A rapid marine biodiversity assessment of the coral reefs of northwest Madagascar

Alison Simmance

impetus and direction for the protection of our oceans. In this article we review progress since the Congress and set out some key actions to help the world community pick up the pace to meet commitments and adequately protect some of the largest ecosystems on our planet. THE VTH IUCN WORLD PARKS CONGRESS, a premier international meeting dealing with protected areas, was held in Durban, South Africa in September 2003. The Congress marked a paradigm shift from parks as ‘islands of conservation ’ to ensuring that parks and protected areas provide benefits ‘beyond boundaries’, while also ensuring that such areas continue to serve as the best means globally to contribute to the conservation and sustainable use of biodiversity. The Durban Congress was a significant opportunity to move the marine conservation agenda forward. Whilst only two of the 33 recommendations focused on marine protected areas (MPAs), they were major milestones in re-setting the overall direction and levels of ambit...

31 Participants ..................................................................................... 4 Organizational Profiles ................................................................. 5 Acknowledgments ......................................................................... 7 Executive Summary ....................................................................... 8 Overview ....................................................................................... 11 Map and Photos ........................................................................... 19 Technical Reports ....................................................................... 23 Appendices ................................................................................... 72 Conservation International 1919 M Street, NW Suite 600 Washington, DC 20036 USA TELEPHONE: 202-912-1000 FAX: 202-912-0772 WEB: www.conservation.org www.biodiversityscience.org A Rapid Marine Biodiversity Assessment of the Coral Reefs of Northwest Madagascar A Rapid Marine Biodiversity Assessment of the Coral Reefs of Northwest Madagascar Rapid Assessment Program A Rapid Marine Biodiversity Assessment of the Coral Reefs of Northwest Madagascar Editors Sheila A. McKenna and Gerald R. Allen RAP Bulletin Biological Assessment of 31 Center for Applied Biodiversity Science (CABS) Conservation International Australian Institute of Marine Science 9 0 0 0 0> 9 781881 173847 C ONSERVATION I NTERNATIONAL IS B N 1-881173-84-4 Western Australian Museum Food & Agriculture Organization of the United Nations Protocole of Agreement CI & FAO no PO 1081138 Additional Published Reports of the Rapid Assessment Program South America * Bolivia: Alto Madidi Region. Parker, T.A. III and B. Bailey (eds.). 1991. A Biological Assessment of the Alto Madidi Region and Adjacent Areas of Northwest Bolivia May 18 - June 15, 1990. RAP Working Papers 1. Conservation International, Washington, DC. * Bolivia: Lowland Dry Forests of Santa Cruz. Parker, T.A. III, R.B. Foster, L.H. Emmons and B. Bailey (eds.). 1993. The Lowland Dry Forests of Santa Cruz, Bolivia: A Global Conservation Priority. RAP Working Papers 4. Conservation International, Washington, DC. † Bolivia/Perú: Pando, Alto Madidi/Pampas del Heath. Montambault, J.R. (ed.). 2002. Informes de las evaluaciones biológicas de Pampas del Heath, Perú, Alto Madidi, Bolivia, y Pando, Bolivia. RAP Bulletin of Biological Assessment 24. Conservation International, Washington, DC. * Bolivia: South Central Chuquisaca Schulenberg, T.S. and K. Awbrey (eds.). 1997. A Rapid Assessment of the Humid Forests of South Central Chuquisaca, Bolivia. RAP Working Papers 8. Conservation International, Washington, DC. * Bolivia: Noel Kempff Mercado National Park. Killeen, T.J. and T.S. Schulenberg (eds.). 1998. A biological assessment of Parque Nacional Noel Kempff Mercado, Bolivia. RAP Working Papers 10. Conservation International, Washington, DC. * Bolivia: Río Orthon Basin, Pando. Chernoff, B. and P.W. Willink (eds.). 1999. A Biological Assessment of Aquatic Ecosystems of the Upper Río Orthon Basin, Pando, Bolivia. Bulletin of Biological Assessment 15. Conservation International, Washington, DC. § Brazil: Rio Negro and Headwaters. Willink, P.W., B. Chernoff, L.E. Alonso, J.R. Montambault and R. Lourival (eds.). 2000. A Biological Assessment of the Aquatic Ecosystems of the Pantanal, Mato Grosso do Sul, Brasil. RAP Bulletin of Biological Assessment 18. Conservation International, Washington, DC. § Ecuador: Cordillera de la Costa. Parker, T.A. III and J.L. Carr (eds.). 1992. Status of Forest Remnants in the Cordillera de la Costa and Adjacent Areas of Southwestern Ecuador. RAP Working Papers 2. Conservation International, Washington, DC. * Ecuador/Perú: Cordillera del Condor. Schulenberg, T.S. and K. Awbrey (eds.). 1997. The Cordillera del Condor of Ecuador and Peru: A Biological Assessment. RAP Working Papers 7. Conservation International, Washington, DC. § Guyana: Kanuku Mountain Region. Parker, T.A. III and A.B. Forsyth (eds.). 1993. A Biological Assessment of the Kanuku Mountain Region of Southwestern Guyana. RAP Working Papers 5. Conservation International, Washington, DC. * Guyana: Eastern Kanuku Mountains. Montambault, J.R. and O. Missa (eds.). 2002. A Biodiversity Assessment of the Eastern Kanuku Mountains, Lower Kwitaro River, Guyana. RAP Bulletin of Biological Assessment 26. Conservation International, Washington, DC. * Paraguay: Río Paraguay Basin. Chernoff, B., P.W. Willink and J. R. Montambault (eds.). 2001. A biological assessment of the Río Paraguay Basin, Alto Paraguay, Paraguay. RAP Bulletin of Biological Assessment 19. Conservation International, Washington, DC. Cover photos Top: Nosy Kivinjy Photo by G. Allen Center: New species of damselﬁsh (Pomacentrus caeruleopunctatus) Photos by G. Allen Bottom: Coral reef Photo by J. Veron Central America § Belize: Columbia River Forest Reserve. Parker, T.A. III. (ed.). 1993. A Biological Assessment of the Columbia River Forest Reserve, Toledo District, Belize. RAP Working Papers 3. Conservation International, Washington, DC. * Guatemala: Laguna del Tigre National Park. Bestelmeyer, B. and L.E. Alonso (eds.). 2000. A Biological Assessment of Laguna del Tigre National Park, Petén, Guatemala. RAP Bulletin of Biological Assessment 16. Conservation International, Washington, DC. Asia-Pacific * Indonesia: Wapoga River Area. Mack, A.L. and L.E. Alonso (eds.). 2000. A Biological Assessment of the Wapoga River Area of Northwestern Irian Jaya, Indonesia. RAP Bulletin of Biological Assessment 14. Conservation International, Washington, DC. * Indonesia: Togean and Banggai Islands. Allen, G.R., and S.A. McKenna (eds.). 2001. A Marine Rapid Assessment of the Togean and Banggai Islands, Sulawesi, Indonesia. RAP Bulletin of Biological Assessment 20. Conservation International, Washington, DC. * Indonesia: Raja Ampat Islands. McKenna, S.A., G.R. Allen and S. Suryadi (eds.). 2002. A Marine Rapid Assessment of the Raja Ampat Islands, Papua Province, Indonesia. RAP Bulletin of Biological Assessment 22. Conservation International, Washington, DC. * Indonesia: Yongsu - Cyclops Mountains and the Southern Mamberamo Basin. Richards, S.J. and S. Suryadi (eds.). 2002. A Biodiversity Assessment of Yongsu - Cyclops Mountains and the Southern Mamberamo Basin, Papua, Indonesia. RAP Bulletin of Biological Assessment 25. Conservation International, Washington, DC. * Papua New Guinea: Lakekamu Basin. Mack, A.L. (ed.). 1998. A Biological Assessment of the Lakekamu Basin, Papua New Guinea. RAP Working Papers 9. Conservation International, Washington, DC. † Papua New Guinea: Milne Bay Province. Werner, T.B. and G. Allen (eds.). 1998. A Rapid Biodiversity Assessment of the Coral Reefs of Milne Bay Province, Papua New Guinea. RAP Working Papers 11. Conservation International, Washington, DC. * Papua New Guinea: Southern New Ireland. Beehler, B.M. and L.E. Alonso (eds.). 2001. Southern New Ireland, Papua New Guinea: A Biodiversity Assessment. RAP Bulletin of Biological Assessment 21. Conservation International, Washington, DC. * Papua New Guinea: Milne Bay Province. Allen, G.R., J.P. Kinch, S.A. McKenna and P. Seeto (eds.). 2003. A Rapid Marine Biodiversity Assessment of Milne Bay Province, Papua New Guinea - Survey II (2000). RAP Bulletin of Biological Assessment 29. Conservation International, Washington, DC. † Philippines: Palawan Province. Werner, T.B. and G. Allen (eds.). 2000. A Rapid Marine Biodiversity Assessment of the Calaminanes Islands, Palawan Province, Philippines. RAP Bulletin of Biological Assessment 17. Conservation International, Washington, DC. Africa & Madagascar * Perú: Tambopata-Candamo Reserved Zone. Foster, R.B., J.L. Carr and A.B. Forsyth (eds.). 1994. The Tambopata-Candamo Reserved Zone of southeastern Perú: A Biological Assessment. RAP Working Papers 6. Conservation International, Washington, DC. * Botswana: Okavango Delta. Alonso, L.E. and L. Nordin (eds.). 2003. A Rapid Biological Assessment of the aquatic ecosystems of the Okavango Delta, Botswana: High Water Survey. RAP Bulletin of Biological Assessment 27. Conservation International, Washington, DC. * Perú: Cordillera de Vilcabamba. Alonso, L.E., A. Alonso, T. S. Schulenberg and F. Dallmeier (eds.). 2001. Biological and Social Assessments of the Cordillera de Vilcabamba, Peru. RAP Working Papers 12 and SI/MAB Series 6. Conservation International, Washington, DC. † Côte d’Ivoire: Marahoué National Park. Schulenberg, T.S., C.A. Short and P.J. Stephenson (eds.). 1999. A Biological Assessment of Parc National de la Marouhe, Côte d’Ivoire. RAP Working Papers 13. Conservation International, Washington, DC. * Venezuela: Caura River Basin. Chernoff, B., A. Machado-Allison, K. Riseng and J.R. Montambault (eds.). 2003. A Biological Assessment of the Aquatic Ecosystems of the Caura River Basin, Bolívar State, Venezuela. RAP Bulletin of Biological Assessment 28. Conservation International, Washington, DC. * Guinea: Pic de Fon. McCullough, J. (ed.). 2004. A Rapid Biological Assessment of the Foret Classée du Pic de Fon, Simandou Range, Southeastern Republic of Guinea. RAP Bulletin of Biological Assessment 35. Conservation International, Washington, DC. * Venezuela: Orinoco Delta and Gulf of Paria. Lasso, C.A., L.E. Alonso, A.L. Flores and G. Love (eds.). 2004. Rapid assessment of the biodiversity and social aspects of the aquatic ecosystems of the Orinoco Delta and the Gulf of Paria, Venezuela. RAP Bulletin of Biological Assessment 37. Conservation International, Washington, DC. * Madagascar: Ankarafantsika. Alonso, L.E., T.S. Schulenberg, S. Radilofe and O. Missa (eds). 2002. A Biological Assessment of the Réserve Naturelle Intégrale d’Ankarafantsika, Madagascar. RAP Bulletin of Biological Assessment 23. Conservation International, Washington, DC. * Available through the University of Chicago Press. To order call 1-800-621-2736; www.press.uchicago.edu † Available only through Conservation International. To order call 202-912-1249. § Out of Print Rapid Assessment Program A Rapid Marine Biodiversity Assessment of the Coral Reefs of Northwest Madagascar Editors Sheila A. McKenna and Gerald R. Allen RAP Bulletin Biological Assessment of 31 Protocole of Agreement CI & FAO no PO 1081138 Center for Applied Biodiversity Science (CABS) Conservation International Australian Institute of Marine Science Western Australian Museum The RAP Bulletin of Biological Assessment is published by: Center for Applied Biodiversity Science Conservation International 1919 M St., N.W., Suite 600 Washington, DC 20036 USA 202-912-1000 202-912-0773 fax www.conservation.org Editors: Sheila A. McKenna and Gerald R. Allen Design: Glenda Fábregas Conservation International is a private, non-proﬁt organization exempt from federal income tax under section 501 c (3) of the Internal Revenue Code. ISBN 1-881173-84-4 ©2005 by Conservation International and FAO All rights reserved. Library of Congress Catalog Card Number: 2005923043 The designations of geographical entities in this publication, and the presentation of the material, do not imply the expression of any opinion whatsoever on the part of Conservation International or its supporting organizations concerning the legal status of any country, territory, or area, or of its authorities, or concerning the delimitation of its frontiers or boundaries. Any opinions expressed in the RAP Bulletin of Biological Assessment are those of the writers and do not necessarily reﬂect those of CI. Printed on recycled paper RAP Bulletin of Biological Assessment was formerly RAP Working Papers. Numbers 1-13 of this series were published under the previous title. Suggested citation: McKenna S.A. and G.R Allen, eds. 2003. A Rapid Marine Biodiversity Assessment of Northwest Madagascar. Bulletin of the Rapid Assessment Program 31, Conservation International, Washington, DC. Table of Contents Participants ................................................................................ 4 Chapter 6 ................................................................................... 68 Organizational Proﬁles ............................................................. 5 Community use and conservation of marine resources at northwest Madagascar Salimo and Bakary Gisele Acknowledgments .................................................................... 7 Appendices Executive Summary .................................................................. 8 Appendix 1 ................................................................................ 72 Overview ................................................................................... 11 Map and Photos ....................................................................... 19 Technical Reports Chapter 1 ................................................................................... 23 Part 1: Zooxanthellate Scleractinia of Madagascar Part 2: Reef corals of Northwest Madagascar J.E.N. Veron and Emre Turak Total inventory of species recorded during the present study and previously recorded for the whole of the SW Indian Ocean Appendix 2 ................................................................................ 83 Coral species recorded at individual sites at northwest Madagascar Appendix 3 ................................................................................ 90 Molluscs recorded at northwest Madagascar Appendix 4 .............................................................................. 102 List of the reef ﬁshes of Madagascar Chapter 2 ................................................................................... 32 Molluscs of northwest Madagascar Fred E. Wells Chapter 3 ................................................................................... 39 Reef ﬁshes of northwest Madagascar Gerald R. Allen Chapter 4 ................................................................................... 49 Fisheries of northwest Madagascar Jean Maharavo Chapter 5 ................................................................................... 54 Condition of coral reefs of northwest Madagascar Sheila A. McKenna Coral Reefs of Northwest Madagascar 3 Participants Gerald R. Allen, Ph.D. (Ichthyology and Science Team Leader) Conservation International 1919 M. St., N.W. Suite 600 Washington, DC 20036 USA Mailing address: 1 Dreyer Road Roleystone, WA 6111 Australia Fax: (618) 9397 6985 Email: tropical_reef@bigpond.com Giséle Bakary (Community liaison team) Centre National de Recherches Océanographiques (CNRO) B.P. 68 CNRO Hell-Ville/Nosy Be (207) Madagascar Jean Maharavo, Ph.D. (Reef ﬁsheries) Centre National de Recherches sur l’Énvronment (CNRE) B.P. 1739 39, Rue Rasamimanana Fiadanana 101 Antananarivo, Madagascar Sheila McKenna, Ph.D. (Reef ecology, RAP Survey Team Leader) Director, Marine Biodiversity Analysis Program Center for Applied Biodiversity Science Conservation International 1919 M. St., N.W. Suite 600 Washington, DC 20036 USA Fax: (1) (202) 912 1217 Email: s.mckenna@conservation.org 4 Rapid Assessment Program Bemahafaly Randriamanantsoa (Reef ecology) Institut Halieutique et des Sciences Marines (IHSM) PO Box 141- Route du Port, Mahavatse II - TULEAR 601 Tel/Fax.: (261) 94 435 52; Email: ihsm@syfed.refer.mg Rémi Ratsimbazafy, Ph.D. (Reef ecology) WWF Madagascar B.P. 738 (101) Antananarivo, Madagascar Email: wwfrep@dts.mg Salimo (Community liaison team) GTZ-Madagascar Deutsche Gesellschaft für Technische Zusammenarbeit Emre Turak (Reef corals) CORMEC 1 rue Francois Villon Cergy, France 9500 Email: emreturak@wanadoo.fr John E. N. Veron, Ph.D. (Reef corals) Australian Institute of Marine Sciences P.M.B. No. 3 Townsville, Queensland 4810 Australia Email: j.veron@aims.gov.au Fred E. Wells, Ph.D. (Malacology) Department of Aquatic Zoology Western Australian Museum Francis Street Perth, WA 6000 Australia Email: molluscau@yahoo.com.au Organizational Proﬁles CONSERVATION INTERNATIONAL Conservation International (CI) is an international, non-proﬁt organization based in Washington, DC. CI acts on the belief that the Earth’s natural heritage must be maintained if future generations are to thrive spiritually, culturally, and economically. Our mission is to conserve biological diversity and the ecological processes that support life on earth and to demonstrate that human societies are able to live harmoniously with nature. 1919 M St., N.W., Suite 600 Washington D.C. 20036 USA (202) 912-1000 (telephone) (202) 912-0773(fax) http://www.conservation.org CONSERVATION INTERNATIONAL (MADAGASCAR) 6 Rue Razaﬁndratandra, Ambohidahy B.P. 5178 Antananarivo 101 Madagascar (261) 20 22 60979 (telephone) (261) 20 22 25029 (fax) AUSTRALIAN INSTITUTE OF MARINE SCIENCE The mission of the Australian Institute of Marine Science (AIMS) is to generate the knowledge to support the sustainable use and protection of the marine environment through innovative, world-class scientiﬁc and technological research. It is an Australian Commonwealth Statutory Authority established by the Australian Institute of Marine Science Act of 1972 in recognition of a national need to manage Australia’s marine environment and marine resources. Cape Ferguson, Queensland PMB No 3, Townsville MC QLD 4810 (61-7) 4753-4444 (telephone) (61-7) 4772-5852 (fax) http://www.aims.gov.au Coral Reefs of Northwest Madagascar 5 Organizational Proﬁles WESTERN AUSTRALIAN MUSEUM The Western Australian Museum was established in 1891 and its initial collections were geological, ethnological and biological specimens. The 1960s and 1970s saw the addition of responsibility to develop and maintain the State’s anthropological, archaeological, maritime archaeological and social and cultural history collections. The collections, currently numbering over two million specimens/artefacts, are the primary focus of research by the Museum’s own staff and others. The aim is to advance knowledge on them and communicate it to the public through a variety of media, but particularly a program of exhibitions and publications. Francis Street Perth, WA 6000 Australia (61-8) 9427-2716 (telephone) (61-8) 9328-8686 http://www.museum.wa.gov.au 6 Rapid Assessment Program Acknowledgments This Marine RAP survey was jointly ﬁnanced by Conservation International and United Nation Trust Funds subcontract UNTS/MAG/001/GEF (FAO) to Conservation International (Protocole of Agreement CI & FAO no PO 1081138). CI funding was made possible by generous donations from John McCaw. We are very grateful to Léon Rajaobelina and Sahondra Radilofe, Director and Scientiﬁc Research Ofﬁcer respectively, for CI-Madagascar. We are also thankful to the entire CIMadagascar staff for their generous assistance. We thank Patrick Tesha (former) and M. Martin Smith (current), FAO Representatives for Madagascar, Comores, Mauritius, and Seychelles for their expert guidance and support. Andrew Cooke provided a wealth of information concerning the background of marine conservation in Madagascar. He also played an important role in the formative stage of the RAP survey and helped to prepare the proposal for Madagascar funding from FAO. We also extend our sincere thanks to the captain and crew of Inga Viola for providing logistic assistance. Mark Allen assisted with color scanning and prepared the layout for the color pages appearing in this report. Coral Reefs of Northwest Madagascar 7 Executive Summary INTRODUCTION Madagascar is the beneﬁciary of a multi-donor national environmental action program (PNAE), founded on the Charter for the Environment of 1990 (Law 90-033). The program consists of three ﬁve-year phases: 1992-1996, 1997-2001, and 2002-2006. Phase Two of this program includes a marine and coastal component due to its considerable importance to an integrated national environmental plan, as well as increasing concern for the rapid degradation of certain coastal areas. Moreover, there is also a critical need for more information concerning coastal ecosystems, which hopefully can be used to formulate sound management policies. The goal of the coastal and marine environmental component of Phase Two is to lay the groundwork for the development of a comprehensive conservation and development strategy to achieve the following objectives: 1) Develop and implement effective mechanisms for the establishment of an integrated plan of coastal zoning (GIZC); 2) Initiate actions for integrated management at the local, regional, and national levels, which would be ultimately integrated into a single, national plan; and 3) Obtain the necessary knowledge base for marine and coastal systems that is required to realize Objectives 1 and 2. A national committee, the Group for Marine and Coastal Conservation Initiatives (AMCC), was formed to implement a network of marine and coastal protected areas. The principal members of this group are the National Ofﬁce of the Environment (ONC), the National Association for Management of Protected Areas, the Minister of Fisheries and Marine Resources (MPRH), the Minister of the Environment, UNESCO, and several international NGOs, including Conservation International, World Wide Fund for Nature, and the Wildlife Conservation Society. This group has begun the process of formulating a national strategy and action plan for Marine Protected Areas. One of its major objectives is to gather biological and other relevant information for the prioritization of potential marine and coastal reserves. Marine Protected Areas (MPAs) are a well known, important component for the management and conservation of sustainable resources and general biodiversity. As well as their management beneﬁts, MPAs may serve as critical sanctuaries for a variety of marine organisms. They also form a buffer for maintaining an area’s biodiversity and encourage the development of low-impact economic activities such as ecotourism. Additionally, they are priority sites for scientiﬁc research. Presently there are only two national MPAs in Madagascar: the reefs around Nosy Atafana in the Biosphere Reserve of Mananara-Nord and the three marine reserves at Masoala National Park. Both of these parks are situated in the northeastern region. Although they have been the focus of signiﬁcant biological research, further studies are needed. In the southeastern part of the country, the coral reefs around Toliara have been the focus of important marine biological studies since the 1960s. The other major reef area on the western coast is situated in the far north and, in contrast to the southeast, is poorly known. Despite its reputation for possessing some of the country’s ﬁnest coral reefs, few studies have been done in this area. Many conservationists feel that the Northwest may have the greatest potential for the development of MPAs and concomitant development of marine ecotourism. Basic biological knowledge of this region is therefore essential to facilitating the MPA implementation process. 8 Rapid Assessment Program Executive Summary Conservation International’s RAP survey is intended to assess the biological importance of the Northwest and make recommendations concerning the best areas for conserving general marine biodiversity. This project was assigned by FAO Madagascar via a sub-contract (FAO project code UNTS/MAG/001/GEF) to Conservation International (Protocole of Agreement CI & FAO no PO 1081138). The objectives of this project were to carry out a rapid evaluation of the coral reefs of northwest Madagascar in terms of their biodiversity, health status, ﬁshing resources and community use, and conservation of marine resources in order, among other things, to: 1) identify the priority sites for marine biodiversity conservation; 2) strengthen national capacities in the coral reefs rapid evaluation techniques; 3) provide speciﬁc support to development projects in the new marine protected areas of the region and, 4) carry out joint publications with the national teams on the project results so as to promote international knowledge of the marine biodiversity of Madagascar. The survey area (see map) covered an approximate linear distance of 220 km along the northwestern coast, encompassing Nosy Iranja, Ambavatoby Bay, vicinity of Nosy Be, Mitsio Islands, Cape St. Sebastien, and Andranomaimbo Bay as far north as Nosy Hao. The selected survey sites were accessed by auxiliary motor boats operating from a liveaboard charter vessel. SUMMARY OF RESULTS The reefs of northwestern Madagascar are relatively remote; hence the shortage of basic biological information on them. Nevertheless, the survey indicated that the area has good potential for conservation initiatives, including the establishment of MPAs. Notable results from the RAP survey include: • Overview of Marine RAP The goal of Marine RAP is to rapidly generate and disseminate information on coastal and near-shore shallowwater marine biodiversity for conservation purposes, with a particular focus on recommending priorities for conservation area establishment and management. Marine RAP deploys multi-disciplinary teams of marine scientists and coastal resource experts to determine the biodiversity signiﬁcance and conservation opportunities of selected areas. Through underwater inventories generally lasting three weeks, Marine RAP surveys produce species lists that serve as indicators of overall biological richness. Surveys also record several measurements to assess overall ecosystem health. During each survey, RAP supports parallel assessments of local human community needs and concerns, which are incorporated into the ﬁnal recommendations. By comparing the results obtained from many surveys, Marine RAP focuses on ensuring that a representative sample of marine biodiversity is conserved within protected areas and through other conservation measures. The Northwest Madagascar Survey The Marine RAP survey of northwestern Madagascar assessed 30 sites over a 16-day period (11-26 January 2002). General site areas were selected prior to the actual survey in order to maximize the diversity of habitats visited, thus facilitating a species list that incorporates maximum biodiversity. These areas were also recommended by a national committee, the Group for Marine and Coastal Conservation Initiatives (AMCC), as potential areas for marine conservation initiatives. At each site, an underwater inventory was made of three faunal groups selected to serve as indicators of overall coral reef biodiversity: scleractinian corals, molluscs, and reef ﬁshes. Additional observations were made on the environmental condition of each site, including evaluation of various threat parameters. Observations and data on reef ﬁsheries were also gathered. • • • • Corals: 318 coral species were recorded at sites surveyed, including seven new species and many range extensions. This total greatly exceeds previous estimates and approximates the predicted total of 340 species for the entire western Indian Ocean. Molluscs: 525 molluscs were recorded, which is similar to areas that are much closer to the “Coral Triangle” center of marine diversity in the Indo-Australian region. Reef Fishes: A total of 463 species were recorded including two new species of damselﬁshes (family Pomacentridae). A review of the literature, combined with results of the current RAP survey, reveal a total reef ﬁsh fauna for Madagascar of 752 species. Reef Fisheries: About 55 reef ﬁsh species are targeted by local ﬁshers. The most common families include Serranidae, Lutjanidae, Caesionidae, Lethrinidae, Haemulidae, and Scaridae. Shark stocks appear to be declining, a trend that has been observed over several years. Stocks of commercially important holothurians also appear to be scarce. Reef Condition: Based on CI’s comparative reef condition rank (RCR), the majority of the reef sites surveyed were in relatively good condition. The most frequently observed damage was from the coral predator Acanthaster plancii and appeared to be localized. CONSERVATION RECOMMENDATIONS Madagascar’s marine fauna has been overshadowed by its wealth of unique terrestrial wildlife. Consequently most conservation efforts to date have focused on the important terrestrial aspect of its natural heritage. The relatively recent attention to Madagascar’s marine ecostystems is timely because of the increasing need to protect the island nation’s diverse coastline and the biodiversity it contains. In order to achieve this goal there remains considerable scope for biological assessment surveys that can assist with the process of establishing a network of MPAs. As a result of our RAP survey we make the following speciﬁc recommendations. Coral Reefs of Northwest Madagascar 9 Executive Summary 1) Pursue a vigorous program of biological assessment. The coastline extends for approximately 5,000 km, and the estimated length of all reefs (including fringing reefs, islands, platforms, and both emergent and submerged barrier reefs) is 3,459 km (Cooke et al. 2000). Such an extensive area necessitates a series of biological assessment surveys to aid the process of establishing the best possible network of Marine Protected Areas. Minimally, this would entail at least six surveys, covering the primary coral reef areas. It is important to assess all reef habitats including coastal bays and headlands, near-shore islands, offshore islands, platform reefs, and outer oceanic banks. Eventual reserves should be designed to embrace representatives of all reef types and exposure situations, thus ensuring that the full range of coral reef biodiversity is conserved. 2) Review and consolidate previous knowledge of Madagascar’s marine biota. Considerable research has been conducted involving the marine fauna and ﬂora, particularly at ﬁeld stations located at Toliara and Nosy Be. This knowledge should be compiled and consolidated to avoid a duplication of effort. Moreover, these previous studies will greatly compliment future ﬁeld assessments for conservation purposes. 3) Collection of additional data and long-term monitoring programs essential for marine conservation planning and implementation. Marine biological studies, particularly those with conservation implications (including taxonomic studies) need to be encouraged, possibly through grants to Madagascar university students or by inviting foreign experts to tackle problems of local signiﬁcance. Long-term monitoring programs that include oceanographic and biological data should be set up to follow the status of the marine resources and provide a guide to their use (e.g., ﬁsh stocks and subsequent ﬁshing quota) as well as an early warning system of threats to their existence (e.g., sea surface temperature and bleaching). Studies involving the connectivity of reef communities around the island would be especially useful in designing a system of MPAs. 4) Establish a network of marine protected areas. The establishment of a sound network of MPAs should remain a prime objective. The location of protected areas should follow scientiﬁc guidelines to ensure that they cover a representative sample of the region’s marine biodiversity. Target areas should include those that are exceptionally diverse, physically unique, and known to protect currently or potentially rare or endangered species as well as Madagascar endemics. MPAs should be established on the basis of both local and regional distributions of marine biodiversity. Furthermore, areas known to be resilient to certain stresses such as bleaching should be included in MPAs. 5) Enact more effective laws to regulate ﬁshing activities. More precise laws may need to be enacted, particularly at the 10 Rapid Assessment Program local level, to cover all aspects of ﬁshing and environmental destruction. This legislation should be sensitive to cultural values of local people, including their system of taboos related to the natural environment. The efﬁcacy of existing laws also needs to be examined. For example, the socioeconomic team identiﬁed the opening and closing dates of the shrimp ﬁshing season in Ambaro Bay as an issue of concern. 6) Promotion and development of conservation-oriented marine tourism for the beneﬁt of local communities. Madagascar is relatively unknown to the global tourist diving market and has considerable potential for promotion and development. Dive tourism and marine conservation are compatible and complimentary activities. As the country establishes a network of MPAs, an effort should be made to involve the dive industry and encourage the development of new resorts and dive concessions within them. Any such endeavor should also involve local communities in the form of employment opportunities and the payment of nominal fees for the privilege of diving in traditional ﬁshing grounds. 7) Provide dive training for staff of local universities and conservation organizations. In order to encourage local biologists to do more to promote marine conservation values, more of them should be trained to dive. This will foster a greater appreciation for the undersea environment. 8) Set up outreach and public awareness programs on marine resources to all local stakeholders. The importance of protecting and sustaining the marine environment needs to be taught in villages, schools, and other venues. Raising the awareness of the community as to what makes a reef healthy, what damages a reef, and what the beneﬁts of protection are will lead people to take action. 9) Promote community participation in conservation planning and management. Local communities need to be involved in the implementation and ongoing management of MPAs. Community participation could be encouraged by establishing and helping to ﬁnance (via government, private corporations, and NGOs) various outreach programs that involve educational assistance, health care, and a general raising of living standards. This type of aid would provide real rewards for villages that successfully implement reef management programs. 10) Strengthen species conservation programs for rare and endangered marine wildlife. The status of rare, endangered, and endemic species needs to be assessed to ensure that adequate habitat for their continued survival is protected. Their direct exploitation and trade must also be controlled. Marine species that appear to be particularly vulnerable include sharks, sea turtles, dugong, giant clams, and certain ﬁshes (e.g., Napoleon wrasse and large groupers). Overview INTRODUCTION Madagascar is the world’s fourth largest island, covering 587,045 square kilometers. It is sometimes described as a living laboratory of evolution in reference to its incredible abundance of endemic wildlife. Indeed, 91% of its reptiles, 80% of its ﬂowering plants, 74% of its mammals, and 46% of its birds are unique to the island (Mittermeier et al. 1999). Among its most outstanding biological attributes are its 51 species of endemic lemurs and more than 60 species of chameleons. Although overshadowed by this treasure trove of terrestrial wildlife, the marine environment of Madagascar also deserves attention. The extensive coastline, spanning more than 5,000 km, harbors a wealth of biodiversity. Reef building corals are particularly well represented. The present RAP survey documented more than 300 species, surpassing the previous total for the entire western Indian Ocean. Coastal waters also abound with ﬁshes, including approximately 750 reef-associated species and perhaps another 250 species that occur in mangroves, trawling grounds, and deep offshore habitats. Terrestrial conservation has deservedly been the main concern for many decades. In spite of its abundance of unique wildlife, Madagascar’s remaining natural habitat is under severe threat due to a host of factors of which the most prominent are deforestation, overgrazing, and severe soil erosion. Considering this background, marine issues have taken a back seat to the more immediate concern of terrestrial conservation. However, there has been a gradual realization over the past decade that marine habitats are also under increasing threat. Thus far only two national marine protected areas (MPAs) have been established in Madagascar. Both are in the northeastern province—one at Mananara-Nord and another at Masoala. Mananara-Nord was Madagascar’s ﬁrst marine national park, having been gazetted by Presidential Decree (89/216) in 1989. It extends along a 50-km stretch of coast and includes two strict conservation zones with national park status, a circular 1,000 ha marine zone centered on Nosy Atafana, and a considerably larger terrestrial forest zone. The marine park is partially protected, with ﬁshing access for local residents on alternate days. A recent survey conﬁrmed that this protected area is in better condition than adjacent non-protected areas, and is characterized by higher ﬁsh diversity (Randrianamantsoa & Brand 2000). The MPA at Masoala consists of three marine reserves totalling 10,000 ha of coral reef, lagoon, and open water habitats. Although not yet fully functional, zoning plans have been negotiated with local stake holders, including small “no take”zones totalling about 0.75 ha for each reserve. Two locally protected marine areas have also been established. Nosy Tanikely, near Nosy Be, was declared a marine reserve in 1968. The island is surrounded by a ﬂourishing coral reef. It has a total reserve area of about 100 ha and a no-ﬁshing zone that extends for 500 m from the low-tide line. The second reserve is at Nosy Ve in the Toliara region. It consists of a small coral cay with a land area of about 24 ha. The island is surrounded by a kidney-shaped ring of reef, which along with the lagoon is designated as a controlled ﬁshing area that includes a 4 ha no-take zone. The area has been protected since July 1999. In 1999, an ecoregional approach was proposed by the Association Nationale pour la Gestion des Aires Protégées (ANGAP 2001) under the auspices of the Minister of the EnviCoral Reefs of Northwest Madagascar 11 Overview ronment. Six principal marine/coastal eco-regions were identiﬁed (north, northeast, east, southeast, southwest, and west) with the possibility of reﬁnement as additional data becomes available. In addition, a new Marine Protected Area (MPA) initiative was launched in late 1999 by ANGAP, the Ofﬁce of the Environment (ONE), and various NGOs (CI, WWF, and WCS). Under this plan 20 potential MPAs were identiﬁed based on their extraordinary biodiversity or the presence of special organisms such as turtles, lemurs, and sea birds. In spite of a considerable body of marine biological knowledge (particularly for the Toliara and Nosy Be areas) that has already been gathered, basic biodiversity assessment is urgently needed. The present RAP represents a ﬁrst response to this need, which is critical in order to capture and preserve the island’s outstanding biodiversity by means of an eventual network of MPAs. The RAP survey was conducted entirely within the northern region, which extends from Cap St. André on the northwest coast to Cap du Diable in the northeast. According to the Plan de Gestion du Reseau National de Aires Protégées de Madagascar (ANGAP 2001), the islands, coral reefs, and lagoons have the highest conservation priority within this region. This plan includes a number of locations that have been targeted for further conservation assessment including Nosy Hara island complex, Mitsio Islands, Nosy Tanikely, Nosty Sakatia, Nosy Iranja, Radama Islands, and the Sahmalaza Peninsula. This report presents the results of a Conservation International Marine RAP (Rapid Assessment Program) survey of marine biodiversity of northwestern Madagascar with emphasis on selected faunal groups, speciﬁcally reef-building (scleractinian) corals, molluscs, and ﬁshes. Additional chapters present the results of ﬁsheries and reef condition surveys, as well as a study of marine resource use by local communities. This project was assigned and partially ﬁnanced by FAO Madagascar via a sub-contract (FAO project code UNTS/MAG/001/GEF) to Conservation International (Protocole of Agreement CI & FAO no PO 1081138). The purpose of this report is to document local marine biodiversity and to assess the condition of coral reefs and the current level of marine resource use and conservation in order to guide regional planning, marine conservation, and the use of sustainable marine resources. Marine RAP Areas of global importance for wildlife conservation obviously need to be identiﬁed, but the data required for this purpose are often unavailable, considering that many of the world’s remote regions are inadequately surveyed. Scarcity of data, especially in the form of basic taxonomic inventories, is a problem particularly for tropical ecosystems. Hence, Conservation International has developed a technique for rapid biological assessment. The method essentially involves sending a team of taxonomic experts into the ﬁeld for a brief period, often 2-4 weeks, in order to obtain an overview of the area’s ﬂora and fauna. Although most surveys to date have involved terrestrial systems, the method is equally applicable for marine and freshwater environments. 