Chaetodontidae Ophichthidae Carapidae Sphyraenidae Balistidae Clupeidae Ephippidae Bothidae Fistulariidae 48 Pempherididae 100.00 €0.01

Totals 100.00 4,989.92 1,681.10 338.48 2,227.19 843.16