12 Rapid Assessment Program A key difference in the way terrestrial and tropical marine localities are evaluated for conservation potential is the relative emphasis placed on species endemism. Terrestrial conservation initiatives frequently focus on localities or regions with high rates of species endemism. Other aspects need to be addressed, of course, but endemism is often treated as one of the most important criteria for assessing a terrestrial area’s conservation worth. Indeed, it has become a universal measure for evaluating and comparing conservation “hot spots.” In contrast, coral reefs and other tropical marine ecosystems frequently exhibit relatively low levels of endemism. The considerable homogeneity found in tropical inshore communities is in large part due to the pelagic larval stage typical of most marine organisms. For example, reef ﬁsh larvae are commonly pelagic for periods ranging from 9 to 100 days (Leis 1991). A general lack of physical isolating barriers and numerous island “stepping stones” have facilitated the wide dispersal of larvae throughout the Indo-Paciﬁc. Thus, the most important feature to assess in determining the conservation potential of a marine location devoid of signiﬁcant endemism is overall species richness or biodiversity. Additional data relating to relative abundance are also important. Other factors requiring assessment are more subjective and depend largely on the observer. Obviously, extensive biological survey experience over a broad geographic range yields the best results, enabling the observer to recognize any unique assemblages within the community, unusually high numbers of normally rare taxa, or the presence of unusual environmental features. Finally, imminent threats such as explosive ﬁshing, use of cyanide, over-ﬁshing, and nearby logging activities need to be considered. Reef corals, ﬁshes, and molluscs are the primary biodiversity indicator groups used in Marine RAP surveys. Corals provide the major environmental framework for ﬁshes and a host of other organisms. Without reef-building corals, there is limited biodiversity. This is dramatically demonstrated in areas consisting primarily of sand, rubble, or seaweeds. Fishes are an excellent survey group as they are the most obvious inhabitants of the reef and account for a large proportion of the reef ’s overall biomass. Furthermore, ﬁshes depend on a huge variety of plants and invertebrates for their nutrition. Therefore, areas rich in ﬁshes invariably have a wealth of plants and invertebrates. Molluscs represent the largest phylum in the marine environment, the group is relatively well known taxonomically, and they are ecologically and economically important. Mollusc diversity is exceedingly high in the tropical waters of the Indo-Paciﬁc, particularly in coral reef environments. Gosliner et al. (1996) estimated that approximately 60% of all marine invertebrate species in this extensive region are molluscs. Molluscs are particularly useful as a biodiversity indicator for ecosystems adjacent to reefs where corals are generally absent or scarce (e.g., mud, sand, and rubble bottoms). Overview PHYSICAL ENVIRONMENT Cooke et al. (2000) summarized physical aspects of the Madagascar coast. The northwestern coast is a scenic region with forest-clad hills rising behind predominantly mangrove-lined embayments. The latter range in size from the broad gulf south of Nosy Be (Baie d’Ampaindava) to narrow indentations near Cap d’Ambre at the extreme northern tip. The continental shelf is relatively broad along most of the west coast, but abruptly narrows north of Cap Saint Sébastien. For example, in the vicinity of the Mitsio Islands the shelf extends seaward for approximately 70 km, in vivid contrast to Cap d’Ambre, where it is nearly absent. Although there is limited coral reef development along the coast, the best examples of this habitat lie offshore, mainly around islands and cays or on numerous shallow banks. The largest of these is Banc de Leven, which lies about 40 km west-northwest of the Mitsio Islands. The larger islands such as Nosy Be and the Mitsio Group have volcanic origins, but there are also a number of weather-sculptured, scenic limestone islets north of Cap Saint Sébastien. Some of the least explored reefs are situated on oceanic banks near the edge of the continental shelf. Most of the reefs that were surveyed during the current RAP were of the fringing variety, but we also dived on three platform reefs (sites 3, 10, and 30), and a submerged shoal (site 28) near the edge of the continental shelf. Madagascar experiences a monsoon regime of winds and rainfall. Winds are predominantly from the southeast, but the central mountains have a moderating effect so that winds tend to be variable and generally milder on the west coast than on the east coast. The Nosy Be area, which is protected by the Tsarantana Massif, is the least windy. The windiest period in that area is generally between September and December (Cooke et al. 2000). The rainy season or Northeast Monsoon extends from November to March, with relatively dry conditions prevailing during the Southeast Monsoon (April to October). Madagascar’s largest rivers drain along the western seaboard, delivering massive amounts of sediment into the sea during the rainy season. The island is also strongly affected by cyclones at this time, with an average of 1.3 per year between 1920 and 1972 (Cooke et al. 2000). The predominant sea current along the northwestern coast is a northward-ﬂowing, counter clock-wise gyre that occupies the northern one-third of the Mozambique channel and rotates around the Comores. It contacts the Madagascar coast north of Cap St. André and is eventually deﬂected towards the African coast at the northern tip of the island, where it contacts a north-ﬂowing branch of the South Equatorial Current. This current pattern is no doubt responsible for the great similarity in marine biota between Madagascar and the Comores. Tidal ranges are greatest between Toliara and Nosy Be, both on the west coast, with a mean spring range of 3.8 m. Periodic strong currents are common throughout the area, especially in channels between islands and around offshore shoals. Sea temperatures on Madagascar exhibit relatively wide variation due to the effects of season and latitudinal gradient (spanning almost 14° C). Mean annual surface temperatures range between 22°C and 28°C, with the warmest temperatures being recorded in the northwestern region during the Northeast Monsoon (Cooke et al. 2000). During the present survey the average sea temperatures was 29°C. SOCIOECONOMIC ENVIRONMENT Madagascar is truly a racial melting pot. The population consists of a unique blend of Malay-Indonesian (Merina and related Betsileo), Cotiers (mixed African, MalayoIndonesian, and Arab ancestry, i.e., Betsimisaraka, Tsimihety, Antaisaka, Sakalava), French, Indian, Creole, and Comoran. The ofﬁcial languages are Malagasy and French. Fifty-two percent of the people hold indigenous beliefs (including a signiﬁcant ancestor worship component), 41% are Christians, and 7% are Muslims. Almost half the population is under the age of 15, and it is estimated that the population will double by the year 2025. Unfortunately, the island’s population dynamics are placing a severe strain on the economy as well as the environment, with about 75% of the people living in rural areas. Madagascar is one of the poorest countries in the world with a yearly per-capita income of about US $230. Child malnutrition is a major problem to the degree that approximately half the children under the age of three have growth abnormalities due to lack of proper food. Agriculture is the mainstay of the population, yet the intense slash-andburn methods are destroying the environment, particularly with regard to ﬂooding and soil erosion. Human settlement in the survey area is relatively sparse, consisting of widely scattered, small villages. Fishing is invariably important at all the villages with an estimated 6075% of adult males engaged in this activity. Women also ﬁsh from the beach and participate in ﬁsh drying. About 80% of the ﬁshing is conducted by seasonal immigrant ﬁshermen using mainly dugout canoes (with sails) and small motor boats. The primary methods include hook and line, ﬁsh traps, nets, and beach seines. SURVEY SITES AND METHODS A general lack of biological knowledge for the extreme northwestern coast of Madagascar (with the exception of Nosy Be) was the primary basis for our selection of this area. Dive sites were generally located in areas of potential conservation signiﬁcance as previously determined by ANGAP (see above). We also consulted pertinent nautical charts (particularly British Admiralty charts 706, 3872, and 3873 and French Hydrographic Service charts 4379, 5476, and 5704) for details of reef topography. Actual site selection was accomplished upon arrival at the general area, and was further inﬂuenced by weather and sea conditions. Coral Reefs of Northwest Madagascar 13 Overview At each site, the biological team conducted underwater assessments that produced species lists for key coral reef indicator groups. General habitat information was also recorded, as was the extent of live coral cover and other benthic substrata/biota at various depths. The main survey method consisted of direct underwater observations by scientists, who recorded species of corals, molluscs, and ﬁshes. Visual transects were the main method for recording ﬁshes and corals in contrast to molluscs, which relied primarily on collecting live animals and shells (most released or discarded after identiﬁcation). Relatively few specimens were preserved for later study, and these were invariably species that were either too difﬁcult to identify in the ﬁeld or were undescribed. Further collecting details are provided in the chapters dealing with corals, molluscs, and ﬁshes. Concurrently, the ﬁsheries and reef condition team used a 100 m-line transect placed on top of the reef to record substrate details and approximate biomass of commercially important (target) species, as well as observations on key indicator species (for assessing ﬁshing pressure) such as groupers and snappers. Additional information on marine resources utilization was obtained from informal interviews with villagers conducted by a community liaison team. The expedition was conducted aboard the live-aboard diving vessel Inga Viola, based at Nosy Be. It was equipped with scuba tanks and an air compressor, as well as two small auxiliary motor boats, used for transport to nearby dive sites. Table A. Summary of survey sites for Marine RAP survey of northwestern Madagascar. 14 No. Date Location Latitude Longitude 1 11/1/02 S Tsarabajina 13°02.18’S 48°32.69’E 2 11/1/02 NW Tsarabajina 13°01.83’S 48°32.41’E 3 12/1/02 Marie Bank 12°56.09’S 48°32.28’E 4 12/1/02 Maromby Village 12°52.90’S 48°32.60’E 5 13/1/02 Ankarea 12°50.78’S 48°34.39’E 6 13/1/02 Nosy Fisaka 12°49.37’S 48°35.43’E 7 13/1/02 NW corner Mitsio 12°49.35’S 48°36.63’E 8 14/1/02 Kalomisampa Rocks 12°25.34’S 48°39.45’E 9 14/1/02 Ambariokisimany 12°26.90’S 48°42.12’E 10 15/1/02 Magnet Platform 12°10.73’S 48°55.16’E 11 15/1/02 SE Nosy Hao 12°07.39’S 49°03.42’E 12 16/1/02 NE Nosy Hao 12°05.87’S 49°03.63’E 13 16/1/02 Nosy Hao channel 12°07.17’S 49°03.28’E 14 17/1/02 Andavakalovo 12°14.23’S 48°58.82’E 15 17/1/02 Lakandava (N beach) 12°15.74’S 48°57.77’E 16 17/1/02 Ambatomaranitra 12°15.26’S 48°57.84’E 17 18/1/02 Lakandava (S wall) 12°15.58’S 48°57.27’E 18 19/1/02 Nosy Fanihy 13°10.89’S 48°14.05’E 19 19/1/02 Befotaka Bay 13°15.13’S 48°12.80’E 20 20/1/02 NE Sakatia 13°17.93’S 48°10.65’E 21 20 /1/02 SE Sakatia 13°19.22’S 48°09.42’E 22 22/1/01 W Ambavatoby Bay 13°33.18’S 47°59.44’E 23 22/1/01 Makamby Point 13°31.47’S 48°00.72’E 24 22/1/01 E Ambavatoby Bay 13°33.10’S 48°01.38’E 25 23/1/01 S Nosy Iranja 13°36.66’S 47°49.81’E 26 24/1/01 Nosy Kivinjy 13°30.25’S 47°57.12’E 27 24/1/01 Nosy Kisimany 13°34.10’S 48°06.06’E 28 25/1/02 5 Metre Bank 13°23.59’S 48°03.28’E 29 26/1/02 W Nosy Tanikely 13°29.08’S 48°14.39’E 30 26/1/02 Souzy Bank 13°25.28’S 48°12.66’E Rapid Assessment Program Overview Details for individual sites are provided in the reef condition section (Chapter 5). Table A provides a summary of sites. Their location is also indicated on the accompanying map. RESULTS Biological diversity Our brief RAP survey indicates a relatively rich and varied marine fauna associated with the northwestern reefs. Totals for the three major indicator groups are presented in Table B. Table B. Summary of fauna recorded during the RAP survey. Faunal Group No. Families No. Genera No. Species Reef corals 17 62 318 Molluscs 92 193 525 Fishes 65 191 463 Highlights of the RAP survey are presented below. Detailed results are included in separate chapters for corals, molluscs, ﬁshes, reef ﬁsheries, reef condition, and community use of marine resources. Corals – A diverse coral fauna was recorded at sites surrveyed consisting of 318 species, which approximates the predicted total of 340 species for the entire East African/ Western Indian Ocean region (Veron 2000). This remarkable total greatly exceeds the previous estimate of 200 scleractinian species (Cooke et al. 2000). It is comparable to certain areas close to the center of the Indo-Paciﬁc region, such as the northern Great Barrier Reef of Australia. Two-thirds of the known genera and all of the known families from the Indo-Paciﬁc were recorded during this survey. In addition, seven new species were collected and many new range extensions were noted. More than 100 coral species were recorded at most sites, with an average of 103 species per site. The richest sites included Nosy Ankarea (143 species), Nosy Fisaka (129), Nosy Sakatia NE (125), Befotaka Bay (123), and Andavakalolo (122). The Mitsio Islands had the highest overall diversity. Sites on reefs with extensive or well-developed reef ﬂats had the highest coral diversity, while submerged reef banks had the lowest. Molluscs – A total of 525 species of molluscs were identiﬁed. This diversity is consistent with that recorded on similar surveys in areas much closer to the recognized center of Indo-Paciﬁc marine diversity (i.e., the area known as the “Coral Triangle”). For example, 313 species were recorded at Christmas Island, 380 at the Cocos-Keeling Islands, and 433 species at Ashmore Reef off northwestern Australia. Clearly, a short survey will not detect all species in a region, and the actual diversity is greater. Cooke (pers. comm.) reports that French surveys in the 1960s and 1970s recorded a total of 1400 species of molluscs in the region of Toliara. Mollusc species recorded at each site ranged from 49 to 93, with a mean of 72.3. Most sites had totals exceeding 60 species. Marie Bank (site 3) was the richest site with 93 species. Southeastern Sakatia (site 21), Nosy Kisimany (site 27), and Nosy Fanihy (site 18) were nearly as rich with 89-91 species. Fishes – A total of 463 species were recorded during the survey. An extrapolation method utilizing six key index families (Chaetodontidae, Pomacanthidae, Pomacentridae, Labridae, Scaridae, and Acanthuridae) indicates a total fauna for northwestern Madagascar consisting of at least 576 species. Species numbers at visually sampled sites ranged from 33 to 166, with an average of 117 per site. The richest sites for ﬁshes were Nosy Hao (sites 12-13), Nosy Tanikely (site 29), Nosy Ankarea (site 5), and Nosy Kivinjy (site 26), with species totals ranging between 152 and 166. A review of the literature, combined with results of the current RAP survey, reveal a total reef ﬁsh fauna for Madagascar consisting of 752 species. A comprehensive checklist is included with this report. In terms of number of species and general faunal composition, Madagascar is similar to other insular localities in the western Indian Ocean including Mauritius, Seychelles, Chagos, and Maldives. Fisheries – Approximately 55 reef ﬁsh species in 18 families are targeted by local ﬁshers. Estimates were made of their abundance and biomass at the various survey sites. The most common families include Serranidae, Lutjanidae, Caesionidae, Lethrinidae, Haemulidae, and Scaridae. The richest sites for target species included Nosy Tanikely (site 29), Sarabajina (sites 1-2), and Cape Saint Sebastian (site 8). Shark stocks are apparently declining, a trend that has been observed over several years. Only a single individual was observed during the reef ﬁsh assessment transects. Stocks of commercially important holothurians were also scarce. Only two individuals (both Holothuria nobilis) of the four exploited species were observed during the entire survey. Reef condition – According to CI’s comparative reef condition rating (RCR), the majority of sites were in relatively good condition. Reef survey sites with the highest RCR included Anakarea (site 5), Northeast Nosy Hao (site 12), Nosy Fanihy (site 18), and Nosy Tanikely (site 20). The most commonly observed disturbance, at 20% of the reef sites surveyed, was from the coral predator, Acanthaster plancii. The extent of damage from this coral predator ranged from light to extensive. A very low incidence of coral colonies (one to three per site) exhibiting early stages of bleaching was observed on 20% of the reefs surveyed. Evidence indicating possible stress from light sedimentation and ﬁshing activity was noted during the survey. Further long term Coral Reefs of Northwest Madagascar 15 Overview monitoring and study over a year or more would be needed to determine if these observations are signiﬁcant and the extent of the impact. No excessive sedimentation stress was observed (i.e., dead reef substrate covered in silt) on any of the reefs surveyed. Community issues A strong community-based approach is the only way to effectively protect and manage reefs in a sustainable manner. Accordingly, the community liaison team visited 15 villages in the survey area. Initial contacts were made with the head of the village, and three-four ﬁshermen from each village were then interviewed. Fishermen generally felt that ﬁsh stocks are declining both in terms of the size and abundance of individual ﬁshes. Migrant ﬁshermen also pose a problem because they seldom respect local beliefs and taboos related to the natural environment. This lack of respect results in frequent conﬂicts. Other major problems facing most villages include poor health standards, few educational opportunities (90% of ﬁshermen are illiterate), a shortage of potable water, erosion due to mangrove destruction (particularly acute in far northern areas), and a lack of economic resources. Economic problems are exacerbated by the practice among itinerant ﬁshermen of selling their catches and taking the money elsewhere rather than investing in the local economy. Financial aid is acutely needed to improve living standards. Many wish that the government would do more to manage local marine resources. Outstanding sites Several sites were observed to have exceptional biodiversity, outstanding or unusual reefs or structure, attractive scenery (both above and below the sea surface), or a combination of these features. These sites have considerable potential as possible MPAs. At the very least, they should be included within larger marine reserves. and coral, and good visibility. The reef next to shore on the southern side of the island is particularly scenic and is ideal for snorkelling tourists. The island and surrounding reefs in combination with the Nosy Fisaka reef complex (site 6) and the northwestern tip of Nosy Mitsio (site 7) seem like an excellent candidate for an MPA. The area is also a nesting ground for the rare and endangered Madagascar Fish Eagle, and several of these birds were observed. Nosy Kivinji (site 26) – This is a small mound-shaped islet, perhaps 150-200 m in diameter and rising to an elevation of about 100 m. It is characterized by steep cliffs and relatively sparse vegetation, mainly on the upper half. The cliffs continue below the sea surface to a depth of approximately 15-25 m. It has an abundance of diverse ﬁshes and corals, particularly on the eastern side. Much of the island is surrounded by rubble bottom, which provides good habitat for a variety of molluscs. Further investigations are required around nearby rocky islets and Nosy Ankazoberavina, as well as the rocky shoreline of Point d’Angadoka. This entire area may well be deserving of MPA status. Nosy Tanikely (site 29) - The best reef in the Nosy Be area was situated at Nosy Tanikely, a small coral cay surrounded by excellent coral gardens with an abundance of ﬁsh life. The island incorporates a good tourist beach, is well vegetated, and is inhabited by one lemur species. Although it does not have ofﬁcial status as a reserve, it is “semi-protected” by local regulations. No ﬁshing is allowed within 500 m of the island. Although this regulation is not strictly enforced, it has been incredibly successful. The density of ﬁsh around the island was the best observed at any locality during the RAP survey, providing an excellent indication of the results that can be expected if certain key reef areas are set aside as reserves and are properly managed. CONSERVATION RECOMMENDATIONS Nosy Hoa (sites 11-13) - Although only three sites on the extensive Nosy Hoa reef complex were surveyed, the area obviously has exceptional marine biodiversity. It is characterized by an extensive shallow reef ﬂat that surrounds a low, sparsely vegetated island, with a deep (to about 55 m) channel on the eastern side. There is also an exposed seaward reef on the northern side with breaking surf. Our survey activities were conﬁned to the edge of the deep channel, where impressive coral and ﬁsh diversity and an above-average mollusc count were recorded along the edge of a steep drop-off. Visibility was particularly good at the northern (seaward) end of the channel, and several ﬁshes recorded here were not seen at any other site. Nosy Ankarea (site 5) – This roughly circular island is approximately 1.2 km in diameter and rises spectacularly to an elevation of about 230 m. It features steep cliffs, forested slopes, and white sand beaches. Underwater scenery is equally good with excellent reef development, plentiful ﬁshes 16 Rapid Assessment Program Unlike many terrestrial areas of Madagascar, much of the country’s reef habitat remains intact in its natural state. Thus, a real opportunity exists to protect these reefs and insure that high levels of biodiversity are maintained. However, in view of its rapidly increasing human population, with a projected 100% increase by the year 2025, Madagascar faces a huge challenge in preserving its biological integrity. As a result of the RAP survey we make the following speciﬁc recommendations. 1) Pursue a vigorous program of biological assessment. The necessary governmental and NGO infrastructure for marine conservation is already in place, and a marine program has already been created, setting the stage for the next step in conserving Madagascar’s unique marine environment: establishing a network of Marine Protected Areas (MPAs). However, before this can be done, more comparative data is needed in the form of biological Overview assessment surveys such as this RAP. The most effective and representative reserve system cannot be designed until all potential areas have been surveyed. Minimally, this would likely require about six surveys covering the primary coral reef areas. All reef habitats must be assessed, including coastal bays and headlands, near shore islands, offshore islands, platform reefs, and outer oceanic banks. Eventual reserves should be designed to embrace representatives of all reef types and exposure situations, thus ensuring that the full range of coral reef biodiversity is conserved. 2) Review and consolidate previous knowledge of Madagascar’s marine biota. Considerable research has already been conducted involving Madagascar’s marine fauna and ﬂora. The Toliara and Nosy Be areas in particular have been relatively well studied and were formerly or presently are the sites for marine research stations. Much of the research results appear in various French publications. Unpublished Masters and Ph.D. theses on Madagascar’s marine biodiversity can also be found at various French universities. A concerted effort should be made to consolidate this knowledge in order to avoid a duplication of effort. Additionally, these previous studies will greatly compliment future ﬁeld assessments for conservation purposes. With this goal in mind, the present RAP report includes a list of the reef ﬁshes thus far known from Madagascar. It represents a consolidation of all previous taxonomic studies beginning with Sauvage (1891). 3) Collect the additional data that is essential for marine conservation planning. Marine biological studies, particularly those with conservation implications (including taxonomic studies), need to be encouraged, possibly through grants to Madagascar university students or by inviting foreign experts to tackle problems of local signiﬁcance. Studies involving the connectivity of reef communities around the island would be especially useful in designing a system of MPAs. A good body of knowledge concerning current patterns around the island already exists and will be useful in this regard, but more information is needed about larval duration and behavior. Detailed studies are also needed on endangered marine wildlife such as sharks, endemic reef organisms, dugongs, and sea turtles. Financial support of university students would provide an incentive for these studies. Biological data are not the only type of information that is important for conservation planning—additional layers of geophysical, political, ecological, cultural, and socioeconomic information are also essential. All these factors need to be considered in deﬁning a local conservation strategy. 4) Establish a network of marine protected areas. A solid infrastructure already exists for the establishment of MPAs, and much has already been accomplished with respect to drafting a national plan for marine conservation and coastal zoning. A new MPA initiative was launched in late 1999 by ANGAP, ONE, and various international NGOs (WWF, WCS, and CI). It included the preparation of a joint action plan identifying numerous potential MPAs. However, no new MPAs have been designated yet. The establishment of a sound network of MPAs, therefore, should remain a prime objective. The location of protected areas should follow scientiﬁc guidelines to ensure that they cover a representative sample of the region’s marine biodiversity. Target areas should include those that are exceptionally diverse, physically unique, and known to protect currently or potentially rare or endangered species as well as Madagascar endemics. Identifying the location of marine conservation areas should be guided by an understanding of the distribution of marine biodiversity, not just locally but also on a regional scale. Other factors also need to be considered, especially the degree of local interest and support. Whatever mechanisms are used, they should be in tune with the needs of coastal communities. In particular, the term “protected area” must be clariﬁed to show that it does not necessarily mean that an area’s resources are denied to a local population. 5) Enact more effective laws to regulate ﬁshing activities. More precise laws may need to be enacted, particularly at the local level, to cover all aspects of ﬁshing and environmental destruction. One of villagers’ main concerns is that itinerant ﬁshermen are ignoring local taboos related to the environment. If practical, and particularly if they support conservation management, local environmental taboos should be given legal backing. Legislation along these lines should have the principal goal of sustaining natural resources and conserving the natural environment. Laws may also be needed that deal with type and quantity of ﬁshing gear and catch quotas for various species, based on sound biological information. In addition, long-term catch-monitoring programs would provide essential information for ﬁshery management. 6) Develop and promote conservation-oriented marine tourism for the beneﬁt of local communities. Dive tourism and marine conservation are complimentary activities because tourism only thrives when reefs are maintained in good condition. Moreover, diving tourists function as “watchdogs” for frequently visited sites. Madagascar is lagging behind the Seychelles and Maldives as a dive tourist destination, and there is tremendous scope for improvement. As the country establishes a network of MPAs, an effort should be made to involve the dive industry and encourage the development of new resorts and dive concessions within them. Any such enterprises should also involve local communities in the form of employment opportunities and the payment of nominal fees in exchange for the privilege of diving in traditional village ﬁshing grounds. Local communities will take more pride in the reef environment and show more enthusiasm for conservation if they are able to realize commercial advantages. On a global scale, there are numerous examples of dive tourism industries that are compatible with both conservation needs and the needs of local communities. Coral Reefs of Northwest Madagascar 17 Overview 7) Provide dive training for staff of local universities and conservation organizations. During the preparatory phase of the marine RAP, it became evident that trained divers working for NGOs and other marine-related organizations in Madagascar are in short supply. Consequently, local enthusiasm for marine conservation is limited, and much of the impetus for it is currently provided by foreign NGOs. Training local biologists and others to dive will help remedy this situation by promoting marine conservation values and fostering a greater appreciation for the undersea environment. 8) Promote community participation in conservation planning and management. Local communities need to be involved in the implementation and ongoing management of MPAs. The challenge is raising the level of conservation awareness in local communities so that people will want to get involved. Poster campaigns and special educational initiatives have proved successful in other countries. However, considering the poverty and high levels of illiteracy in most coastal villages, it may prove more effective to encourage community participation by establishing and helping to ﬁnance (via government, private corporations, and NGOs) various outreach programs that involve educational assistance, health care, and a general raising of living standards. This type of aid would provide real rewards for villages that implement reef management programs. 9) Strengthen species conservation programs for rare and endangered marine wildlife. Conservation authorities have vast experience in maintaining conservation programs for Madagascar’s rare and endangered terrestrial wildlife. The same level of expertise should be applied to the marine environment. First, the status of rare, endangered, and endemic species must be assessed to ensure adequate habitat protection for these species’ continued survival. The direct exploitation of and trade in these species must also be controlled. Species that seem particularly vulnerable include sharks, sea turtles, dugong, giant clams, and certain ﬁshes (e.g., Napoleon wrasse and large groupers). REFERENCES ANGAP. 1999. Plan Strategique du Réseau des Aires Protégées de Madagascar. Report of a scientiﬁc workshop, Antananarivo, 28-29 July 1999. Association Nationale pour la Gestion des Aires Protégées: 1-61. ANGAP. 2001. Plan de Gestion du Réseau National des Aires Protégées de Madagascar. Parcs Nationaux Madagascar, Antananarivo. Cooke, A., O. Ratomahenina, E. Ranaivoson, and H. Razaﬁndrainibe. 2000. Chapter 60. Madagascar. In: Sheppard, C. R. C. (ed.). Seas at the millennium: an environmental evaluation. Volume 2. Regional chapters: the Indian Ocean to the Paciﬁc. Pergamon, Amsterdam. Pp. 113-131. 18 Rapid Assessment Program Gosliner, T. M., D. W. Behrens, and G. C. Williams. 1996. Coral reef animals of the Indo-Paciﬁc. Sea Challengers, Monterey California. Leis, J. M. 1991. Chapter 8. The pelagic stage of reef ﬁshes: The larval biology of coral reef ﬁshes. In: Sale, P. F. (ed.). The ecology of ﬁshes on coral reefs. Academic Press, San Diego. Pp. 183-230. Mittermeier, R. A., N. Myers, P. R. Gil, and C. G. Mittermeier. 1999. Hotspots. Cemex, Mexico City. ONE. 1999. Document d’orientation pour une politique nationale de developpment durable des zones cotieres de Madagascar. Résumé. Ofﬁce de l’Environment, Antananarivo. Randrianamantsoa, B. J. and J. Brand. 2000. Etude de reconnaissance et zonage des écosystèms marins. Rapport d’étude. Development Evironment Consultants. Report prepared for Projet UNESCO-MAB Mananara-Nord. Antananarivo. 66 pp. Sauvage, H. E. 1891. Historie Naturelle de Poissons. In: Histoire Physique, Naturelle et Politique de Madagascar. Vol. 16. Paris: L’Imprimerie Nationale. Veron, J. E. N. 2000. Corals of the World. Australian Institute of Marine Science, Townsville, Australia. Coral Reefs of Northwest Madagascar 19 Maps and Photos Nudibranch, Glossodoris pallida. (G. Allen) 20 Soft coral, Cespitularia sp.. (G. Allen) Nosy Kivinjy. (G. Allen) Charter vessel Inga Viola. (G. Allen) Coral reef at site 5. (J. Veron) Rapid Assessment Program Maps and Photos Madagascar Anemoneﬁsh, Amphiprion latifasciatus. (G. Allen) Starﬁsh, Protoreaster lincki. (G. Allen) New species of damselﬁsh (Pomacentrus caeruleopunctatus). (G. Allen) Coralliomorpharians, Amplexidiscus fenestrafer. (G. Allen) Science team recording data at site 12. (J. Veron) Coral Reefs of Northwest Madagascar 21 Maps and Photos Ornate lobster, Panulirus ornatus. (G. Allen) 2002 Madagascar Marine RAP Science Team. Back row (L to R): F. Wells, R. Ratsimbazafy, G. Allen, and B. Randriamanantsoa. Front row (L to R): Salimo, S. McKenna, J. Maharavo, E. Turak, G. Bakary, and J. Veron. 22 Rapid Assessment Program Chapter 1 Part 1: Zooxanthellate Scleractinia of Madagascar J.E.N. Veron and Emre Turak SUMMARY • Madagascar has the highest recorded coral diversity of the Western Indian Ocean and Red Sea. • A total of 323 species of zooxanthellate Scleractinia were recorded by the present authors from northwest Madagascar during this study. • An additional 57 species not found during the present study have previously been recorded from Madagascar. This brings the total species complement of Madagascar to 380. • Twelve species were not identiﬁed in this study. Of these, seven are known to be new to science, and six were studied in detail and will be described elsewhere. • One of the new species necessitates distinction as a new genus. Two species currently placed in the genus Platygyra will be included in this genus. METHODS The coral survey team consisted of two people. One individual (Veron) was principally concerned with compiling a species list for the entire area studied (Appendix 1), while the other (Turak) concentrated on making site-by-site comparisons (see Turak, Chapter 1, Part 2). The coral survey team collected species of interest as well as species requiring further study. Part of the substantial though not comprehensive collection made during the survey was sent to the Australian Institute of Marine Science (AIMS), where the survey team was able to study it further. The results of this study have contributed to this chapter. The collection will continue to be housed at AIMS except for type specimens that will be deposited in a still-tobe-determined museum in Madagascar. Field work was carried out over the full depth range where corals occurred and in as wide a range of environmental conditions as possible. Most of the unidentiﬁed species were photographed in situ. RESULTS A total of 323 species of zooxanthellate Scleractinia were recorded during this study1. An additional 57 species have previously been recorded from Madagascar but were not found during the present study (Appendix 1). Of these 57 species, all but seven were recorded from the southwest coast (north of Tuléar) by Veron (Veron 2000). The remainder have been The number and identity of species recorded in this chapter and Turak (Chapter 1, Part 2) differ because this chapter includes work undertaken at the Australian Institute of Marine Science on collected specimens. 1 Coral Reefs of Northwest Madagascar 23 Chapter 1 recorded in earlier French reports, and are not detailed here. The majority of the 57 species not found during the present study are usually found in protected lagoons and estuaries. Two species believed to be endemic to Madagascar (Pocillopora fungiformis Veron 2000 and Stylophora madagascarensis Veron 2000), both of which occur in shallow habitats exposed to strong wave action, were not found during the present study. Conversely, Madagascar’s one endemic genus (Horastrea indica Pichon 1971) was not found by Veron in the Tuléar region (Veron 2000), yet was seen frequently during the present study. These records bring the total number of known coral species in Madagascar to 380, which is the highest number of species recorded in the western Indian Ocean (Figure 1.1), exceeding the combined total for the Red Sea and Socotra (340 species). It also substantially exceeds the 336 species previously recorded from the entire southwest Indian Ocean (the area extending north to and including the Seychelles, Appendix 1). Appendix 1 differentiates between undescribed species and unidentiﬁed species. The former are a group of species that are known to be new to science. Except for the Turbinaria these were studied in detail and will be described as new species (Veron and Turak in prep). The group of undescribed species includes: 1. a Seriatopora that is distinctive and common in some turbid habitats; 2. an Acropora that is common and forms large colonies in shallow exposed habitats; 3. a Coscinaraea that is uncommon; 4. a Turbinaria that is rare but also known from Indonesia; 5. a Blastomussa that is very distinctive; 6. a Platygyra-like faviid that warrants separation as a new genus—this genus is not endemic to Madagascar and will contain two described species currently placed in Platygyra; and 7. an Alveopora that is rare and restricted to muddy substrates. Except for the Turbinaria, none of the above species is known to occur in other countries. 24 Rapid Assessment Program “Unidentiﬁed” species are species that were collected but not studied in the ﬁeld sufﬁciently to be described or identiﬁed reliably. The authors’ reservations about the identities of some of the species in Appendix 1 are indicated by a “cf ” before the species name. This chapter does not provide a detail of biogeographic patterns revealed by the present study, but it is noteworthy that, for the ﬁrst time, a total of 61 species were recorded from the southwest Indian Ocean. A high proportion (30%) of these belong to the genus Porites and are poorly known and largely unstudied. A total of 16 species have not been previously recorded anywhere in the Indian Ocean. Nine species have not been recorded closer to Madagascar than Sri Lanka. CONCLUSION The results of this survey lead to a number of conclusions. First and most generally, this survey underscores the fact that Madagascar is a major center of both endemism and species diversity. More speciﬁcally, since most of the western Indian Ocean and Red Sea has been studied as comprehensively as Madagascar, this survey indicates that Madagascar has more species of zooxanthellate Scleractinia than any country within the region, with the only possible exception of the Comoros, where the corals are still too poorly known to evaluate. Finally, Madagascar’s level of coral species endemism is high, primarily because the total complement of species is high. Madagascar has one endemic genus (Horastrea) and at least eight species that have not been recorded elsewhere. REFERENCES Veron, J.E.N. 2000. Corals of the World. Vols 1-3. El Cajon, CA: Odyssey Publishing. Zooxanthellate Scleractinia of Madagascar Figure 1.1. Total number of species recorded at sites sampled in the Indo-Paciﬁc ocean. Darker areas indicate higher numbers of species. Coral Reefs of Northwest Madagascar 25 Chapter 1 Part 2: Reef Corals of Northwest Madagascar E. Turak SUMMARY • Previously, Madagascar was not considered to contain signiﬁcant coral diversity. This study revealed a rich coral fauna despite the fact that a limited amount of sites were visited in a relatively small area with limited habitat diversity. • Site diversity of coral species was high (average 106 species) and is comparable to near shore reefs of the northern Great Barrier Reef of Australia. Of a total of 340 species predicted to occur in the region, 323 species were recorded during this study, including at least seven undescribed species. For 318 of these species detailed site-speciﬁc data was collected. • Species distribution was relatively homogeneous, mainly due to the low variability of habitat types. Approximately one-third of the total number of species for the area were found at ~60% of the sites. • Reef development in general was limited. With the exception of the Nosy Hao area, most of the sites visited had corals growing on bedrock with limited reef accretion. • Very large coral formations were encountered at some sites, suggesting an absence of severe disturbances in the area. For example, there was no evidence of the 1998 bleaching event that killed most corals in the Central Indian Ocean. INTRODUCTION Coral reefs are one of the most signiﬁcant and extensive marine habitats of Madagascar. Cooke et al. (2000) reported a total of 3,540 linear km of coral reefs, consisting of the following components: 1,130 km of fringing reef; 557 km of reef around islets, islands, and patch reefs; 52 km of true barrier reef; and 1,711 km of submerged coral banks and shoals. The best known reefs are those of the southwestern part of the country, particularly the Toliara region, which were reported in various publications during the 1960s and 1970s (e.g., Clausade et al. 1971 and Thomassin 1971). One of the most intensive studies of coral diversity was that of Pichon (1978), who reported 130 species of scleractinians for the Grand Récif at Toliara. More recently, McClanahan and Obura (1998) recorded at least 164 species for the Masaola area in the Northeast. A total of 200 scleractinian species has been estimated for the entire island (Gabrié et al. 2000 and Cooke et al. 2000). Although very limited in area and duration, the current survey is the ﬁrst detailed study of the northwest region and reveals a much richer fauna for Madagascar than was previously suspected. 26 Rapid Assessment Program Reef Corals of Northwest Madagascar Part 2 METHODS Coral surveys were conducted using the method described in DeVantier et al. (1998). A complete inventory of coral species was compiled per site, giving each species a relative abundance score. Most corals were recorded on the basis of visual observations, but samples were collected of problematical species for shipboard identiﬁcation, primarily using the work of Veron (2000). In addition, a large number of coral samples was collected and sent to Australia for further comparison with reference collections and positive identiﬁcation. A total of 29 sites were surveyed for corals during the RAP. Seven of these were sampled at two depths: shallow (surface to 8-10 m) and deep (>12 m, maximum to 44 m). RESULTS AND DISCUSSION A total of 323 species (see Chapter 1, Part 1) in 62 genera and 17 families of scleractinian corals was recorded during the survey (Appendix 2), including at least seven new species. This ﬁgure is close to the predicted total of 340 species for the entire East Africa and West Indian Ocean region as indicated by species distribution maps in Veron (2000). Detailed site-speciﬁc data was collected for 318 of these species (Appendix 2) and all following analysis is based on this total. Site diversity of coral species was high (average 106 species) and is comparable to areas near the central Indo-Paciﬁc (e.g., northern Great Barrier Reef, Australia). A total of 62 genera, or about two-thirds of the known 93 genera and all of the known families from the Indo-Paciﬁc, was recorded during the RAP survey. Species distribution was relatively homogenous throughout the survey area, which covered an approximate linear distance of 220 km. One-third of all recorded species occurred at over 60% of the sites. This is considerably more than northern Sulawesi, Indonesia, but less than Milne Bay, Papua New Guinea (Table 1.1). Most likely, the relatively high homogeneity in the distribution of coral species in northwestern Madagascar is mainly due to the limited variety of habitats that were visited. However, northwestern Madagascar in general has a lower diversity of coral and reef habitats in comparison to North Sulawesi. More than 100 species were recorded at most sites (Table 1.2). The top ten sites with the highest coral species diversity are listed in Table 1.3. Most of these sites were located in the northernmost section and at northern Nosy Be. The Mitsio Archipelago had the highest overall diversity, followed by Nosy Be and the far northern section (Table 1.4). The area south of Nosy Be had the lowest site diversity. Sites on reefs with extensive or well-developed reef ﬂats had the highest diversity, while submerged reef banks had the lowest (Table 1.5). Acroporidae was by far the richest family (91 species), which is usually the case in the Indo-Paciﬁc. Other major groups included Faviidae and Poritidae, which in combination with Acroporidae accounted for 62% of the species total. The families Fungiidae, Mussidae, and Agariciidae were also well represented, collectively comprising 20% of the species total (Table 1.6). The most speciose genera included Acropora, Montipora, Porites, Favia, Fungia, Favites, Goniopora, and Pavona, accounting for half of the recorded species (Table 1.7). The species grouping referred to as “Porites massive” includes a number of similar species that are not easily identiﬁed underwater. This was the most commonly observed “species.” The 12 most common (i.e., seen at the most sites) species were encountered at over 90% of the sites (Table 1.8). They represent a diversity of families, although half are faviids. The 12 most abundant (i.e., most individual colonies seen over the entire survey) coral species were found at over 80% of the sites, with faviids again accounting for half the species (Table 1.9). The contribution of each family per site in relation to the total number of records gives a somewhat different perspective. Considering families with more than just a few species, Faviidae, Pocilloporidae, and Merulinidae had the highest proportion and Dendrophylliidae, Siderastreidae, and Poritidae the lowest. At sites that had reef or corals deeper than about 20 m, a signiﬁcant difference was recorded between shallow (1-10m) and deep (>10m) zones in terms of species composition and abundance (Figure 1.2). Shallow areas were dominated by encrusting, massive, or branching forms, whereas foliaceous, encrusting, or solitary forms were more common Table 1.1. Comparison of some reef characteristics of northwestern Madagascar with other regions in the Indo-Paciﬁc. Results are from all surveys conducted using the same methods and by the same observer. NW Madagascar N Sulawesi Indonesia Milne Bay PNG North GBR Australia Total number of species 318 445 379 318 Average no. of species per site 103 100 147 100* % of sites with over 1/3 species 60 8 82 - Number of sites surveyed 29 52 28 26 Area covered (x1000 km ) 1.2 23 15 0.8 Average % hard coral cover 35.1 21.3 33.3 34.8 2 * Estimate based on a combination of values for two depths per site. Coral Reefs of Northwest Madagascar 27 Chapter 1 Table 1.2. Number of scleractinian corals recorded for each site. Seven of the sites include a combined total for two depth zones (shallow and deep) that were surveyed separately. Site Species Site Species Site Species 1 111 11 103 21 125 2 111 12 125 22 88 3 104 13 110 23 104 4 109 14 122 24 71 5 143 15 92 25 30 6 129 16 107 26 103 7 109 17 119 27 121 8 90 18 120 28 75 9 88 19 123 29 110 10 46 20 110 Total 318 Table 1.3. The top ten sites with highest coral diversity and the percent of total species for each site. Table 1.6. Number of species recorded for each family and the percent of total species recorded during the survey. Site Family Percent of total spp. Acroporidae 91 29 Faviidae 72 23 Poritidae 33 10 Fungiidae 23 7 Mussidae 22 7 Agariciidae 19 6 Pocilloporidae 14 4 Siderastreidae 13 4 37.42 Pectinidae 9 3 34.91 Dendrophylliidae 8 3 Merulinidae 4 1 Euphyllidae 3 1 Astrocoeniidae 2 1 Oculinidae 2 1 Meandrinidae 1 <1 Trachyphylliidae 1 <1 Caryophilliidae 1 <1 Species Percent Ankarea 5 143 44.97 Nosy Fisaka 6 129 40.57 NE Nosy Hao 12 125 39.31 SE Sakatia 21 125 39.31 Befotaka Bay 19 123 38.68 Andavakalovo 14 122 38.36 Nosy Kisimany 27 121 38.05 Nosy Fanihy 18 120 37.74 Lakandava S wall 17 119 S Tsarabanjina 1 111 Table 1.4. The average species number species per site at four different localities. Locality Avg. no spp. Mitsio Archipelago 117 Far northern section 100 Nosy Be and surounds 111 Southern section 86 Table 1.5. The average number of species per site at four different major reef habitat types. Habitat types 28 Number of species Site no. Avg. no spp. Fringing reefs with moderate ﬂats 107 Submerged banks 75 Sheltered bays 98 Extensive reef ﬂats 115 Rapid Assessment Program Reef Corals of Northwest Madagascar Part 2 Table 1.7. Number of species in the 20 most diverse genera and the percent of total species recorded. Table 1.9. The 12 most abundant coral species found at over 80% of the sites. Family Genus Species Acroporidae No. spp. % total Acropora 63 20 Acroporidae Montipora 22 7 Poritidae Porites 16 5 Faviidae Favia 15 5 Fungiidae Fungia 11 3 Faviidae Favites 11 3 Poritidae Goniopora 10 3 Agariciidae Pavona 10 3 Faviidae Goniastrea 10 3 Faviidae Platygyra 10 3 Mussidae Acanthastrea 8 3 Siderastreidae Psammocora 7 2 Agariciidae Leptoseris 7 2 Dendrophylliidae Turbinaria 7 2 Pocilloporidae Seriatopora 6 2 Acroporidae Astreopora 6 2 Poritidae Alveopora 6 2 Pocilloporidae Pocillopora 5 2 Faviidae Montastrea 5 2 Faviidae Leptastrea 5 2 Average abundance Number of sites Porites massive 2.62 29 Acropora divaricata 2.44 27 Goniastrea peresi 2.22 27 Platygyra daedalea 2.21 24 Favia favus 2.15 27 Leptoria phrygia 2.12 25 Hydnophora exesa 2.11 28 Galaxea fascicularis 2.07 27 Goniastrea pectinata 2.04 26 Astreopora myriophthalma 2.00 28 Echinopora hirsutissima 2.00 28 Pocillopora damicornis 1.96 28 Table 1.8. The 12 most common coral species, found at over 90% of the sites. Total abundance Number of sites Porites massive 76 29 Hydnophora exesa 59 28 Species Linkage Distance 0 25 50 12_s 13_s Astreopora myriophthalma 56 28 Echinopora hirsutissima 56 28 Pocillopora damicornis 55 28 9_s Acropora divaricata 66 27 17_s Goniastrea peresi 60 27 Favia favus 58 27 Galaxea fascicularis 56 27 Goniastrea pectinata 53 26 Favia matthaii 50 26 Diploastrea heliopora 44 26 Shallow 11_s 6_s 5_s 9_d 12_d Deep 17_d 13_d 11_d 6_d 5_d Figure 1.2. Cluster analysis of species presence and abundance on seven of the sites where two depths were surveyed. The X_s indicates shallow sites, and X_d indicates deep sites. This and the following dendrograms were created using Wards Amalgamation schedule and a Squared Euclidean Distance measurement. Coral Reefs of Northwest Madagascar 29 Chapter 1 Linkage Distance 0 Mitsio Archipelago High energy Nosy Hao area Submerged and low diversity Ambavatoby Bay Nosy Hara area Nosy Be area S1 S2 S4 S7 S6 S5 S8 S23 S26 S14 S18 S3 S9 S11 S13 S12 S10 S25 S28 S22 S24 S15 S16 S17 S19 S27 S29 S20 S21 20 40 60 80 North South Figure 1.3. A dendrogram based on a cluster analysis of species presence and abundance at all sites indicates a strong geographical and to a lesser extent environmental separation of sites. in greater depth. Deeper reefs were found mostly in the northern section. A cluster analysis of species presence and abundance separates the northern and southern sites (Figure 1.3) into two discrete groupings. The northern grouping includes the Mitsio Archipelago, Nosy Hao reefs, and a group of high-energy sites, which are subjected to strong wave action or currents. The southern group contains sites of lower diversity including submerged banks, Ambavatoby Bay, and sites in the Nosy Be area. Although located in the north, the Nosy Hao sites are nested in the less diverse southern group. At least nine previously unknown species were discovered during the survey. They were mainly found in protected bays with high turbidity or in deeper water. These will eventually be described by the second author. This present survey increases the total species for Madagascar to well over 300 species, a notable increase to the previous estimate of 200 species (Gabrié et al. 2000 and Cooke et al. 2000). The azooxanthelate scleractinian Tubastrea was not common, but usually found in areas with periodic strong currents. Of the non-scleractinian hard corals the hydrocoral (ﬁre coral) Millepora was the most common. Four species were recorded during the survey: M. tenella, M. exesa, M. platyphylla, and M. intricata. Millepora was particularly abundant at site 21 on the southeastern side of Nosy Sakatia and appeared to be more common in the southern section of the survey area. The other hydrocoral, Stylaster, was common and sometimes abundant on deep vertical walls of submerged banks, with some colonies reaching 20-30 cm in height. A total of 22 genera in ﬁve families of soft corals (excluding gorgonians) were recorded during the survey. They were generally common and were the dominant bottom cover at a few sites. Xeniidae, Nephtheidae, and 30 Rapid Assessment Program the alcyoniid coral Rhytsima were the most abundant representatives. Soft corals were particularly abundant at sites 3 and 17, where they formed 60-80% of the bottom cover and were common at sites 9, 11, and 12. The organpipe coral Tubipora musica was found at most sites, but was always rare or only a few colonies were seen, with the exception of site 11 where it was common. The blue coral Heliopora coerulea was not seen during the survey. With the exception of reefs in the far northern section of the survey area and occasional reef ﬂats in other areas, reef development was generally limited. However, coral communities on basal rock with limited accretion sometimes give the appearance of true reef development. Large “massive” formations were rare, with the majority of non-plating corals being less than 50 cm in diameter. However, few sites contained a number of very large (45 m diameter) plate-forming corals. The scarcity of large formation indicates a recent (within the past 20-30 years) major disturbance. The region is in a major cyclone pathway and destructive storms no doubt periodically alter the seascape. There is minor evidence of current bleaching, but no apparent associated mortality. There was virtually no evidence of the effects from the 1998 bleaching event that caused widespread severe mortality in other parts of the Indian Ocean, including the Seychelles. Although there were reports (in 1998) of bleaching along the central western coast of Madagascar, no mortality was indicated. CONSERVATION RECOMMENDATIONS Coral areas of northwestern Madagascar, although exhibiting only limited reef development, are in relatively good shape compared to many other parts of the world. In terms of diversity and abundance the most interesting area is the Mitsio Archipelago. However, in terms of health, diversity, abundance of corals, and presence of new species, Nosy Hara and Nosy Hao are exceptional. Less accessible areas such as Cape d’Ambre and large protected bays such as Befotaka Bay (site 19) and Ambavatoby Bay (sites 22 and 24) remain largely unknown and require further surveys. They may well harbor additional species, including undescribed forms. Although they are not presently threatened, they should be considered for part of an eventual reserve system to safeguard against future threats. REFERENCES Clausade, M., N. Gravier, J. Picard, M. Pichon, M-L. Roman, B. Thamassin, P. Vasseur, M. Vivien, and P. Weydert. 1971. Morphologie des récifs coralliens de la région de Tuléar (Madagascar): Eléments de terminologie récifale. Téthys, Supplement 2: 1-74. Cooke, A., O. Ratomahenina, E. Ranaivoson, and H. Razaﬁndrainibe. 2000. Chapter 60. Madagascar. In: C. R. C. Sheppard (ed.). Seas at the millennium: an environmental evaluation. Volume 2. Regional chapters: Reef Corals of Northwest Madagascar Part 2 the Indian Ocean to the Paciﬁc. Pergamon, Amsterdam. Pp. 113-131. DeVantier, L. M., G. De’ath, T. J. Done, and E. Turak. 1998. Ecological assessment of a complex natural system: a case study from the Great Barrier Reef. Ecological Applications 8: 480-496. Gabrié, C., P. Vasseur, J. Maharavo, H. Randriamiarana, and E. Mara. 2000. The coral reefs of Madagascar. In: T. R. McClanahan, D. O. Obura, and C. R. C. Sheppard (eds.). Coral Reefs of the Western Indian Ocean. Oxford: Oxford University Press. McClanahan, T. and D. Obura. 1998. Monitoring, training and assessement of the coral reefs of the Masoala Peninsula. Wildlife Conservation Society, June 1998. Pichon, M. 1979. Recherches sur les peuplements à dominance d’Anthozoaires dans les récifes coralliens de Tuléar (Madagascar). Atoll Research Bulletin 222: 1-147. Thomassin, B. 1971. Révue bibliographique des travaux de la Station Marine de Tuléar (Madagascar) 1961-1970. Téthys, Supplement 1: 3-49. Veron, J. E. N. 2000. Corals of the World. Australian Institute of Marine Science, Townsville, Australia. Coral Reefs of Northwest Madagascar 31 Chapter 2 Molluscs of Northwestern Madagascar Fred E. Wells SUMMARY 32 Rapid Assessment Program • Molluscs were collected at 30 sites in northern Madagascar, 11-26 January 2002. As many habitats as possible were examined at each site to develop as comprehensive a species list of the molluscs present as possible in the limited time available. • A total of 525 species of molluscs were collected: 382 gastropods, 139 bivalves, 1 scaphopod, 2 cephalopods, and 1 chiton. The collection included 92 families and 193 genera. • Recorded diversity of marine molluscs in Madagascar was higher than expected considering Madagascar’s distance from the widely acknowledged center of marine biodiversity in the coral triangle region (Indonesia, Philippines, and New Guinea). • A mean of 72.3±2.4 species was recorded per site. The range was from 49 to 93 species. • The most abundant species at each site were generally arcid bivalves and Lithophaga spp., which live in the coral, and Lopha cristagalii. Pedum spondyloidaeum was common at several sites, but was not as abundant as on previous Marine RAP surveys. • The least and most diverse sites for molluscs were widely spread over the study area. Sites with the greatest diversity of molluscs were SE Sakatia (site 21, 93 species), S Tsarabanjina (site 1, 91 species), Marie Bank (site 3, 91 species), Nosy Kisimany (site 27, 89 species), and NE Sakatia (site 27, 89 species). Sites with the lowest diversity of molluscs were Souzy Bank (site 30, 49), Ambariokisimany (site 9, 51 species), NW corner Mitsio Island (site 7, 51), and Kalomisampa Rocks (site 8, 55 species). • The geographical distributions of 203 species are analyzed. The vast majority (93.6%) are widespread Indo-West Paciﬁc forms. Only 11 species (5.4%) are restricted to the Indian Ocean. • A number of commercially important mollusc species (Tridacna, Strombus, Pinna, Pinctada, and Lambis) occurred widely at the sites surveyed. Populations of all of these groups were small, and commercial quantities were never found. • No clear pattern of regional hotspots of molluscan diversity was evident among the stations examined in Madagascar. More detailed surveys would be necessary to develop a reserve zoning pattern that adequately protects a representative series of habitats in the islands. Molluscs of Northwestern Madagascar INTRODUCTION The present chapter examines molluscs collected during the Marine RAP survey of Madagascar. As the basic building blocks of the reef, corals are an obvious group to include in the Marine RAP surveys. Fish are also an important component because of their ecological and economic importance in the system. The rationale for including molluscs is that they dominate diversity in many marine systems. Molluscs can be used as a surrogate for measuring the diversity of invertebrates other than corals in the reefs. With their high diversity, often high density, and variety of lifestyles, molluscs are ecologically important to trophic ﬂows within the system. Many groups, such as giant clams, spider shells, trochus, cephalopods, and others, are economically important. For all of these reasons, molluscs have been included in the Marine RAP surveys. METHODS The survey was conducted from 11 to 26 January 2002, with a total of 30 sites being examined; an additional site (Site 0) was examined at Nosy Be before the survey commenced. All sites were surveyed by scuba diving. Dive times ranged from 50 to 122 minutes; 26 of the 30 dives were between 80 and 105 minutes. Each site was examined by starting at depths of 10-25 m and working up the reef slope. Most of the time was spent in shallow (<6 m) water, as the greatest diversity of molluscs occurs in this region; the shallow depth also maximizes diving time. To obtain as many species as possible, all habitats encountered at each site were examined for molluscs, including living coral, the upper and lower surfaces of dead coral, shallow and deep sandy habitats, and intertidal habitats. For the same reason, no differentiation was made between species collected alive or as dead shells, as the dead shells would have been living at the site. Beach drift collections were made at sites 18 and 27. The collection at site 18 was made by limiting the dive time to one hour. The beach collection at site 27 was made after the dive was completed. These types of collections can signiﬁcantly increase the number of species recorded at a site. Unfortunately, time available for these collections is very limited during Marine RAP surveys. While this collecting approach allows the rapid assessment of a variety of mollusc species, it is not complete. For example, no attempt was made to break open the corals to search for boring species, such as Lithophaga spp. Similarly, arcid bivalves burrowing into the corals were not thoroughly examined. Furthermore, micro molluscs were not sampled. However, I undertook the sampling of molluscs on all six Conservation International Marine RAP trips done to date, as well as on many surveys by the Western Australian Museum, and used the same techniques on all of these expeditions. Therefore, the sampling results provide a good indication of diversity relative to other Marine RAP surveys. A variety of standard shell books and ﬁeld guides were available for reference during the expedition. Most species were identiﬁed according to the following texts: Cernohorsky (1972); Springsteen and Leobrera (1986); Lamprell and Whitehead (1992); Lamprell and Healy (1998); and Coleman (2001). Specimens of small species were retained in plastic vials or bags, and tissue was removed with bleach. These were taken to the Western Australian Museum, where they were identiﬁed using the reference collections of the Museum and specialist texts and papers on particular groups (listed below under References). Representatives of these species were deposited in the WA Museum and in the CNRE in Madagascar. Additionally, a collection of 70 identiﬁed species was left in Madagascar at CNRE at the end of the survey. RESULTS A total of 525 species of molluscs were collected: 382 gastropods, 139 bivalves, 1 scaphopod, 2 cephalopods, and 1 chiton (Table 2.1; Appendix 3). There were 92 families and 193 genera collected. Recorded diversity of marine molluscs in Madagascar was higher than expected given the distance of Madagascar from the center of diversity in the coral triangle between the Philippines, Malaysia, and New Guinea. The greatest diversity recorded in previous Marine RAP surveys was in the Raja Ampats, where 699 species were recorded in 45 dives (Table 2.2). The TogeanBanggai expedition included 31 dives, the same number as in Madagascar. The 501 species recorded in Madagascar are very close to the 541 recorded on the Togean expedition. Diversity of Madagascar marine molluscs is greater than surveys of Christmas Island (313 species), Cocos (Keeling) Islands (380), Ashmore Reef (433), and surveys undertaken on the west coast of Australia (Table 2.2). As with previous surveys, the most abundant species at each site were generally arcid bivalves and Lithophaga spp., which live in the coral, and Lopha cristagalii. Pedum spondyloidaeum was common at several sites, but was not as abundant as on previous surveys. A mean of 72.3±2.4 species was recorded per site; the range was from 49 to 93 species (Table 2.3). Higher diversity was recorded at sites with more variable habitat types. The sites with the greatest diversity of molluscs (Table 2.4) were those with the greatest habitat diversity. In particular these sites had shallow sand in addition to the subtidal corals and intertidal rocks. Shallow sand is important both because of the species which live within it and because dead shells are washed in from adjacent coral habitats and accumulate in the sand. The mean of 72.3 species per site compares very favorably with previous Marine RAP surveys, again emphasizing the relatively high diversity of Madagascar molluscs compared to areas in the coral triangle. The distributions of molluscs in the various areas of northern Madagascar were variable, with no apparent pattern. The most diverse sites for molluscs (Table 2.4) and Coral Reefs of Northwest Madagascar 33 Chapter 2 the least diverse sites (Table 2.5) were widely spread over the study area. Not enough is known about the distributions of many mollusc species collected in Madagascar to place them within a geographical context. Table 2.6 shows the distributions of 203 species, as recorded in other literature (See References). The species were gastropods of the families Haliotidae, Neritidae, Cerithiidae, Planaxidae, Littorinidae, Strombidae, Cypraeidae, Bursidae, Ranellidae, Tonnidae, Muricidae, Nassariidae, Olividae, Harpidae, Mitridae, Costellariidae, Terebridae, Conidae, and Phyllidiidae. The vast majority Table 2.1. Taxonomic composition of mollusc species collected in northwestern Madagascar. Class Families Genera Species Polyplacophora 1 1 1 Gastropoda 61 120 382 Bivalvia 27 69 139 Scaphopoda 1 1 1 Cephalopoda 2 2 2 Totals 92 193 525 Table 2.2. Numbers of mollusc species collected during previous Marine RAP surveys undertaken by Conservation International and similar surveys by the Western Australian Museum. Location Collecting days Mollusc species Reference Northern Madagascar 16 525 Present Survey Raja Ampat Islands 15 665 Wells 2002 Togean-Banggai Islands, Indonesia 11 541 Wells 2001 Calamian Group, Philippines 16 651 Wells 2000 Milne Bay, Papua New Guinea 19 638 Wells 1998 Milne Bay, Papua New Guinea 11 643 Wells & Kinch 2003 20 380 on survey; total known fauna of 610 species Abbott 1950, Maes 1967, Wells 1994 12 plus accumulated data 313 on survey; approx. 520 total Iredale 1917, Wells et al. 1990, Wells & Slack-Smith 2000 Ashmore Reef 12 433 Wells 1993, Willan 1993 Cartier Island 7 381 Wells 1993 Hibernia Reef 6 294 Willan 1993 Scott/Seringapatam Reef 8 279 Wilson 1985, Wells & Slack-Smith 1986 Rowley Shoals 7 260 Wells & Slack-Smith 1986 Montebello Islands 19 633 Preston 1914, Wells et al. 2000 Muiron Islands and Exmouth Gulf 12 655 Slack-Smith & Bryce 1995 Bernier and Dorre Islands, Shark Bay 12 425 Slack-Smith & Bryce 1996 Accumulated data 492 Wells & Bryce 1997 Accumulated data 384 Shepherd 1984 CI Marine RAP surveys Western Australian Museum Surveys Cocos (Keeling) Islands Christmas Island (Indian Ocean) Abrolhos Islands Other surveys Chagos Islands 34 Rapid Assessment Program Molluscs of Northwestern Madagascar Table 2.3. Total number of mollusc species collected at each site in northwestern Madagascar. Site Number of species Site Number of species Site Number of species 1 91 11 78 21 93 2 85 12 71 22 68 3 91 13 85 23 77 4 77 14 69 24 58 5 80 15 66 25 70 6 76 16 61 26 68 7 51 17 57 27 89 8 55 18 81 28 73 9 51 19 60 29 81 10 69 20 89 30 49 Mean = 72.3±2.4 (93.6%) of the species are widespread Indo-West Paciﬁc forms. Only 11 species (5.4%) are restricted to the Indian Ocean. Due to the shell trade, a number of new species have been described from Madagascar in recent years. Only one of these, Bursa nigrita, was collected during the present survey. In a recent revision of the genus Cerithium, Houbrick (1992) described Cerithium africanum from Madagascar and the nearby region. This species was widespread in the survey area, being recorded at a total of 17 sites. A number of commercially important edible mollusc species occurred widely at the sites surveyed. These include spider shells (Lambis), conchs (Strombus), Murex ramosus, pen shells (Pinna and Atrina), pearl oysters (Pinctada), and giant clams (Tridacna). Populations of all of these groups were small, and commercial quantities were never found. Many species were found at only a few sites, with a high proportion of the records being only dead shells. Table 2.4. Ten sites observed to have the richest mollusc diversity among the 30 sites surveyed in northern Madagascar. Site Number of species Location 21 SE Sakatia 93 1 S Tsarabanjina 91 3 Marie Bank 91 20 NE Sakatia 89 27 Nosy Kisimany 89 2 NW Tsarabanjina 83 13 Nosy Hao channel 85 18 Nosy Fanihy 81 29 W Nosy Tanikely 81 5 Ankarea 80 DISCUSSION The results of the Marine RAP of northwestern Madagascar reveal a diverse molluscan biota on the coral reefs of the study area. In a survey that lasts only 16 days, all species that occur in northwestern Madagascar clearly cannot be collected or even recorded. However, the survey demonstrates that a wide range of mollusc species occur in the area, making it well worth conserving. No clear pattern of hotspots for molluscan diversity exists in the region. Rather, high species diversity sites are intermingled with sites of much lower diversity. Consequently, the best strategy for protecting marine biodiversity in northwestern Madagascar is to focus on the entire archipelago. More detailed surveys are needed to develop a zoning pattern that would adequately protect a representative series of habitats in the islands. Cooke’s summary (in press) of knowledge about the marine and coastal ecosystems of Madagascar includes cur- Table 2.5. Ten sites observed to have the lowest mollusc diversity among the 30 sites surveyed in northern Madagascar. Number of species Site Location 30 Souzy Bank 49 9 Ambariokisimany 51 7 NW corner Mitsio 51 8 Kalomisampa Rocks 55 17 Lakandava S wall 57 24 E Ambavatoby Bay 58 19 Befotaka Bay 60 16 Ambatomaranitra 61 15 Lakandava N beach 66 22 W Ambavatoby 68 Coral Reefs of Northwest Madagascar 35 Chapter 2 Table 2.6. Geographical distributions of selected species recorded in Madagascar. Number of species Percentage of species Southwestern Indian Ocean 2 1.0 Western Indian Ocean 4 2.0 Indian Ocean 5 2.5 Indian Ocean-Western Paciﬁc 2 1.0 Indo-West Paciﬁc 190 93.6 Totals 203 100.1 Geographical Range rent understandings of the region’s marine biodiversity. In the western Indian Ocean, 10,627 species of invertebrates have been recorded (Richmond in press), and at least 3,500 of these have been described (Gabrié et al. 2000). Moreover, the marine fauna of Madagascar is the most diverse in the region. Knowledge of diversity is based principally on studies of reefs in the Toliara region undertaken by scientists at Site d’Endoume, Universite of Aix-Marseille, during the 1960s and early 1970s. Less detailed studies undertaken in other areas of Madagascar indicate the same general patterns, but report lower numbers of species due to less intensive sampling. For example, Cooke (in press) records the known marine macrofauna in the region of Nosy Be as >600 species. As of yet, no comprehensive list of the molluscs of Madagascar has been compiled. The body of literature on the subject is divided between publications from the late nineteenth and early twentieth centuries (Dautzenberg 1923, 1929, 1932, Hoffman 1877, Odhner 1919, Rost & Soot-Ryen 1955) and recent single-species descriptions (e.g., Cosel & Blöcher 1976). Thomassin (1978) reported at least 1,400 species of molluscs from the Toliara region. Laboute and Maharavo (personal communication cited in Cooke in press) reported 64 species from Nosy Be, including 38 opisthobranchs, but the species were not identiﬁed. This Marine RAP study recorded essentially the same number of opisthobranchs, but increased by an order of magnitude the known number of species in other groups. FAO (1998) reported the primary commercial food molluscs in the Toliara region to be Octopus cyanea, O. aegina, and O. macropus as well as pelagic squid (Loligo species). In contrast, the most important commercial species in Nosy Be are Sepioteuthis lessoniana and cuttleﬁsh, particularly Sepia zanzibarica. These species, however, were not sampled during the Marine RAP. Madagascar has a large ornamental shell trade industry, centered on the Toliara region. In a 1993 study, 82 species of gastropods were recorded in the ornamental shell markets in Toliara (WWF 1993). This total increased to 138 species in 1997. The rarest and most valuable species include Lambis truncata, Cypraea mauritania, Cassis cornuta, Cypraecassis rufa, Charonia tritonis, Tonna canaliculata, and species of Conus (Romaine 1997). The shell trade, which has probably increased since 1997, is a major conservation concern in the 36 Rapid Assessment Program Toliara region. This Marine RAP recorded numerous species of Conus available in the Toliara markets, including a number of fully adult Lambis truncata and one huge C. cornuta. In Nosy Bé, the few shells available for purchase included Triton Trumpet (Charonia tritonis). REFERENCES Abbott, R. T. 1950. Molluscan fauna of the Cocos-Keeling Islands. Bulletin of the Rafﬂes Museum 22: 68-98. Cernohorsky, W. O. 1972. Marine shells of the Paciﬁc. Volume 2. Sydney, New South Wales: Paciﬁc Publications. Coleman, N. 2001. 1001 nudibranchs. Catalogue of IndoPaciﬁc sea slugs. Springwood, Queensland: Neville Coleman. Cooke, A. In press. Marine and coastal ecosystems of Madagascar. Cosel, R. von and M. Blöcher. 1976. Eine neue Lyria aus Madagaskar (Prosobranchia: Volutidae). Archives Molluskunde 107: 195-201. Dautzenberg, P. 1923. Liste préliminaire des mollusques marins de Madagascar et description de deux espèces nouvelles. Journal de Conchyliologie 67: 21-74. Dautzenberg, P. 1929. Mollusques testacés marins de Madagascar. Fauna des Colonies Francaises. Volume 3. Dautzenberg, P. 1932. Mollusques testacés marins de Madagascar. Supplément. Journal de Conchyliologie 76: 5-119. Hoffman, C. K. 1877. Recherches sur le faune de Madagascar et de ses dependances, d’après les découvertes de Dr Francois P.L. Pollen et D.C. van Dam. Leiden, The Netherlands: E. J. Brill. FAO (Food and Agriculture Organization of the United Nations). 1998. Aperçu sur les Zones de Pêche Traditionelle Maritime dans la Region de Toliara. Programme Sectiorel Pêche PNUD/FAO-MAG/92/004 – DT/10/98. 14 pp. Gabrié, C., J. Maharavo, H. Randriamiarana, and E. Mara. 2000. The coral reefs of Madagascar. In: McClanahan, T. R., D. O. Obura, and C. R. C. Shepherd (eds.). Coral reefs of the West Indian ocean. Oxford: Oxford University Press. Houbrick, R. S. 1992. Monograph of the genus Cerithium Bruguière in the Indo-Paciﬁc (Cerithiidae: Prosobranchia). Smithsonian Contributions to Zoology 510: 1-211. Iredale, T. 1917. On some new species of marine molluscs from Christmas Island, Indian Ocean. Proceedings of the Malacological Society of London 12: 331-334. Lamprell, K. and J. M. Healy. 1998. Bivalves of Australia. Volume 2. Leiden, The Netherlands: Backhuys Publishers. Lamprell, K. and T. Whitehead. 1992. Bivalves of Australia. Volume 1. Bathurst, New South Wales: Crawford House Press. Molluscs of Northwestern Madagascar Maes, V. O. 1967. The littoral marine molluscs of CocosKeeling Islands (Indian Ocean). Proceedings of the Academy of Natural Science of Philadelphia 119: 93-217. Odhner, N. J. 1919. Contribution a la faune malacologique de Madagascar. Arkiv för Zoologi utgifvet af K. Svenska Vetenskapsakademien 12(6): 1-52. Preston, H. B. 1914. Description of new species of land and marine shells from the Montebello Islands, Western Australia. Proceedings of the Malacological Society of London 11: 13-18. Richmond, M. D. In press. The marine biodiversity of the western Indian ocean and its biogeography. How much do we know? In: Richmond, M. D. and J. Francis (eds). Proceedings of the Twentieth Anniversary Conference on Marine Science in Tanzania, June 2001. Western Indian Ocean Marine Science Association, Institute of Marine Sciences, Zanzibar, Tanzania. Romaine, T. E. 1997. Localized marine resource exploitation in Toliara, Madagascar. Project assignment report, School for International training, Vermont, USA. 30 pages. Rost, H. and T. Soot-Ryen. 1955. Notes on a small collection of pelecypods from Madagascar. Astarte 11: 1-5. Sheppard, A. L. S. 1984. The molluscan fauna of Chagos (Indian Ocean) and an analysis of its broad distribution patterns. Coral Reefs 3: 43-50. Slack-Smith, S. M. and C. W. Bryce. 1995. Molluscs. In: Hutchins, J. B., S. M. Slack-Smith, L. M Marsh, D. S. Jones, C. W. Bryce, M. A. Hewitt, and A. Hill (eds.). Marine biological survey of Bernier and Dorre Islands, Shark Bay. Western Australian Museum and Department of Conservation and Land Management, manuscript report. Pp. 57-81. Slack-Smith, S. M. and C. W. Bryce 1996. Molluscs. In: Hutchins, J. B., S. M. Slack-Smith, C. W. Bryce, S. M. Morrison, and M. A. Hewitt (eds.). Marine biological survey of the Muiron Islands and the eastern shore of Exmouth Gulf, Western Australia. Western Australian Museum and Department of Conservation and Land Management, manuscript report. Pp. 64-100. Springsteen, F. J. and F. M. Leobrera. 1986. Shells of the Philippines. Manila, Philippines: Carfel Seashell Museum. Thomassin, B. 1978. Les peuplements des sédiments coralliens de la région de Tuléar (SW de Madagascar). Leur insertion dans le contexte côtier indo-paciﬁque. Thèse Doct. Ès Sci., Univ Aix Marseille. Wells, F. E. 1993. Part IV. Molluscs. In: Berry, P. F. (ed.). Faunal Survey of Ashmore Reef, Western Australia. Records of the Western Australian Museum, Supplement 44: 25-44. Wells, F. E. 1994. Marine Molluscs of the Cocos (Keeling) Islands. Atoll Research Bulletin 410: 1-22. Wells, F. E. 1998. Marine Molluscs of Milne Bay Province, Papua New Guinea. In: Werner, T. and G. R. Allen. (eds.). A rapid biodiversity assessment of the coral reefs of Milne Bay Province, Papua New Guinea. RAP Working Papers Number 11. Washington, DC: Conservation International. Pp. 35-38. Wells, F. E. 2001. Molluscs of the Calamianes Islands, Palawan Province, Philippines. In: Werner, T. B. and G. R. Allen (eds.). A rapid marine biodiversity assessment of the Calamianes Islands, Palawan Province, Philippines. RAP Bulletin of Biological Assessment 17. Washington, DC: Conservation International. Pp. 27-30; 81-94. Wells, F. E. 2002. Molluscs of the Gulf of Tomini, Indonesia. In: Allen, G. R. and S. A. McKenna (eds.). A rapid biodiversity assessment of the coral reefs of the Togean and Banggai Islands, Sulawesi, Indonesia. RAP Bulletin of Biological Assessment 20. Washington, DC: Conservation International. Wells, F. E. 2002. Molluscs of the Raja Ampat Islands, Papua Province, Indonesia. In: McKenna, S.A., G. R. Allen and S. Suryadi. 2002. A marine rapid assessment of the Raja Ampat Islands, Papua Province, Indonesia. RAP Bulletin of Biological Assessment 22. Washington, DC: Conservation International. Wells, F. E. and C. W. Bryce. 1997. A preliminary checklist of the marine macromolluscs of the Houtman Abrolhos Islands, Western Australia. In: Wells, F. E. (ed). Proceedings of the seventh international marine biological workshop: The marine ﬂora and fauna of the Houtman Abrolhos Islands, Western Australia. Western Australian Museum, Perth. Pp. 362-384. Wells, F. E., C. W. Bryce, J. E. Clarke, and G. M. Hansen. 1990. Christmas Shells: The Marine Molluscs of Christmas Island (Indian Ocean). Christmas Island: Christmas Island Natural History Association. Wells, F. E. and J. Kinch, 2003. Molluscs of Milne Bay Province, Papua New Guinea . In: Allen, G. R., J. P. Kinch, S. A. McKenna and P. Seeto (eds.). A rapid marine biodiversity assessment of Milne Bay Province, Papua New Guinea – Survey II (2000). RAP Bulletin of Biological Assessment 29. Washington, DC: Conservation International. Wells, F. E. and S. M. Slack-Smith. 1986. Part IV. Molluscs. In: Berry, P. F. (ed.). Faunal Survey of the Rowley Shoals and Scott Reef, Western Australia. Records of the Western Australian Museum, Supplement 25: 41-58. Wells, F. E. and S. M. Slack-Smith. 2000. Molluscs of Christmas Island. In: Berry, P. F. and F. E. Wells (eds.). Survey of the marine fauna of the Montebello Islands, Western Australia and Christmas Island, Indian Ocean. Records of the Western Australian Museum, Supplement 59: 103-116. Wells, F. E., S. M. Slack-Smith, and C. W. Bryce. 2000. Molluscs of the Montebello Islands. In: Berry, P. F. and F. E. Wells (eds.). Survey of the marine fauna of the Montebello Islands, Western Australia and Christmas Island, Indian Ocean. Records of the Western Australian Museum, Supplement 59: 29-46. Coral Reefs of Northwest Madagascar 37 Chapter 2 Willan, R. C. 1993. Molluscs. In: Russell, B. C. and J. R. Hanley (eds.). The marine biological resources and heritage values of Cartier and Hibernia Reefs, Timor Sea. Northern Territory Museum, manuscript report. Wilson, B. R. 1985. Notes on a brief visit to Seringapatam Atoll, North West Shelf, Australia. Atoll Research Bulletin 292: 83-100. World Wildlife Fund. 1993. Coral reefs and coastal zone of Toliara – Conservation and development through ecotourism. Pre-project Report edited by A. J. Cooke. Prepared for WWF Madagascar. 38 Rapid Assessment Program Chapter 3 Reef Fishes of Northwestern Madagascar Gerald R. Allen SUMMARY • A total of 463 species of reef ﬁshes belonging to 191 genera and 65 families were recorded for 30 sites. An extrapolation method utilizing six key index families (Chaetodontidae, Pomacanthidae, Pomacentridae, Labridae, Scaridae, and Acanthuridae) indicates a total reef ﬁsh fauna for northwest Madagascar consisting of at least 576 species. • Species numbers at visually sampled sites ranged from 33 to 166, with an average of 117 per site. • The richest sites for ﬁshes were Nosy Hao, Nosy Tanikely, Nosy Ankarea, and Nosy Kivinjy, with species totals ranging between 152 and 166. • A review of the literature combined with results of the current RAP survey reveal a total reef ﬁsh fauna for Madagascar consisting of 788 species. A comprehensive checklist is included with this report. • In terms of number of species and general faunal composition, Madagascar is similar to other insular localities in the western Indian Ocean including Mauritius, Seychelles, Chagos, and Maldives. • The majority of Madagascar ﬁshes have broad distributions in the Indo-West and central Paciﬁc. Approximately 20% of the species range widely over the entire tropical Indian Ocean or are restricted to the western Indian Ocean. • About 30 species are conﬁned to Madagascar and adjacent regions including Mozambique, East Africa, Comoro Islands, St. Brandon’s Shoals, and the Mascarene Group. • Eight species are presently known only from the seas of Madagascar, including two new damselﬁshes (Pomacentridae: Pomacentrus) collected for the ﬁrst time during the current RAP survey. Another four species are known only from Madagascar and the Comoros/ Saint Brandon’s Shoals. Coral Reefs of Northwest Madagascar 39 Chapter 3 INTRODUCTION Comprehensive documentation of the reef ﬁsh fauna based on results of Conservation International’s RAP survey during January 2002 are presented in this chapter. The background of this project and detailed descriptions of the 30 survey sites are provided elsewhere in this report. The principal aim of the ﬁsh survey was to provide a comprehensive inventory of the reef ﬁshes along the northwestern coast of Madagascar. This segment of the fauna includes ﬁshes living on or near coral reefs down to the limit of safe sport-diving or approximately 40 m depth. Survey activities therefore excluded estuary species, deepwater ﬁshes, and offshore pelagic species such as ﬂyingﬁshes, tunas, and billﬁshes. An effort was also made to review previous knowledge of Madgascar reef ﬁshes. Considerable research has been done on Madagascar reef ﬁshes, resulting in a number of publications. The areas around Nosy Bé and Toliara (also known as Tuléar) have been especially well studied. However, much of the work is now outdated, and previously published species lists contain many misidentiﬁcations. Therefore, a review of previous taxonomic work was made and incorporated into the present survey results, resulting in a comprehensive list of Madagascar species. The results of this survey facilitate a comparison of the reef ﬁsh fauna of northwestern Madagascar, as well as the entire island, with other parts of the tropical IndoPaciﬁc. However, the list of ﬁshes from the survey area is still incomplete, due to the time restriction and the cryptic nature of many small reef species. Nevertheless, a basic knowledge of the cryptic component of the fauna in other areas and an extrapolation method that utilizes key “index” families can be used to accurately estimate the overall species total. Moreover, a wealth of supplemental information concerning the west Madagascar ﬁsh fauna has been gathered by previous researchers at Nosy Bé and Toliara. HISTORICAL BACKGROUND The earliest ﬁshes collected and described from the tropical southwest Indian Ocean were generally procured on various French expeditions in the early part of the nineteenth century. Several famous naturalists, including Commerson, Desjardins, Dussumier, Gaimard, Leschenault, Lesson, and Quoy, were among the most active collectors. The numerous new species they brought back to Paris were primarily described by Cuvier and Valenciennes in their Histoire Naturelle des Poissons, published in 22 volumes between 1828 and 1850. Although this work provided a useful reference for tropical Indo-Paciﬁc ﬁshes in general, it contained very little information speciﬁc to Madagascar as the early expeditions mainly visited Mauritius and Réunion. The ﬁrst major work dealing with Madagascar ﬁshes appeared in 1891. Histoire Naturelle des Poissons de Madagascar (Sauvage 1891) contained detailed descriptions, 40 Rapid Assessment Program illustrations, and a checklist of 1,238 species. However, the coverage is not limited to Madagascar, but also includes the Mascarene Islands (Mauritius, Réunion, and Rodriques) and Seychelles. It also includes many non-reefal species (e.g., freshwater, deepsea, and pelagic). Although approximately 950 reef species were listed from the region, the nomenclature is seriously outdated. The list contains numerous species now regarded as junior synonyms, and single species frequently have multiple listings under a host of invalid names. For example, of the 45 pomacentrids listed, only 14 are now considered valid. The remaining names are primarily misidentiﬁcations, using either names of species not occurring in the region or names that are invalid junior synonyms. The most useful publications that deal speciﬁcally with Madagascar ﬁshes were produced by French investigators working from research stations at Nosy Bé and Tuléar between 1950 and 1975. These include the papers by Fourmanoir (1957), Maugé (1967), and Vivien (1973ab, 1974, and as Harmelin-Vivien 1977 and 1979). The combined efforts of these authors resulted in a list of approximately 600 coral reef species. In addition, Bauchot and Bianchi (1984) published a guide to approximately 350 commercial ﬁsh species of Madagascar, including many reef inhabitants. Fricke (1999) presents the most recent summary of the Madagascar fauna. Although the main focus of this work is the Mascarene Group, Fricke also provides a lengthy table of species for key locations in the western Indian Ocean, including Madagascar. Approximately 700 species are indicated for Madagascar, of which about 590 are associated with coral reefs. However, the accuracy of this list is compromised by the inclusion of certain species that are presumed to occur on the island, but have not actually been recorded there. The United States Museum of Natural History, Washington, DC, currently maintains the world’s largest collection of ﬁsh specimens, including approximately 900 lots from Madagascar. Much of the Museum’s extensive Madagascar collection remains unreported in the literature. METHODS The ﬁsh portion of this survey involved approximately 45 hours of scuba diving by G. Allen to a maximum depth of 40 m. A list of ﬁshes was compiled for 30 sites. The basic method consisted of underwater observations, in most cases during a single, 60-90 minute dive at each site. The name of each species encountered was written on a plastic sheet attached to a clipboard. The technique usually involved a rapid descent to the maximum depth (usually 15-40 m) and then a slow, zigzag path was traversed on the ascent back to the shallows. The majority of time was spent in the 2-12 m depth zone, which consistently contains the largest number of species. The visual transect at each site included Reef Fishes of Northwestern Madagascar a representative sample of all bottom types and habitat situations. Only the names of ﬁshes whose identiﬁcation was absolutely certain were recorded. However, less than two percent of the species could not be identiﬁed to species level. The visual survey was supplemented with seven small collections procured with the use of a commercial ichthyocide (rotenone) and several specimens that were collected with a rubber-sling propelled multi-prong spear. The purpose of the rotenone collections was to ﬂush out small crevice- and sand-dwelling ﬁshes (e.g., eels and tiny gobies) that are seldom recorded with the visual technique. A total of 45 species were added by using this method. Coral and rocky reefs were by far the richest habitat in terms of ﬁsh biodiversity. The best sites for ﬁshes (Table 3.2) were invariably locations where coral reef was the dominant substratum, although there was usually a mixture of other bottom types, particularly sand or rubble. Seaweed and pure sand-rubble substrates were comparatively poor for ﬁshes. Silty bays and harbors also had impoverished ﬁsh communities, although several species were unique to this environment. In terms of general areas, the reef surrounding Nosy Hoa and those of the Mitsio Islands were the richest with an average of 154.3 and 129.2 species respectively. Table 3.2. Eight sites observed to have the highest diversity of ﬁsh among the 30 sites surveyed in northern Madagascar. SURVEY RESULTS A total of 463 species belonging to 191 genera and 65 families were recorded during the present survey. A review of previous work on Madagascar coral reef ﬁshes revealed an additional 325 species, 138 genera, and 26 families. Thus the total coral reef ﬁsh fauna of Madagascar as determined by this study consists of 788 species in 329 genera and 91 families (Appendix 4). The present report includes at least 75 new records for the island. Detailed accounts and illustrations of most Madagascar reef ﬁshes are included in Smith and Heemstra (1986), Allen and Steene (1987), and Lieske and Myers (1994). Further coverage will be provided in a treatise of western Indian Ocean ﬁshes currently in preparation by Heemstra and Randall. ANALYSIS OF SITE DATA The number of species found at each site is indicated in Table 3.1. The number of species at each site ranged from 33 to 166, with an average of 116.8 per site. Table 3.1. Number of species observed at each site. Site Species Site 1 132 11 141 21 107 2 111 12 166 22 119 3 145 13 156 23 119 4 118 14 82 24 89 5 152 15 114 25 33 6 133 16 101 26 152 7 114 17 117 27 123 8 104 18 140 28 122 9 106 19 91 29 154 10 98 20 92 30 73 Species Site Species Site No. Location Total ﬁsh spp. 12 Nosy Hao (northeastern corner) 166 13 Nosy Hao (channel on E. side) 156 29 Nosy Tanikely (west side) 154 5 Nosy Ankarea 152 26 Nosy Kivinjy 152 3 Marie Bank 145 11 Nosy Hao (southeastern corner) 141 18 Nosy Fanihy 140 CORAL FISH DIVERSITY INDEX (CFDI) Allen (1998) devised a convenient method for assessing and comparing overall reef ﬁsh diversity. The technique essentially involves an inventory of six key families: Chaetodontidae, Pomacanthidae, Pomacentridae, Labridae, Scaridae, and Acanthuridae. The number of species in these families is totaled to obtain the Coral Fish Diversity Index (CFDI) for a single dive site, relatively restricted geographic areas (e.g., Raja Ampat Islands), or countries and large regions (e.g., Indonesia). CFDI values can be used to make a reasonably accurate estimate of the total coral reef ﬁsh fauna of a particular locality by means of regression formulas. The latter were obtained after analysis of 35 Indo-Paciﬁc locations for which reliable, comprehensive species lists exist. The data were ﬁrst divided into two groups: those from relatively restricted localities (surrounding seas encompassing less than 2,000 km2) and those from much larger areas (surrounding seas encompassing more than 50,000 km2). Simple regression analysis revealed a highly signiﬁcant difference (p = 0.0001) between these two groups. Therefore, the data were separated and subjected to additional analysis. The Macintosh program Statview was used to perform simple linear regression analyses on each dataset in order to determine a predictor formula, using CFDI as the predictor variable (x) for estimating the independent variable (y) Coral Reefs of Northwest Madagascar 41 Chapter 3 or total coral reef ﬁsh fauna. The resultant formulae were obtained: (1) total fauna of areas with surrounding seas encompassing more than 50,000 km2 = 4.234(CFDI) - 114.446 (d.f = 15; R2 = 0.964; p = 0.0001) and (2) total fauna of areas with surrounding seas encompassing less than 2,000 km2 = 3.39 (CFDI) - 20.595 (d.f = 18; R2 = 0.96; p = 0.0001). The CFDI regression formula is particularly useful for large regions, such as the Philippines, where reliable totals are lacking. Moreover, the CFDI predictor value can be used to gauge the thoroughness of a particular short-term survey that is either currently in progress or already completed. For example, the CFDI obtained for northwestern Madagascar during the present survey is 176, and the appropriate Table 3.3. Coral ﬁsh diversity index (CFDI) values for restricted localities, number of coral reef ﬁsh species as determined by surveys to date, and estimated numbers using the CFDI regression formula (refer to text for details). 42 Locality CFDI No. reef ﬁshes Estim. reef ﬁshes Milne Bay, Papua New Guinea 337 1109 1313 Maumere Bay, Flores, Indonesia 333 1111 1107 Raja Ampat Islands, Indonesia 326 972 1084 Togean and Banggai Islands, Indonesia 308 819 1023 Komodo Islands, Indonesia 280 722 928 Madang, Papua New Guinea 257 787 850 Kimbe Bay, Papua New Guinea 254 687 840 Manado, Sulawesi, Indonesia 249 624 823 Capricorn Group, Great Barrier Reef 232 803 765 Ashmore/Cartier Reefs, Timor Sea 225 669 742 Kashiwa-Jima Island, Japan 224 768 738 Scott/Seringapatam Reefs, Western Australia 220 593 725 Samoa Islands 211 852 694 Chesterﬁeld Islands, Coral Sea 210 699 691 Sangalakki Island, Kalimantan, Indonesia 201 461 660 Bodgaya Islands, Sabah, Malaysia 197 516 647 Pulau Weh, Sumatra, Indonesia 196 533 644 Izu Islands, Japan 190 464 623 Christmas Island, Indian Ocean 185 560 606 Sipadan Island, Sabah, Malaysia 184 492 603 Northwestern Madagascar 176 463 576 Rowley Shoals, Western Australia 176 505 576 Cocos-Keeling Atoll, Indian Ocean 167 528 545 North-West Cape, Western Australia 164 527 535 Tunku Abdul Rahman Is., Sabah, Malaysia 139 357 450 Lord Howe Island, Australia 139 395 450 Monte Bello Islands, W. Australia 119 447 382 Bintan Island, Indonesia 97 304 308 Kimberley Coast, Western Australia 89 367 281 Cassini Island, Western Australia 78 249 243 Johnston Island, Central Paciﬁc 78 227 243 Midway Atoll 77 250 240 Rapa (S.E. Polynesia) 77 209 240 Norfolk Island, Australia 72 220 223 Rapid Assessment Program Reef Fishes of Northwestern Madagascar regression formula (3.39 x 176 - 20.595) predicts an approximate total of 576, indicating that approximately 113 additional species can be expected or that only 80 percent of the fauna was actually surveyed. On a much larger scale the CFDI can be used to estimate the reef ﬁsh fauna of the entire Indo-west Paciﬁc region, a frequent subject of conjecture. This method estimates a faunal total of 3,764 species, a ﬁgure that compares favorably with the estimated total of 3,950 proposed by Springer (1982). Moreover, Springer’s ﬁgure covers shore ﬁshes rather than reef ﬁshes and therefore includes species not always associated with reefs (e.g., estuarine ﬁshes). The total CFDI for northwestern Madagascar has the following components: Labridae (59), Pomacentridae (44), Acanthuridae (25), Chaetodontidae (20), Scaridae (20), and Pomacanthidae (8). Table 3.3 presents a ranking of IndoPaciﬁc areas that have been surveyed to date based on CFDI values. It also includes the number of reef ﬁshes thus far recorded for each area, as well as the total fauna predicted by the CFDI regression formula. Table 3.4 presents CFDI values, number of shallow reef ﬁshes recorded to date, and the estimated number of species based on CFDI data for selected countries or regions in the Indo-Paciﬁc. In most cases the predicted number of species is either similar to or lower than the number actually recorded, and is thus indicative of the level of knowledge. For example, an actual number that is substantially lower than the estimated total (e.g., Sabah) indicates incomplete sampling. The total number of species for the Philippines is yet to be determined and is therefore excluded. The CFDI total (200) used for Madagascar is based on the total number of species recorded to date (i.e., RAP survey totals plus additional species previously recorded) for the six indicator families: Labridae (75), Pomacentridae (49), Acanthuridae (27), Chaetodontidae (22), Scaridae (22), and Pomacanthidae (9). ANALYSIS OF THE MADAGASCAR REEF FISH FAUNA The following analysis is based on the present RAP survey and a review of past collections as reported in published literature and records of the United States National Museum of Natural History. The most abundant families in terms of number of species are gobies (Gobiidae), wrasses (Labridae), damselﬁshes (Pomacentridae), groupers (Serranidae), cardinalﬁshes (Apogonidae), blennies (Blenniidae), moray eels (Muraenidae), surgeonﬁshes (Acanthuridae), trevallies (Carangidae), parrotﬁshes (Scaridae), and soldierﬁshes (Holocentridae). These 11 families collectively account for about 56 percent of the total reef fauna (Table 3.5). Table 3.4. Coral ﬁsh diversity index (CFDI) for regions or countries with ﬁgures for total reef and shore ﬁsh fauna (if known), and estimated fauna from CFDI regression formula. Locality CFDI No. reef ﬁshes Estim. Reef ﬁshes Indonesia 504 2056 2019 Australia (tropical) 401 1627 1584 Philippines 387 ? 1525 Papua New Guinea 362 1494 1419 S. Japanese Archipelago 348 1315 1359 Great Barrier Reef, Australia 343 1325 1338 Taiwan 319 1172 1237 Micronesia 315 1170 1220 New Caledonia 300 1097 1156 Sabah, Malaysia 274 840 1046 Northwest Shelf, Western Australia 273 932 1042 Mariana Islands 222 848 826 Marshall Islands 221 795 822 Ogasawara Islands, Japan 212 745 784 French Polynesia 205 730 754 Maldives 219 894 813 Madagascar 204 790 749 Seychelles 188 765 682 Society Islands 160 560 563 Tuamotu Islands 144 389 496 Hawaiian Islands 121 435 398 Marquesas 90 331 267 Table 3.5. Largest ﬁsh families of Madagascar. Rank Family Species % of total species 1 Gobiidae 76 9.62 2 Labridae 74 9.37 3 Pomacentridae 49 6.20 4 Serranidae 41 5.19 5 Apogonidae 36 4.56 6 Blenniidae 35 4.43 7 Muraenidae 33 4.18 8 Acanthuridae 27 3.42 9 Carangidae 25 3.16 10 Scaridae 22 2.78 10 Holocentridae 22 2.78 Coral Reefs of Northwest Madagascar 43 Chapter 3 The relative abundance of Madagascar ﬁsh families is similar to other reef areas in the Indo-Paciﬁc, although the ranking of individual families is variable (Table 3.6). Zoogeographic afﬁnities Madagascar belongs to the western Indian Ocean faunal community, which forms an integral part of the greater Indo-West and central Paciﬁc biotic province. Reef ﬁshes of Madagascar are very similar to those inhabiting other areas within this vast region (from East Africa and the Red Sea to the islands of Micronesia and Polynesia). Although most families and many genera and species are consistently present across the region, the species composition varies greatly according to locality. Dispersal capabilities and the larval lifespan of a given species are usually reﬂected in its geographic distribution. Most reef ﬁshes have a relatively long pelagic stage, hence a disproportionate number of wide-ranging species inhabit tropical seas. Figure 3.1 presents the major zoogeographic categories of Madagascar reef ﬁshes. There are basically two major groups. The vast majority of species (about 72%) range widely across the Indo-Paciﬁc region, with distributions extending to either the western or central Paciﬁc. Aside from a small number of circumtropical ﬁshes, the remaining species are mainly conﬁned to the Indian Ocean, although distribution patterns within this large region are variable. Table 3.6. Fish family rankings in terms of species number for various localities in the Indo-Paciﬁc region. Data for the Chagos Archipelago is from Winterbottom et al. (1989), for northwestern Sumatra (Weh Island) from Allen (unpublished), for east-central Sulawesi (Togean and Banggai islands) from Allen (in press), for Milne Bay from Allen (1998), and for the Marshall Islands from Randall and Randall (1987). Madagascar Chagos NW Sumatra E. Cen. Sulawesi Milne Bay, PNG Marshall Islands Gobiidae 1st 1st 3rd 3rd 2nd 1st Labridae 2nd 2nd 1st 2nd 3rd 2nd Pomacentridae 3rd 5th 2nd 1st 1st 4th Serranidae 4th 3rd 5th 5th 4th 3rd Apogonidae 5th 8th 7th 4th 4th 8th Blenniidae 6th 9th 8th 7th 6th 6th Muraenidae 7th 4th 12th 15th 16th 5th Acanthuridae 8th 8th 5th 8th 8th 7th Carangidae 9th 16th 44th 13th 12th 14th Scaridae 10th 11th 8th 9th 10th 10th Holocentridae 10th 11th 10th 11th 11th 11th Family Figure 1. Zoogeographic Analysis of Madagascar Fishes Mad. & St. Brandon's Mad. & Comoros Mad., Mascar. & S. Africa Mad. & Mascarenes Madagascar Circumtropical Madagascar - E Africa Undetermined Indo-Pacific to Americas W. Ind. - Red Sea Indian Ocean W. Indian O. Indo-W. Pacific Indo-W. & C. Pacific 0 50 100 150 # species Figure 3.1. Zoogeographic analysis of the ﬁsh of Madagascar. 44 Rapid Assessment Program 200 250 300 Reef Fishes of Northwestern Madagascar Approximately 23 percent of the species range widely over the entire tropical Indian Ocean or are restricted to the western Indian Ocean (with many species entering the Red Sea). About 38 species (or 4.8%) are conﬁned to Madagascar and immediately adjacent regions including Mozambique, East Africa, Comoro Islands, St. Brandon’s Shoals, and the Mascarene Group (Mauritius, Réunion, and Rodriquez Island). A select group containing 12 species is conﬁned either to Madagascar alone or to Madagascar and the Comoro Group or St. Brandon’s Shoals (Table 3.7). All of the families represented by these 12 species, with the exception of Labridae, exhibit parental care of eggs, and either have a reduced pelagic larval stage or may lack it entirely. These families, particularly Pseudochromidae and Pomacentridae, exhibit considerable local and regional endemism throughout the Indo-Paciﬁc region. An additional seven species are mainly conﬁned to Madagascar and the Mascarene Islands, although three of these also range to northern South African waters: Cristatacirrhitus punctatus (Cirrhitidae), Pomacentrus agassizi and Stegastes limbatus (Pomacentridae), Thalassoma genivittatum (Labridae), Chlorurus cyanescens (Scaridae), Entomacrodus lemuria (Blenniidae), and Zebrasoma gemmatum (Acanthuridae). Comparison of faunal diversity from it. This phenomenon is well illustrated by numerous groups of reef ﬁshes. It is particularly evident in the damselﬁsh family Pomacentridae, as shown for the following localities (number of species in parentheses): Indonesia (138), Philippines (122), Papua New Guinea (109), northern Australia (95), Fiji (60), western Thailand (60), Madagascar (49), Red Sea (34), Society Islands (30), and Hawaiian Islands (15). Although Madagascar is approximately one-third as rich for reef ﬁshes as the “coral triangle” center of diversity, its fauna presents a unique blend of elements from the Indian and Paciﬁc oceans. It is not as isolated as the Mascarenes, and therefore does not have as many endemic species. The size and composition of its fauna is similar to other insular locations in the western Indian Ocean, including Mauritius, Seychelles, Chagos, and the Maldives (Table 3.8). Table 3.8. Comparison of reef-ﬁsh faunas of selected locations in the western Indian Ocean. Location No. species Maldives 894 Mauritius 800 Madagascar 790 Seychelles 765 Chagos 716 The general pattern of faunal diversity in the Indo-Paciﬁc region is well documented (Briggs 1999; Randall 1998). The center of distribution or “coral triangle” consists of Indonesia, Philippines, and Papua New Guinea. Marine biodiversity reaches its highest levels in this area, and a general faunal attenuation proceeds in any direction away Table 3.7. Endemic and near-endemic ﬁshes of Madagascar. Species Family Distribution Chlidichthys abruptus Pseudochromidae Madagascar & St. Brandon’s Chlidichthys sp. Pseudochromidae Madagascar & Comoros* Halimuraena lepopareia Pseudochromidae Madagascar & Comoros Halimuraenoides isostigma Pseudochromidae Madagascar Pseudochromis sp. 1 Pseudochromidae Madagascar Pseudochromis sp. 2 Pseudochromidae Madagascar Plesiops malalaxus Plesiopidae Madagascar Amphiprion latifasciatus Pomacentridae Madagascar & Comoros Pomacentrus atriaxillaris Pomacentridae Madagascar Pomacentrus caeruleopunctatus Pomacentridae Madagascar Paracheilinus hemitaeniatus Labridae Madagascar Flabellogobius fourmanoiri Gobiidae Madagascar * also at Aldabra Coral Reefs of Northwest Madagascar 45 Chapter 3 New and rare taxa Several undescribed and rarely encountered species were recorded during the current RAP survey. These are discussed in more detail in the following paragraphs. Chlidichthys abruptus Lubbock, 1977 (Pseudochromidae) – This species was previously known on the basis of seven specimens collected from St. Brandon’s (Cargados Carajos) Shoals in 1971 and 1976. It was thought to be restricted to this locality, but during the current survey four specimens were collected with rotenone at sites 13 and 9. The species is a secretive inhabitant of coral crevices that is difﬁcult to detect unless collected with chemical ichthyocides. Caesio sp. 1 (Caesionidae) – The fusilier family Caesionidae was reviewed by Carpenter (1987 and 1988). Considering the comprehensive nature of this recent work and the relatively wide distributional ranges of most species, the occurrence of three apparently undescribed species on reefs of northwestern Madagascar is surprising. Unfortunately, no specimens were collected, but all three species were photographed. The ﬁrst of these species generally resembles Caesio caerulaurea in shape and color, but possesses a black tip instead of a dark longitudinal streak on each caudal-ﬁn lobe. It has a thin yellow stripe immediately above the lateral line, which is similar to that of C. caerulaurea, but also has a faint suggestion of a second yellow stripe, just below the lateral line. It was common in the survey area, forming large mixed aggregations with other Caesio and Pterocaesio. Caesio sp. 2 (Caesionidae) – This species also resembles C. caerulaurea in general shape and color. It was illustrated by Allen and Steene (1987, plate 45-8) as Pterocaesio sp. However, the characteristic yellow stripe is wider than in C. caerulaurea, has dark margins, and extends slightly below the lateral line. The species forms aggregations and is often found in association with C. caerulaurea. Aware of the possibility that this ﬁsh might represent a temporary color-change variant of the latter species, the author watched several aggregations for extended periods. No color change was observed, providing convincing evidence that the pattern of this ﬁsh is a permanent feature. Pterocaesio sp. (Caesionidae) – This species is similar in shape and general coloration to Pterocaesio trilineata of the western Paciﬁc. However, in contrast to trilineata, which has a trio of bluish-white stripes on the upper back, it has approximately six pale stripes that become progressively thinner and less deﬁned towards the dorsal-ﬁn base. The species was common, forming mixed aggregations with various Caesio species. It was previously reported and illustrated (Randall and Kunzmann 1988) from western Sumatra and is therefore broadly distributed in the Indian Ocean. 46 Rapid Assessment Program Pomacentrus atriaxillaris (Pomacentridae) – This recently described species (see Allen, 2002) was discovered during the current RAP survey. It superﬁcially resembles Pomacentrus reidi from the Indo-Australian Archipelago. Both species have a similar shape and are overall light gray. They also possess 14 dorsal spines, a feature that, although not unusual, is shared by only a relatively small number of species in this large Indo-Paciﬁc genus. The Madagascar ﬁsh differs from reidi in lacking small blue spots on the head, and it has a much larger dark spot on the upper pectoral-ﬁn base, which also covers the entire axil. It was relatively rare in the survey area, with a group of about 10 adults seen at site 3 and single subadult ﬁsh observed at sites 14 and 17. Two specimens were collected. Pomacentrus caeruleopunctatus (Pomacentridae) – This is another recently described species (see Allen, 2002) that was discovered during the survey. It was common in mixed live coral and rubble areas at depths between about 5 and 15 m. It is similar to P. caeruleus of the Indian Ocean, and the two species are sometimes found in close proximity at the same sites. Both are generally blue with yellow pelvic ﬁns. However, caeruleus is generally more brilliant and has yellow on the anal and caudal ﬁns, whereas these ﬁns are blue in the new species. Close inspection of the pattern of the latter species reveals a pair of small blue dots and blue margin on most of the body scales. Several specimens were collected. Chrysiptera sp. (Pomacentridae) – A species of damselﬁsh (Pomacentridae) usually identiﬁed as Chrysiptera unimaculatus most likely represents an undescribed species. Allen (1991) reported that C. unimaculatus ranged from East Africa and the Red Sea to Fiji and illustrated several geographical color variants. The illustrated varieties from Mauritius and South Africa are similar to the Madagascar ﬁsh, and although further investigation is required, the southwestern Indian Ocean population is probably distinct. Adults are generally dark brown with a large, often poorly deﬁned black splotch on the rear portion of the dorsal ﬁn, in contrast to the pale-edged ocellus found in other populations. The species was common at ﬁve sites in shallow water inﬂuenced by moderate surge activity. Sites of special signiﬁcance for ﬁshes An assessment of the 2002 survey area presents special difﬁculties, primarily for the following reasons: (1) this was the ﬁrst Marine RAP survey for Madagascar or any place in the western Indian Ocean, hence there is little local or regional basis for comparison, and (2) coverage of the area was incomplete due to bad weather and mechanical problems associated with the charter vessel. Despite these limitations, the following areas were noteworthy for their overall ﬁsh diversity: Nosy Hao, northwestern Mitsio Islands in the vicinity of Nosy Ankarea, Nosy Tanikely, and Nosy Kivinjy. Reef Fishes of Northwestern Madagascar Nosy Hao – This island is surrounded by a good mix of habitat types, including an extensive reef ﬂat, exposed outer reef slope, and a vast area of reef ﬂat and patch reefs lying immediately to the east and south. The latter is bisected by a deep channel, where three dives were made (sites 11, 12, and 13). The channel has rich coral gardens along its edge and harbors an abundance of ﬁshes. A number of species associated with steep dropoffs and not recorded elsewhere during the survey were found here including Myripristis vittata, Pseudanthias evansi, P. pulcherrimus, Forcipiger ﬂavissimus, Hemitaurichthys zoster, Centropyge bispinosa, Chromis nigrura, Labropsis xanthonota, Nemateleotris magniﬁca, Acanthurus thompsoni, Melichthys indicus, and Arothron mappa. Mitsio Islands – The best reefs for ﬁshes were located off the northern tip of Mitsio Island (sites 6 and 7) and near Nosy Ankarea (site 5). The latter site, which was particularly noteworthy, featured a large patch reef in about 10 m, gradually sloping to about 25 m depth, and a good mix of both corals and ﬁshes. An impressive reef is also located adjacent to the southern side of Nosy Ankarea. Although it was not an ofﬁcial survey site, we made brief snorkelling observations, which revealed extensive stands of live coral and a rich ﬁsh assemblage. The above-water scenery at Nosy Ankarea was also impressive, with the jungle-clad island rising abruptly to a elevation of about 240 m above sea level. Nosy Tanikely – This site (29) had the most impressive ﬁsh community in terms of numbers of individuals. It also had the third highest species count for the survey. The habitat consisted of broad sandy areas interspersed with variable sized coral formations that gradually sloped to about 25 m depth. Among the many ﬁsh attractions were dense schools of Lethrinus lentjan and Lutjanus lutjanus. The island, which is situated near Nosy Bé, is a popular tourist destination. Although it has no ofﬁcial national conservation status, it is semi-protected with a locally-enforced no-ﬁshing zone. Therefore, the island serves as a good example of the beneﬁts of protective status. Nosy Kivinjy – This is a spectacular, small beehive-shaped island that rises abruptly from the sea to an elevation of 110 m. It requires about one hour to circumnavigate its base while scuba diving at depths varying from about 10 to 20 m. It contains a good mixture of bottom types including steep walls, rubble, and isolated coral patches. A well-developed reef of live coral is also found on the southeastern side. This was the ﬁfth richest ﬁsh site surveyed during this RAP. Impact of ﬁshing pressure Fishes were generally abundant throughout the survey area, although no remarkable concentrations of large commercially-important species such as groupers, jacks, or scombrids were noted. Snappers and the closely related (but less important for human consumption) fusiliers were generally common. The only species besides fusiliers that were seen in dense shoals were the emperor Lethrinus lentjan and the snapper Lutjanus lutjanus. It is extremely difﬁcult to assess the impact of local ﬁshing without having made prior visits to the area or having access to catch records. However, our community liaison team interviewed a number of ﬁshermen. Their general opinion was that stocks of commercially important ﬁshes had declined in recent years and maximum sizes had decreased. The Napoleon Wrasse ranges throughout the Indo-West and central Paciﬁc and commands a huge price in the Asian live-ﬁsh trade. It is one of the largest reef ﬁshes, reaching a length of 230 cm and weight in excess of 190 kg. Sadly, it has been ﬁshed to dangerously low levels in the Philippines and Indonesia, where it was formerly common. The species is an excellent indicator of ﬁshing pressure in the IndoAustralian Archipelago. However, it is difﬁcult to assess its use for this purpose in Madagascar without any knowledge of its previous abundance. During the current RAP survey it was observed on only three occasions (one small adult and two juveniles). Table 3.9 presents a comparison of sightings for various RAP surveys between 1997 and 2002. The highest concentrations of this ﬁsh have been noted at Milne Bay Province in Papua New Guinea, where there is a small human population and vast areas of nearly pristine reefs. Table 3.9. Frequency of Napoleon Wrasse (Cheilinus undulatus) sightings during Marine RAPs. No. sites where seen % of total sites Approx. no. seen Milne Bay, PNG – 2000 28 49.12 90 Milne Bay, PNG – 1997 28 52.83 85 Raja Ampat Islands – 2001 7 15.55 7 Togean/Banggai Islands – 1998 6 12.76 8 Northwest Madagascar 3 10.0 3 Calamianes Is., Philippines – 1998 3 7.89 5 Weh Island, Sumatra – 1999 0 0.00 0 Location Sharks appeared to be in short supply throughout the survey area, although observations were restricted due to low visibility at most dive sites. Five individuals belonging to three species (Reef Whitetip, Gray Reef, and Tawny Nurse) were seen at ﬁve sites. Apparently sharks have been heavily exploited for the Asian shark-ﬁn trade, a practice that still continues. This problem is prevalent throughout the IndoWest Paciﬁc. Coral Reefs of Northwest Madagascar 47 Chapter 3 REFERENCES Allen, G. R. 1991. Damselﬁshes of the world. Mentor, Ohio: Aquarium Systems. Allen, G. R. 1998. Reef Fishes of Milne Bay Province, Papua New Guinea. In: Werner, T. B. and G. R. Allen (eds.). A Rapid Marine Biodiversity Assessment of the coral reefs of Milne Bay Province, Papua New Guinea. Bulletin of the Rapid Assessment Program 11. Washington, DC: Conservation International. Allen, G.R. 2002. Descriptions of two new species of damselﬁshes (Pomacentridae: Pomacentrus) from Madagascar. Aqua, Journal of of Ichthyology and Aquatic Biology 6(2): 45-52. Allen, G. R. 2002. Reef ﬁshes of the Togean and Banggai Islands, Central Sulawesi, Indonesia. In: G. R. Allen and S. A. McKenna (eds.). A rapid biodiversity assessment of the coral reefs of the Togean and Banggai Islands, Sulawesi, Indonesia. RAP Bulletin of Biological Assessment. Washington, DC: Conservation International. Allen, G. R. and R. C. Steene. 1987. Reef ﬁshes of the Indian Ocean. Neptune City, New Jersey: T. F. H. Publications. Bauchot, M. L. and G. Bianchi. 1984. Guide des poissons commerciaux de Madagascar. Rome: Food and Agricultural Organization of the United Nations. Briggs, J. C. 1999. Coincident biogeographic patterns: Indowest Paciﬁc Ocean. Evolution 53(2): 326-335. Carpenter, K. E. 1987. Revision of the Indo-Paciﬁc ﬁsh family Caesionidae (Lutjanoidea) with descriptions of ﬁve new species. Indo-Paciﬁc Fishes 15: 1-56. Carpenter, K. E. 1988. FAO species catalogue. Vol. 8. Fusilier ﬁshes of the world. An annotated and illustrated catalogue of caesionid species known to date. FAO Fish. Synop. 125, Vol. 8: 1-75. Fourmanoir, P. 1957. Poissons téléostéens de eaux malgaches du Canal de Mozambique. Mémoires de l’Institut Sciences de Madagascar, série F, 1: 1-316. Fricke, R. 1999. Fishes of the Mascarene Islands (Réunion, Mauritius, Rodriquez). An annotated checklist, with descriptions of new species. Theses Zoologicae 31. Koenigstein, Germany: Koeltz Scientiﬁc Books. Harmelin-Vivien, M. L. 1977. Ecological distribution of ﬁshes on the outer slope of Tulear Reef (Madagascar). Proceedings of the Third International Coral Reef Symposium, Miami, Florida: 289-295. Harmelin-Vivien, M. L. 1979. Ichthyofaune des recifs coralliens de Tuléar (Madagascar): ecologie et relations trophiques. Unpublished Doctor of Sciences thesis. University of Marseille. Lieske, E. and R. F. Myers. 1994. Coral reef ﬁshes. Indo-Paciﬁc and Caribbean. London: Harper Collins. Lubbock, R. 1977. Fishes of the family Pseudochromidae (Perciformes) in the western Indian Ocean. Ichthyological Bulletin J. L. B. Smith Institute of Ichthyology 35: 1-28. 48 Rapid Assessment Program Maugé, L. A. 1967. Contribution préliminaire à l’inventaire ichtyologique de la région de Tulear. Travaux de la Station Marine d’Endoume, Marseille, fascicule hors série, Supplement 7: 101-132. Randall, J. E. 1998. Zoogeography of shore ﬁshes of the Indo-Paciﬁc region. Zoological Studies 37(4): 227-268. Randall, J.E. and A. Kunzmann. 1998. Seven new records of ﬁshes from Indonesia, with discussions of western Indian Ocean ﬁshes in southwestern Indonesia. Rafﬂes Bulletin of Zoology 46: 477-485. Randall, J. E. and H. A. Randall. 1987. Annotated checklist of ﬁshes of Enewetak Atoll and Other Marshall Islands. In: Vol 2. The natural History of Enewetak Atoll. Ofﬁce of Scientiﬁc and Technological Information U.S. Dept. of Energy: 289-324. Sauvage, H. E. 1891. Historie Naturelle de Poissons. In: Histoire Physique, Naturelle et Politique de Madagascar. Vol. 16. Paris: L’Imprimerie Nationale. Smith, M. M. and P. C. Heemstra (eds.). 1986. Smith’s sea ﬁshes. Johannesburg: Macmillan South Africa. Springer, V. G. 1982. Paciﬁc plate biogeography with special reference to shoreﬁshes. Smithsonian Contributions to Zoology 367: 1-182. Vivien, M. L. 1973a. Ecology of the ﬁshes of the inner coral reef ﬂat in Tuléar (Madagascar). Journal of the Marine Biological Association of India 15(1): 20-45. Vivien, M. L. 1973b. Contribution á la connaisance de l’éthologie alimentaire de l’ichtyofaune du platier interne des récifs coralliens de Tuléar (Madagascar). Téthys Supplement. 5(1): 221-308. Vivien, M. L. 1974. Ichtyofaune des herbiers de Phanérogames marines du Grand Récif de Tuléar (Madagascar). I: Les peuplements et leur distribution écologique. Téthys 5(2-3): 425-436. Winterbottom, R., A. R. Emery, and E. Holm. 1989. An annotated checklist of the ﬁshes of the Chagos Archipelago, Central Indian Ocean. Royal Ontario Museum Life Sciences. Contributions 145: 1-226. Chapter 4 A Basic Stock Assesment of Coral Reef Fishes for the Northwest Coast of Madagascar Jean Maharavo SUMMARY • A marine rapid assessment survey (RAP) was undertaken along the northwest coast of Madagascar in January 2001 by Conservation International and the Centre National de Recherche sur l’Environnment (CNRE), Madagascar. • A total of 55 species representing 18 families were classiﬁed as target species for reef ﬁsheries. Fusiliers (Pterocaesio spp.: Caesionidae) and snapper (Lutjanus lutjanus: Lutjanidae) were particularly abundant. • Counts of target species at individual sites ranged from 2 to 36 (mean = 13.41 ± 1.54). Counts of individual target ﬁshes at a single site ranged between 6 and 894 (mean = 189.50 ± 5.80). Estimated target ﬁsh biomass at a single site ranged between 3.11 and 1720.17 ton/km2 (mean = 182.30 ± 5.69 ton/km2). • In comparison with the results of past marine RAPs in other regions of the world, the mean total biomass for the northwest coast of Madagascar is the highest recorded with the exception of the Raja Ampat Islands, Indonesia, which exhibited slightly higher biomass levels. • Although fusiliers (Caesionidae) were the most abundant group of reef ﬁshes in terms of number of individuals, they are not targeted by local ﬁshers. INTRODUCTION Madagascar is one of the largest islands of the world. It is surrounded by about 5,600 km of coast and more than 1,200 km of fringing and barrier coral reefs that play a key role in the health of marine and terrestrial ecosystems as well as local human communities. Reefs form a barrier that protects coastal ecosystems against oceanic waves, and they provide breeding grounds for numerous marine animals. Coral reefs are also vital resources for Malagasy communities because much of the population lives in close proximity to the sea and depends on marine resources for a livelihood. Finally, reefs and coral sand beaches are an important resource for all of Madagascar because they have major ecotourism potential. Unfortunately, reefs have been severely degraded at many locations due to human activities, especially near the main coastal settlements. The primary objective of the present study was to collect information concerning commercial reef ﬁsh stocks and to provide tools for achieving an acceptable compromise between the needs of conservation and resource users. Coral Reefs of Northwest Madagascar 49 Chapter 4 MATERIAL AND METHODS Observations were carried out in January 2001 at 25 of 30 sites along the northwest coast of Madagascar. Data were collected visually while scuba diving and recorded with pencil on waterproof plastic paper. The visual census methodology outlined by Dartnal and Jones (1986) was employed with some modiﬁcations. Observations were made while slowly swimming along a 5 m wide corridor centered on a 100 m tape measure that was placed on the bottom along a predetermined depth contour, forming a survey area of approximately 500 m2 per transect. The time spent on each transect ranged from 40 to 45 minutes. Data were collected for two transects at each site, one at 10-13 m depth and the other at 4-6 m depth. Target species are deﬁned as ﬁshes that are important in traditional and artisanal catches. The average size and the numbers of individuals were recorded for every species observed. In the case of large schools, estimates were made to the nearest 50-100 ﬁshes. Estimation of the average size was made to the nearest ﬁve centimeters (5 cm). The cubic law Weight = 0.05(length)3 was used in the calculation of the biomass (expressed as ton/km2, weight in grams and length in centimeters). Bibliographic sources used included Lieske and Myers (1994) for the identiﬁcation of species and Elliot (1977) and Snedecor and Cochran (1984) for statistical analysis (calculation of the mean, conﬁdence limits, etc.). RESULTS AND DISCUSSION A total of 55 species representing 18 families were recorded. The most speciose families were Serranidae, Lethrinidae, Lutjanidae, Carangidae, Scaridae, and Caesionidae. Figure 4.1 illustrates the relative importance of different families. The category Other refers to those families with only one species recorded. In terms of abundance of individuals, the most important families were Caesionidae (18%), Lutjanidae (14%), Lethrinidae (9%), Scaridae (8%), Haemulidae (6%), and Serranidae (4%). Families with a single representative species accounted for 27% of the total. The high abundance of Casionidae and Lutjanidae was due to the proclivity of these families to produce large aggregations. This was especially true for Pterocaesio spp. and Lutjanus lutjanus. The results of the data analysis are presented in Table 4.1. Counts of target species ranged from 2 to 22 (mean = 10.33 ± 1.35) for 10-13 m transect and from 3 to 30 (mean 9.15 ± 1.73) for the shallow transect at 4-6 m depth. The data show that for target species, there is no signiﬁcant difference between the 10-13 m and 4-6 m transects. If we combine the count for target species from the various depth zones, the numbers range from 2 to 36 (mean 13.41 ± 1.54). The ﬁsh counts ranged from 5 to 600 (mean 125.91 ± 4.73) for 10-13 m transects and from 19 to 617 (mean 119.15 ± 6.26) for shallow transects. There was no signiﬁ- 50 Rapid Assessment Program cant difference between the two transect depths concerning ﬁsh counts of target species. The total ﬁsh count per site (for combined deep and shallow transects) ranged from 6 to 894 (mean 189.50 ± 5.80). The richest site was Tanikely with 894 ﬁsh counted for the two transects. The estimated biomass of target ﬁsh ranged from 3.11 ton/km2 to 1255.85 ton/km2 (mean 170.12 ± 5.53) for 1013 m transects and from 12.44 ton/km2 to 2184.49 ton/km2 (mean 270.73 ± 9.43) for shallow transects. Therefore, the data show there was a greater biomass of target ﬁsh in shallow water. The total biomass for each site ranged from 3.11 to 1720.17 ton/km2. The highest-ranking sites for the three data categories are indicated in Table 4.2. The data show that site number 29 (Tanikely Island) was by far the best site for the three values (target species, ﬁsh count, and estimated biomass). Biomass is the most important parameter from a ﬁsheries perspective. The high ranking of Tanikely is no doubt due to the reduced ﬁshing pressure because the island is a locally decreed marine reserve. Tanikely was followed by Tsarabanjina (site 1) and Ankarea (site 5). It is interesting to compare the “mean site total biomass” measurements for past RAP surveys conducted by Conservation International. This comparison provides a measure by which Madagascar can be ranked with other regions in the Indo-Paciﬁc. The mean total biomass per site for the current survey was second only to the Raja Ampat Islands, eastern Indonesia (Table 4.3). Caesionid ﬁshes are particularly abundant at both Madagascar and the Raja Ampats, where they comprise 18 and 52% of the total biomass, respectively. However, in contrast to Indonesia, these plankton-feeding midwater ﬁshes are seldom used for food in Madagascar. Figure 4.1. Percentage of target species belonging to different families. Other refers to families with only one species recorded. A Basic Stock Assesment of Coral Reef Fishes for the Northwest Coast of Madagascar Table 4.1. Summary table of coral reef ﬁsh stock assessment. 116.95 Site total No. Approx. target ﬁsh count species 20 324 354.80 26 21.24 22 385 189.95 3 112 49.49 20 418 120.92 119.27 ----- ----- ------- 6 51 119.27 224 350.26 ----- ----- -------- 16 224 350.26 -----11 ----54 ------126.40 ----6 -----20 -------206.02 ----13 ----74 -------166.21 8 7 71 12.42 11 617 505.66 16 688 259.04 9 12 179 163.84 7 23 20.77 16 179 92.30 10 11 8 3 83 5 205.26 6.15 ---10 ----160 -----162.84 8 11 83 165 205.26 84.49 12 13 66 59.05 4 33 24.35 17 99 41.70 13 5 60 67.91 13 130 176.49 17 190 122.2 14 2 6 3.11 ------ ------- ------- 2 6 3.11 15 2 28 7.18 ------ ------- ------- 2 28 7.18 16 12 133 180.20 ------ ------- ------- 12 133 180.20 17 6 43 23.03 ----- ------- -------- 6 43 23.30 18 19 20 21 9 ---------------- 83 --------------- 12.24 ---------------------- 3 ------------- 22 ------------------ 12.44 ------------------- 11 ------------- 105 ------------- 12.34 --------------------- 22 11 67 44.55 ----- -------- ------- 11 67 44.55 23 9 24 11.55 ------ -------- ------- 9 24 11.55 24 13 66 31.19 9 36 25.03 19 102 28.11 25 26 ------------ ----------- ------------- -------- ------------- ------------- ------- --------- ------------- 27 11 92 56.88 6 19 13.83 14 111 35.35 28 6 99 79.86 ------- ------ ------- 6 99 79.86 29 22 600 1255.85 30 294 2184.49 36 894 1720.17 30 n min max 12 24 2 22 56 24 5 600 123.16 24 3.11 1255.85 ----13 3 30 -----13 19 617 --------13 12.44 2184.49 12 24 2 36 56 24 6 894 123.16 24 3.11 1720.17 Avg 10.33 125.91 170.12 9.15 119.15 270.73 13.41 189.50 182.30 Std. E. 1.35 4.73 5.53 1.73 6.26 9.43 1.54 5.80 5.69 Approx. ﬁsh count Biomass (ton/km2) 1 Transect No. target species 16 Approx. ﬁsh count Biomass (ton/km2) 592.66 Transect 2 No target species 10 267 57 2 19 359 358.66 7 3 17 306 192.36 4 6 51 5 16 6 7 Sites Biomass (ton/km2) Most abundant species (% of total ﬁsh count) Pterocaesio spp. (32.40%) Lutjanus lutjanus (41.85%) Lethrinus mahsena (23.92%) Lethrinus mahsena (78.43%) Pterocaesio pisang (44.64%) ------------------------Naso unicornis (14.86%) Lutjanus lutjanus (29.06%) Pterocaesio marri (27.93%) Naso unicornis (26.50%) Lutjanus gibbus (24.24%) Pterocaesio pisang (24.24%) Caranx melampygus (25.26%) Epinephelus sp (33.33%) Lutjanus lutjanus (89.28%) Pterocaesio spp. (37.59%) Lutjanus kasmira (34.88%) Pterocaesio spp. (33.33%) -------------------------------------------------------------------------------Scolopsis bimaculatus (29.85%) Siganus argenteus (25%) Plectorhinchus sordidus (11.76%) ----------------------------------------------------Pterocaesio pisang (34.69%) Pterocaesio pisang (57.57%) Lutjanus lutjanus (36.35%) Pterocaesio pisang (89.28%) Ranking of most abundant species 1-Scolopsis bimaculatus (12 sites) 2-Plectorhinchus gaterinus (10 sites) 3-Pterocaesio pisang (8 sites) Coral Reefs of Northwest Madagascar 51 Chapter 4 CONCLUSIONS REFERENCES • Sharks were rarely observed during the present survey, which is not surprising considering the intensive shark ﬁshing in recent years that various ﬁsheries projects have actively encouraged. Sharks have been ﬁshed out in large numbers to supply the demand for shark ﬁn, mainly in the Asian market. A similar situation (i.e., overﬁshing) was noted for sea cucumbers. • Several sites have considerable potential as conservation reserves, notably Tanikely, Tsarbanjina, and Ankarea. • The CI-RAP provided an excellent opportunity for capacity building for the national participants. For example, a variety of new survey techniques were introduced. • The results of the survey will prove useful in developing guidelines for working out a compromise between the needs of conservation and resource users. • Future marine surveys are recommended for Madagascar, especially in the Southwest and Northeast. Allen, M., J. Kinch, and T. Werner. 2003. Coral reef resources of Milne Bay Province, Papua New Guinea. In: G. R. Allen, J. Kinch, S. A. McKenna, and P. Seeto (eds.). A Rapid Marine Biodiversity Assessment of Milne Bay Province, Papua New Guinea. RAP Bulletin of Biological Assessment. Washington, DC: Conservation International. Dartnall, H. J. and M. Jones. 1986. A manual of survey methods of living resources in coastal areas. ASEANAustralia Cooperative Programme on Marine Science Hand Book. Townsville: Australian Institute of Marine Science. Elliot, J. M. 1977. Some methods for statistical analysis of benthic invertebrates. Freshw. Biol. Assoc. Sci. Publ., UK., 25: 1-156. Ingles, J. 2000. Fisheries of the Calalmianes Islands, Palawan Province, Philippines. In: Werner, T. B. and G. R. Allen (eds.). A Rapid Marine Biodiversity Assessment of the Calamianes Islands, Palawan Province, Philippines. RAP Bulletin of Biological Assessment 17. Washington, DC: Conservation International. Table 4.2. Best sites for target ﬁshes in three data categories for the northwest coast of Madagascar. Number of target species Approximate ﬁsh count Estimated biomass (ton/km2) Site Value Site Value Site Value 1 29 36 29 894 29 1720.17 2 2 22 8 688 1 354.80 3 1, 3 20 3 418 5 350.26 4 24 19 2 385 8 259.04 5 12, 13 17 1 324 10 205.26 6 5, 8, 9 16 5 224 2 189.95 7 27 14 13 190 16 180.20 8 7 13 9 179 7 166.21 9 30 12 11 165 30 123.6 10 11,18, 22 11 16 133 13 122.2 Rank Table 4.3. Comparison of mean site total biomass for this survey and past CI RAP surveys. CI-RAP Mean total biomass (ton/km2) per site Raja Ampat Is., Indonesia 2001 208.97 Northwest Madagascar 2002 182.30 Milne Bay Province, Papua New Guinea 2000 125.56 Togean and Banggai Is., Indonesia 1998 66.49 Calamianes Is., Philippines 1998 16.94 Source : Allen et al. 2003, La Tanda 2002 a and b, Ingles 2000, and data from present survey. 52 Rapid Assessment Program A Basic Stock Assesment of Coral Reef Fishes for the Northwest Coast of Madagascar La Tanda. 2002a. Chapter 4. A basic stock assessment of economically important coral reef ﬁshes of the Raja Ampat Islands, Papua Province, Indonesia. In: S. A. McKenna, G. R. Allen, and S. Suryadi (eds.). A Marine Rapid Assessment of the Raja Ampat Islands, Papua Province, Indonesia. RAP Bulletin of Biological Assessment 22. Washington, DC: Conservation International. La Tanda. 2002b. Coral reef ﬁsh stock assessment in the Togean and Banggai Islands, Sulawesi, Indonesia. In: G. R. Allen and S. A. McKenna (eds.). A Rapid Biodiversity Assessment of the Coral Reefs of the Togean and Banggai Islands, Sulawesi, Indonesia. RAP Bulletin of Biological Assessment. Washington, DC: Conservation International. Lieske, E. and R. Myers. 1994. Coral Reef Fishes – Indo-Paciﬁc & Caribbean. London: Harper Collins. Snedecor, G. W. and W. G. Cochran. 1984. Methodes statistiques. Traduit par H. Boelle et Camahji. Paris: Association de Coordination Technique Agricole. Coral Reefs of Northwest Madagascar 53 Chapter 5 The Condition of Coral Reefs in Northwest Madagascar Sheila A. McKenna SUMMARY 54 Rapid Assessment Program • Reef condition refers to the general “health” of a particular site as determined by assessment of key variables including natural and human-induced damage or stress and general biodiversity as deﬁned by major indicator groups (corals and ﬁshes). • Reef condition was assessed at 30 sites in northwest Madagascar and included reefs in the vicinity of Mitsio Island, Cape Saint Sebastian, and Nosy Bé. • A Reef Condition Rating (RCR) was calculated for each site to indicate its general diversity and health. Essentially, the RCR is a semi-quantitative measure that combines the total species number of reef ﬁsh and hard corals with the percent of live hard coral coverage. Points are subtracted from this sum according to evidence of threat or stress and its relative extent or impact on the reef. RCRs allow for comparison of reefs surveyed in the region. • Reef survey sites with the highest RCR included Anakarea (site 5), Northeast Nosy Hao (site 12), Nosy Fanihy (site 18), and Nosy Tanikely (site 29). These sites are strong candidates for ofﬁcial protection and management. • The most frequently observed damage was from the coral predator Acanthaster plancii and occurred on 20% of the reef sites surveyed. The extent of impact on those sites ranged from light to excessive. • Evidence of threat or stress from bleaching and coral pathogens was low for the sites surveyed. A very low incidence of coral bleaching was noted and appeared to be in the early stages at 20% of the reef sites surveyed. Coral pathogens were seen only on two coral colonies at one reef site. • Damage resulting from anchors and ﬁshing nets was infrequently observed with only three (or 10%) of the reef sites surveyed having entangled ﬁshing nets and only one site having anchor damage. No evidence of physical damage from severe storms or cyclones was observed on the reef sites surveyed. • Evidence indicating possibly light siltation and ﬁshing pressure was noted during the survey. Further long-term monitoring and study would be needed to determine quantitatively the extent of activity and impact. No excessive sedimentation stress was observed (i.e., dead reef substrate covered in silt) on any of the reefs surveyed. • Charismatic marine fauna observed during the survey were manta rays, sharks, and turtles. The Condition of Coral Reefs in Northwest Madagascar INTRODUCTION Surrounding Madagascar’s 4,828 km coastline are 2,230 km2 of coral reef area (Spalding et al. 2001). The majority of reef studies in Madagascar have focused on the reefs in the southwest or Toliara (Tulear) region followed by reefs in the Nosy Bé area. This survey focused on the coral communities and reefs lying off the northwest coast of Madagascar, stretching approximately 200 km from Nosy Hao to Nosy Iranja. The main areas included Andramaimba Bay, Cape Saint Sebastian, Mitsio Islands, Nosy Bé (including Sakatia and Nosy Tanikely), Ambavatoby Bay, and Nosy Iranja. The conditions of 30 reef sites are described here to provide a snapshot of reef “health” as observed during the time period of January 6–26, 2002. Several sources of damage or stress to the coral reefs of Madagascar and the Indian Ocean have been documented. Although a major bleaching event occurred in the Indian Ocean with the 1998 El Niño, Madagascar was not as heavily impacted as other islands of the Indian Ocean (Quod & Bigot 2000). As a result of this severe bleaching event, the CORDIO (Coral Reef Degradation in the Indian Ocean) project was formed to increase knowledge and monitor reef status and recovery in the region. For the Madagascar reefs that were monitored following the bleaching event, positive signs of recovery are reported (Mahavaro, pers. comm.). In addition to bleaching, other observed disturbances and threats to the coral reefs of Madagascar include over-exploitation of marine resources (e.g., ﬁshing and coral extraction), run-off from poor land-use practices (e.g., deforestation), point sources of pollution (e.g., sewage and sugar cane efﬂuent), destructive ﬁshing practices, and predation by the Crown of Thorns starﬁsh (COTS). A global risk assessment of coral reefs found Madagascar’s reefs to be at medium to high risk (Bryant et al. 1998). MATERIALS AND METHODS At each survey site the reef condition team collected substrata/biota data of the benthos. Transects were used to sample the benthos as described in English et al. (2000). The following is a brief description of the transects: a 100 meter transect tape was placed along the bottom of the reef as close as possible to the biota/substrata. Depending on the topography of each reef, transects were placed at two depth zones (4-6 m and 12-16 m). The biota/substrata was sampled at selected 0.5 m intervals (for 40 sample points) along 20 m segments of the 100 m transect. Below each sampling point, the type of substrata/biota is identiﬁed or characterized as follows: hard coral (hc), soft coral (sc), sponge (sp), macro algae (ma), turf algae (ta), calcareous algae (ca), rubble (rb), other, mud, and bare substrata (bs). The “other” category includes invertebrates such as tunicates, sea stars, sea cucumbers, etc. After the ﬁrst 20 of the 100 m was sampled, the diver would skip 5 m and then continue sampling another 20 m (40 points) along the transect. This allowed for rep- lication during sampling, with four 20 m segments of each transect sampled at half-meter intervals per depth. Because of their limited reef structure, only one depth was surveyed at sites 2, 10, 14-17, 22, 23, 28, and 30. Weather and dive conditions did not permit transects to be surveyed at reef sites 25 and 26. Prior to surveying the benthos/substrata, ﬁsh counts were performed along each transect to assess ﬁsh biomass (see Chapter 4). The abundance of commercially and artisanally important ﬁshes such as groupers (Serranidae) and snappers (Lutjanidae) was recorded. Charismatic marine fauna was also noted at each reef site. These include sharks, manta rays, turtles, dugongs, etc. The reef condition team also recorded any visible signs of damage, threats, or disturbance at each reef site. Evidence of disturbance, damage, or threat was rated according to the relative amount or level of impact/frequency (none, light, moderate, and excessive). The divers looked for evidence of damage from ﬁshing (nets, ﬁsh pots, lines, explosives, cyanide, nets), boating activities (anchor damage, grounding scars, ﬁn marks from snorkelers), and storms or cyclones. Damage from the coral predators Acanthaster plancii and Drupella cornus on the reef was detected by the presence and number of individuals seen or by feeding scars on the coral. The team documented substantial reef damage caused by the Crown of Thorns starﬁsh (A. plancii). The coral-eating mollusc Drupella cornus also can damage coral, but usually not as severe as A. plancii. Population outbreaks of Drupella cornus have caused severe damage in some cases (e.g., Ningaloo Reef in Western Australia). The reef condition teams’ observations were supplemented by other divers of the RAP team after the site survey dive had been completed. Bleaching refers to the discoloration of coral tissue color—the more discolored the coral tissue, the more severe the bleaching. Light (or early stages of ) bleaching is indicated by a slight discoloration of the coral tissue. Moderate or extreme bleaching is usually indicated by the coral tissue being transparent, opaque, or clear in color with the coral skeleton visible. The number of colonies showing signs of bleaching and the level of tissue discoloration indicates the extent of the bleaching on the reef. In addition to bleaching, coral pathogens or diseases on the reef were noted. Several coral diseases have been identiﬁed to occur on hard and soft coral. Diseases are identiﬁable by a distinctive banding or pattern of discoloration on the surface of hard and soft coral. For example, black band disease on hard corals is evident by an obvious black band across the coral head—behind the band the coral skeleton is visible and the coral tissue is dead and gone. On the other side of the band the coral surface looks normal. Descriptions of coral disease have been well documented and can be found on the Web at http://ourworld.compuserve.com/ homepages/mccarty_and_peters/coraldis.htm. Other forms of threat or pressure on the reef cannot be diagnosed equivocally without testing, monitoring, or experimentation. Such types of threat or pressure include pollu- Coral Reefs of Northwest Madagascar 55 Chapter 5 tion/eutrophication, ﬁshing pressure, siltation, and freshwater runoff. In some cases the source of the damage (e.g., sewage outfall pipe, deforested area along the shoreline, coastal development, and river outfall) can be seen from the reef site, thereby providing qualitative evidence. An abundance of algae with low coral cover can be an anecdotal indicator of pollution/eutrophication on reefs. However, the population of herbivores and type of algae need to be considered. The presence of ﬁshers actively ﬁshing or a low abundance of target biota (e.g., sea cucumbers or groupers) on the reef site may indicate ﬁshing pressure, but the frequency and extent of marine resource use and abundance of stocks need to be further investigated and monitored to obtain quantitative data to provide empirical evidence. High percent cover of mud or silt on the reef benthos indicates siltation stress. These types of threats or disturbance are better characterized by direct measurements of speciﬁc parameters (e.g., nutrients in the water column, stock abundance and ﬁshers activity, sediments, and percent cover of biota/substrata) over a long sampling period of at least one year or more. The nature of the rapid assessment only allows for observation of what may under further investigation prove to be eutrophication/pollution, ﬁshing pressure, siltation, or runoff on the reef site and provides an important ﬁrst step in determining reef health and the need for follow up. A reef condition rating (RCR) was calculated for each site and was modiﬁed from CI’s previous reef condition index (McKenna et al. 2002). The RCR is calculated by combining data collected from each reef site surveyed. This includes the summation of the total number of species from hard corals and reef ﬁshes observed and the average percent of hard coral cover. In cases where only one depth (i.e., one transect) was surveyed, that percentage is used in the RCR calculation. The relative amount of damage or stress expressed as penalty points at each site is then subtracted from the sum of the hard coral and reef ﬁsh and percent hard coral cover to determine the ﬁnal RCR. The number of penalty points to be subtracted is determined by the relative amount of damage or stress observed on the reef. The four levels of damage or threat include none, light, moderate, and extensive for the following factors: predation from Acanthaster plancii and Drupella, bleaching, coral pathogens, and physical damage from anchoring, explosives, and nets. If no damage is noted, no points are subtracted. For each factor and its corresponding level, points are deducted as follows: for light damage, two points are subtracted from the sum of the number of total reef ﬁsh and coral species and percent live coral cover; for moderate damage, four points; and for excessive damage, six points. No points were subtracted for anecdotal evidence of siltation or ﬁshing pressure. However, sites where these were noted are indicated under site description and in summary Table 5.2. The resultant rating is a semi-quantitative tool for comparing sites within the survey region to identify reefs of high diversity and good health for protection as well as reefs that have been damaged or are under threat or are in need of further study and mitigation. 56 Rapid Assessment Program INDIVIDUAL SITE DESCRIPTIONS Mitsio Island 1. South end of Tsarabajina Time: 2430 hours, dive duration 90 minutes; depth range 1-11 m; visibility 5-7 m at 0-4 m depth and 10 m at 5-19 m depth; temperature 30º C; current none. Site description: Small rocky islet with scattered reef that is moderately exposed; dominant hard coral included mostly Favids with Acropora and Pocillopora; dominant soft coral included Sinularia and Rhytisima. Other noted biota observed included the coral predators Acanthater plancii (approximately 50 individuals) and Drupella cornus. The boring sponge Cliona was also observed. Also noted were cyanobacteria, a slight incidence of coral bleaching, one manta ray, Manta birostris, and one hawksbill turtle Erectmochelys imbricata. RCR = 270. 2. Inshore northwest corner of Tsarabajina Time: 1600 hours, dive duration 90 minutes; depth range 1-16 m; visibility 8-10 m across depths; temperature 30º C, exposure variable with some current, protected in some areas. Site description: Fringing reef with dense coral coverage 313 m; sparse hard coral with some soft coral at base of reef slope; dominant hard coral included species of Favids and Acroporids; most abundant soft coral was Rhytisimia. Acanthaster plancii feeding scars observed on hard coral, although no individuals were observed. RCR = 232.4. 3. Marie Bank Time: 0800 hours, dive duration 90 minutes; depth range 1-13 m; visibility 10-12 m across depth; temperature 30º C; current none. Site description: Platform reef with moderate wave exposure; dominant hard coral taxa were Favids and foliose Tubinaria; dominant soft coral was Sinularia and Rhytisima. Approximately 30 individuals of Acanthaster plancii were observed, and the damage from this predator was classiﬁed as moderate. Other common biota included Diadema and star ﬁsh. Net damage with some light incidence of coral bleaching was noted on the reef. Fisherman huts were seen on the shore. RCR = 253.5. 4. Maromby Village Time: 1630 hours, dive duration 46 minutes; depth range 1-10 m; visibility 8 m across depth; temperature 30º C; current none. Site description: Fringing reef with light wave exposure; dominant hard coral taxa were Favids and Acropora, with a The Condition of Coral Reefs in Northwest Madagascar good variety of coral species noted (approximately 119 species). Dominant soft corals were Sinularia and Rhytisima. Coral garden type reef with some sections having nearly 100 percent hard coral cover, although transect surveys averaged 18.7% + 6.33 for deep depth, with 57.5 % + 2.5 for shallow depth. Moderate ﬁshing activities likely to occur on the reef from ﬁshers in the village. Very light siltation observed that may be natural or attributable to runoff from the village. Both the ﬁshing and siltation observations would need further monitoring and study to determine extent. RCR = 284.5. 5. Anakarea Time: 0830 hours, dive duration 90 minutes; depth range 124.3 m; visibility 7-8 m at 0-11 m depth and 12 m visibility below 12 m depth; temperature 30º C; current none. Site description: Fringing reef with moderate exposure; dominant hard coral taxa were Favids, Acroporids, and Mussids; distinct zonation of Favids in deeper depths. Dominant soft corals were Cespitularia and Pachyclavularia. Slight coral bleaching noted on A. variolosa. No coral trout observed, which may indicate some light ﬁshing pressure. RCR = 348.5. 6. Nosy Fisaka Time: 1400 hours, dive duration 100 minutes; depth range 1-18 m; visibility 6-7 m across depth; temperature 30º C; current none. Site description: Fringing reef with light to moderate exposure; in shallow depths (<6m) Favids, Acroporids, and Pocilliporids were dominant hard corals while Sinularia and Nepthea were dominant soft corals; in deeper depths (> 6m) Favids, Poritids, and Goniopora spp. were dominant hard corals with Rhytisima and Cespitularia the dominant soft corals. Two to three coral colonies were observed to have some kind of coral pathogen. Slight incidence of coral bleaching; excessive damage likely from A. plancii as evidenced by dead Acropora plates covered with turf algae. This is consistent with other observed remnants of feeding activity by A. plancii on reefs. RCR = 283.1. 7. Northwest corner of Mitsio Island Time: 1700 hours, dive duration 90 minutes; depth range 1-12 m; visibility 6 m across depth; temperature 30º C; current none. Site description: Fringing reef with light to moderate exposure; Favids, Acroporids, and Poritids dominant hard coral taxa across depth, with Sinularia and Nepthea dominant soft corals. Village located onshore approximately < 1 km from survey site. RCR = 268.8. Cape Saint Sebastian 8. Kalomisampa Rocks Time: 1030 hours, dive duration 90 minutes; depth range 1-19 m; visibility 7 m across depths; temperature 30º C, exposure variable, moderate to heavy depending on position relative to rocks. Site description: Fringing reef surrounding rocky islets, complex topography in < 6m depth with simple topography at approximately 10 m depth. Favids, Acroporids, and Pocilliporids dominant hard coral across depth with Xenia and Lemnalia dominant soft corals across depth. Interesting site for commercial ﬁsh. No observed threat or damage noted. RCR = 244. 9. Ambariokisimany Time: 1615 hours, dive duration 75 minutes; depth range 1-25 m; visibility 3.5 m across depths; temperature 27.8º C, exposure moderate to heavy with current. Site description: No reef development, patchy rocky area covered with corals; in shallow depths (< 6m) Favids and Pocilliporids dominant hard corals while Sinularia and Cespitularia dominant soft corals; in deeper depths (> 6m) Favids and Pocilloporids dominant hard corals with Cespitularia dominant soft coral. Two hawksbill turtles (Erectmochelys imbricata) and one shark (unidentiﬁed taxonomically) were observed. RCR = 223.4. 10. Magnet Platform Time: 1045 hours, dive duration 65 minutes; depth range 1-40 m; visibility 10 m at 0-19 m depth and 15 m visibility below 19 m depth; temperature 30º C; exposure oceanic. Site description: Rock platform with no extensive reef development; top of platform 15 m depth with scant patches of soft coral and sponges; dropoff vertical with wall having soft and hard coral with sponges; overall most dominant hard coral was Stylastera followed by Agaricids and Fungids, with dominant soft corals Lithophyton and Gorgonia. Large coral trout approximately 1.8 m observed off wall; one white tip reef shark (Triaenodon obesus) observed. No threat or damage observed. RCR = 146.5. Northern Reefs 11. Southeast Nosy Hao Time: 1530 hours, dive duration 90 minutes; depth range 1-30 m; visibility 8 m at 0-19 m depth and 15 m below 19 m depth; temperature 27.8º C; sheltered from sea with potential of exposure to strong current. Site description: Reef on edge of passage (channel) off of ﬂat island, gradual slope (about 17º) with extensive reef ﬂat Coral Reefs of Northwest Madagascar 57 Chapter 5 till 17 m then drops off to 30 m. Large hard coral colony, approximately 3 m across, of Alveopora cf. spongiosa. Favids and Agaricids dominant hard corals while Gorgonids and Cespitularia dominant soft corals across depth. Interesting commercial ﬁsh noted. No threat or damage observed, although ﬁshing boats seen on beach. Mangroves are reported in the center of the island. RCR = 282.7. 12. Northeast Nosy Hao Time: 0830 hours, dive duration 75 minutes; depth range 126 m; visibility 15 m at 0-19 m depth and an average of 20 m visibility below 19 m depth; temperature 27.8º C; moderate exposure with current. Site description: Fringing reef occurring at opening of channel between two islands. Hard coral species mixed in deeper depths (+15m) with most dominant Poritids and Favids, while dominant soft coral was Cespitularia. In shallow depths (< 15m) dominant hard corals were Acroporids, Poritids, and Favids, with dominant soft coral Xenia. Other than discarded ﬁshing line observed in shallows, no other damage or threat observed. Charismatic marine fauna included Napoleanﬁsh, Cheilinus undulatas, gray reef shark, Carcharinus amblyrhynchos, and dogtooth tuna (Gymnosarda unicolor). RCR = 328.5. RCR = 328.5. 13. Nosy Hao channel (east side) Time: 1330 hours, dive duration 90 minutes; depth range 1-30 m; visibility 8-10 m across depth; temperature 27.8º C; sheltered and exposed to the sea depending on location, exposure to current at times. Site description: Reef on edge of passage, similar to site 11 with slope 10-15º that goes into a sheer drop at 16 m and ﬂattens to a sand and rubble bottom at 30 m. In shallow depths (<6 m) Poritids, Acroporids, and Pocilliporids dominant hard corals while Lemnalia dominant soft coral; in deeper depths (> 6 m) Favids and Acroporids dominant hard corals with Cespitularia dominant soft corals. More ﬁsh observed in shallow than deeper depths. No damage or threat observed. RCR = 295.4. 14. Andavakalovo Island Time: 0755 hours, dive duration 70 minutes; depth range 1-19 m; visibility 3-4 m across depth; temperature 27.8º C; moderate exposure to the sea with wave surge. Site description: Fringing reef off of rugged cliff face of island. Across depth, Acroporids, Favids, and Pocilloporids dominant hard corals while Rhytisima and Capnella dominant soft corals. No damage or threat observed. RCR = 235.9. 15. Lakandava, North Beach Time: 1300 hours, dive duration 60 minutes; depth range 1- 58 Rapid Assessment Program 13 m; visibility 4 m across depth; temperature 27.8º C; light to sheltered exposure. Site description: Fringing reef in bay surrounded by karst islets. Across depth, Favids, Mussids, and Acroporids dominant hard corals with Cestipularia and Gorgonids dominant soft corals. Anecdotal evidence that Holothurians are harvested by Chinese ﬁshers. Very slight siltation observed that is likely to be natural. RCR = 229. 16. Ambatomaranitra Time: 1600 hours, dive duration 75 minutes; depth range 1-14 m; visibility 5-8 m across depth; temperature 27.8º C; moderate exposure with waves shoaling in shallows. Site description: Submerged coral community on rock (similar to platform reef ). Across depth, Favids, Pectinids, Pocilloporids, and Acroporids dominant hard corals, with Capnella and Sinularia dominant soft corals. No damage or threat observed. RCR = 230.5. 17. Lakandava, south wall Time: 0830 hours, dive duration 75 minutes; depth range 1-31 m; visibility 3-4 m across depth; temperature 28º C; moderate exposure with surge in shallows. Site description: Fringing reef off of cliff face of island, slope approximately 30o then increasing to 70o in shallows (0-4m). In shallow depths (<6m) Poritids, Agaricids, and Favids dominant hard corals, with Sinularia and Sacrophyta dominant soft corals; in deeper depths (> 6m) Favids, Pectnids, and Agaricids dominant hard corals with Ctenocella and Cespitularia dominant soft corals. Reef depth at 25 m plus characterized by sand and sediment with patches of coral. No damage or threat observed. RCR = 264.1. Reefs in vicinity of Nosy Bé 18. Nosy Fanihy Time: 0930 hours, dive duration 75 minutes; depth range 1-16 m; visibility 6 m across depth; temperature 29º C. Site description: Fringing reef with moderate exposure. Across depth, Acroporids and Favids dominant hard corals, with Rhytisima and Sinularia dominant soft corals. In shallow depths (< 8m) plates and branching Acropora colonies noted; in deeper depths (approximately > 8m) mainly sand with patches of coral outcrops. Feeding scars from Acanthaster plancii observed, some scars measuring 22 cm in diameter. Early signs of bleaching indicated by light tissue color of Acropora colonies. RCR = 326.6. 19. Befotaka Bay Time: 1158 hours, dive duration 65 minutes; depth range 113 m; visibility 0.5 to 2 m across depth; temperature 29º C. The Condition of Coral Reefs in Northwest Madagascar Site description: Fringing reef with good reef development; gradual slope in shallows with shoals. Sheer wall begins at depth of approximately 4 m and ends at approximately 13 m depth, then ﬂattens or bottoms out with sand-dominant substrata, nooks, and crannies with some caves occuring on wall. Across depth, Favids, Poritids, and Acroporids dominant hard corals, with Rhytisima and Paralemnalia dominant soft corals. Seagrass observed in lagoonal area of reef. No damage or threat observed. RCR = 258.4. 20. Sakatia (northeast side of channel) Time: 0826 hours, dive duration 75 minutes; depth range 1-6.9 m; visibility 4 m at 0-4 m depth and 2 m visibility below 5 m depth; temperature 29º C. Site description: Fringing reef with boulders inshore. Across depth, Poritids, Acroporids, Favids, Milleporids, and Tubastrea sp. dominant hard corals, with Rhytisima and Sinularia dominant soft corals. Seagrass observed. Coral predator Drupella cornus seen in shallows on colonies of Acropora. Debris in form of plastic litter noted on reef. No other damage or threat observed. RCR = 253.7. 21. Sakatia (southeast side of channel) Time: 1100 hours, dive duration 60 minutes; depth range 17 m; visibility 4 m at 0-4 m depth and 2 m visibility below 5 m depth; temperature 29º C. Site description: Fringing reef with variable slope depending on location; large coral balmies near shore with scattered coral outcrops starting in approximately 4 m with sand, patches of Halimeda and sand dominant from 7 m+. Across depth, Favids, Agaricids, Fungids, and Milliporids dominant hard corals, with Rhytisima and Nephthea dominant soft corals. Seagrass observed. Coral predator Drupella cornus seen in shallows on colonies of Seriatopora. No other damage or threat observed. RCR = 257.5. 22. Ambavatoby Bay (west) Time: 0840 hours, dive duration 104 minutes, depth range 1-10 m; visibility 5 m across depth, temperature 29º C. Site description: Patch reef in sheltered bay with variable slope depending on location. Porites and Diploastrea colonies or balmies in shallow 2-4 m; at 10 m depth mostly dominated by sand with scattered patches of coral with soft coral. Across depth, Poritids, Fungids, and Favids dominant hard corals, with Xenia and Gorgonids dominant soft corals. Coral predator Drupella cornus observed. Approximately 12+ sea urchins observed with consequential bioerosion evident. Slight siltation observed that is likely to be natural. No other damage or threat observed. RCR = 229.4. Time: 1215 hours, dive duration 115 minutes, depth range 1-10 m, visibility 4 m across depth, temperature 29º C. Site description: Fringing reef off of channel entrance to Baie de Russes. Across depth, Favids and Acropora dominant hard corals with Rhytisima and Nephthea dominant soft corals. No evidence of threats noted. RCR = 238. 24. Ambavatoby Bay (east) Time: 1630 hours, dive duration 75 minutes, depth range 110 m, visibility 2 m across depth, temperature 29º C. Site description: Fringing reef with only light exposure to wave energy. Across depth, Porites and Favids dominant hard corals, with Gorgonians and Lemnalia. Damage from shark net observed with some slight natural siltation noted. No evidence of other threats or damage noted. RCR = 194.2. 25. Nosy Iranja (south) Time: 930 hours, dive duration 60 minutes, depth range 1-6 m, visibility 2 m across depth, temperature 29º C. Site description: Sand with boulders containing sea fans, hard corals and algae, and some balmies of Porites. Medium to high exposure to wave energy. Across depth, Favids dominant hard coral with Rhytismia dominant soft coral. Abundant sea urchins observed embedded in rock boulders. Seagrass, Thalassia present. No evidence of other threats or damage noted. 26. Nosy Kivindry Time: 1030 hours, dive duration 60 minutes, depth range 120 m, visibility 6 m across depth, temperature 29º C. Site description: Fringing reef surrounding a rock island containing some vegetation, depending on area. Slope gradual 10º to sheer 90º. Intermittent, well-developed reef areas interspersed with sand. Across depth, Favids, Poritids, and Acroporids dominant hard corals, with Asterospicularia, Xenia, and Cespitula dominant soft corals. Feeding scars from Acanthaster plancii noted. However, only two individuals seen. Light incidence of coral bleaching; no other damage/ threats noted. Two hawksbill turtles and 2 rays observed. 27. Nosy Kisimany Time: 1530 hours, dive duration 90 minutes, depth range 110 m, visibility 1-2 m across depth, temperature 29º C. Site description: Fringing reef with good development. Across depth, Acropora, Favia, and Pectinids dominant hard corals, with Rumphellia and Nepthea dominant soft corals. Numerous sea urchins, Diadema observed on reef. Mangroves present. No observed threats or damage noted on reef. RCR = 279. 23. Makamby Point Coral Reefs of Northwest Madagascar 59 Chapter 5 28. Five Metre Bank Time: 930 hours, dive duration 90 minutes, depth range 140 m, visibility 20 m across depth, temperature 29º C. Site description: Platform reef with oceanic exposure and approximately 30º slope, mostly sand with a few scarce patches of hard and soft coral. Halimeda and Diadema noted on top of reef. Across depth, Favidae and Agaricidae dominant hard corals, with Nepthea and Cespitularia dominant soft corals. No observed threats or damage on reef. RCR = 194.2. 29. Tanikely (west) Time: 700 hours, dive duration 100 minutes, depth range 120 m, visibility 20 m across depth, temperature 29º C. Site description: Fringing reef with variable slope 5-25º, light to moderate exposure to wave energy. Across depth, acroporids, fungids, and favids dominant hard corals, with Nepthea and Sacrophyton dominant soft corals. Beds of the algae Halimeda pelarus present. Only one Acanthaster plancii seen. Very light bleaching observed and some anchor damage noted. No other damage or threat visible. Tourist area. Fish plentiful. RCR = 320. 30. Banc Souzy Time: 1100 hours, dive duration 90 minutes, depth range 112 m, visibility 5 m across depth, temperature 29º C. Site description: Patch reef offshore with moderate exposure to wave energy. Across depth, Echinopora, Leptoseris, and Goniopora dominant hard corals, with gorgonids dominant soft coral. Light net damage. Reef used for artisinal ﬁshers. No other observed damage or threat. cornus was observed on reef sites S. Tsarabajina (site 1), NE Sakatia (site 20), SE Sakatia (site 21), and W. Ambavatoby Bay (site 22). Pathogens Coral diseases or pathogens were noted only on Nosy Fisaka (site 6) and were of very low frequency (i.e., 2-3 corals) on the reef. Coral bleaching Bleaching was light and appeared to be in the early stages (evidenced by slight discoloration of coral tissue) on six (or 20%) of the reefs surveyed. These included the sites of Tsarabajina (site 1), Marie Bank (site 3), Anakarea (site 5), Fisake (site 6), Nosy Fanihy (site 18), and Nosy Tanikely (site 29). Damage from ﬁshing, boating, and storms Damage resulting from anchors and ﬁshing nets was infrequently observed. Three (or 10%) of the reef sites surveyed had ﬁshing net entangled on the reef. These sites included Marine Bank (site 3), Ambavatoby Bay E (site 24), and Banc Souzy (site 30). Anchor damage was observed only on one site, Nosy Tanikely (site 29). No evidence of physical damage from severe storms or cyclones was observed on the reef sites surveyed. Siltation Five reef sites were noted to have slight sedimentation that may need further monitoring to determine signiﬁcance. These included Maromby Village (site 4), Lakandava (site 15), Ambavatoby Bay W (site 22), Ambavatoby Bay E (site 24). No excessive sedimentation stress was observed, i.e., dead reef substrate covered in silt. Fishing pressure RESULTS Benthic cover (substrata/biota) Hard coral coverage ranged approximately from 2.5 to 70.6% (Table 5.1). Soft coral dominated the substrata at some sites (e.g., Ambariokisimany, site 9) and ranged from 0 to 63.7%. Three reef sites had evidence of light ﬁshing pressure. These included Nosy Fisaka (site 4), Northeast Nosy Hao (site 12), and Lakandava, North Beach (site 15). At site 15 there was evidence of sea cucumber harvesting by Chinese ﬁshers. Further long-term monitoring would be needed to determine quantitatively the level of activity and impact. For example, stock assessments of sea cucumber species and catch-per-unit effort by the ﬁshers would help determine the extent and effect of the sea cucumber ﬁshery. Predators Disturbance from the coral predator Acanthaster plancii was localized and observed on six (or 20%) of the reefs surveyed. The amount of damage from this coral predator ranged from excessive to light. Excessive damage was noted on the fringing reef at Nosy Fisake (site 6). Moderate damage from A. plancii was noted on reef sites at Tsarabanjina (site 1) and Marie Bank (site 3). Light damage from A. plancii was noted on the three sites: Tsarabanjina (site 2), Nosy Fanihy (site 18), and Nosy Kivindry (site 26). One individual was seen on Tanikely (site 29), but no damage was observed. Light to negligible damage from the other coral predator Drupella 60 Rapid Assessment Program Overall reef condition The reef condition rating (Table 5.2) indicated that several sites had comparatively higher amounts of biodiversity and better reef health. Sites comparatively rated with the highest RCR for the sites surveyed included Anakarea (site 5), Northeast Nosy Hao (site 12), Nosy Fanihy (site 18), and Nosy Tanikely (site 29). Sites 25, 30, 26, 20, 21, and 6 were missing data. Although missing benthic data, Nosy Kivindry (site 26) had a well-developed reef structure and was a notable site. The Condition of Coral Reefs in Northwest Madagascar Charismatic marine fauna sightings Sharks, turtles, and rays were observed at a few sites; however, no duogongs were seen. A total of three sharks were seen during the survey at sites 9, 10, and 12. At sites 1 and 26, one hawksbill turtle (Erectmochelys imbricata) was noted while two hawksbill turtles were seen at site 9. A second unidentiﬁed turtle was also seen at site 26. One manta ray (Manta birostris) was seen at site 1, and two were observed at site 26. DISCUSSION AND RECOMMENDATIONS For the reef sites surveyed, the reef condition rating ranged from 146.5 to 348.5 (Table 5.2). Reef survey sites with the highest RCR included Anakarea (site 5), Northeast Nosy Hao (site 12), Nosy Fanihy (site 18), and Nosy Tanikely (site 29). These sites are strong candidates for ofﬁcial protection. A network of marine protected areas set up next to terrestrially protected areas would be ideal given the deadly effects of poor land-use practices on coral reefs. It is important to note that this comparative rating is only for the reefs that were surveyed for this assessment. The reef condition rating (RCR) is a synthesis of the biodiversity data (number of species per site for corals and ﬁsh), percent hard coral cover, and relative amount of threat or stress observable at each site during the survey. Therefore differences are expected and do occur when the results of the combined rank are compared to that of only one parameter (e.g., number of coral species). Incorporating the commercially important ﬁsh data of approximate count and biomass with the RCR values (Table 5.2), it is clear that Nosy Tanikely (site 29) off of Nosy Bé is by far the best site. Although Nosy Tanikely is not ofﬁcially a marine protected area (MPA), local management by the community (e.g., tour operators, ﬁshers) effectively regulates activity, affording a level of protection. This illustrates that MPAs are already an effective tool for conserving marine biodiversity and resources in Madagascar. Further tourism can play a positive role in protecting marine resources. Interestingly, sites off of Nosy Tanikely have been monitored since 1998 as part of the CORDIO and GCRMN programs (Bigot et al. 2000). The Nosy Tanikely survey site assessed during this RAP was at a different locality then the ones monitored as part of CORDIO and GCRM (Mahavaro, pers. comm.). Nevertheless, percent hard coral cover at the CORDIO/GCRM site as well as the RAP survey site were similar, at approximately 60-65.7% (Bigot et al. 2000). For the reefs surveyed, the most frequently observed disturbance was from Acanthaster plancii. Quod and Bigot noted COTS on two reef sites in the Toliara region during post bleaching surveys. The incidence of coral bleaching, while very low and apparently only in the early stages, should continue to be monitored, as is done through CORDIO/ GCRM. Sedimentation has been a major cause of degradation to many of the reefs of Madagascar. However, only light and potentially natural sedimentation was observed for the reef sites surveyed. This may partly be due to the fact that northwest Madagascar’s coastline is less developed and less populated than other Madagascar coastlines. For instance, reefs lying off the more populated and developed southwest coastline near cities of Toliara (Tulear) and Mahajanga suffer from high levels of sedimentation and degradation. In other regions of Madagascar, reefs are reported to be under major threat from coral extraction for building material and tourist curios. This activity is reported to be a major problem near Toliara, Dzamanadjar, and Foulpointe (Bigot et al. 2000). However, it was not observed in the areas covered during this survey. The extraction of ﬁsh and other coral reef invertebrates in large quantities has taken its toll on the reefs of Madagascar as well. Forty-three percent of the ﬁshery (or 65,090 tons) is based on coral reefs (FAO 1999). Fishing is an important source of food and to a lesser extent a source of employment for the people of Madagascar (Westmacott et al. 2000). As ﬁshing pressure is measured by catch-per-unit effort over time, and marine RAP surveys are only designed to be a present snapshot of an area’s conservation status, it is hard to deﬁnitively diagnose ﬁshing pressure on the reefs. However, proof of the effectiveness of no take zones such as Tanikely, where ﬁsh biomass and count is an order of magnitude greater than all other sites, strongly suggests that ﬁshing pressure is a problem in Madagascar and that MPAs are an effective tool to address it. In conclusion, setting up an ofﬁcial network of MPAs in northwest Madagascar, as is currently planned by Madagascar ofﬁcials in various sectors, would contribute substantially to conserving the country’s marine resources. Based on results of this survey, the areas highly recommended for inclusion in this network of MPAs, given their biodiversity and level of threat, include but are not limited to Tanikely, Anakarea, Northeast Nosy Hao, and Nosy Fanihy. Socioeconomic and biophysical factors, such as current patterns and watershed (i.e., land use practices) in the region, need to be considered for the overall design and implementation as well. The establishment of marine protected areas is best accomplished by engaging all stakeholders in the process of setting up and running them. REFERENCES Bigot, L., L. Charpy, J. Maharavo, F. Abdou Rabi, N. Ppaupiah, R. Aumeeruddy, C. Viledieu, and A. Lietaud. 2000. Status of coral reefs of the Southern Indian Ocean: The Indian Ocean Commission Node for Comoros, Madagascar, Mauritius, Reunion and Seychelles. In: C. R. Wilkinson (ed.), Status of coral reefs of the world: 2000. GCRMN Report, Australian Institute of Marine Science, Townsville. Chapter 5, pp. 77-94. Bryant, D., L. Burke, J. McManus, and M. D. Spalding. 1998. Reefs at Risk. World Resource Institute. 56 pp. Coral Reefs of Northwest Madagascar 61 Chapter 5 English S., C. Wilkinson, and V. Baker (eds.). 1997. Survey manual for Tropical Marine Resources (2nd Edition). Australian Institute of Marine Science. ASEAN-Australia Marine Science Project. 390 pp. McKenna, S. A., G. R. Allen and S. Suryadi. 2002. A marine rapid assessment of the Raja Ampat Islands, Papua Province, Indonesia. RAP Bulletin of Biological Assessment 22. Washington, DC: Conservation International. Quod, J. P. and L. Bigot. 2000. Coral bleaching in the Indian Ocean islands: Ecological consequences and recovery in Madagascar, Comoros, Mayotte and Reunion. In: D. Souter, D. Obura, and O. Linden (eds.). Coral reef degradation in the Indian Ocean: Status reports and project presentations. Spalding, M. D., C. Ravilious, and E. P. Green. 2001. World Atlas of Coral Reefs. Prepared at the UNEP World Conservation Monitoring Centre, University of California Press, Berkeley, U.S.A. Westmacott S., H.S. J. Cesar, L. Pet-Soede, and O. Linden. 2000. Coral Bleaching in the Indian Ocean: Socio-Economic Assessment of Effects. In: H. S. J. Cesar (ed.). Essays on the Economics of Coral Reefs. CORDIO, Kalmar University, Sweden. pp. 94-106. 62 Rapid Assessment Program 4 4 4 3 6 6 7 7 4 4 5 10 4 5 4 2 4 9 4 4 4 4 3 9 4 3 4 4 2 8 2 1 4 4 1 8 n site deep shallow deep shallow deep shallow deep shallow deep shallow deep shallow deep shallow deep shallow shallow deep depth 1.2 (1.25) 3.1 (0.63) 1.3 (0.72) 4.38 (2.95) 10.6 (8.20) 6.9 (2.13) 5.8 (3.63) 10.6 (4.25) 17.5 (1.77) 28.7 (3.31) 63.7 (5.82) 4.4 (2.14) 1.2 (0.72) 0.6 (0.63) 3.1 (1.20) 1.9 (0.63) 10 (3.68) 0 1.2 (0.72) 0 0.6 (1.25) 0 0 13.7 (6.25) 8.7 (2.16) 21.9 (2.13) 47.5 (2.70) 7.5 (3.06) 35 (15) 14.4 (3.87) 12.5 (0.77) 11.2 (2.16) 18.7 (6.33) 57.5 (2.5) 43.1 (8.68) 55.6 (4.00) 33.1 (3.13) 21.8 (3.59) 42.5 (2.82) 45.8 (7.41) 11.3 (5.25) 50 (10.21) 29.4 (4.61) 23.1 (4.72) 2.5 (1.02) SP 3.7 (0.7) 3.8 (1.3) 1.2 (1.25) 0.6 (0.62) 1.2 (0.72) 0.6 (0.62) 0 2.5 (1.0) SC 10.6 (1.2) HC 6.9 (3.12) 0 23.7 (5.82) 14.4 (2.77) 20 (4.89) 41.9 (9.76) 8.1 (4.49) 19.2 (7.95) 3.12 (2.37) 3.1 (2.37) 1.9 (1.87) 0 2.5 (1.44) 0 MA 1.2 (0.72) 6.2 (3.7) 3.1 (1.19) 21.2 (4.39) TA 13.1 (3.3) 26.2 (11.2) 12.5 (1.02) 4.4 (1.57) 16.3 (4.62) 12.5 (5.20) 28.7 (11.25) 6.2 (2.98) 20.6 (2.58) 6.2 (1.25) 10.6 (5.44) 6.2 (2.16) 17.5 (15.07) 16.2 (5.25) 13.1 (2.37) 10.6 (2.13) 6.2 (0.72) 10.0 (1.02) 2.5 (0.96) 0.8 (0.83) 0.2 (0.25) 2.5 (1.77) 1.0 (0.71) 1.2 (1.25) 4.0 (0.41) 0 5 (0.71) 0 0.2 (0.25) 0 3.1 (1.88) 3.25 (1.89) 1.9 (1.87) 2.4 (1.22) 0 CA 0.5 (0.3) 16.2 (5.4) 0 RB 21.8 (5.9) 5 (0) 18.4 (3.75) 1.9 (1.19) 4.4 (1.57) 11.2 (4.15) 7.5 (2.50) 7.5 (3.06) 2.5 (2.50) 11.2 (2.16) 6.9 (2.77) 7.5 (1.02) 5.8 (3.63) 33.1 (4.13) 1.9 (1.87) 9.4 (3.29) 40 (4.21) 0 SD 45 (8.3) 5 (0) 34.4 (1.57) 21.9 (3.29) 11.3 (2.16) 38.7 (16.38) 6.3 (6.25) 13.7 (8.00) 4.4 (2.14) 10.6 (2.13) 7.5 (6.69) 19.4 (3.87) 3.3 (1.67) 22.5 (1.77) 8.1 (4.13) 8.7 (5.54) 0 0 0 0 5.6 (1.57) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Mud 0 1.2 (0.72) 0 1.9 (1.19) 0 1.2 (0.72 0 0.6 (0.63) 0 1.9 (1.87) 0 Other 0.6 (0.6) 5 (0) 3.7 (2.98) 6.9 (2.77) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 BS The n represents the number of 20 m transects surveyed along each 100 m transect at shallow and/or deep depths (shallow = 4-6 m, deep = 12-16m). Biota/ substrata were categorized as: Hard Coral (HC), Soft Coral (SC), Sponge (SP), Macro Algae (MA), Turf Algae (TA), Calcareous Algae (CA), Rubble (RB), Sand (SD), and Bare Substrate (BS). The “Other” category represents invertebrates such as echinoderms, tunicates, etc. Percent mean is given with standard error below in parentheses. In cases where transects were not done, the transect/site has been omitted from the table. Table 5.1. Summary of benthic data for reef sites surveyed in northwest Madagascar. The Condition of Coral Reefs in Northwest Madagascar Coral Reefs of Northwest Madagascar 63 64 n 4 2 4 4 4 4 4 2 4 4 4 4 3 4 4 4 4 4 4 2 2 site 11 11 12 Rapid Assessment Program 12 13 13 14 15 16 17 18 18 19 19 20 20 21 21 22 23 24 deep shallow deep shallow deep shallow deep shallow deep shallow deep deep deep deep deep shallow deep shallow deep shallow deep depth 22.5 (5.10) 28.1 (3.87) 26.2 (2.60) 70.6 (6.16) 42.5 (6.61) 44.4 (4.49) 40 (3.95) 53.7 (4.39) 26.2 (5.25) 27.5 (3.95) 24.4 (3.29) 15.0 (2.50) 36.2 (3.75) 25 HC 3.1 (1.20) 38.7 (6.25) 37.5 (5.40) 26.9 (3.12) 8.7 (0.72) 29.4 (11.01) 31.9 (7.39) SC 20.6 (2.58) 25 (5.0) 21.9 (3.12) 43.1 (2.77) 36.9 (3.87) 33.75 (3.61) 11.2 (4.39) 26.2 (1.25) 32.5 (6.69) 42.5 (4.89) 3.7 (1.61) 0.6 (0.62) 10.0 (1.44) 11.8 (3.59) 3.1 (2.37) 0.6 (0.62) 9.4 (2.13) 7.5 (3.23) 14.4 (1.57) 17.5 (7.50) 3.7 (1.25) 1.9 (1.20) 0.6 (0.62) 4.4 (0.62) 5.0 (5.0) 20 (2.50) 0 7.5 (3.82) 5.0 (2.70) 3.1 (1.19) 0 1.9 (1.19) 1.2 (1.25) 0 3.1 (1.20) 6.2 (6.25) 0 5 (1.44) 0 0 0 SP 10.6 (2.13) 0.6 (0.63) 12.5 (5.0) 1.2 (1.25) 0 0.6 (0.62) 18.1 (0.63) 2.5 (2.50) 1.3 (1.25) 5.0 (1.77) 20 (5.86) 7.5 (3.68) 10.8 (3.63) 13.1 (1.87) 3.7 (2.39) 10 (5.40) 1.9 (0.63) 0 4.4 (1.20) 3.1 (1.57) 0 MA 6.2 (3.75) TA 7.5 (2.70) 1.2 (1.25) 6.88 (2.77) 18.1 (3.12) 6.2 (2.60) 0.6 (0.62) 3.1 (1.87) 5.0 (2.50) 16.3 (1.61) 5.0 (1.02) 21.3 (2.60) 20.6 (5.62) 22.5 (10.41) 18.7 (3.75) 26.9 (5.72) 32.5 (4.45) 20.6 (2.58) 17.5 (3.68) 15.6 (3.59) 11.2 (8.75) 16.2 (1.25) 0 2.5 (2.50) 0 0.3 (0.33) 3.7 (0.72) 0.2 (0.25) 3.1 (1.87) 1.0 (0.41) 1.2 (1.25) 0 4.5 (0.29) 0.5 (0.50) 0 0 0 0 0.2 (0.25) 2.5 (1.02) 0.7 (0.25) 0 0 CA 0 9.4 (2.77) 8.1 (1.87) 2.5 (1.44) 1.2 (1.25) 35 (25.0) 20 (2.50) 0 25 (7.36) 26.2 (3.61) 0 0.6 (0.62) 13.8 (9.82) 0.6 (0.62) 2.5 (1.02) 0 0 0 0 21.8 (6.16) 3.8 (2.98) 0 0 10 (5.40) 0 SD 29.4 (3.73) 13.7 (3.75) 14.4 (5.53) 2.5 (1.77) 15.6 (3.29) 20 (7.57) 18.7 (2.98) 0 RB 21.2 (8.81) 17.5 (2.50) 14.4 (2.37) 3.8 (2.39) 20 (2.70) 12.5 (3.54) 13.1 (7.17) 3.7 (1.25) 13.1 (2.14) 2.5 (2.50) 1.9 (1.20) 10.6 (3.12) 0.6 (0.62) 0 0 0.6 (0.62) 5.8 (3.00) 1.8 (0.62) 6.3 (2.39) 0 4.4 (1.20) 5.6 (1.20) 2.5 0 0 3.1 (2.37) 0 0 0 Other 1.2 (0.72) 0 0 37.5 (6.45) 0 1.3 (1.25) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.6 (0.63) 0 0 0 27.5 (2.50) 0 0 0 0 0 0 1.3 (1.25) 0 BS 0 0 2.0 (2.5) 0 0 0 0 Mud Chapter 5 n 4 4 4 4 4 1 4 site 24 27 27 28 29 29 30 deep shallow deep deep shallow deep shallow depth HC 23.1 (7.24) 21.2 (6.65) 35 (7.86) 8.1 (2.58) 35.6 (4.13) 60 10.6 (1.87) SC 0.6 (0.62) 4.4 (1.57) 2.5 (1.02) 5.6 (2.37) 2.5 (1.02) 0 3.7 (1.61) SP 2.5 (1.02) 1.3 (0.72) 1.9 (1.20) 3.1 (1.87) 3.1 (1.20) 0 5.6 (0.63) 0 MA 0.6 (0.62) 0.6 (0.62) 1.25 (0.7) 17.5 (4.21) 3.1 (1.57) 0 TA 13.7 (5.25) 8.8 (3.75) 18.1 (5.14) 13.8 (3.61) 15.6 (3.87) 12.5 12.5 (1.77) 0 0 0 CA 0.6 (0.62) 0.8 (0.45) 1.2 (1.25) 3.3 (1.49) RB 5.0 (2.28) 10.6 (2.77) 16.9 (10.07) 17.5 (3.68) 11.9 (2.1) 7.5 13.8 (3.89) SD 51.9 (12.26) 45.6 (6.64) 19.4 (9.86) 23.7 (4.62) 23.1 (3.87) 0 33.1 (6.07) Other 1.9 (1.87) 5.6 (3.29) 3.7 (2.39) 2.5 (1.77) 5.0 (1.02) 20 20.6 (5.81) 0 0 0 0 0 0 0 Mud 0 0 0 0 0 0 0 BS The Condition of Coral Reefs in Northwest Madagascar Coral Reefs of Northwest Madagascar 65 66 Rapid Assessment Program 111 111 104 109 143 129 109 90 88 46 103 125 110 122 92 107 119 120 123 110 125 88 104 71 12 13 14 15 16 17 18 19 20 21 22 23 24 Coral 1 2 3 4 5 6 7 8 9 10 11 Site 89 119 119 117 140 91 92 107 101 114 156 82 166 132 111 145 118 152 133 114 104 106 98 141 Fish 58 77 68 57 81 60 89 93 61 66 85 69 71 91 85 91 77 80 76 51 55 51 69 78 Mollusc Number of species 218 300 275 293 341 274 291 325 269 272 351 273 362 334 307 340 304 375 338 274 249 245 213 322 Total 102 24 67 43 105 na na na 133 28 190 6 99 324 385 418 51 224 na 74 688 179 83 165 Approx. ﬁsh count 28.1 11.5 44.5 23.3 12.3 na na na 180.2 7.2 122.2 3.1 41.7 354.8 189.9 120.9 119.3 350.3 na 166.2 259.0 92.3 205.26 84.5 Biomass (ton/km2) 29.65 15.0 24.4 28.1 48.4 43.45 46.85 26.85 22.5 25 19.05 31.9 32.2 22.8 14.4 11.85 38.1 49.35 27.45 44.15 30.65 26.25 2.5 20.9 Percent hard coral 194.2 238 229.4 264.1 326.6 258.4 253.7 257.5 230.5 229 295.4 235.9 328.5 270 232.4 253.5 284.5 348.5 283.1 268.8 244 223.4 146.5 282.7 Reef Condition Rating (RCR) D(P) D(P) D(P) A(L) L A(E) L L L L Bleaching L Coral disease A(M), D(P) A(L) A(M) Predators shark net (L) ﬁshing line in shallows Physical damage from ﬁshing or boating activity siltation present, appears natural siltation present, appears natural debris siltation (L), evidence of sea cucumber harvest by Chinese ﬁshing (L), siltation (L) ﬁshing (L) Observations indicating possible threat from siltation, ﬁshing activity, other source This table lists syntheses of species diversity, approximate counts and biomass quantities of commercially and artisanally important ﬁsh, percents of hard coral cover, reef condition ratings, and damage and threat observations for each reef site surveyed. Where values are not available due to missing data, “na” is used. For predators, A = Acanthaster planci, D = Drupella cornus. Threats or levels of damage are rated as L = light, M = moderate, and E = excessive (listed next to predators and damage or threat observations). In cases where no damage was observed but a predator was seen, the letter P for “present” follows the predator notation. If no threat or damage was observed the corresponding cell for that site is empty. The last column contains observations on siltation and ﬁshing pressure and other comments on the reef site. Table 5.2. Summary of reef condition. Chapter 5 30 103 121 75 110 na 26 27 28 29 30 Coral 25 Site 152 123 122 154 73 33 Fish 68 89 73 81 49 70 Mollusc Number of species 323 333 270 345 122 133 Total na 111 99 894 56 na Approx. ﬁsh count na 35.3 79.9 1720.2 123.2 na Biomass (ton/km2) na 28.1 8.1 47.8 10.6 na Percent hard coral na 279 194.2 320 na na Reef Condition Rating (RCR) A(P) A(L) Predators Coral disease L Bleaching anchor net (L) Physical damage from ﬁshing or boating activity artisanal ﬁshers on reef Observations indicating possible threat from siltation, ﬁshing activity, other source The Condition of Coral Reefs in Northwest Madagascar Coral Reefs of Northwest Madagascar 67 Chapter 6 Survey Mission Report Salimo and Bakary Gisèle INTRODUCTION At the request of Conservation International and within the framework of the 2002 RAP (Rapid Assessment Program) project, we have conducted a socioeconomic survey designed to gather a number of data speciﬁc to the following village communities: Nosy Faly, Ampampamena, Ankazomborona, a few villages in the Ambaro Bay region in northwestern Madagascar and a little further to the south, and a number of Grande Terre villages and villages on neighboring islets such as the Radama Islands. This survey was carried out 11-26 January 2002 for the purpose of assessing the health of the coral reefs of these regions by gathering data on the quantities and types of species as well as the impact of local human communities. METHODOLOGY AND MATERIALS We adopted a participative approach in our interactions with the PCLS and village notables. We traveled from village to village aboard an Argos 700 boat to collect our data. RESULTS Main activities Overall, the main economic activity of the communities surveyed is traditional ﬁshing, but men also dedicate time to secondary activities to meet their consumption needs, including small-scale crop and livestock agriculture (rice, zebu, poultry, etc.). While women sometimes go ﬁshing with their husbands, most of their time is spent drying and curing unsold output, including chavaquines and bivalves. These products are then taken to Diego, Ambilobe, Ambanha, and Nosy Ne by collectors and wholesale ﬁsh merchants. Only 3% of the output produced by local communities is consumed locally. 68 Rapid Assessment Program Survey Mission Report Table 6.1: Percent of village populations that are ﬁshermen. Commercialization The prices of the products vary according to quantity and quality. The following constitute the average prices of the different products at the time of the survey: Small pelagics Rock ﬁsh and big pelagics Shrimp Camaron Lobster Octopus Trépang (depending on size and quality) Mangrove green crabs Moray eel Aileron or shark ﬁn 1st choice 2nd choice 3rd choice Chevaquine 1,000 - 1,500 fmg /kg 3,500 fmg/kg 5,000 10,000 fmg/kg 15,000 fmg/kg 15,000-25,000fmg/kg 1,000-2,000 fmg / kg 1,000-45,000fmg/kg 2,000-2500 fmg/kg 500-1,000 fmg/kg 450,000 fmg /kg 350,000 fmg/kg 50,000 fmg/kg 15,000-35,000 fmg/kg Fishing output sales Fishermen with their own production means such as pirogues and nets retain all the proﬁts of their sales. Those who do not own their production means, however, rely on a system, common in the villages, that is based on the principle of sharecropping. Ethnicity and migration In the villages there are fewer native ﬁshermen than migrant ones. The latter come from the northwest, southwest, and southeast regions of Madagascar. Education Many ﬁshermen have a very low level of education. School is poorly attended because of the distances between villages and the lure of the earnings to be made ﬁshing. Beliefs and customs Each village has its own “fady” or taboos. New generations and migrants have perpetuated some taboos and abandoned others. Examples of current taboos include: - working on Tuesdays (Nosy Faly) killing dolphins (in almost all of the regions) using ﬁshing equipment such as ﬁsh traps (Ampampamena) Health Most landlocked villages have practically no basic medical care infrastructure. Most of these villagers have to walk several kilometers to reach a clinic (CSB I). Village Ampampamena Ankazomborona Andavanemboka Mitsio Marotogny Iranja Total Population 510 4,000 80 700 320 60 Number of Fishermen 90 1,400 21 205 25 12 Percentage 17.64 35 26.25 29.28 7.81 20 Dialogue According to the villagers, the coral reefs of the Amparo Bay are still intact. The villagers of Grande Terre (Marotogny, Agnorontsanga) and of its neighboring islets (Iranja, Radama islands), however, have noted advanced damages most likely caused by destructive equipment, possibly bâtons sticks, and the phenomenon of coral bleaching. Almost all of the villagers are aware that the size and quantity of products ﬁshed throughout the region have decreased. This growing scarcity of ﬁsh prompts ﬁshermen to travel farther and farther to improve their catch. The chemical pollution emitted by SIRAMA, which is dumped into the inlets, has a negative impact on marine animals (Andavanemboka). The landlocked villages have no shopping centers; as a result, the price of materials remains high even as the sales price of ﬁshing outputs is decreasing. RECOMMENDATIONS Based on our preliminary ﬁndings, we recommend that the following actions be taken: - A public awareness campaign for the preservation of the coral reefs should be carried out. - Oceanographic and biological data should be periodically gathered to evaluate the state of health of the coral reefs and the consequences of ﬁshing activities and bad environmental protection. - A drinking water conveyance should be implemented to offset the problem of water shortages during the dry season. - Basic schooling and basic health care centers should be established. - The opening and closing dates of the shrimp ﬁshing season in the Ambaro Bay should be re-examined. - With the aim of conserving stocks, traditional ﬁshing practices should be improved through training and information campaigns. - The acquisition of adequate ﬁshing equipment should be facilitated. Coral Reefs of Northwest Madagascar 69 Chapter 6 Table 6.2. Fishing methods, market price, and species captured by geographic area. AREA: Ambaro Bay intertidal zone (sandy bottom) Equipment Amount - High season 700 kg (Mar-Apr) Gill net: 12–15 to 8–15 m deep, 100 m long (kaokobe); nonmotorized outrigger pirogue: 7–8 m long with 4 men aboard Valakira (coastal dam); nonmotorized outrigger pirogue: 7–8m long with 1 or 2 men aboard Amount - Low season 40-100 kg Species captured Shrimp Upenaeus indicus Upenaeus semisulcatus Metapenaeus monoceros Upenaeus monodon Fishes Rastrelliger kanaguria Selar crumenophthalmus Chirocentrus dorrab Leiognathus ecuulus Secutor incidiator Sphyraena obtusata Sphyraena foresteri Psettodes erumei Scomberoides tol Liza melinoptera Mugil esphalus 100-500 kg (Jan-Feb) 10-15 kg Crab Scylla serrata 120 kg (Mar-Apr) 60 kg Shrimp Upenaeus indicus Upenaeus semisulcatus Metapenaeus monoceros Upenaeus monodon 200-300 kg 50-80 kg Fishes Rastrelliger kanaguria Selar crumenophthalmus Chirocentrus dorrab Leiognathus ecuulus Secutor incidiator Sphyraena obtusata Sphyraena foresteri Psettodes erumei Scomberoides tol Liza melinoptera Mugil esphalus 50-100 kg 60 kg Crab Scylla serrata AREA : Grande Terre and its neighboring islets (reef zone) Equipment Amount - High season Amount - Low season Species captured Sea turtle Chelonia mydas Gill net: 50 (jarifa) from motorized boat 7–9 m long, with 4–5 men aboard 70 Rapid Assessment Program 90 kg (year round) 10 kg Cartilage ﬁsh Rhincobatus dijddensis Sphyrna lewini Pliotrema warreni Cephalopholus paquicentron Miliobatis punctatus Survey Mission Report AREA : Grande Terre and its neighboring islets (reef zone) Equipment Line ﬁshing: line strength 6–9 to 36 kg from non-motorized outrigger pirogue 7–8 m long with 2–3 men aboard Dragnet ﬁshing from sailboat or other motorized type of boat with 2–3 persons aboard (men or women) Fish trap: non-motorized outrigger pirogue, 3–6 m long with 1–2 men aboard Scuba diving: non-motorized outrigger pirogue, 7–9 m long with 4–5 men aboard Amount - High season 70-100 kg (Dec-Jan) 5-10 pieces, 80-100 kg each (year round) 5-20 kg (year round) Amount - Low season Species captured 30 kg Big pelagic ﬁsh Somberomurus commerson Spyraena barracuda Sphyraena jello Lethrinus lentjan Lenthrius elongates Llenthrinus harak Scomberoides tol Thurnus alabacares Monotaxis grandoculis Canrangoides fulvogutatus Lutjanus bohar 1-2 pieces Big pelagic ﬁsh Scomberomorus commersoni Xiphias gladius Istiophorus platypterus Scomberoides tol Thunnus albacares Thunnus obesus Gnathanodon speciosus 0-2 kg Big pelagic ﬁsh Siganus stellatus Siganus oramin Lethrinus mahsena Lethrinus elongatus Lutjanus argentimaculatus Sea cucumber Holothuria ananas Sticopus chloronatus Holoturia amamelle Holothuria scabra 2-4 pieces Beach seine on foot, 2–4 women 5-10 buckets (Jan) 2-3 buckets Small crustaceans Acetes erhutraeus Non-motorized outrigger pirogue, 3–5m long with 1–2 men aboard 10-15 kg 1-2 kg Small crustaceans Sepia latimanus Rock ﬁsh Lethrinus lentjean Caranx ignobilis Caranx sexfasciatus Lutjanus argentimaculatus Spyraena barracuda Epinephelus posteli Underwater hunting 2-20 kg (year round) Motorized or non-motorized boat, 7–8 m long, with 2–4 men aboard 10-20 kg 3-5 kg Lobster Panulirus versicolor Panulirus ornatus Self consumption 5-10 kg Octopus macropus Harpoon on foot 1 person with buoy Gathering on foot, alone or with several people 50 kg Self consumption Scylla serrata Anadara natalensis Coral Reefs of Northwest Madagascar 71 Appendix 1 Total inventory of species recorded during the present study and previously recorded for the whole of the SW Indian Ocean Zooxanthellate Scleractinia Family Astrocoeniidae Koby, 1890 Genus Stylocoeniella Yabe and Sugiyama, 1935 Stylocoeniella armata (Ehrenberg, 1834) Stylocoeniella guentheri Bassett-Smith, 1890 Genus Madracis Milne Edwards and Haime, 1849 Madracis kirbyi Veron and Pichon, 1976 Madracis sp. Family Pocilloporidae Gray, 1842 Genus Pocillopora Lamarck, 1816 Pocillopora ankeli Scheer and Pillai, 1974 Pocillopora damicornis (Linnaeus, 1758) Pocillopora eydouxi Milne Edwards and Haime, 1860 Pocillopora fungiformis Veron, 2000 Pocillopora indiania Veron, 2000 Pocillopora cf. kelleheri Veron, 2000 Pocillopora verrucosa (Ellis and Solander, 1786) Genus Seriatopora Lamarck, 1816 Seriatopora undescribed species Seriatopora aculeata Quelch, 1886 Seriatopora caliendrum Ehrenberg, 1834 Seriatopora guttatus Veron, 2000 Seriatopora hystrix Dana, 1846 Genus Stylophora Schweigger, 1819 Stylophora madagascarensis Veron, 2000 Stylophora pistillata Esper, 1797 Stylophora subseriata (Ehrenberg, 1834) Stylophora wellsi Scheer, 1964 Family Acroporidae Verrill, 1902 Genus Montipora Blainville, 1830 Montipora aequituberculata Bernard, 1897 Montipora australiensis Bernard, 1897 Montipora calcarea Bernard, 1897 Montipora caliculata (Dana, 1846) Montipora danae (Milne Edwards and Haime, 1851) Montipora digitata (Dana, 1846) Montipora efﬂorescens Bernard, 1897 72 Rapid Assessment Program Recorded in SW Indian Ocean (Veron 2000) X X Recorded in Madagascar √ √ X √ X X X X X X X X √ √ √ P √ √ √ √ √ √ √ X X X X √ √ √ P X X X √ P √ √ √ √ √ X X X Total inventory of species recorded during the present study and previously recorded for the whole of the SW Indian Ocean Zooxanthellate Scleractinia Montipora effusa Dana, 1846 Montipora ﬂoweri Wells, 1954 Montipora foliosa (Pallas, 1766) Montipora friabilis Bernard, 1897 Montipora grisea Bernard, 1897 Montipora hispida (Dana, 1846) Montipora informis Bernard, 1897 Montipora kellyi Veron, 2000 Montipora lobulata Bernard, 1897 Montipora meandrina (Ehrenberg, 1834) Montipora millepora Crossland, 1952 Montipora mollis Bernard, 1897 Montipora monasteriata (Forskäl, 1775) Montipora cf. nodosa (Dana, 1846) Montipora orientalis Nemenzo, 1967 Montipora peltiformis Bernard, 1897 Montipora spongodes Bernard, 1897 Montipora spumosa (Lamarck, 1816) Montipora stilosa (Ehrenberg, 1834) Montipora tuberculosa (Lamarck, 1816) Montipora turgescens Bernard, 1897 Montipora undata Bernard, 1897 Montipora venosa (Ehrenberg, 1834) Montipora verrucosa (Lamarck, 1816) Genus Anacropora Ridley, 1884 Anacropora forbesi Ridley, 1884 Genus Acropora Oken, 1815 Acropora undescribed species Acropora abrotanoides (Lamarck, 1816) Acropora aculeus (Dana, 1846) Acropora acuminata (Verrill, 1864) Acropora anthocercis (Brook, 1893) Acropora appressa (Ehrenberg, 1834) Acropora arabensis Hodgson and Carpenter, 1995 Acropora aspera (Dana, 1846) Acropora austera (Dana, 1846) Acropora bifurcata Nemenzo, 1971 Acropora branchi Riegl, 1995 Acropora brueggemanni (Brook, 1893) Acropora cerealis (Dana, 1846) Acropora clathrata (Brook, 1891) Acropora copiosa Nemenzo, 1967 Acropora cuneata (Dana, 1846) Acropora cytherea (Dana, 1846) Acropora digitifera (Dana, 1846) Acropora divaricata (Dana, 1846) Acropora donei Veron and Wallace, 1984 Acropora echinata (Dana, 1846) Acropora ﬂorida (Dana, 1846) Recorded in SW Indian Ocean (Veron 2000) X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X Recorded in Madagascar P √ √ √ √ P P √ P √ √ √ P √ P √ √ √ √ √ P √ √ √ √ √ √ √ √ √ √ P √ √ √ P √ √ √ √ P √ Coral Reefs of Northwest Madagascar 73 Appendix 1 Zooxanthellate Scleractinia Acropora formosa (Dana, 1846) Acropora forskali (Ehrenberg, 1834) Acropora gemmifera (Brook, 1892) Acropora glauca (Brook, 1893) Acropora grandis (Brook, 1892) Acropora granulosa (Milne Edwards and Haime, 1860) Acropora haimei (Milne Edwards and Haime, 1860) Acropora hemprichii (Ehrenberg, 1834) Acropora horrida (Dana, 1846) Acropora humilis (Dana, 1846) Acropora indonesia Wallace, 1997 Acropora inermis (Brook, 1891) Acropora insignis Nemenzo, 1967 Acropora irregularis (Brook, 1892) Acropora lamarcki Veron, 2000 Acropora latistella (Brook, 1891) Acropora longicyathus (Milne Edwards and Haime, 1860) Acropora loripes (Brook, 1892) Acropora massawensis Marenzeller, 1906 Acropora macrostoma (Brook, 1891) Acropora microphthalma (Verrill, 1859) Acropora millepora (Ehrenberg, 1834) Acropora mirabilis (Quelch, 1886) Acropora monticulosa (Brüggemann, 1879) Acropora nana (Studer, 1878) Acropora nasuta (Dana, 1846) Acropora natalensis Reigl, 1995 Acropora nobilis (Dana, 1846) Acropora ocellata (Klunzinger, 1879) Acropora palifera (Lamarck, 1816) Acropora paniculata Verrill, 1902 Acropora papillarae Latypov, 1992 Acropora parilis (Quelch, 1886) Acropora pharaonis (Milne Edwards and Haime, 1860) Acropora pinguis Wells, 1950 Acropora plantaginea (Lamarck, 1816) Acropora polystoma (Brook, 1891) Acropora prostrata (Dana, 1846) Acropora pulchra (Brook, 1891) Acropora rambleri (Bassett-Smith, 1890) Acropora retusa (Dana, 1846) Acropora robusta (Dana, 1846) Acropora rosaria (Dana, 1846) Acropora roseni Wallace, 1999 Acropora samoensis (Brook, 1891) Acropora secale (Studer, 1878) Acropora selago (Studer, 1878) Acropora squarrosa (Ehrenberg, 1834) Acropora stoddarti Pillai and Scheer, 1976 Recorded in SW Indian Ocean (Veron 2000) X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X Recorded in Madagascar √ √ √ P √ P √ √ √ √ P √ √ P √ √ √ √ √ √ P P P √ √ √ √ √ √ √ √ P √ P √ √ √ √ √ P P √ √ √ √ √ continued 74 Rapid Assessment Program Total inventory of species recorded during the present study and previously recorded for the whole of the SW Indian Ocean Zooxanthellate Scleractinia Acropora striata (Verrill, 1866) Acropora subulata (Dana, 1846) Acropora tenuis (Dana, 1846) Acropora cf. valenciennesi (Milne Edwards and Haime, 1860) Acropora valida (Dana, 1846) Acropora variabilis (Klunzinger, 1879) Acropora variolosa (Klunzinger, 1879) Acropora vermiculata Nemenzo, 1967 Acropora verweyi Veron and Wallace, 1984 Acropora willisae Veron and Wallace, 1984 Acropora yongei Veron and Wallace, 1984 Genus Astreopora Blainville, 1830 Astreopora expansa Brüggemann, 1877 Astreopora listeri Bernard, 1896 Astreopora myriophthalma (Lamarck, 1816) Astreopora ocellata Bernard, 1896 Astreopora suggesta Wells, 1954 Family Euphilliidae Veron, 2000 Genus Catalaphyllia Wells, 1971 Catalaphyllia jardinei (Saville-Kent, 1893) Genus Nemenzophyllia Hodgson and Ross, 1981 Euphyllia glabrescens (Chamisso and Eysenhardt, 1821) Genus Physogyra Quelch, 1884 Physogyra lichtensteini (Milne Edwards and Haime, 1851) Genus Plerogyra Milne Edwards and Haime, 1848 Plerogyra sinuosa (Dana, 1846) Family Oculinidae Gray, 1847 Genus Galaxea Oken, 1815 Galaxea astreata (Lamarck, 1816) Galaxea fascicularis (Linnaeus, 1767) Family Meandrinidae Gray, 1847 Genus Ctenella Matthai, 1928 Ctenella chagius Matthai, 1928 Family Siderasteridae Vaughan and Wells, 1943 Genus Horastrea Pichon, 1971 Horastrea indica Pichon, 1971 Genus Anomastraea Marenzeller, 1901 Anomastraea irregularis Marenzeller, 1901 Genus Gyrosmilia Gyrosmilia interrupta (Ehrenberg, 1834) Genus Coscinaraea Milne Edwards and Haime, 1848 Coscinaraea undescribed species Coscinaraea sp. Coscinaraea columna (Dana, 1846) Coscinaraea crassa Veron and Pichon, 1980 Coscinaraea monile (Foskål, 1775) Coscinaraea wellsi Veron and Pichon, 1980 Genus Psammocora Dana, 1846 Psammocora contigua (Esper, 1797) Recorded in SW Indian Ocean (Veron 2000) X X X X X X X X X X X Recorded in Madagascar P √ √ √ √ √ √ √ √ √ √ X X X X X P √ √ √ P X √ X P X √ X √ X X √ √ X X √ X P X √ X X X X √ √ √ P √ √ X √ Coral Reefs of Northwest Madagascar 75 Appendix 1 Zooxanthellate Scleractinia Psammocora digitata Milne Edwards and Haime, 1851 Psammocora explanulata Horst, 1922 Psammocora haimeana Milne Edwards and Haime, 1851 Psammocora nierstraszi Horst, 1921 Psammocora obtusangula (Lamarck, 1816) Psammocora profundacella Gardiner, 1898 Psammocora superﬁcialis Gardiner, 1898 Psammocora cf. vaughani Yabe and Sugiyama, 1936 Genus Pseudosiderastrea Yabe and Sugiyama, 1935 Pseudosiderastrea tayami Yabe and Sugiyama, 1935 Genus Siderastrea Blainville, 1830 Siderastrea savignyana Milne Edwards and Haime, 1850 Family Agariciidae Gray, 1847 Genus Coeloseris Vaughan, 1918 Coeloseris mayeri Vaughan, 1918 Genus Gardineroseris Scheer and Pillai, 1974 Gardineroseris cf. planulata Dana, 1846 Genus Leptoseris Milne Edwards and Haime, 1849 Leptoseris explanata Yabe and Sugiyama, 1941 Leptoseris foliosa Dineson, 1980 Leptoseris hawaiiensis Vaughan, 1907 Leptoseris incrustans (Quelch, 1886) Leptoseris mycetoseroides Wells, 1954 Leptoseris papyracea (Dana, 1846) Leptoseris scabra Vaughan, 1907 Leptoseris solida (Quelch, 1886) Leptoseris yabei (Pillai and Scheer, 1976) Genus Pachyseris Milne Edwards and Haime, 1849 Pachyseris rugosa (Lamarck, 1801) Pachyseris speciosa (Dana, 1846) Genus Pavona Lamarck, 1801 Pavona bipartita Nemenzo, 1980 Pavona cactus (Forskål, 1775) Pavona clavus (Dana, 1846) Pavona decussata (Dana, 1846) Pavona duerdeni Vaughan, 1907 Pavona explanulata (Lamarck, 1816) Pavona frondifera (Lamarck, 1816) Pavona maldivensis (Gardiner, 1905) Pavona varians Verrill, 1864 Pavona venosa (Ehrenberg, 1834) Family Fungiidae Dana, 1846 Genus Cycloseris Milne Edwards and Haime, 1849 Cycloseris costulata (Ortmann, 1889) Cycloseris curvata (Hoeksema, 1989) Cycloseris cyclolites Lamarck, 1801 Cycloseris erosa (Döderlein, 1901) Cycloseris patelliformis (Boschma, 1923) Recorded in SW Indian Ocean (Veron 2000) X X X X X X X Recorded in Madagascar P √ √ P P √ √ √ X P X P X P X √ X X X X X X X X X √ √ √ √ √ P √ P P X X √ √ X X X X X X X X X X √ P √ √ √ √ √ √ √ √ X X X X X √ √ √ √ P continued 76 Rapid Assessment Program Total inventory of species recorded during the present study and previously recorded for the whole of the SW Indian Ocean Zooxanthellate Scleractinia Cycloseris sinensis Milne Edwards and Haime, 1851 Cycloseris somervillei (Gardiner, 1909) Cycloseris tenuis (Dana, 1846) Cycloseris vaughani (Boschma, 1923) Genus Diaseris Milne Edwards and Haime, 1849 Diaseris distorta (Michelin, 1843) Diaseris fragilis Alcock, 1893 Fungia concinna Verrill, 1864 Fungia corona Döderlein, 1901 Fungia danai Milne Edwards and Haime, 1851 Fungia fungites (Linneaus, 1758) Fungia granulosa Klunzinger, 1879 Fungia horrida Dana, 1846 Fungia klunzingeri Döderlein, 1901 Fungia moluccensis Horst, 1919 Fungia paumotensis Stutchbury, 1833 Fungia puishani Veron and DeVantier, 2000 Fungia repanda Dana, 1846 Fungia scabra Döderlein, 1901 Fungia scruposa Klunzinger, 1879 Fungia scutaria Lamarck, 1801 Fungia seychellensis Hoeksema, 1993 Genus Halomitra Dana, 1846 Halomitra pileus (Linnaeus, 1758) Genus Herpolitha Eschscholtz, 1825 Herpolitha limax (Houttuyn, 1772) Herpolitha weberi Horst, 1921 Genus Podabacia Milne Edwards and Haime, 1849 Podabacia crustacea (Pallas, 1766) Podabacia motuporensis Veron, 1990 Genus Polyphyllia Quoy and Gaimard, 1833 Polyphyllia talpina (Lamarck, 1801) Genus Cantharellus Hoeksema and Best, 1984 Cantharellus doederleini (Marenzeller, 1907) Genus Fungia Lamarck, 1801 Family Pectinidae Vaughan and Wells, 1943 Genus Echinophyllia Klunzinger, 1879 Echinophyllia aspera (Ellis and Solander, 1788) Echinophyllia echinata (Saville-Kent, 1871) Echinophyllia orpheensis Veron and Pichon, 1980 Genus Mycedium Oken, 1815 Mycedium elephatotus (Pallas, 1766) Mycedium mancaoi Nemenzo, 1979 Genus Oxypora Saville-Kent, 1871 Oxypora crassispinosa Nemenzo, 1979 Oxypora glabra Nemenzo, 1959 Oxypora lacera Verrill, 1864 Recorded in SW Indian Ocean (Veron 2000) X X X X Recorded in Madagascar √ √ √ X X X X X X X X X X X X X X X X X P P √ X √ X X √ √ X X √ √ X √ √ √ √ √ √ √ √ √ √ √ √ √ X X X X √ √ √ X X √ √ X √ √ √ Coral Reefs of Northwest Madagascar 77 Appendix 1 Zooxanthellate Scleractinia Genus Pectinia Oken, 1815 Pectinia africanus Veron, 2000 Pectinia lactuca (Pallas, 1766) Family Merulinidae Verrill, 1866 Genus Hydnophora Fischer de Waldheim, 1807 Hydnophora exesa (Pallas, 1766) Hydnophora microconos (Lamarck, 1816) Hydnophora rigida (Dana, 1846) Genus Merulina Ehrenberg, 1834 Merulina ampliata (Ellis and Solander, 1786) Merulina scabricula Dana, 1846 Family Dendrophylliidae Gray, 1847 Genus Turbinaria Oken, 1815 Turbinaria undescribed species Turbinaria frondens (Dana, 1846) Turbinaria irregularis, Bernard, 1896 Turbinaria mesenterina (Lamarck, 1816) Turbinaria peltata (Esper, 1794) Turbinaria reniformis Bernard, 1896 Turbinaria stellulata (Lamarck, 1816) Genus Heteropsammia Milne Edwards and Haime, 1848 Heteropsammia cochlea (Spengler, 1781) Family Caryophylliidae Gray, 1847 Genus Heterocyathus Milne Edwards and Haime, 1848 Heterocyathus aequicostatus Milne Edwards and Haime, 1848 Family Mussidae Ortmann, 1890 Genus Blastomussa Wells, 1961 Blastomussa undescribed species Blastomussa merleti Wells, 1961 Genus Micromussa Veron, 2000 Micromussa cf. amakusensis (Veron, 1990) Genus Acanthastrea Milne Edwards and Haime, 1848 Acanthastrea brevis Milne Edwards and Haime, 1849 Acanthastrea echinata (Dana, 1846) Acanthastrea hemprichii (Ehrenberg, 1834) Acanthastrea hillae Wells, 1955 Acanthastrea ishigakiensis Veron, 1990 Acanthastrea lordhowensis Veron and Pichon, 1982 Genus Cynarina Brüggemann, 1877 Cynarina lacrymalis (Milne Edwards and Haime, 1848) Genus Lobophyllia Blainville, 1830 Lobophyllia corymbosa (Forskål, 1775) Lobophyllia hataii Yabe and Sugiyama, 1936 Lobophyllia hemprichii (Ehrenberg, 1834) Lobophyllia robusta Yabe and Sugiyama, 1936 Genus Scolymia Haime, 1852 Scolymia cf. vitiensis Brüggemann, 1877 Recorded in SW Indian Ocean (Veron 2000) Recorded in Madagascar X X √ P X X X √ √ P X X √ √ X X X X X X √ √ √ √ √ √ √ X √ √ √ X √ X X X X X X √ √ √ √ √ √ X √ X X X X √ √ √ √ X √ continued 78 Rapid Assessment Program Total inventory of species recorded during the present study and previously recorded for the whole of the SW Indian Ocean Zooxanthellate Scleractinia Genus Symphyllia Milne Edwards and Haime, 1848 Symphyllia agaricia Milne Edwards and Haime, 1849 Symphyllia erythraea (Klunzinger, 1879) Symphyllia recta (Dana, 1846) Symphyllia valenciennesii Milne Edwards and Haime, 1849 Family Faviidae Gregory, 1900 Genus Caulastrea Dana, 1846 Caulastrea connata (Ortmann, 1892) Caulastrea furcata Dana, 1846 Caulastrea tumida Matthai, 1928 Genus Favia Oken, 1815 Favia danae Verrill, 1872 Favia favus (Forskål, 1775) Favia helianthoides Wells, 1954 Favia laxa (Klunzinger, 1879) Favia lizardensis Veron and Pichon, 1977 Favia maritima (Nemenzo, 1971) Favia matthai Vaughan, 1918 Favia pallida (Dana, 1846) Favia rotumana (Gardiner, 1899) Favia speciosa Dana, 1846 Favia stelligera (Dana, 1846) Favia truncatus Veron, 2000 Favia veroni Moll and Borel-Best, 1984 Genus Barabattoia Yabe and Sugiyama, 1941 Barabattoia amicorum (Milne Edwards and Haime, 1850) Genus Favites Link, 1807 Favites abdita (Ellis and Solander, 1786) Favites chinensis (Verrill, 1866) Favites complanata (Ehrenberg, 1834) Favites ﬂexuosa (Dana, 1846) Favites halicora (Ehrenberg, 1834) Favites micropentagona Veron, 2000 Favites paraﬂexuosa Veron, 2000 Favites pentagona (Esper, 1794) Favites russelli (Wells, 1954) Favites spinosa (Klunzinger, 1879) Favites vasta (Klunzinger, 1879) Genus Goniastrea Milne Edwards and Haime, 1848 Goniastrea aspera Verrill, 1905 Goniastrea australensis (Milne Edwards and Haime, 1857) Goniastrea columella Crossland, 1948 Goniastrea edwardsi Chevalier, 1971 Goniastrea minuta Veron, 2000 Goniastrea palauensis (Yabe and Sugiyama, 1936) Goniastrea pectinata (Ehrenberg, 1834) Goniastrea peresi (Faure and Pichon, 1978) Recorded in SW Indian Ocean (Veron 2000) X X X Recorded in Madagascar √ √ √ P X X X √ P X X X X X X X X X X X X X √ √ √ √ √ √ √ √ √ √ √ √ √ √ X X X X X √ √ √ √ √ √ P √ √ √ √ X X X X X X X X √ √ P √ P P √ √ X X X X X Coral Reefs of Northwest Madagascar 79 Appendix 1 Zooxanthellate Scleractinia Goniastrea thecata Veron, DeVantier and Turak, 2000 Goniastrea retiformis (Lamarck, 1816) Genus undescribed Species undescribed Genus Platygyra Ehrenberg, 1834 Platygyra acuta Veron, 2000 Platygyra carnosus Veron, 2000 Platygyra crosslandi Matthai, 1928 Platygyra daedalea (Ellis and Solander, 1786) Platygyra lamellina (Ehrenberg, 1834) Platygyra pini Chevalier, 1975 Platygyra ryukyuensis Yabe and Sugiyama, 1936 Platygyra sinensis (Milne Edwards and Haime, 1849) Platygyra verweyi Wijsman-Best, 1976 Genus Oulophyllia Milne Edwards and Haime, 1848 Oulophyllia bennettae (Veron, Pichon, 1977) Oulophyllia crispa (Lamarck, 1816) Genus Leptoria Milne Edwards and Haime, 1848 Leptoria phrygia (Ellis and Solander, 1786) Genus Montastrea Blainville, 1830 Montastrea annuligera (Milne Edwards and Haime, 1849) Montastrea colemani Veron, 2000 Montastrea curta (Dana, 1846) Montastrea magnistellata Chevalier, 1971 Montastrea serageldini Veron, 2000 Montastrea valenciennesi (Milne Edwards and Haime, 1848) Genus Plesiastrea Milne Edwards and Haime, 1848 Plesiastrea devantieri Veron, 2000 Plesiastrea versipora (Lamarck, 1816) Genus Leptastrea Milne Edwards and Haime, 1848 Leptastrea aequalis Veron, 2000 Leptastrea bottae (Milne Edwards and Haime, 1849) Leptastrea pruinosa Crossland, 1952 Leptastrea purpurea (Dana, 1846) Leptastrea transversa Klunzinger, 1879 Genus Cyphastrea Milne Edwards and Haime, 1848 Cyphastrea chalcidium (Forskål, 1775) Cyphastrea microphthalma (Lamarck, 1816) Cyphastrea serailea (Forskål, 1775) Cyphastrea sp. Genus Diploastrea Matthai, 1914 Diploastrea heliopora (Lamarck, 1816) Genus Echinopora Lamarck, 1816 Echinopora forskaliana (Milne Edwards and Haime, 1850) Echinopora fruticulosa (Ehrenberg, 1834) Echinopora gemmacea Lamarck, 1816 Echinopora hirsutissima Milne Edwards and Haime, 1849 Echinopora lamellosa (Esper, 1795) Echinopora mammiformis (Nemenzo, 1959) Recorded in SW Indian Ocean (Veron 2000) X Recorded in Madagascar √ √ √ X X X X X X X X √ P √ √ √ √ √ √ √ X √ √ X √ X X X X X X P P √ √ √ √ X √ √ X X X X √ √ √ √ √ X X X √ √ √ √ X √ X X X X X X √ P √ √ √ P continued 80 Rapid Assessment Program Total inventory of species recorded during the present study and previously recorded for the whole of the SW Indian Ocean Zooxanthellate Scleractinia Family Trachyphyllidae Verrill, 1901 Genus Trachyphyllia Milne Edwards and Haime, 1848 Trachyphyllia geoffroyi (Audouin, 1826) Family Poritidae Gray, 1842 Genus Alveopora Blainville, 1830 Alveopora undescribed species Alveopora allingi Hoffmeister, 1925 Alveopora daedalea (Forskål, 1775) Alveopora excelsa Verrill, 1863 Alveopora fenestrata (Lamarck, 1816) Alveopora cf. spongiosa Dana, 1846 Alveopora tizardi Bassett-Smith, 1890 Genus Goniopora Blainville, 1830 Goniopora albiconus Veron, 2000 Goniopora burgosi Nemenzo, 1955 Goniopora columna Dana, 1846 Goniopora djiboutiensis Vaughan, 1907 Goniopora lobata Milne Edwards and Haime, 1860 Goniopora minor Crossland, 1952 Goniopora planulata (Ehrenberg, 1834) Goniopora savignyi Dana, 1846 Goniopora somaliensis Vaughan, 1907 Goniopora stokesi Milne Edwards and Haime, 1851 Goniopora stutchburyi Wells, 1955 Goniopora cf. tenella (Quelch, 1886) Goniopora tenuidens (Quelch, 1886) Genus Porites Link, 1807 Porites annae Crossland, 1952 Porites attenuata Nemenzo 1955 Porites australiensis Vaughan, 1918 Porites columnaris Klunzinger, 1879 Porites cumulatus Nemenzo, 1955 Porites cylindrica Dana, 1846 Porites deformis Nemenzo, 1955 Porites densa Vaughan, 1918 Porites echinulata Klungzinger, 1879 Porites evermanni Vaughan, 1907 Porites ﬂavus Veron, 2000 Porites horizontalata Hoffmeister, 1925 Porites latistellata Quelch, 1886 Porites lichen Dana, 1846 Porites lobata Dana, 1846 Porites lutea Milne Edwards and Haime, 1851 Porites mayeri Vaughan, 1918 Porites monticulosa Dana, 1846 Porites murrayensis Vaughan, 1918 Porites napopora Veron, 2000 Porites negrosensis Veron, 1990 Porites nigrescens Dana, 1846 Recorded in SW Indian Ocean (Veron 2000) X X X X X X X X X X X X X X X X Recorded in Madagascar √ √ √ √ √ √ √ √ √ √ √ √ √ √ √ √ √ √ √ √ √ √ X X X X X X X X X X X √ √ √ √ √ √ √ √ √ √ √ √ √ √ √ √ √ √ √ √ Coral Reefs of Northwest Madagascar 81 Appendix 1 Zooxanthellate Scleractinia Porites ornata Nemenzo, 1971 Porites profundus Rehberg, 1892 Porites rugosa Fenner & Veron, 2000 Porites rus (Forskål, 1775) Porites sillimaniana Nemenzo, 1976 Porites solida (Forskäl, 1775) Porites somaliensis Gravier, 1911 Porites stephensoni Crossland, 1952 Porites tuberculosa Veron, 2000 Porites vaughani Crossland, 1952 Porites sp. 1 Porites sp. 2 Genus Poritipora Veron, 2000 Poritipora paliformis Veron, 2000 Genus Stylaraea Milne Edwards and Haime, 1851 Stylaraea punctata (Linneaus, 1758) TOTALS Recorded in SW Indian Ocean (Veron 2000) X X X X X Recorded in Madagascar √ √ √ √ √ √ √ √ √ √ √ √ X √ X 336 P 380 = Recorded during the present trip (323 species). P = Previous record but not recorded during the present trip (57 species). X = Recorded in the SW Indian Ocean previous to the present trip (336 species). undescribed species = species known to be new. sp. = unidentiﬁed species. √ continued 82 Rapid Assessment Program Appendix 2 Site records of scleractinian coral species of Northwest Madagascar by Turak and Veron. This working checklist includes only the species found in designated study sites. It does not include the results of subsequent laboratory study. See full checklist by Veron and Turak, Appendix 1. Species Astrocoeniidae Stylocoeniella armata Stylocoeniella guentheri Pocilloporidae Pocillopora damicornis Pocillopora eydouxi Pocillopora indiania Pocillopora verrucosa Pocillopora cf. kellerheri Seriatopora aculeata Seriatopora caliendrum Seriatopora guttatus Seriatopora hystrix Seriatopora sp. Seriatopora sp. new Stylophora madagascarensis Stylophora pistillata Stylophora subseriata Acroporidae Montipora aequituberculata Montipora calcarea Montipora caliculata Montipora cf. confusa Montipora danae Montipora efﬂorescens Montipora effusa Montipora ﬂoweri Montipora cf. foveolata Montipora grisea Montipora informis Montipora millepora Montipora mollis Montipora meandrina Montipora monasteriata Montipora peltiformis Site records 18, 22 2, 5, 6, 8, 9, 13-17, 21, 23 1-14, 16-29 1-4, 6, 8, 9, 11-14, 16-18, 21, 23, 26, 28 1-7, 12, 14, 16-18 1, 5, 8, 9, 11-14, 16-18, 21, 26 3, 8, 9, 11, 18, 26 1, 18-22, 27 1, 3-8, 10, 11, 13, 14, 16, 22, 23, 27, 28, 29 8, 27 1, 2, 5-7, 10-23, 27, 28, 29 5, 7, 11-14, 17 15, 16, 21, 23, 27, 29 5, 7, 9, 11, 20 5-7, 9, 11-17, 19-21, 27 3, 8 5, 6, 8, 14, 20 11-14, 16-18, 20-24 9, 11, 14, 16, 18, 19, 21, 26, 28 5, 6, 8 4-7, 10, 12, 14, 15, 17, 28 4, 11, 13 14, 26 24 2, 7, 13 11, 12, 14, 15, 17, 19 3-5, 7, 11, 15, 17-22, 27, 29 6, 12, 19, 21, 27 25, 26 17, 18 2-6, 8, 11-13, 17-22, 24, 27, 28 12, 14, 17-19 Coral Reefs of Northwest Madagascar 83 Appendix 2 84 Species Site records Montipora spumosa Montipora tuberculosa Montipora turgescens Montipora undata Montipora venosa Montipora verrucosa Acropora abrotanoides Acropora aculeus Acropora anthocercis Acropora appressa Acropora arabensis Acropora aspera Acropora austera Acropora bifurcata Acropora cerealis Acropora clathrata Acropora copiosa Acropora cuneata Acropora cytherea Acropora digitifera Acropora divaricata Acropora donei Acropora eurystoma Acropora ﬂorida Acropora formosa Acropora forskali Acropora gemmifera Acropora cf. gemmifera Acropora grandis Acropora granulosa Acropora haimei Acropora hemprichii Acropora horrida Acropora humilis Acropora hyacinthus Acropora insignis Acropora irregularis Acropora latistella Acropora loripes Acropora lutkeni Acropora microphthalma Acropora mirabilis Acropora nasuta Acropora natalensis Acropora nobilis Acropora ocellata Acropora palifera Acropora paniculata Acropora papillarae Acropora pinguis 24, 26, 27 1-8, 10-14, 18-21, 23, 25-29 12, 15, 17, 18, 20, 21, 27 5, 6, 10-18, 21-23, 27 3-8, 11-13, 18, 21, 26 18, 21 1-6, 9, 11, 16 1, 2, 4-17, 19, 20, 21, 27, 29 1-3, 8, 15, 16, 20, 21, 26, 29 2, 8, 13, 16-20, 23, 26, 27 1-9, 11-13, 18, 19, 21-23, 26, 29 4, 11, 18, 20, 27 1, 2, 4-8, 11-15, 17-21, 26 12, 18, 27-29 1, 3, 5-7, 11, 14, 16, 17, 21-26 1, 2, 5-9, 11-14, 16-18, 21, 26, 28, 29 8, 13, 14, 18 18, 27 1, 2, 4-9, 11-14, 16-20, 23, 27, 29 2-6, 9, 11, 14, 18, 20, 24 1-24, 26, 27, 29 21, 22 2-5, 7, 8, 13-15, 19, 20, 22-24, 27, 29 6, 13 2, 4-8, 11-14, 16, 18-20, 26, 27, 29 13, 14 1-9, 11-14, 16-18, 20, 24, 26, 28, 29 18, 19, 22, 27, 29 14, 18, 26, 27 17, 18 14 4, 5, 7, 13-15, 18, 27, 29 5, 6, 15, 16, 19, 26 1-5, 8, 9, 11, 12, 14-27, 29 2, 4, 5, 7, 9, 11, 13 8, 10, 13, 25, 26, 29 2, 5-8, 12-14, 18, 26, 29 1, 2, 4, 15, 17-21, 27, 29 4, 5, 8, 9, 12, 14, 16, 17, 19, 21, 28 1, 4, 6, 11, 14, 16, 18, 19 1, 2, 4-6, 8, 14, 16-20, 26, 27, 29 3 3, 5, 12, 15, 19, 29 12, 23 1-8, 12, 14, 18-20, 27, 29 3, 6, 9, 12-14, 17, 26, 29 1, 2, 4-7, 11-14, 18-21, 23, 26-29 21 2, 3, 6, 7 16, 17 Rapid Assessment Program continued Site records of scleractinian coral species of northwest Madagascar Species Acropora plantaginea Acropora prostrata Acropora pulchra Acropora robusta Acropora rosaria Acropora samoensis Acropora secale Acropora selago Acropora squarrosa Acropora subulata Acropora tenuis Acropora valida Acropora variabilis Acropora variolosa Acropora verweyi Acropora willisae Acropora yongei Acropora sp. new Acropora sp. Astreopora expansa Astreopora cf. incrustans Astreopora listeri Astreopora myriophthalma Astreopora ocellata Astreopora suggesta Euphyllidae Catalophyllia jardinei Plerogyra sinuosa Physogyra lichtensteini Oculinidae Galaxea astreata Galaxea fascicularis Merulinidae Hydnophora exesa Hydnophora microconos Hydnophora rigida Merulina ampliata Siderastreidae Horastrea indica Psammocora contigua Psammocora explanulata Psammocora haimeana Psammocora nierstraszi Psammocora obtusangula Psammocora profundacella Psammocora superﬁcialis Coscinaraea columna Coscinaraea crassa Coscinaraea monile Site records 3 19, 22, 27 3 2, 5, 9, 14, 17 29 17, 18, 21, 25-27 1-8, 11-18, 21, 22, 25, 29 1-4, 6, 9, 10, 17, 21, 23, 26, 27, 29 29 2, 4-8, 11-20, 22, 23, 26, 27, 29 1, 2, 4-8, 10-23, 26, 29 1, 4-6, 11-13, 18, 20-23, 27, 29 3, 11, 12, 27 2, 4-6, 8, 12, 17, 22, 24 26, 29 7, 9, 12, 14, 17, 20, 21, 23, 27, 29 1, 2, 4, 7, 13, 17, 19, 27, 29 5, 15, 29 5 7 5, 6, 8, 12, 14-16, 18-20, 23, 26 1, 5, 14-19, 27 1-28 1, 15, 23, 27 2, 4-6, 16, 23, 25, 26 17 3-6, 11-13, 15, 17-24, 27-29 2-8, 10-12, 15, 19-24, 27-29 1, 5, 13, 16, 19-21, 26, 27, 29 1-8, 10-24, 26-29 1-24, 26-29 1, 2, 4-8, 11, 12, 14, 19-21, 23, 26-28 6 1-7, 11-24, 26-29 6, 14, 16 1, 17, 20, 22, 24, 27 15, 17, 27 1, 5, 6, 12, 14, 18, 20, 21, 28 11, 21 1, 27, 29 1, 2, 5, 11-14, 17, 18, 21, 23, 25, 26, 28 19 11, 17, 18, 19, 29 19, 22, 26-28 1-6, 8-21, 23, 24, 27-29 Coral Reefs of Northwest Madagascar 85 Appendix 2 86 Species Site records Coscinaraea cf. monile Coscinaraea wellsi Agariciidae Pavona bipartita Pavona clavus Pavona decussata Pavona difﬂuens Pavona duerdeni Pavona explanulata Pavona frondifera Pavona maldivensis Pavona varians Pavona venosa Leptoseris explanata Leptoseris foliosa Leptoseris hawaiiensis Leptoseris incrustans Leptoseris mycetoseroides Leptoseris scabra Leptoseris yabei Gardineroseris planulata Pachyseris speciosa Fungiidae Cycloseris costulata Cycloseris curvata Cycloseris cyclolites Cycloseris patelliformis Cantharellus sp. Cantharellus doederleini Fungia concinna Fungia danai Fungia fungates Fungia granulosa Fungia horrida Fungia klunzingeri Fungia paumotensis Fungia repanda Fungia scutaria Fungia seychellensis Fungia sp. Herpolitha limax Herpolitha weberi Polyphyllia talpina Halomitra pileus Podabacia crustacea Lithophyllon mokai Pectiniidae Echinophyllia aspera Echinophyllia echinata Echinophyllia echinoporites 3, 5-7, 11, 13-15, 19, 21, 23 17 Rapid Assessment Program 19 3, 13, 21, 29 3, 5, 6, 12, 13, 17, 20-22, 24, 29 29 1, 11-13, 17, 20, 27, 29 1, 3-7, 9, 11, 13, 17-19, 21-23, 28, 29 5, 18-20, 22, 29 4, 12, 13, 17 1-5, 7-13, 17-23, 25, 27, 28 21, 25, 28 5, 9, 28 10, 19, 28 10, 11, 13 11, 13, 16, 17, 19, 21 9, 11-13, 16, 17, 19, 20, 24, 27 10, 21, 28 7 1, 6, 12-14, 17, 19-21, 23, 29 2, 5, 9-13, 15-24, 27-29 3, 8, 10, 14, 16, 18 7 1-3, 5, 9, 10, 15, 16, 21, 24, 26, 28 14, 15 29 12 2, 4, 5, 7, 12-14, 16-24, 26, 27, 29 1, 4, 12, 29 1, 2, 5, 6, 12-22, 24, 26, 27, 29 1, 2, 9, 13-23, 26-29 1, 2, 4, 5-7, 10, 12-14, 16-22, 26-29 8, 12, 15, 22, 27-29 1, 3, 5-7, 10, 12, 14, 16, 17, 19, 21, 22, 27-29 20 3, 6, 9-13, 18 5, 18, 20, 22, 27, 28 29 7, 10, 12, 16-19, 21, 22, 24, 27, 29 16, 21 1, 5, 14, 16, 17, 21 5, 22, 29 2, 5, 10, 12, 13, 17, 19, 21, 27, 29 10 3-7, 9, 11, 13-17, 19-21, 23, 24, 27 11 4, 5, 7, 19 continued Site records of scleractinian coral species of northwest Madagascar Species Echinophyllia orpheensis Oxypora crassispinosa Oxypora lacera Mycedium elephantotus Mycedium mancaoi Pectinia africanus Meandrinidae Gyrosmillia interrupta Dendrophylliidae Turbinaria frondens Turbinaria irregularis Turbinaria mesenterina Turbinaria peltate Turbinaria reniformis Turbinaria stellulata Turbinaria sp. Heteropsammia cochlea Caryophylliidae Heterosyathus aequicostatus Mussidae Blastomussa merleti Blastomussa sp. Micromussa amakusensis Acanthastrea brevis Acanthastrea echinata Acanthastrea hillae Acanthastrea hemprichii Acanthastrea ishigakiensis Acanthastrea lordhowensis Acanthastrea rotandoﬂora Acanthastrea subechinata Lobophyllia corymbosa Lobophyllia hataii Lobophyllia hemprichii Lobophyllia robusta Symphyllia agaricia Symphyllia erythraea Symphyllia hassi Symphyllia recta Symphyllia valenciennesi Scolymia vitiensis Cynarina lacrymalis Faviidae Caulastrea connata Favia danae Favia favus Favia helianthoides Favia cf. lacuna Favia laxa Favia lizardensis Site records 4, 7, 9, 14-18, 21, 23, 24, 27, 29 5, 22 2, 4, 5, 11, 13-19, 22, 23, 27-29 1, 4-7, 10-24, 27-29 12, 17, 23 5, 6, 17 1, 3-10, 12, 14, 17-19, 21, 23 26 4, 6, 11, 19, 22, 24, 26, 27 3-5, 7, 9, 12, 15-18, 20, 21, 27, 28 2-7, 9, 19, 20 6 4, 6-9, 16, 19-22, 26-28 17, 26 8 8 2-6, 8, 9, 15-17 8, 14, 15, 23, 28 3, 9, 21 1, 5, 6, 12, 15, 16, 20, 21, 26, 27 1, 4-7, 9, 16, 18-24, 27, 28 2, 14-17, 21, 23, 24 1-19, 23, 24, 27 4, 8, 9, 12-14, 19, 20, 26-29 26 1, 4, 5, 7, 15, 28 1, 2 4-7, 12, 14-16, 21-24, 29 2, 4, 5, 8, 14-17, 19-22, 24, 26, 29 1-9, 12, 14-23, 26-29 8, 9, 14-17, 19, 22, 27, 29 2, 9, 24, 27 16, 17, 26, 28 2 1 9, 14, 26, 27 13 5, 15, 23 3-9, 15, 16, 23, 26, 27 2, 3, 7, 9, 11, 15, 22 1-7, 9-20, 22-29 12, 17, 22 2, 6, 7, 25 24 1-5, 7, 9, 13-21, 23, 26, 27, 29 Coral Reefs of Northwest Madagascar 87 Appendix 23 Appendix Species Favia maritima Favia matthaii Favia pallida Favia cf. rosaria Favia rotumana Favia speciosa Favia stelligera Favia truncatus Favia veroni Barabattoia amicorum Favites abdita Favites chinensis Favites complanata Favites ﬂexuosa Favites halicora Favites micropentagona Favites paraﬂexuosa Favites pentagona Favites russelli Favites spinosa Favites vasta Goniastrea aspera Goniastrea australiensis Goniastrea columella Goniastrea edwardsi Goniastrea favulus Goniastrea palauensis Goniastrea pectinata Goniastrea peresi Goniastrea retiformis Goniastrea thecata Platygyra acuta Platygyra carnosus Platygyra contorta Platygyra crosslandi Platygyra daedalea Platygyra lamellina Platygyra pini Platygyra ryukyuensis Platygyra sinensis Platygyra cf. verweyi Leptoria phrygia Oulophyllia bennette Oulophyllia crispa Montastrea annuligera Montastrea colemani Montastrea curta Montastrea magnistellata Montastrea serageldini Plesiastrea versipora 88 Rapid Assessment Program Site records 4, 5, 8, 14-16, 18-21, 23, 26 1, 3-15, 17-27, 29 1-3, 7, 11-16, 18, 19, 21, 23, 29 24 1, 2 1-3, 5-9, 11-16, 19-24, 26-29 1-7, 9, 11-14, 16, 18-21, 24, 26-28 1, 2, 4-7, 12, 13, 20-23, 24, 26 8 1, 22, 24 1, 2, 5, 9, 12-14, 16-18, 20, 24 9, 11, 16 1, 2, 4-9, 12, 14-16, 18-21, 25-27 1-7, 9, 11-24, 26-28 2, 6, 11, 17, 23 7, 11, 12, 20, 26, 29 7 1, 2, 4-6, 8-14, 16, 17, 19-21, 23, 26-29 1, 2, 4-9, 11-14, 17, 22, 23, 26 1, 12, 21, 23 1-8, 12-23, 26-28 12, 13, 18, 19, 21, 22, 28 1, 3, 6, 18, 24 3, 24, 26 5-7, 11, 13, 14, 18-21, 25-29 3 23 1, 2, 4-7, 10-24, 26-29 1-24, 26-28 1-3, 5-7, 9, 11-14, 17-27, 29 2, 7, 9 1-6, 8, 9, 14-16, 18-24, 26, 27, 29 2, 7 1, 3-5, 9, 12, 19, 23, 24, 27 5-7, 11-13, 16, 17, 19, 22, 26, 27 1-12, 14-22, 24, 26, 27 1, 2, 4-7, 9, 14-20, 22, 23, 24, 29 1, 2, 4-9, 11-15, 17-19, 22, 23, 25-29 2, 3, 7, 16, 19, 22, 23, 27 2, 4-7, 9, 13, 16, 17, 19-24, 26, 27, 29 1, 4 1-9, 11-14, 16-21, 23, 24, 26-29 2, 6, 7, 9 1-3, 5-7, 9-18, 20 23 26, 27, 28 1-6, 12, 14-17, 19, 24-26, 28 1, 2, 4-7, 13, 15, 16, 19-24, 27, 29 2-8, 11, 12, 14, 18-20, 25, 26, 28 1, 3, 4, 6, 8-13, 15, 18, 20-23, 26, 29 continued Site records of scleractinian coral species of northwest Madagascar Species Plesiastrea devantieri Plesiastrea sp. Diploastrea heliopora Leptastrea bottae Leptastrea pruinosa Leptastrea purpurea Leptastrea transversa Leptastrea aequalis Cyphastrea chalcidium Cyphastrea microphthalma Cyphastrea serailia Echinopora gemmacea Echinopora hirsutissima Echinopora lamellosa Unidentiﬁed genus Trachyphylliidae Trachyphyllia geoffreyi Poritidae Porites massive Porites attenuata Porites cylindrica Porites echinulata Porites evermani Porites latistella Porites lichen Porites mayeri Porites monticulosa Porites nigrescens Porites cf. nodifera Porites profundus Porites rus Porites solida Porites vaughani Porites ﬂat branches Poritopora paliformis Site records 5-8, 11-14, 18-21, 23, 25-28 8, 23 1-10, 14-29 9, 21 2, 5, 6, 9, 11-14, 16-18, 20, 22-29 1, 4-9, 11-13, 16-22, 24 2, 3, 8-13, 15, 19, 21, 22, 24, 25, 27-29 7, 11, 12, 14, 20, 23 3, 5-7, 12-14, 17-21, 23, 24, 26-29 1-3, 5, 8, 9, 11-16, 18, 20, 21, 23, 25-29 3, 5-9, 11, 15, 16, 19, 28, 29 1-9, 11, 12, 18 1-4, 6-29 3, 6, 20, 21 3 Goniopora albiconus Goniopora ciliatus Goniopora columna Goniopora djiboutiensis Goniopora lobata Goniopora minor Goniopora somaliensis Goniopora stokesi Goniopora stutchburyi Goniopora tenuidens Alveopora allingi Alveopora daedalea Alveopora fenestrata Alveopora spongiosa Alveopora tizardi Alveopora sp. 2, 6, 15, 20, 21, 23, 26, 27, 29 14, 16, 20 1-29 1-3, 5, 6, 19, 22, 23, 29 4, 5, 7, 12-16, 18-21, 23, 24, 26, 27, 29 25, 26 1 8, 12, 13, 17, 19, 23 12, 21 1 12, 19 12, 13, 21 1, 8 2, 3, 5-7, 11-15, 17-21, 23, 24, 26-28 2, 6, 12, 17, 19-24, 27, 29 18 1, 19 11, 13, 17, 18, 26 22 1, 3, 9 1, 3-6, 8, 15, 21, 29 1, 4, 6, 23, 24, 29 7, 10, 11, 18, 20, 23 2-5, 16, 18, 26, 29 1, 3-8, 14-16, 18-23, 26-29 1-6, 8, 14, 15, 18, 20-24, 26, 28, 29 3 10, 27 8 8, 9, 12, 15, 23, 26 3 1, 4-6, 9, 11-14, 16, 17, 19, 21, 23, 26, 27 5, 6, 8, 13, 14, 17, 28, 29 4-6, 11, 14, 15 Coral Reefs of Northwest Madagascar 89 Appendix 3 List of molluscs collected Species CLASS POLYPLACOPHORA Family Chitonidae Acanthopleura gemmata (Blainville, 1825) CLASS GASTROPODA Family Patellidae Cellana rota (Gmelin, 1791) Patelloida striata (Quoy & Gaimard, 1834) Family Haliotidae Haliotis cf. crebrisculpta Sowerby, 1914 Family Fissurellidae Diodora sp. Emarginula sp. Family Turbinidae Monodonta labio (Linnaeus, 1758) Turbo argyrostomus (Linnaeus, 1758) Turbo cinereus Born, 1778 Turbo coronatus (Gmelin, 1791) Turbo petholatus Linnaeus, 1758 Family Trochidae Calliostoma sp. Cantharidus sp. Clanculus pharaonis (Linnaeus, 1758) Herpetopoma sp. Tectus maculatus Linnaeus, 1758 Tectus pyramis Born, 1778 Tectus virgatus (Gmelin, 1791) Thalotia sp. Stomatella varia (A. Adams, 1850) Family Neritidae Nerita albicilla Linnaeus, 1758 Nerita plicata Linnaeus, 1758 Nerita polita Linnaeus, 1758 Nerita undata Linnaeus, 1758 Family Cerithiidae Cerithium africanum Houbrick, 1992 Cerithium balteatum Philippi, 1848 Cerithium caeruleum Sowerby, 1855 Cerithium citrinum Sowerby, 1855 Cerithium echinatum (Lamarck, 1822) Cerithium egenum Gould, 1849 90 Rapid Assessment Program Site numbers 1, 2, 5, 7, 9, 20, 27B 1, 5, 7, 9, 20, 27B 5, 7, 9, 20, 27B 22 15 20 5, 27B 2, 3, 5, 6, 7, 9, 14, 15, 18, 23, 25, 26, 28 27B 27 23 11 21 3, 10, 18 6 1, 18, 22, 29 2, 6, 22, 23, 25, 27 13, 18, 19, 23 11 3 0, 5, 7, 9, 18, 20 0, 18, 20, 27B 2, 18, 20, 27B 18, 20, 27B 0, 2-9, 15, 16, 18B, 19, 21, 22, 24, 27 4-7, 18, 27 25 1, 3-7, 9, 11, 12, 14, 16-18, 21, 25-27B 2 List of molluscs collected Species Cerithium nesioticum Pilsbry & Vanetta, 1906 Cerithium nodulosus (Bruguière, 1792) Cerithium ruppellii Philippi, 1848 Cerithium salebrosum Sowerby, 1855 Clypeomorus batillariaeformis Habe & Kosuge, 1966 Clypeomorus bifasciatus (Sowerby, 1855) Clypeomorus petrosus (Wood, 1828) Clypeomorus purpurastoma Houbrick, 1985 Rhinoclavis articulata (Adams & Reeve, 1850) Rhinoclavis aspera (Linnaeus, 1758) Rhinoclavis fasciatus (Bruguière, 1792) Rhinoclavis kochi (Philippi, 1848) Rhinoclavis sinensis (Gmelin, 1791) Family Turritellidae Turritella sp. Family Planaxidae Planaxis sulcatus (Born, 1780) Family Potamididae Terebralia palustris (Linnaeus, 1758) Family Modulidae Modulus tectum (Gmelin, 1791) Family Littorinidae Littoraria scabra (Linnaeus, 1758) Littorina cf. kraussi Rosewater, 1970 Family Strombidae Lambis chiragra (Linnaeus, 1758) Lambis lambis (Linnaeus, 1758) Lambis scorpius (Linnaeus, 1758) Lambis truncatus (Humphrey, 1786) Strombus decorus (Röding, 1798) Strombus dentatus Linnaeus, 1758 Strombus fusiformis Sowerby, 1842 Strombus gibberulus Linnaeus, 1758 Strombus labiatus Röding, 1798 Strombus lentiginosus Linnaeus, 1758 Strombus luhuanus Linnaeus, 1758 Strombus mutabilis Swainson, 1821 Strombus terebellatus Sowerby, 1842 Terebellum terebellum (Linnaeus, 1758) Family Hipponicidae Hipponix conicus (Schumacher, 1817) Family Vanikoridae Vanikoro cancellata (Lamarck, 1822) Family Capulidae Cheilea equestris (Linnaeus, 1758) Family Vermetidae Serpulorbis colubrina (Röding, 1798) Family Cypraeidae Cypraea annulus Linnaeus, 1758 Cypraea arabica Linnaeus, 1758 Cypraea argus Linnaeus, 1758 Cypraea asellus Linnaeus, 1758 Cypraea caputserpentis Linnaeus, 1758 Cypraea carneola Linnaeus, 1758 Cypraea caurica Linnaeus, 1758 Cypraea cicercula Linnaeus, 1758 Cypraea clandestina Linnaeus, 1767 Site numbers 2, 4, 5, 11, 12, 15, 18, 25, 26 1-5, 18, 21, 24-26 0, 4, 12, 14-17, 19, 20, 22-26, 30 2, 5, 12-14, 21, 23, 24 20, 24 27 27B 22, 24, 27 12 1-8, 11-15, 18, 19, 21-28, 30 12 16, 22, 25 12 10 20 27B 2, 6, 7, 15, 18, 25, 26, 28 27B 21 19 1, 4 3, 5, 11, 13, 18, 19, 21, 23-26, 28, 29 1, 2, 5, 6, 13, 14, 21, 23, 25, 28 4, 12, 13, 18B, 21-23, 25, 26, 28 10, 11 10, 14, 20, 24, 29 0, 15, 18, 20, 22, 29 1-5, 13, 20, 23-27 4, 18B, 21, 23-25 4, 10, 11 5, 11, 18B, 19, 24, 25, 28 13, 14, 19 1, 12, 19, 26, 28 1, 3, 6-8, 12, 21 8 1, 3, 6, 12, 18, 25, 26, 28, 29 1-4, 6-9, 16, 18, 20, 21, 23, 25, 27, 28 0, 6, 15, 18B, 20, 27B 9 18 14, 17, 27 1, 2, 5, 6, 9 3-9, 13, 14, 16-18B, 23-26, 30 14, 24, 27, 29 11, 19, 21, 29 3, 7, 20 Coral Reefs of Northwest Madagascar 91 Appendix 3 Species Cypraea cribraria Linnaeus, 1758 Cypraea contaminata Sowerby, 1832 Cypraea diliculum (Reeve, 1845) Cypraea erosa Linnaeus, 1758 Cypraea ﬁmbriata Gmelin, 1791 Cypraea helvola Linnaeus, 1758 Cypraea hirundo Linnaeus, 1758 Cypraea histrio (Gmelin, 1791) Cypraea isabella Linnaeus, 1758 Cypraea kieneri Hidalgo, 1906 Cypraea limacina Lamarck, 1810 Cypraea lynx Linnaeus, 1758 Cypraea mappa Linnaeus, 1758 Cypraea microdon Gray, 1828 Cypraea owenii (Sowerby, 1837) Cypraea punctata Linnaeus, 1758 Cypraea scurra Gmelin, 1791 Cypraea staphylaea Linnaeus, 1758 Cypraea stolida Linnaeus, 1758 Cypraea talpa Linnaeus, 1758 Cypraea teres Gmelin, 1791 Cypraea tigris Linnaeus, 1758 Family Ovulidae Calpurneus verrucosus (Linnaeus, 1758) Family Triviidae Trivia oryza (Lamarck, 1810) Family Velutinidae Coriocella nigra (Blainville, 1824) Family Naticidae Natica euzona Rècluz, 1844 Natica gualteriana (Rècluz, 1844) Natica onca (Röding, 1798) Natica sp. Natica violacea Sowerby, 1825 Polinices didyma (Röding, 1798) Polinices tumidus (Swainson, 1840) Polinices simae (Dehsayes, 1838) Polinices tumidus (Swainson, 1840) Sinum sp. Family Bursidae Bursa cruentata (Sowerby, 1835) Bursa granularis (Röding, 1798) Bursa rhodostoma (Sowerby, 1835) Bursa nigrita Mulhauser & Blocker, 1979 Tutufa bubo Linnaeus, 1758 Family Cassidae Casmaria erinaceus (Linnaeus, 1758) Cassis cornuta (Linnaeus, 1758) Family Ficidae Ficus subintermedia (d’Orbigny, 1852) Family Ranellidae Cymatium exile (Reeve, 1844) Cymatium ﬂaveolum (Röding, 1798) Cymatium hepaticum (Röding, 1798) Cymatium labiosum (Wood, 1828) Cymatium lotorium (Linnaeus, 1758) Cymatium pileare (Linnaeus, 1758) 92 Rapid Assessment Program Site numbers 1, 5, 10, 12-14, 25, 30 15, 16, 18B 6, 14, 28, 29 0-2, 9-11, 13-15, 18B-22, 25, 27-29 1, 3-6, 8, 10-12, 15, 17, 28-30 0, 3, 4, 9, 11-15, 18B, 19, 21, 26, 28 17 30 0, 3, 6, 8-11, 14, 16, 18B, 19, 22, 25, 29, 30 0, 3, 20 1, 11 19, 27B 21 4 16 14, 17, 22, 29 19 1, 2, 6, 8, 16-18B, 21, 22, 25, 27 5 4, 16, 27-30 9, 11 0, 1, 3, 6, 9, 11, 16, 24, 27 3 11, 13, 17, 29 9 3 2, 20 1, 4, 14, 15, 21, 24, 26, 29 22 3, 10, 29 10 23 1-3, 18B, 23, 28 0-2, 5, 10, 11, 13, 15, 19, 21, 23-27, 29 21 13 2, 4-6, 10, 11, 13, 16, 18, 28, 30 10, 13, 28 2 3, 8, 14 2, 4, 18B, 23, 25, 29 23 18B, 22, 23 21, 27 11,12 19 3, 22 3, 5, 21, 28 2 List of molluscs collected Species Cymatium rubeculum (Linnaeus, 1758) Distorsio reticulata (Röding, 1798) Gyrineum pusillum (A. Adams, 1854) Family Tonnidae Malea pomum (Linnaeus, 1758) Tonna perdix (Linnaeus, 1758) Family Triphoridae Triphorid sp. 1 Triphorid sp. 2 Family Epitoniidae Epitonium sp. Family Eulimidae Hypermastus sp. Family Muricidae Aspella sp. Chicoreus torrefactus Sowerby, 1840 Chicoreus microphyllus (Lamarck, 1816) Chicoreus trigonulus (Lamarck, 1816) Coralliophila erosa (Röding, 1798) Coralliophila neritoidea (Lamarck, 1816) Cronia contracta (Reeve, 1846) Cronia fenestrata (Blainville, 1832) Drupa morum (Röding, 1798) Drupa ricinus (Linnaeus, 1758) Drupa rubusidaeus (Röding, 1798) Drupella cornus (Röding, 1798) Drupella rugosa (Born, 1778) Favartia minatauros Radwin & D’Attilio, 1976 Homalocantha anatomica (Perry, 1811) Latiaxena ﬁmbriata (Hinds, 1844) Morula anaxeres (Kiener, 1835) Morula biconica (Blainville, 1832) Morula granulata (Duclos, 1832) Morula margariticola (Broderip, 1832) Morula nodulifera Pease, 1860 Morula spinosa (H. & A. Adams, 1855) Morula uva (Röding, 1798) Murex ramosus (Linnaeus, 1758) Naquetia triqueter (Born, 1778) Nassa francolina (Bruguière, 1789) Quoyola madreporarum (Sowerby, 1832) Rapa rapa (Gmelin, 1791) Stramonita muricina (Blainville, 1832) Thais alouina (Röding, 1798) Thais savignyi (Deshayes, 1844) Thais tuberosa (Röding, 1798) Family Turbinellidae Vasum ceramicum (Linnaeus, 1758) Vasum turbinellus (Linnaeus, 1758) Family Buccinidae Colubraria nitidula (Sowerby, 1833) Cantharus fumosus (Dillwyn, 1817) Cantharus iostomus (Gray in Grifﬁth & Pidgeon, 1834) Cantharus pulcher (Reeve, 1846) Cantharus undosus (Linnaeus, 1758) Engina concinna (Reeve, 1846) Engina incarnata (Deshayes, 1834) Site numbers 17, 19, 30 28, 30 3, 4, 5, 11 0, 29 11, 13 1, 29 15 26 11 19, 26 0-3, 6, 7, 22, 23, 27 7, 17, 19, 24, 26, 27 4, 16 23 1, 7, 8, 17, 21, 24, 28, 29 22 0, 19 9 6, 9 6, 9, 13 1-3, 5, 8, 11, 13, 17-22, 27-30 6, 13, 15, 19 1 1, 3 22 0, 2, 9 1, 5, 19 0-2, 5, 7, 9, 18, 20, 26 1, 9, 18B, 19, 27 3, 6, 12, 26 1, 5, 28, 29 1, 5-9, 11, 15, 17, 28, 20, 25, 29 3 1, 4 5, 18B, 21 11, 17, 30 15, 16 1 0-2, 6, 8, 9, 17, 19, 20, 26, 28 9 9 7, 19 0-9, 12, 16-19, 21-27B, 29 11, 12 1-3, 5-7, 16-20 11, 12, 15 13 1, 8, 11, 21, 23 28 10, 11 Coral Reefs of Northwest Madagascar 93 Appendix 3 Species Engina zatricum Melvill, 1893 Pollia fragaria (Wood, 1828) Phos cyanostoma A. Adams, 1850 Phos textum (Gmelin, 1791) Family Columbellidae Mitrella albina (Kiener, 1841) Mitrella marquesa (Gaskoin, 1852) Pyrene ﬂava (Bruguière, 1789) Pyrene punctata (Bruguière, 1789) Pyrene scripta (Lamarck, 1822) Pyrene testudinaria (Link, 1807) Pyrene turturina (Lamarck, 1822) Pyrene varians (Sowerby, 1832) Family Nassariidae Nassarius albescens (Dunker, 1846) Nassarius arcularia (Linnaeus, 1758) Nassarius castus (Gould, 1850) Nassarius coronatus (Bruguière, 1798) Nassarius gemmulifera (A. Adams, 1852) Nassarius glans (Linnaeus, 1758) Nassarius granifer (Kiener, 1834) Nassarius luridus (Gould, 1850) Nassarius oneratus (Deshayes, 1863) Nassarius papillosus (Linnaeus, 1758) Nassarius pauperus (Gould, 1850) Nassarius siquijorensis (A. Adams, 1852) Family Fasciolariidae Fusinus colus (Linnaeus, 1758) Latirus fastigium (Reeve, 1847) Latirus gibbulus (Gmelin, 1791) Latirus nodatus (Gmelin, 1791) Latirus polygonus (Gmelin, 1791) Peristernia incarnata (Deshayes, 1830) Peristernia nassatula (Lamarck, 1822) Peristernia forskalli (Tapparone-Canefri, 1879) Peristernia pulchella (Reeve, 1847) Peristernia sp. Pleuroploca trapezium (Linnaeus, 1758) Family Olividae Oliva annulata (Gmelin, 1791) Oliva caerulea (Röding, 1798) Oliva paxillus Reeve, 1855 Family Harpidae Harpa amouretta Röding, 1798 Family Mitridae Mitra ambigua Swainson, 1829 Mitra coffea Schubert & Wagner, 1829 Mitra eremitarum Röding, 1798 Mitra imperialis Röding, 1798 Mitra mitra (Linnaeus, 1758) Mitra unifascialis Kiener, 1838 Mitra vexillum Reeve, 1844 Neocancilla circula (Kiener, 1838) Neocancilla pretiosa (Reeve, 1844) Pterygia sinensis (Reeve, 1844) Subcancilla abyssicola (Schepman, 1813) 94 Rapid Assessment Program Site numbers 18 29 0, 29 29 1-5, 7, 8, 10-14, 16, 21, 22, 25, 26, 28 10, 11, 17 1-2, 6, 10, 11, 14, 19, 21, 27 0, 6 2 26 1-3, 5-8, 10-14, 16-18B, 21, 22 1, 3 2, 4, 14, 15, 20, 23, 25 27 20, 22 15, 22, 27 15 22, 24 1, 2, 10-13, 15, 18, 21 5 11, 12 1, 2, 7, 8, 26, 28 2, 11-13 20, 22 11 13, 16, 26, 28 3, 21, 26 3 1, 5, 8 26 6, 9-12, 15, 18, 26-30 2, 3, 18, 19, 22, 23 1, 2, 13, 18 6, 11, 15, 19 18, 25, 27B 1, 3-5, 10, 12, 13 0, 4, 23 13, 18, 23, 26 10 20, 22 20 19, 24 22 29 0, 14 1 22, 24 14, 15, 17, 22 1, 2, 9 17, 24 List of molluscs collected Species Family Costellariidae Vexillum bernhardina (Röding, 1798) Vexillum cadaverosum (Reeve, 1844) Vexillum cancellaroides (Anton, 1839) Vexillum citrinum (Gmelin, 1791) Vexillum consanguineum (Reeve, 1845) Vexillum crocatum (Lamarck, 1811) Vexillum echinatum (A. Adams, 1853) Vexillum exasperatum (Gmelin, 1791) Vexillum lautum (Reeve, 1844) Vexillum leucodesmum (Reeve, 1845) Vexillum lucidum (Reeve, 1845) Vexillum cf. militaris (Reeve, 1845) Vexillum mirabile (A. Adams, 1853) Vexillum osridis (Issel, 1869) Vexillum paciﬁcum (Reeve, 1845) Vexillum plicarium (Linnaeus, 1758) Vexillum regina (Sowerby, 1828) Vexillum semisculptum (Adams & Reeve, 1850) Vexillum sp. Vexillum speciosum (Reeve, 1844) Vexillum tankervillei (Melvill, 1888) Vexillum turrigerum (Reeve, 1845) Vexillum unifascialis (Lamarck, 1811) Vexillum zelotypum (Reeve, 1845) Family Turridae Clavus lamberti (Montrouzier, 1860) Clavus pica (Reeve, 1843) Clavus unizonalis (Lamarck, 1822) Lophiotoma acuta (Perry, 1811) Lophiotoma indica (Röding, 1798) Lophiotoma sp. 1 Lophiotoma sp. 2 cf. Tritonoturris cumingii (Powys, 1835) Turridrupa bijubata (Reeve, 1843) Turridrupa cerithina (Anton, 1839) Turris crispa (Lamarck, 1816) Turris spectabilis (Reeve, 1843) Xenoturris cingulifera (Lamarck, 1822) Family Terebridae Duplicaria raphanula (Lamarck, 1822) Hastula albula (Menke, 1843) Hastula anomala (Gray, 1834) Hastula penicillata (Hinds, 1844) Hastula solida (Deshayes, 1857) Terebra afﬁnis Gray, 1834 Terebra amanda Hinds, 1844 Terebra areolata (Link, 1807) Terebra argus Hinds, 1844 Terebra babylonia Lamarck, 1822 Terebra cerithina Lamarck, 1822 Terebra crenulata (Linnaeus, 1758) Terebra dimidiata (Linnaeus, 1758) Terebra felina (Dillwyn, 1817) Terebra funiculata Hinds, 1844 Terebra guttata (Röding, 1798) Terebra insalli Bratcher & Burch, 1976 Terebra laevigata Gray, 1834 Site numbers 1 2, 28 2 29 9 6, 13, 17, 21 1, 2 1, 15, 19, 23 4 6, 18 4, 17, 22 0 28 28 1, 4, 5, 8, 13, 16, 18 17 29 0, 21 23 13 1, 17 15 13 21 9 8 30 11 4, 12 10, 28 30 13 2, 16 21, 25, 28 4, 22 11, 13 1, 2, 10, 11, 14, 23, 28, 30 10 4 0 12 29 0, 1, 3-5, 11-13, 15, 20, 23-26, 28-30 22, 27 24, 29 11, 15 0, 13, 23, 29 12 4, 18 5, 29 5, 12, 18, 19, 21, 25 26 4 13, 29 13 Coral Reefs of Northwest Madagascar 95 Appendix 3 Species Terebra maculata (Linnaeus, 1758) Terebra nebulosa Sowerby, 1825 Terebra cf. parkinsoni Cernohorsky & Bratcher, 1976 Terebra pertusa (Born, 1778) Terebra quoygaimardi Cernohorsky & Bratcher, 1976 Terebra subulata (Linnaeus, 1767) Terebra undulata Gray, 1834 Family Conidae Conus achatinus Gmelin, 1791 Conus arenatus Hwass in Bruguière, 1792 Conus auger Lightfoot, 1786 Conus boeticus Reeve, 1844 Conus canonicus Hwass in Bruguière, 1792 Conus capitaneus Linnaeus, 1758 Conus catus Hwass in Bruguière, 1792 Conus chaldeus (Röding, 1798) Conus consors Sowerby, 1833 Conus coronatus (Gmelin, 1791) Conus distans Hwass in Bruguière, 1792 Conus eburneus Hwass in Bruguière, 1792 Conus ebraeus Linnaeus, 1758 Conus episcopatus da Motta, 1982 Conus emaciatus Reeve, 1849 Conus ﬂavidus Lamarck, 1810 Conus ﬂoccatus Sowerby, 1839 Conus generalis Linnaeus, 1767 Conus litoglyphus Röding, 1798 Conus litteratus Linnaeus, 1758 Conus lividus Hwass in Bruguière, 1792 Conus magniﬁcus Reeve, 1843 Conus marmoreus Linnaeus, 1758 Conus miles Linnaeus, 1758 Conus miliaris Hwass in Bruguière, 1792 Conus musicus Hwass in Bruguière, 1792 Conus nussatella Linnaeus, 1758 Conus omaria Hwass in Bruguière, 1792 Conus pertusus Hwass in Bruguière, 1792 Conus planorbis Born, 1778 Conus quercinus Lightfoot, 1786 Conus rattus Hwass in Bruguière, 1792 Conus sanguinolentus Quoy & Gaimard, 1834 Conus scabriusculus Dillwyn, 1817 Conus spiculum Reeve, 1849 Conus sponsalis Hwass in Bruguière, 1792 Conus striatellus Link, 1807 Conus striatus Linnaeus, 1758 Conus suratensis Hwass in Bruguière, 1792 Conus terebra Born, 1778 Conus tessellatus Born, 1778 Conus textile Linnaeus, 1758 Conus tulipa Linnaeus, 1758 Conus varius Linnaeus, 1758 Conus vexillum Gmelin, 1791 Conus virgo Linnaeus, 1758 Conus vulpinus Hwass in Bruguière, 1792 Conus zelanicus Gmelin, 1791 Family Architectonicidae Architectonica sp. 96 Rapid Assessment Program Site numbers 12, 15, 19, 23, 25, 29, 30 0, 3, 4, 10, 11, 13, 15, 19, 21, 23 12 29 15, 22, 27, 29 24, 29 12, 21, 22, 27, 29, 30 20 1, 4, 5, 12, 13, 23-25, 28, 29 2, 6, 19 3 3, 5, 6, 8, 10, 13, 18B, 23-25, 28 20 0, 9, 18B, 20 21, 28 6, 23 4, 29 5 0, 18B, 20 21 25 0-2, 25, 29 18B 4, 8, 21, 25 3, 4, 8 2, 4-6, 10-12, 17, 25, 28 0, 1, 3, 5, 6, 8, 14, 18, 20, 21, 25, 26, 30 21, 26, 28 2, 3, 10, 25 0-7, 9, 12, 13, 18, 20, 21, 28, 30 1, 2, 6, 7, 18B, 20, 25, 28 0, 1, 6-8, 10, 11, 18 6 28 13 3, 16, 23, 27 7 0-2, 5, 6, 19, 20 0-2, 5, 11, 20, 21, 25 19 23 0, 1, 3, 6, 8, 10, 12, 13, 16, 17, 19, 21, 25, 29 4, 12, 13, 21, 28 15, 19, 21, 23, 25, 26 12 18 10, 12, 13, 15, 25, 28, 29 4, 6, 7, 10, 11, 17, 25 18 1, 14, 18B 13, 16 0, 2-4, 11-13, 21, 23, 28, 29 14, 16 5 21, 27 List of molluscs collected Species Architectonica perspectiva (Linnaeus, 1758) Heliacus variegatus (Gmelin, 1791) Philippia radiata (Röding, 1798) Family Pyramidellidae Otopleura auriscati (Holten, 1802) Pyramidella acus (Gmelin, 1791) Pyramidella teres (A. Adams, 1855) Pyramidella ventricosa (Guerin, 1831) Family Acteonidae Pupa sulcata (Gmelin, 1791) Family Cylichnidea Acteocina sp. Family Aglajidae Chelidonure electra Rudman, 1970 Chelidonura pallida Risbec, 1851 Philinopsis sp. Family Haminoeidae Atys cylindricus (Helbling, 1779) Atys naucum (Linnaeus, 1758) Family Bullidae Bulla vernicosa Gould, 1859 Family Elysiidae Elysia sp. Family Aplysiidae Dactylomela sp. Family Pleurobranchidae Berthellina citrina (Ruppell and Leuckart, 1828) Family Dorididae Discodoris fragilis (Alder and Hancock, 1864) Family Polyceridae Nembrotha kubyarana Bergh, 1877 Roboastra gracilis (Bergh, 1877) Family Chromodorididae Chromodoris africana (Eliot, 1904) Chromodoris annulata Eliot, 1904 Chromodoris cf. daphnae (Angas, 1864) Chromodoris sp. 1 Chromodoris sp. 2 Chromodoris sp. 3 Chromodoris cf. strigata Rudman, 1982 Glossodoris cincta (Bergh, 1888) Glossodoris pallida (Ruppell & Leuckart, 1828) Family Dendrodorididae Dendrodoris nigra (Stimpson, 1855) Family Phyllidiidae Phyllidia coelestis Bergh, 1905 Phyllidia elegans Bergh, 1869 Phyllidia ocellata (Cuvier, 1804) Phyllidiella aff. rosans (Bergh, 1873) Phyllidia varicosa Lamarck, 1801 Phyllidiella pustulosa (Cuvier, 1804) Phyllidia zeylanica (Kelaart, 1859) Phyllidiopsis striata (Bergh, 1889) Family Glaucidae cf. Pteraeolidia Pteraeolidia ianthina (Angas, 1864) Site numbers 27 6 21 25, 29 15 13 20, 21, 23 4, 13, 15, 23, 29 23 5-8, 16, 17, 19, 20, 24 22 12 0, 29 0, 10, 27B 18B, 21, 24 19, 29 2 1, 16 2 12 20 6, 7, 19 9 0, 8 29 9 18 6 22 29 10, 27 8, 15, 27 1, 6, 7, 17, 19, 20 8, 11, 13 15-17, 19, 26, 28, 30 9, 20, 22 0, 3, 7, 9, 14, 15, 25 6 21, 26 8, 9 10, 26, 29 Coral Reefs of Northwest Madagascar 97 Appendix 3 Species Family Onchidiidae Onchidium sp. Family Siphonariidae Siphonaria sp. 1 Siphonaria sp. 2 CLASS BIVALVIA Family Nuculidae Nucula sp. Family Mytilidae Lithophaga spp. Modiolus sp. Septifer bilocularis (Linnaeus, 1758) Septifer sp. Family Arcidae Anadara antiquata (Linnaeus, 1758) Anadara maculosa (Reeve, 1844) Anadara sp. Arca navicularis Bruguière, 1798 Arca ventricosa Lamarck, 1819 Barbatia amygdalumtotsum (Röding, 1798) Barbatia foliata Forskål, 1775 Barbatia sp. Family Glycymerididae Tucetona petunculus (Linnaeus, 1758) Family Pteriidae Pinctada margaritifera (Linnaeus, 1758) Pteria avicular (Holten, 1802) Pteria pengiun (Röding, 1798) Family Malleidae Vulsella vulsella (Linnaeus, 1758) Family Isognomonidae Isognomon isognomum (Linnaeus, 1758) Family Pinnidae Atrina pectinata Linnaeus, 1767 Atrina vexillum (Born, 1778) Pinna bicolor (Gmelin, 1791) Streptopinna saccata (Linnaeus, 1758) Family Limidae Ctenoides annulata (Lamarck, 1819) Lima lima (Link, 1807) Limaria orientalis (Adams & Reeve, 1850) Family Gryphaeidae Hyotissa hyotis (Linnaeus, 1758) Family Ostreidae Dendostrea sandvichensis (Sowerby, 1871) Lopha cristagalli (Linnaeus, 1758) Saccostrea cucullata (Born, 1778) Family Pectinidae Bractechlamys nodulifera (Sowerby, 1842) Bratechlamys vexillum (Reeve, 1843) Chlamys senatoria (Gmelin, 1791) Gloripallium pallium (Linnaeus, 1758) Haumea minuta Linnaeus, 1758 Laevichlamys lemniscata (Reeve, 1853) Laevichlamys limatula (Reeve, 1853) Laevichlamys sp. 1 Pallium lentiginosa (Reeve, 1865) 98 Rapid Assessment Program Site numbers 27B 10, 20 18B 0 0, 4, 5, 7-9, 11, 13, 14, 16, 18-23, 26, 28-30 10, 16, 18, 22-25, 27B 0-3, 13, 17, 20, 22, 24, 25, 27B 20, 21 13, 15, 22, 23 0, 1, 14, 15, 18B, 21-24, 26, 28 24 3, 5, 6, 9, 10, 12, 15, 18, 20, 21 0-9, 12-15, 17, 19, 20, 23, 26-30 1, 10, 12, 19, 20, 29, 30 13 1, 10-14, 21, 24-26, 28, 30 1, 3-5, 7, 14, 16, 18, 20-24, 26, 27, 30 0, 1, 3-5, 10, 11, 13, 16, 17, 27, 29, 30 30 27, 29 0, 1, 8, 9, 17, 20-22, 24-27, 29 3-6, 8, 30 0, 2, 3, 4-6, 13, 15-17, 22, 24, 26, 27, 30 11, 14, 20, 21, 24, 27 0, 1, 3, 5, 6, 8-10, 13, 17, 22, 30 16 4, 10, 12-14, 18, 19, 21, 25-27, 29, 30 5, 17, 20, 21, 25 0-9 5, 8, 9, 11, 13-17, 20-30 0-2, 5, 7-9, 20, 26, 27B 3, 10, 11, 16, 17, 21, 26, 28 3 22 2-4, 11-16, 19-23, 25, 27-30 14 3, 12, 13, 27, 29 6, 12, 13 2, 3, 14, 15, 21, 22, 24, 27 21 List of molluscs collected Species Pallium squamata (Gmelin, 1791) Mimachlamys punctata (Gmelin, 1791) Pectinid sp. 1 Pectinid sp. 2 Pectinid sp. 3 Pectinid sp. 4 Pectinid sp. 5 Pectinid sp. 6 Pectinid sp. 7 Pectinid sp. 8 Pectinid sp. 9 Pectum funebris (Reeve, 1853) Pedum spondyloidaeum (Gmelin, 1791) Semipallium luculentum (Reeve, 1853) Decatopecten striatus (Schumacher, 1817) Family Spondylidae Spondylus anacanthus Mawe, 1823 Spondylus barbatus Reeve, 1856 Spondylus butleri Reeve, 1856 Spondylus candidus (Lamarck, 1819) Spondylus imperialis Chenu, 1843 Spondylus multimuricatus Reeve, 1856 Spondylus nicobaricus Schreibers, 1793 Family Chamidae Chama brassica Reeve, 1846 Chama lazarus Linnaeus, 1758 Chama limbula (Lamarck, 1819) Chama paciﬁca (Broderip, 1834) Chama savigni Lamy, 1921 Family Lucinidae Divaricella ornata (Reeve, 1853) Family Fimbriidae Codakia punctata (Linnaeus, 1758) Codakia tigerina (Linnaeus, 1758) Family Carditidae Cardita variegata Bruguière, 1792 Family Cardiidae Acrosterigma alternatum (Sowerby, 1841) Acrosterigma mendanaense (Sowerby, 1896) Acrosterigma reeveanum (Dunker, 1822) Acrosterigma rugosa (Lamarck, 1819) Acrosterigma sp. Acrosterigma transcendens (Melvill & Standen, 1899) Acrosterigma unicolor (Sowerby, 1834) Fragum hemicardium (Linnaeus, 1758) Fragum retusum (Linnaeus, 1767) Lyrocardium lyratum (Sowerby, 1841) Vepricardium multispinosum (Sowerby, 1838) Trachycardium elongatum (Bruguière, 1789) Trachycardium enode (Sowerby, 1841) Trachycardium rubicundum (Reeve, 1844) Family Tridacnidae Tridacna maxima (Röding, 1798) Tridacna squamosa Lamarck, 1819 Family Mactridae Lutraria sp. Site numbers 21, 23 23 6, 20, 21, 23 22 3, 16, 20 3, 4, 16, 17 4 28 20 20 26 2 0-3, 6, 7, 16, 17, 20-23, 26, 28-30 22 10 8 10, 16, 21 18, 22, 28-30 7,30 5 10, 15, 16 18, 20, 21, 23 5, 22 7, 10, 11, 29 0-3, 5, 7, 14, 16, 20, 21, 24, 27 30 6, 8, 26, 29, 30 2, 21 0, 4, 29 0, 4, 18, 20, 25, 27B 25, 28 2, 3, 7, 10, 14-16, 18B, 20-24, 26-28, 30 2, 8, 11, 13, 16 10, 14 13 14, 27 0, 2, 5, 10, 15, 21 1-6, 8, 10-15, 17, 19-23, 25-30 12, 27, 29 14 10, 14, 20, 23 2 10 2, 3, 5, 7, 16, 17, 21 0 0, 2-7, 11, 16-20, 22-29 1-7, 12, 16-21, 23, 29 21, 24, 27, 30 Coral Reefs of Northwest Madagascar 99 Appendix 3 Species Spisula aspersa (Sowerby, 1825) Family Solenidae Solen sp. Family Tellinidae Tellina exculta Gould, 1850 Tellina gargadia Linnaeus, 1758 Tellina linguafelis Linnaeus, 1758 Tellina radians Deshayes, 1854 Tellina rastellum Hanley, 1844 Tellina rostrata Linnaeus, 1758 Tellina serricostata Tokunaga, 1906 Tellina staurella Lamarck, 1818 Tellina tongana Quoy and Gaimard, 1835 Tellina sp. Tellina verrucosa Hanley, 1844 Exotica virgulata (Hanley, 1844) Tellina virgata Linnaeus, 1758 Family Semelidae Semele casta A. Adams, 1853 Semele lamellosa (Sowerby, 1830) Family Psammobiidae Asaphis contraria (Deshayes, 1863) Asaphis violaceans (Forsskal, 1775) Gari inﬂata (Bertin, 1880) Gari maculosa (Lamarck, 1818) Gari pulcherrima (Deshayes, 1855) Gari squamosa (Lamarck, 1818) Family Solecurtidae Solecurtis sp. Family Donacidae Donax sp. Family Trapeziidae Trapezium bicarinatum (Schumacher, 1817) Family Veneridae Antigona chemnitzii (Hanley, 1844) Antigona clathrata (Deshayes, 1854) Antigona purpurea (Linnaeus, 1771) Antigona restriculata (Sowerby, 1853) Antigona reticulata (Linnaeus, 1758) Callista impar (Lamarck, 1818) Dosinia sp. 1 Dosinia sp. 2 Gafrarium pectinatum (Linnaeus, 1758) Gafrarium tumidum Röding, 1798 Globivenus capricorneus (Hedley, 1908) Globivenus toreuma (Gould, 1850) Lioconcha annettae Lamprell & Whitehead, 1990 Lioconcha castrensis (Linnaeus, 1758) Lioconcha fastigiata (Sowerby, 1851) Lioconcha polita (Röding, 1798) Pitar afﬁnis (Gmelin, 1791) Tapes literatus (Linnaeus, 1758) Tapes sulcarius (Lamarck, 1818) Timoclea marica (Linnaeus, 1758) Family Corbulidae Corbula cf tahitensis Lamarck, 1818 100 Rapid Assessment Program Site numbers 3, 8, 10, 14-16, 20, 24-26, 28 20 0 12, 15, 20, 28 15, 22-24 14 4 4 20 9, 11, 14, 23, 25 20 13-15, 20, 24, 29 23, 26 12 0, 14, 16, 19, 23, 24, 27 11 9, 13, 14, 16, 17, 21, 23, 28, 30 10 18B, 27B 27 0, 2-6, 8-12, 14, 16, 20-23, 25-28 0, 4, 16, 19, 23, 24, 27 6 15 20, 29 8, 11, 12, 14, 17, 26 3, 8-10, 13, 14, 20, 26 17, 20, 21, 23, 26 3, 10, 13-15, 21, 29 2, 3, 5, 11, 13, 14, 16, 21, 22, 24 1, 3, 12, 16, 17, 21, 24 15, 20, 24 11, 13, 14 4-6, 10, 11, 13-15, 20 13, 20, 27 20, 27B 3, 16, 25, 27 1-4, 6, 10-15, 17, 18B, 23, 26, 28, 30 10, 11, 13 4, 14, 28 4, 14 10, 11, 13, 15, 17, 21, 22, 24, 27 4, 6, 11, 16, 24 1-17, 20, 21, 23, 25, 27, 30 2 1, 4, 5, 11-14, 24, 26-28 30 List of molluscs collected Species CLASS CEPHALOPODA Family Sepiidae Sepia sp. Family Octopodidae Octopus sp. CLASS SCAPHOPODA Family Dentaliidae Dentalium sp. Site numbers 3, 18B, 21 8 3, 4, 10 Coral Reefs of Northwest Madagascar 101 Appendix 4 List of the reef ﬁshes of Madagascar The following list includes all reef species that have been recorded to date as determined by the RAP survey, collections at the United States National Museum of Natural History, and a search of pertinent literature beginning with Savage’s (1891) Histoire Naturelle des Poissons de Madagascar. The phylogenetic sequence of the families appearing in this list follows Eschmeyer (Catalog of Fishes, California Academy of Sciences, 1998) with slight modiﬁcation (e.g., placement of Mugilidae and Cirrhitidae). Genera and species are arranged alphabetically within each family. Site numbers, relative abundance, and depth distribution are provided for species that were recorded during the current survey. Species recorded previously from Madagascar but not seen during the RAP survey are indicated by “previously recorded” or “USNM collection” in the site records column. Terms relating to relative abundance are as follows: Abundant - Common at most sites in a variety of habitats with up to several hundred individuals being routinely observed on each dive. Common - seen at the majority of sites in numbers that are relatively high in relation to other members of a particular family, especially if a large family is involved. Moderately common - not necessarily seen on most dives, but may be relatively common when the correct habitat conditions are encountered. Occasional - infrequently sighted and usually in small numbers, but may be relatively common in a very limited habitat. Rare - less than 10, often only one or two individuals seen on all dives. Species marked with an asterisk (*) are new records for Madagascar based on a search of scientiﬁc literature, including the items listed in the reference section of Chapter 3. 102 Rapid Assessment Program List of the reef ﬁshes of Madagascar SPECIES STEGOSTOMATIDAE Stegostoma fasciatum (Hermann, 1783) GINGLYMOSTOMIDAE Nebrius ferrugineus (Lesson, 1830) CARCHARHINIDAE Carcharhinus albimarginatus (Rüppell, 1837) C. amblyrhynchos (Bleeker, 1856) C. limbatus (Valenciennes, 1839) C. melanopterus (Quoy & Gaimard, 1824) C. sorrah (Valenciennes, 1839) C. wheeleri Garrick, 1982 Galeocerdo cuvieri (Peron & LeSueur, 1822) Negaprion acutidens (Rüppell, 1837) Triaenodon obesus (Rüppell, 1835) SPHYRNIDAE Sphyrna lewini (Grifﬁth & Smith, 1834) TORPEDINIDAE Torpedo fuscomaculata Peters, 1855 RHINOBATIDAE Rhynchobatus djeddensis (Forsskål, 1775) DASYATIDIDAE Dasyatis kuhlii (Müller & Henle, 1841) D. pastinaca (Linnaeus, 1758) Taeniura lymma (Forsskål, 1775) T. meyeni (Müller & Henle, 1841) MYLIOBATIDAE Aetobatus narinari (Euphrasen, 1790) Manta birostris (Walbaum, 1792) MORINGUIDAE Moringua abbreviata (Bleeker, 1863) M. bicolor Kaup, 1856 M. javanica (Kaup, 1856) M. microchir Bleeker, 1853 MURAENIDAE Anarchias seychellensis Smith, 1962 Echidna nebulosa (Thünberg, 1789) E. polyzona (Richardson, 1845) E. unicolor Schultz, 1953 Enchelycore pardalis (Temminck & Schlegel, 1846) Gymnomuraena zebra (Shaw, 1797) Gymnothorax buroensis (Bleeker, 1857) G. chilospilus Bleeker, 1865 G. elegans Bliss, 1833 G. enigmaticus McCosker & Randall, 1982 G. eurostus (Abott, 1861) G. favagineus (Bloch & Schneider, 1801) G. ﬂavimarginatus (Rüppell, 1828) G. ﬁmbriatus (Bennett, 1831) G. fuscomaculatus (Schultz, 1953) G. javanicus (Bleeker, 1865)* G. johnsoni (Smith, 1962) G. margaritophorus (Bleeker, 1865) G. meleagris (Shaw & Nodder, 1795) G. melatremus Schultz, 1953* SITE RECORDS ABUNDANCE DEPTH (m) Rare, only one seen. 0-70 Rare, only one seen. 0-100 Rare, only 3 seen. 2-100 Rare, only one seen. 1-200 Previously recorded. 5 Previously recorded. 12 Previously recorded. Previously recorded. Previously recorded. Previously recorded. Previously recorded. Previously recorded. 4, 7, 10 22 29 6 Previously recorded. 4, 6, 20, 26, 29 12 Previously recorded. 1, 2 2-40 Rare, only one seen. 2-50 Occasional in sandy areas. Rare, only one seen. 2-30 1-400 Several seen at sites 1-2. 0-100 One collected with rotenone. 1-10 One collected with rotenone. 1-150 Rare, only 3 seen. 0.5-50 A single specimen collected with rotenone. 5-30 Previously recorded. Previously recorded. Previously recorded. Previously recorded. Previously recorded. 27b Previously recorded. Previously recorded. Previously recorded. Previously recorded. Previously recorded. Previously recorded. Previously recorded. Previously recorded. Previously recorded. Previously recorded. 27b Previously recorded. Previously recorded. 3, 13, 21 Previously recorded. Previously recorded. Previously recorded. 13 Coral Reefs of Northwest Madagascar 103 Appendix 4 SPECIES G. picta (Ahl, 1798) G. pindae Smith, 1962 G. richardsoni (Bleeker, 1852) G. rueppelliae (McClelland, 1845) G. undulatus (Lacepède, 1803) SITE RECORDS Previously recorded. Previously recorded. Previously recorded. Previously recorded. 18 G. zonipectus Seale, 1906* 12, 13 Rhinomuraena quaesita Garman, 1888* Scuticara tigrina (Lesson, 1828) Siderea grisea (Lacepède, 1803) Thrysoidea macrura (Bleeker, 1854) Uropterygius concolor Rüppell, 1838 U. marmoratus Lacepède, 1803 U. micropterus (Bleeker, 1852) OPHICHTHIDAE Brachysomophis crocodilinus (Bennett, 1833) Callechelys catostoma (Schneider & Forster, 1801) C. marmorata (Bleeker, 1853) Leiuranus semicinctus (Lay & Bennett, 1839) Myrichthys colubrinus (Boddaert, 1781) M. maculosus (Cuvier, 1817) Myrophis microchir (Bleeker, 1864) M. uropterus (Temminck & Schlegel, 1846) Ophichthus bonaparti (Kaup, 1856) Pisodonophis cancrivorus (Richardson, 1844) Phyllophichthus xenodontus Gosline, 1951 3 Previously recorded. Previously recorded. Previously recorded. Previously recorded. Scolecenchelys gymnotus Bleeker, 1864 6 S. laticaudata (Ogilby, 1897) S. xorae Smith, 1958 Xestochilus nebulosus (Smith, 1962) CONGRIDAE Conger cinereus Rüppell, 1828 Heteroconger haasi (Klausewitz & Eibl-Eibesfeldt, 1959) CLUPEIDAE Herklotsichthys quadrimaculatus (Rüppell, 1835) H. spilurus (Guichenot, 1863) Sardinella jussieui (Valenciennes, 1847) S. melanura (Cuvier, 1829) Previously recorded. Previously recorded. Previously recorded. ABUNDANCE Two specimens collected with rotenone. Rare, only 3 seen. DEPTH (m) 8-45 1-50 Previously recorded. Previously recorded. Previously recorded. Previously recorded. Previously recorded. Previously recorded. Previously recorded. Previously recorded. Previously recorded. Previously recorded. Previously recorded. Previously recorded. A single specimen collected with rotenone. 5-30 One colony of about 20 individuals observed. 3-45 20 Hundreds seen schooling near surface. 0-1 Plotosus lineatus (Thünberg, 1787) 7 One aggregation with about 50 individuals seen. 1-20 SYNODONTIDAE Saurida gracilis (Quoy & Gaimard, 1824) Synodus binotatus Schultz, 1953 5, 6, 9, 10, 19, 29 Previously recorded. Occasional on sand bottoms. 1-30 Spratelloides delicatulus (Bennett, 1832) 13 Previously recorded. Previously recorded. Previously recorded. PLOTOSIDAE 104 S. dermatogenys Fowler, 1912 1-7, 9-13, 15, 17, 1824, 26-30 S. jaculum Russell and Cressy, 1979 17, 26 S. variegatus (Lacepède, 1803) 1, 3, 6, 8, 16, 19, 21, 26, 29 Trachinocephalus myops (Forster, 1801) Previously recorded. Rapid Assessment Program Moderately common, solitary individuals or small groups usually seen resting on dead coral or rubble. Photographed. Rare, only 2 seen. Occasional, solitary individuals or pairs usually seen resting on live coral. 1-25 10-50 5-50 List of the reef ﬁshes of Madagascar SPECIES OPHIDIIDAE Brotula multbarbata Temminck & Schlegel, 1846 CARAPIDAE Carapus mourlani (Petit, 1934) Encheliophis gracilis (Bleeker, 1856) E. homei (Richardson, 1846) Onuxodon fowleri (Smith, 1955) BYTHITIDAE Brosmophyciops pautzkei Schultz, 1960 ABUNDANCE D E P T H (m) 6 One specimen collected with rotenone. ? Antennarius coccineus (Lesson, 1829) 13 One specimen collected with rotenone. 1-40 A. pictus (Shaw & Nodder, 1794) A. striatus (Shaw & Nodder, 1794) Antennatus tuberosus (Cuvier, 1817) Histrio histrio (Linnaeus, 1758) Previously recorded. Previously recorded. Previously recorded. Previously recorded. A. nummifer (Cuvier, 1817) 6 One specimen collected with rotenone. 1-15 GOBIESOCIDAE Lepadichthys coccinotaenia Regan, 1921 MUGILIDAE Previously recorded. Crenimugil crenilabis (Forsskål, 1775) 2, 27b One school of about 30 ﬁsh seen next to rocky shore. Collected. 0-4 Rare, only one seen. 0-4 Rare, only one seen. Occasional, sheltering in caves and under ledges. Rare, aggregation of 8 ﬁsh seen in 15 m. Common, usually in areas affected by silt. Photographed. Occasional, sheltering in reef crevices. Photographed. Several seen sheltering under rocky outcrops. Photographed. Rare, only about 5 seen. 3-30 Moderately common. Rare. Seen at only one site. 3-30 12-80 Rare, only 2 seen. Occasional, usually among branches of staghorn Acropora coral. 3-20 Ogilbia sp. 1 SITE RECORDS Previously recorded. Previously recorded. Previously recorded. Previously recorded. Previously recorded. ANTENNARIIDAE ATHERINIDAE Atherinomorus lacunosus (Forster, 1801) Hypoatherina barnesi Schultz, 1953 BELONIDAE Tylosurus crocodilus (Lesueur, 1821) HEMIRAMPHIDAE Hemiramphus far (Forsskål, 1775) Hyporhamphus afﬁnis (Günther, 1866) H. dussumieri (Valenciennes, 1847) HOLOCENTRIDAE Myripristis adusta Bleeker, 1853 Previously recorded. Previously recorded. 29 Previously recorded. USNM collection. Previously recorded. 26 M. berndti Jordan & Evermann, 1902 3, 18, 28 M. botche Cuvier, 1829* 10 M. hexagona (Lacepède, 1802) 1, 3-11, 21, 22, 27-30 M. kuntee Cuvier, 1831 3, 5, 10-13, 28 M. murdjan (Forsskål, 1775) 13 M. pralinia Cuvier, 1829 M. seychellensis Cuvier, 1829 M. violacea Bleeker, 1851 M. vittata Valenciennes, 1831* Neoniphon aurolineatus (Liénard, 1839) N. opercularis (Valenciennes, 1831) 12, 28 Previously recorded. 6 11 Previously recorded. 18 N. sammara (Forsskål, 1775) 11, 13, 18, 20-22, 24 Plectrypops lima (Valenciennes, 1831) Previously recorded. Sargocentron caudimaculatum (Rüppell, 1835) 3, 6, 11-13, 18, 26, 28 Occasional, sheltering in reef crevices. 8-55 12-240 10-40 3-30 3-40 3-40 2-50 6-45 Coral Reefs of Northwest Madagascar 105 Appendix 4 SPECIES SITE RECORDS S. diadema (Lacepède, 1802) 10-13, 15, 17, 28 S. melanospilos (Bleeker, 1858)* S. praslin (Lacepède, 1802) S. punctatissimum (Cuvier, 1829) S. seychellense (Smith & Smith, 1963) 5, 7, 9, 10, 18, 22, 27 Previously recorded. Previously recorded. Previously recorded. 1, 3-7, 9-13, 16-18, 22, 26, 27, 29 6 S. spiniferum (Forsskål, 1775) S. violaceum (Bleeker, 1853) PEGASIDAE Eurypegasus draconis (Linnaeus, 1758) Moderately common, in caves and under ledges. Rare, only one seen. D E P T H (m) 2-30 10-25 5-122 3-30 Previously recorded. Pegasus volitans Linnaeus, 1758* 8 Only one seen, but relatively cryptic. 2-20 AULOSTOMIDAE Aulostomus chinensis (Linnaeus, 1766) FISTULARIIDAE 11,12 Rare, only 2 seen. 2-122 Fistularia commersoni Rüppell, 1835 10, 26, 29 Rare, except moderately common at site 29. 2-128 One specimen collected. 0-4 Rare, but mainly nocturnal. 1-50 Rare, but mainly nocturnal. Occasional, usually around rocky outcrops with abundant small ﬁshes. 1-15 CENTRISCIDAE Aeoliscus punctulatus (Bianconi, 1855) SOLENOSTOMIDAE Solenostomus cyanopterus Bleeker, 1854 SYNGNATHIDAE Acentronura tentaculata Günther, 1870 Choeroichthys sculptus (Günther, 1870) C. smithi Dawson, 1976 Corythoichthys ﬂavofasciatus (Rüppell, 1838) C. haematopterus (Bleeker, 1851) Cosmocampus dorrosanus (Dawson & Randall, 1975) Hippichthys cyanospilus (Bleeker, 1854) Hippocampus cameleopardalis Bianconi, 1855 H. kuda Bleeker, 1852 Micrognathus andersonii (Bleeker, 1858) Penetopteryx taeniocephalus Lunel, 1881 Phoxocampus belcheri (Kaup, 1856) Syngnathoides biaculaeatus (Bloch, 1785) Trachyrhamphus bicoarctatus (Bleeker, 1857) SCORPAENIDAE Dendrochirus brachypterus (Cuvier, 1829) Parascorpaena mcadamsi (Fowler, 1938) P. mossambica (Peters, 1855) Pterois antennata (Bloch, 1787) P. mombasae (Smith, 1957) P. radiata Cuvier, 1829 106 ABUNDANCE Occasional, in reef crevices or among staghorn Acropora coral. Occasional, in reef crevices. Previously recorded. Previously recorded. Previously recorded. Previously recorded. Previously recorded. Previously recorded. Previously recorded. Previously recorded. Previously recorded. Previously recorded. 29 Previously recorded. Previously recorded. Previously recorded. Previously recorded. Previously recorded. Previously recorded. Previously recorded. Previously recorded. 4 Previously recorded. 2 P. miles (Bennett, 1828) 9, 15, 17, 18, 21, 23, 24 Rhinopias frondosa (Günther, 1891) Scorpaenodes alabaiensis (Evermann & Seale, 1907)* S. guamensis (Quoy & Gaimard, 1824) S. hirsutus (Smith, 1957) S. parvipinnis (Garrett, 1863) S. varipinnis Smith, 1957 Scorpaenopsis diabolus (Cuvier, 1829) S. gibbosa (Bloch & Schnieder, 1801) Previously recorded. Rapid Assessment Program 6 Previously recorded. Previously recorded. Previously recorded. Previously recorded. Previously recorded. Previously recorded. One specimen collected with rotenone. 2-50 0-20 List of the reef ﬁshes of Madagascar SPECIES S. possi Randall & Eschmeyer, 2001 Sebastapistes cyanostigma (Bleeker, 1856) S. mauritiana (Cuvier, 1829) S. strongia (Cuvier, 1829) Taenianotus triacanthus Lacepède, 1802 TETRAROGIDAE Ablabys binotatus (Peters, 1855) SYNANCEIIDAE Inimicus ﬁlamentousus (Cuvier, 1829) Synanceja verrucosa (Bloch & Schneider, 1801) DACTYLOPTERIDAE Dactyloptena orientalis (Cuvier, 1829) PLATYCEPHALIDAE Cociella crocodila (Tilesius, 1812) SITE RECORDS Previously recorded. Previously recorded. Previously recorded. Previously recorded. Previously recorded. Papilloculiceps longiceps (Ehrenberg, 1829) 26 Thysanophrys otaitensis (Parkinson, 1829) CARACANTHIDAE Caracanthus madagascariensis Guichenot, 1869 C. unipinna (Gray, 1831) SERRANIDAE Aporops bilinearis Schultz, 1943 Aethaloperca rogaa (Forsskål, 1775) Anyperodon leucogrammicus (Valenciennes, 1828) Previously recorded. 1-15 Rare, only 4 seen. Rare, only 4 seen. 1-55 5-50 Occasional. 1-40 Previously recorded. Previously recorded. Previously recorded. Previously recorded. C. sexmaculata Rüppell, 1828 11 C. spiloparaea (Valenciennes, 1828)* 12 C. sonnerati (Valenciennes, 1828) Dermatolepis striolatus (Playfair, 1867) Epinephelus andersoni Boulenger, 1903 Previously recorded. Previously recorded. Previously recorded. E. areolatus (Forsskål, 1775) 14, 23, 27 E. fasciatus (Forsskål, 1775) 2, 3, 7, 9, 13, 18, 24, 26, 27b 28 E. ﬂavocaeruleus (Lacepède, 1802) 21, 22, 24 E. fuscoguttatus (Forsskål, 1775) 29 E. hexagonatus (Forster, 1801) E. lanceolatus (Bloch, 1790) E. longispinis (Kner, 1865) E. macrospilus (Bleeker, 1855) E. malabaricus (Bloch & Schneider, 1801) E. melanostigma Schultz, 1953 E. merra Bloch, 1793 E. multinotatus (Peters, 1876) E. rivulatus (Valenciennes, 1830) E. tauvina (Forsskål, 1775) Grammistes sexlineatus (Thünberg, 1792) 18 Previously recorded. 20 Previously recorded. Previously recorded. 18 20 Previously recorded. Previously recorded. Previously recorded. 2, 3 E. caeruleopunctatus (Bloch, 1790) Only one seen, but difﬁcult to detect. Previously recorded. Previously recorded. C. miniata (Forsskål, 1775) C. nigripinnis (Valenciennes,1828) C. boenack (Bloch, 1790) D E P T H (m) Previously recorded. Previously recorded. 2, 9, 23, 24 1, 12, 22, 30 1, 2, 5-7, 9, 13, 26, 27, 29 1-11, 14-18, 20-24, 26-30 11, 13, 21, 26 10, 12, 13 Cephalopholis argus Bloch & Schneider, 1801 ABUNDANCE Common, most abundant serranid species. Rare, only 4 seen. Rare, about 5 seen. Rare, only one seen on steep dropoff. Rare, but its relatively deep habitat not adequately surveyed. Rare, only 3 seen, the largest about 40 cm TL. Occasional, about 5 tiny juveniles seen at site 27b. Rare, only 3 seen. Rare, only 2 seen including one adult and 2 small juveniles. Rare, 2 large adults (about 100 cm TL) seen. Rare, only one seen. 1-20 3-150 1-40 6-140 16-108 6-200 5-25 4-160 2-80 1-60 1-15 Rare, only one juvenile seen. 5-30 Rare, only one seen. Rare, only one seen. 1-10 1-15 Rare, only 2 seen. 0.5-30 Coral Reefs of Northwest Madagascar 107 Appendix 4 SPECIES SITE RECORDS Liopropoma africanum (Smith, 1954)* 12 Liopropoma susumi (Jordan & Seale, 1906) Nemanthias carberryi Smith, 1954* Previously recorded. USNM collection. Plectropomus laevis (Lacepède, 1802) 5, 10, 12, 15, 17, 22, 26 P. punctatus Quoy & Gaimard, 1824 1, 3-5, 8, 11, 13-19, 21, 23, 26, 27, 29 Pseudanthias bimaculatus (Smith, 1955)* 10, 11, 13 P. cooperi (Regan, 1902) Previously recorded. P. evansi (Smith, 1954)* 12, 13 P. pulcherrimus (Heemstra & Randall, 1986)* 11 P. squamipinnis (Peters, 1855) 3, 5, 8-11, 13, 18, 19, 21, 22, 26, 28 Pseudogramma polyacanthum (Bleeker, 1856) 12 Variola albimarginata Baissac, 1953 V. louti (Forsskål, 1775) CIRRHITIDAE Amblycirrhitus bimacula (Jenkins, 1903) 12, 28 13 1, 3, 5, 8, 9, 11, 13, 1517, 22, 30 Cirrhitops fasciatus (Bennett, 1828) Cirrhitus pinnulatus (Schneider, 1801) Cristacirrhitus punctatus (Cuvier, 1829) Cyprinocirrhites polyactis (Bleeker, 1875) Previously recorded. Previously recorded. Previously recorded. Previously recorded. Oxycirrhitus typus Bleeker, 1857* 10 Paracirrhites arcatus (Cuvier, 1829)* P. forsteri (Schneider, 1801) PSEUDOCHROMIDAE Anisochromis kenyae Smith, 1954 11-13 3, 11-13, 25, 28 Previously recorded. Chlidichthys abruptus Lubbock, 1977* 6, 13, 29 Chlidichthys sp. (Gill, pers. comm.) Halimuraena lepopareia Winterbottom, 1980 Halimuraenoides isostigma Maugé & Bardach, 1985 Haliophis guttatus (Forsskål, 1775) Pseudochromis sp. 1 P. sp. 2 Previously recorded. Previously recorded. PLESIOPIDAE Plesiops coeruleolineatus Rüppell, 1835 P. malalaxus Mooi, 1995 P. mystaxus Mooi, 1995 TERAPONTIDAE Terapon jarbua (Forsskål, 1775) T. theraps (Cuvier, 1829) 108 Rapid Assessment Program Occasional. One very large adult (about 150 cm TL) seen at site 29. Moderately common. Most abundant large grouper. Occasional, except common on steep outer slopes below 15 m. Rare, except 2 aggregations seen at sites 12 and 13. Rare, except locally common at one site adjacent to steep dropoff. Moderately common. One specimen collected with rotenone. Rare, only 2 seen. Rare, only one seen. Previously recorded. Cirrhitichthys oxycephalus (Bleeker, 1855) Pseudochromis tauberae Lubbock, 1977 ABUNDANCE One specimen collected with rotenone in 25 m. D E P T H (m) 8-48 4-90 5-65 4-40 20-54 20-50 4-20 1-15 12-90 4-150 5-20 Occasional around live coral outcrops. The most abundant hawkﬁsh. Rare, only one seen in black coral on steep dropoff in 25 m. Rare, only about 5 seen. Occasional. 1-40 10-100 1-35 1-35 Several specimens collected with rotenone. 5-20 Moderately common, but relatively cryptic. Collected and photographed. 2-15 Previously recorded. Previously recorded. Previously recorded. Previously recorded. 3-8, 14, 17, 19, 20, 22, 24, 29 Previously recorded. Previously recorded. Previously recorded. Previously recorded. Previously recorded. List of the reef ﬁshes of Madagascar SPECIES KUHLIIDAE Kuhlia mugil (Forster, 1801) PRIACANTHIDAE Heteropriacanthus cruentatus (Lacepède, 1801) Priacanthus hamrur (Forsskål, 1775) APOGONIDAE Apogon abrogramma (Fraser & Lachner, 1985) A. angustatus (Smith & Radcliffe, 1911) A. apogonides (Bleeker, 1856) Previously recorded. 1, 3, 4, 9, 16-18, 26, 29 A. aureus (Lacepède, 1802) 1, 10, 14, 20-22, 26, 29 A. coccineus Rüppell, 1838 A. cooki Macleay, 1881 Previously recorded. USNM collection. A. cyanosoma (Bleeker, 1853) 4, 5, 15, 18 A. ﬂeurieu (Lacepède, 1802)* 15, 17 A. fraenatus Valenciennes, 1832 5, 6, 10, 14, 15, 17, 2022, 26, 29 A. fragilis Smith, 1961 15, 20, 22 A. guamensis Valenciennes, 1832 Occasional, but nocturnal. 3-35 A. lateralis Valenciennes, 1832 USNM collection. 1, 3-7, 11, 13, 14, 21, 26, 29 Previously recorded. A. leptacanthus Bleeker, 1856 20, 22, 24 Rarely encountered, but locally common. 1-12 A. pharaonis Bellotti, 1874 A. quadrifasciatus Cuvier, 1828 A. savayensis Günther, 1871 Previously recorded. Previously recorded. Previously recorded. A. semiornatus Peters, 1876 10 One specimen collected with rotenone. 8-30 A. talboti Smith, 1961 Previously recorded. A. kallopterus Bleeker, 1856 SITE RECORDS 13, 18, 22, 26 1, 3, 8-12, 18, 26 26 27b A. thermalis Cuvier, 1829* 22, 24 A. timorensis Bleeker, 1854 Previously recorded. Archamia fucata (Cantor, 1850) 1, 3-5, 14, 15, 17, 18, 20-24, 26, 29 Apogonichthys ocellatus (Weber, 1913) A. perdix Bleeker, 1854 Previously recorded. Previously recorded. Cheilodipterus artus Smith, 1961* 13, 16, 20-24 Foa madagascariensis Petit, 1931 1, 3-6, 8, 11-13, 21, 26-29 1, 2, 4-8, 11, 14-24, 26-30 Previously recorded. Fowleria aurita (Valenciennes, 1831) 13 F. variegata (Valenciennes, 1832) Pseudamia gelatinosa Smith, 1955 Pseudamiops pellucidus Smith, 1961 Previously recorded. Previously recorded. Previously recorded. C. quinquelineatus Cuvier, 1828 D E P T H (m) Moderately common. 5-80 Occasional. Rare, about 10 seen at one site. Occasional, forming aggregations with other apogonids. 5-30 3-30 Previously recorded. A. taeniophorus Regan, 1908 C. macrodon Lacepède, 1801* ABUNDANCE Occasional, but common at site 15. Rare, only 2 small aggregations seen. Occasional, but locally common under ledges and in coral crevices. Rarely encountered, but locally abundant. Rare, but occurs in very shallow water and is nocturnal and therefore difﬁcult to accurately survey. Rarely encountered, but locally common. Moderately common, but abundant at sites 3, 4, and 22 around large rocky outcrops. Occasional, often among branching corals. Occasional, in coral crevices or among boulders. Photographed. Common, most frequently seen member of family. One specimen collected with rotenone in 24 m. 10-30 3-35 5-30 3-35 1-15 0.5-2 0-10 3-60 2-20 4-30 1-40 0-15 Coral Reefs of Northwest Madagascar 109 Appendix 4 SPECIES SITE RECORDS Rhabdamia cypselurus Weber, 1909* 1, 4, 18, 21, 26 R. gracilis (Bleeker, 1856) 1, 4, 10, 13, 14, 23 Siphamia mossambica Smith, 1955 24 SILLAGINIDAE Sillago sihama (Forsskål, 1775) MALACANTHIDAE Malacanthus brevirostris Guichenot, 1848 M. latovittatus Lacepède, 1801 ECHENEIDAE 110 D E P T H (m) 2-15 5-20 5-30 Previously recorded. Previously recorded. Previously recorded. Echeneis naucrates Linnaeus, 1758 1, 29 CARANGIDAE Alectis indicus (Rüppell, 1830) Previously recorded. Atule mate (Cuvier, 1833) 20, 22, 28, 30 Carangoides armatus (Rüppell, 1830) C. caeruleopinnatus (Rüppell, 1830) C. chrysophrys (Cuvier, 1833) C. ferdau (Forsskål, 1775) C. fulvoguttatus (Forsskål, 1775) C. malabaricus (Bloch & Schneider, 1801) C. oblongus (Cuvier, 1833) Caranx heberi (Bennett, 1828) C. ignobilis (Forsskål, 1775) Previously recorded. Previously recorded. 27 2 Previously recorded. Previously recorded. Previously recorded. 1, 9 Previously recorded. 1, 10, 11, 13, 16, 18, 23, 26, 28, 29 C. melampygus Cuvier, 1833 ABUNDANCE Occasional, but sometimes in large numbers swarming around rocky outcrops. Occasional, forming large aggregations around coral heads. Photographed. About 100 ﬁsh seen with Didadema urchins in 12 m; several collected. Rare, only 4 seen attached to large ﬁshes. 0-30 Occasional schools seen, including large ones at sites 28 and 30. 5-50 Rare, only one seen. Rare, only 5 seen. 5-90 5-60 Rare, only two seen. 15-60 Occasional, usually seen solitary or in small schools. Rare, but schools containing up to about 20 ﬁsh seen. 1-190 C. papuensis Alleyne & Macleay, 1877 9, 18, 21 C. sexfasciatus Quoy & Gaimard, 1824 C. tille Cuvier, 1833 Elegatis bipinnulata (Quoy & Gaimard, 1824) Previously recorded. 26 Previously recorded. Rare, only one seen. 5-50 Gnathanodon speciosus (Forsskål, 1775) 9, 27 Rare, about 10 small juveniles seen. 1-30 Scomberoides commersonnianus (Lacepède, 1802) S. lysan (Forsskål, 1775) Previously recorded. Previously recorded. S. tol (Cuvier, 1832) 1, 29 Selar crumenophthalmus (Bloch, 1793) 24 Seriolina nigrofasciata (Rüppell, 1829) Trachinotus bailloni (Lacepède, 1801) T. blochii (Lacepède, 1801) LEIOGNATHIDAE Previously recorded. Previously recorded. Previously recorded. Gazza minuta (Bloch, 1797) 22 LUTJANIDAE Aphareus furca (Lacepède, 1802) A. rutilans Cuvier, 1830 Aprion virescens Valenciennes, 1830 Previously recorded. Previously recorded. 3, 9, 12, 13, 29 Lutjanus argentimaculatus (Forsskål, 1775) 23, 29 Rapid Assessment Program Rare, except 2 schools containing 1-30 ﬁsh seen. Rare, except one school of about 50 encountered in 12 m. About 30 ﬁsh seen among large school of apogonids. One collected. Rare, only 5 seen. Rare, but locally common at site 29. Largest seen about 100 cm TL. 1-50 0-10 5-160 2-20 3-40 1-100 List of the reef ﬁshes of Madagascar SPECIES L.bohar (Forsskål, 1775) SITE RECORDS 1-4, 6, 7, 9-19, 21-24, 26, 27, 29 L. ehrenburgi (Peters, 1869)* 8, 29 L. fulviﬂamma (Forsskål, 1775) 2-6, 8, 11, 12, 15-20, 22-24, 26, 27, 29 L. fulvus (Schneider, 1801) L. kasmira (Forsskål, 1775) 13, 18, 29 3, 11-13, 16, 18, 22-24, 26, 27 10, 15 L. lutjanus Bloch, 1790 1, 3, 8, 22, 24, 29 L. gibbus (Forsskål, 1775) L. monostigma (Cuvier, 1828) L. notatus (Cuvier, 1828) L. rivulatus (Cuvier, 1828) L. russelli (Bleeker, 1849) L. sebae (Cuvier, 1816) Macolor niger (Forsskål, 1775) 4, 5, 7-13, 16, 17, 27, 29 6, 12, 16, 24 1, 6, 11, 21, 29 Previously recorded. Previously recorded. 1-3, 5-7, 10-13, 17-24, 26-30 ABUNDANCE Common, especially abundant at site 14. Rare, 2 small schools encountered. Photographed. Common, especially at site 26. Occurs solitarily or in aggregations. Rare, only 5 seen. Occasional, but large aggregation seen at site 22. Rare, only 6 seen. Occasional, but abundant at sites 8 and 29, occurring in large aggregations. Occasional, usually occurring as solitary individuals. Rare, less than 10 seen. Rare, 5 large adults encountered. D E P T H (m) 4-180 1-20 1-35 2-40 6-40 3-265 5-90 5-60 10-40 2-100 Moderately common, but always in low numbers. 3-50 Abundant in variety of habitats. 1-30 Common. 1-35 Common. 1-40 CAESIONIDAE Caesio caerulaurea Lacepède, 1802 C. lunaris Cuvier, 1830 C. xanthonota Bleeker, 1853 1-3, 5-14, 16-18, 2124, 26-30 1-3, 5, 11, 13, 17-19, 22-24, 26-29 1-4, 10-14, 17, 21, 23, 24, 26-28 C. sp. 1* 1, 3, 9, 18, 21, 27 C. sp. 2* 9, 13, 17, 21, 22 1, 6-10, 13, 18, 21, 22, 27, 30 Pterocaesio pisang (Bleeker, 1853)* P. sp.* 9, 13, 16, 17, 21, 22, 24 P. tile (Cuvier, 1830) GERREIDAE Gerres acinaces Bleeker, 1854 G. oblongus Cuvier, 1830 G. oyena (Forsskål, 1775) G. rappi (Barnard, 1927) HAEMULIDAE 27 Diagramma pictum (Thünberg, 1792) Plectorhinchus ﬂavomaculatus P. gaterinus (Forsskål, 1775) Plectorhinchus gibbosus (Lacepède, 1802) P. paulayi Steindachner, 1895 P. plagiodesmus (Fowler, 1935) P. playfairi (Pellegrin, 1914) P. schotaf (Forsskål, 1775) P. sordidus (Klunzinger, 1870) Occasional, usually in mixed schools with C. caerulaurea. An undescribed species. Locally common. Moderately common, sometimes occurring in large schools. Occasional in mixed schools with C. caerulaurea and P. pisang. Possibly an undescribed species. Rare, only one school seen. 1-25 1-25 1-35 1-25 1-60 Previously recorded. Previously recorded. Previously recorded. Previously recorded. 2-4, 7, 9, 15, 17, 20, 21, 29 1, 3, 6, 7, 9, 23, 27 1, 3-7, 11, 16, 18, 2124, 26, 27, 29 Previously recorded. Previously recorded. Previously recorded. Previously recorded. Previously recorded. Previously recorded. Occasional, in silty areas; common at site 29. Occasional. Occasional, except common at site 22. 2-40 3-25 3-30 Coral Reefs of Northwest Madagascar 111 Appendix 4 SPECIES SPARIDAE Acanthopagrus bifasciatus (Forsskål, 1775) Cheimerius nufur (Valenciennes, 1830) LETHRINIDAE Gnathodentex aurolineatus Lacepède, 1802 Previously recorded. Previously recorded. Gymnocranius elongatus 18 G. grandoculus (Valenciennes, 1830) 2, 15 Lethrinus borbonicus Valenciennes, 1830 Previously recorded. l. conchyliatus (Smith, 1959) 10-12 L. enigmaticus Smith, 1959* 12 L. erythracanthus Valenciennes, 1830 22, 28 L. harak (Forsskål, 1775) 2, 4-6, 15, 20, 21, 24, 26, 27, 30 L. lentjan (Lacepède, 1802) 15, 16,18, 19, 29 L. mahsena (Forsskål, 1775) 1-4, 6-13, 20-29 L. microdon Valenciennes, 1830 L. nebulosus (Forsskål, 1775) 18 Previously recorded. L. obsoletus (Forsskål, 1775) 1, 5, 6, 16, 18, 22, 29 L. olivaceous Valenciennes, 1830 L. rubrioperculatus Sato, 1978 L. variegatus Valenciennes, 1830 L. xanthocheilus Klunzinger, 1870 2, 4, 26 Previously recorded. 3, 20, 24 3 4, 5, 7, 9, 10, 12, 13, 15-18, 22-24, 28, 29 Monotaxis grandoculis (Forsskål, 1775) SITE RECORDS 20 ABUNDANCE D E P T H (m) Rare, only 3 seen. Rare, only 2 seen on sandy bottom in 18 m. Rare, only 3 seen on sandy bottom in 1 m. 1-30 Rare, except locally common at sites 11 and 12. Rare, less than 10 seen. Rare, only one juvenile and one adult seen. Moderately common, mainly on sheltered reefs near shore. Occasional, but locally common, especially at site 29. Common and most abundant lethrinid seen. Rare, only one seen. Occasional, and always in low numbers. Most common at site 29. Rare, only 5 seen. 5-40 20-100 5-200 5-40 10-120 1-20 3-50 2-100 5-80 1-25 4-185 Rare, but seagrass is main habitat. Rare, only one seen. 1-10 2-25 Moderately common. 1-100 NEMIPTERIDAE Scolopsis bimaculatus Rüppell, 1828 S. gahnam (Forsskål, 1775) S. vosmeri (Bloch, 1792) 3, 18, 23, 26, 27, 29 1-11, 14-16, 18-24, 26-30 1-3, 5, 6, 8, 9, 14, 17, 21, 23, 24, 26, 27, 29, 30 Occasional, but locally common in sandy areas. Common on sand-rubble bottoms. Photographed. Common on sand-rubble bottoms. Photographed. 3-60 2-30 2-30 MULLIDAE Mulloidichthys ﬂavolineatus (Lacepède, 1802) 22 Parupeneus barberinus (Lacepède, 1801) P. bifasciatus (Lacepède, 1801) 5, 7-27, 29, 30 8, 9, 14, 18, 23 2, 5, 10, 11, 13, 14, 16, 18, 19, 23, 26, 30 10, 18 Previously recorded. 1-13, 15-19, 22, 23, 25, 26, 30 10, 25 3, 4, 30 P. cyclostomus (Lacepède, 1802) P. heptacanthus (Lacepède, 1801) P. indicus (Shaw, 1803) P. macronema (Lacepède, 1801) P. pleurostigma (Bennett, 1830) P. rubescens (Lacepède, 1801) PEMPHERIDAE 112 Parapriacanthus ransonneti Steindachner, 1870 3, 4, 21 Pempheris oualensis Cuvier, 1831 1, 18 Rapid Assessment Program Rare, one small aggregation of 6 ﬁsh seen. Common. Rare, less than 10 seen. 1-40 1-100 1-80 Occasional individuals seen. 2-92 Rare, less than 10 seen. 1-60 Common. 3-30 Rare, only 2 seen. Rare, only 3 seen. 5-46 1-30 Occasional, forming large aggregations in caves and crevices. Occasional, forming large aggregations in caves and crevices. 5-30 5-30 List of the reef ﬁshes of Madagascar SPECIES SITE RECORDS P. vanicolensis Cuvier, 1831 4-6, 8, 9, 18 ABUNDANCE Occasional, forming large aggregations in caves and crevices. D E P T H (m) 3-38 KYPHOSIDAE Kyphosus cinerascens (Forsskål, 1775) 13, 26 K. vaigiensis (Quoy & Gaimard, 1825) MONODACTYLIDAE Previously recorded. Monodactylus argenteus (Linnaeus, 1758) 20 Locally common, but only seen at 2 sites. 1-24 Large aggregation of about 100 ﬁsh seen. 0-5 Common, most abundant chaetodontids. 1-30 Occasional. 5-30 Occasional. Occasional. Occasional. 1-15 1-25 2-45 Occasional. 6-60 Occasional. Moderately common, but always in low numbers at each site. Rare, only seen below about 16 m. 2-170 Moderately common. 2-15 Rare, about 6 seen. Moderately common in areas of tabular Acropora. Common and most abundant species of butterﬂyﬁsh. Rare, about 5 pairs seen. Occasional in rich coral areas. Rare, less than 10 seen. Locally common, but rarely seen. Rare, except about 50 seen at sites 12 and 13. Common, one of 3 most abundant chaetodontids. Often in aggregations. Rare, only 5 seen. 5-25 Occasional. Occasional, and usually in low numbers. Rare, less than 10 seen. Common and most abundant angelﬁsh. Occasional. Moderately common, but always in low numbers. 10-50 CHAETODONTIDAE Chaetodon auriga Forsskål, 1775 C. bennetti Cuvier, 1831 Chaetodon blackburni Desjardins, 1836 C. dolosus Ahl, 1923 C. falcula Bloch, 1793 C. guttatissimus Bennett, 1823 C. interruptus Ahl, 1923 C. kleinii Bloch, 1790 C. lineolatus Cuvier, 1831 C. lunula Lacepède, 1803 C. madagaskariensis Ahl, 1923 C. melannotus Schneider, 1801 C. meyeri Schneider, 1801 C. trifascialis Quoy & Gaimard, 1824 1-9, 11-19, 21-24, 26-30 3, 5-7, 12, 14-17, 19, 22, 27 Previously recorded. Previously recorded. 7, 11-13, 20-24, 27, 29 3, 8-17, 22-24, 27, 29 2, 3, 6, 11-13, 23, 26 1-3, 5-13, 15, 16, 18, 26, 28, 29 1, 3, 5-7, 13, 15, 17, 22 4, 7-9, 11-15, 17-24, 26-29 10-13 3, 4, 6-8, 10, 12-18, 22, 26, 28 11-13, 17 1-9, 11-16, 18-21, 23, 26, 27, 28, 29, 30 C. trifasciatus Park, 1797 1-8, 11-27, 29, 30 C. vagabundus Linnaeus, 1758 C. xanthocephalus Bennett, 1832 C. zanzibarensis Playfair, 1867 Forcipiger ﬂavissimus Jordan & McGregor, 1898 3, 5, 9, 23, 26 11-13, 15, 19, 22, 24 2, 7, 11, 12, 19, 23 11, 12, 13 Hemitaurichthys zoster (Bennett, 1831) 12, 13 Heniochus acuminatus (Linnaeus, 1758) 1-10, 12-30 H. monoceros Cuvier, 1831 POMACANTHIDAE Apolemichthys trimaculatus (Lacepède, 1831) 9, 23, 27 Centropyge acanthops (Norman, 1922) 8, 10, 11, 12, 21, 28 C. bispinosa (Günther, 1860) 12, 13 3-7, 9-16, 18, 19, 22, 26-28 3, 7-9, 12, 13, 17, 28 1-3, 5, 7-9, 11-14, 21, 23, 26-28 Previously recorded. C. multispinis (Playfair, 1867) Pomacanthus chrysurus Cuvier, 1831 P. imperator (Bloch, 1787) P. rhomboides (Gilchrist & Thompson, 1908) P. semicirculatus Cuvier, 1831 8, 11-13, 26, 28 1-3, 6-9, 11-13, 16-18, 23, 24, 26-29 Common, and in much greater numbers compared to most locations in the Indo-Paciﬁc. 1-40 10-60 2-30 1-20 1-30 1-25 3-40 2-114 3-60 2-75 2-25 6-40 10-45 1-30 1-25 3-70 5-40 Coral Reefs of Northwest Madagascar 113 Appendix 4 SPECIES Pygoplites diacanthus (Boddaert, 1772) SITE RECORDS 5-7, 11-13, 16-19, 26-29 ABUNDANCE D E P T H (m) Occasional. 3-50 Rare, but surge zone environment not regularly surveyed. 1-3 POMACENTRIDAE Abudefduf septemfasciatus (Cuvier, 1830) 2, 5, 20, 27b A. sexfasciatus Lacepède, 1802 1, 2, 4-7, 11, 13, 15, 16, 18-20, 22-27, 29, 30 A. sordidus (Forsskål, 1775) 2 A. sparoides (Cuvier, 1830) 2, 4-7, 9-29 A. vaigiensis (Quoy & Gaimard, 1825) 1, 2, 5, 8, 9, 13, 17, 20, 22, 25, 27b, 29 Amblyglyphidodon indicus (Allen & Randall, 2002) Amphiprion akallopisos (Bleeker, 1853) A. latifasciatus Allen, 1972 Chromis atripectoralis Welander & Schultz, 1951* C. dimidiata (Klunzinger, 1871) C. lepidolepis Bleeker, 1877 114 Rare, but surge zone environment not regularly surveyed. Common, most abundant member of the genus. Moderately common, but locally abundant. Spawning observed at site 29. 1-15 1-3 1-15 1-12 1, 2, 4-7, 11-24, 26-30 Abundant on silty reefs. 10-35 1-3, 5, 7-23, 26-30 1, 3-8, 10-12, 15-23, 26-30 1, 2, 4, 5, 8, 11, 13, 15, 16, 18, 19, 29 1, 3, 5, 6, 8-19, 21, 23, 26-28, 30 3, 5-7, 9-13, 16-19, 21, 26, 28 Common. Common, but less abundant than A. akallopisos. Moderately common, but locally abundant. 1-20 Moderately common. 1-35 Moderately common. 2-20 C. nigroanalis Randall, 1989* 12, 28 C. nigrura Smith, 1960 11, 12 C. opercularis (Günther, 1867) 5, 12, 16, 26, 28 C. ternatensis (Bleeker, 1856) 4-7, 11-13, 15-18, 2023, 26-30 C. viridis (Cuvier, 1830) 4-8, 11-17, 29 C. weberi Fowler & Bean, 1928 1-6, 8-24, 26-30 C. xutha Randall, 1989 5, 10-13, 17, 28 Chrysiptera annulata (Peters, 1855) Chrysiptera biocellata (Quoy & Gaimard, 1824) Previously recorded. 20 C. brownriggii (Bennett, 1828) 2, 5, 17, 20, 25 C. glauca (Cuvier, 1830) Previously recorded. C. sp. 5, 7, 20, 25, 27b Dascyllus aruanus (Linnaeus, 1758) 1, 2, 4-7, 14-24, 27, 29, 30 D. carneus Fischer, 1885 1-16, 18, 20, 21, 23, 26, 28-30 D. trimaculatus (Rüppell, 1928) 1-9, 11-30 Rapid Assessment Program Common at variety of sites. Rare, except locally abundant on deep outer reef slopes. Rare, except locally common at 2 sites. Occasional, most common on outer slopes. Common. Moderately common, but locally abundant. Abundant, one of most common damselﬁshes. Occasional, but locally common on steep slopes below 15 m. Rare, about 10 seen at one site. Occasional, but surge environment not regularly surveyed. Occasional, but surge environment not regularly surveyed. Formerly identiﬁed as C. unimaculatus, but is undescribed. Common in sheltered waters, forming aggregations around small coral heads. Abundant, except at sheltered inshore habitats. Photographed. Common. Spawning observed at site 30. Photographed. 1-55 2-15 20-40 1-30 4-40 2-15 1-12 3-25 12-40 0-5 0-2 0-2 1-12 1-10 1-55 List of the reef ﬁshes of Madagascar SPECIES SITE RECORDS Neoglyphidodon melas (Cuvier, 1830) 1-9, 11-30 Neopomacentrus azysron (Bleeker, 1877) 3-9, 14-19, 22-24, 27 N. cyanomos (Bleeker, 1856) 1-9, 14-17, 20-24, 26, 27, 29, 30 N. fuliginosus (Smith, 1960) 24 P. johnstonianus Fowler & Ball, 1924* 1, 4-7, 11-13, 16, 18, 19, 26, 27 6, 11, 12, 26 P. lacrymatus (Quoy & Gaimard, 1824) 1-9, 11-24, 26-30 P. leucozonus (Bleeker, 1859) 1, 2, 9, 12 Pomacentrus agassizi Bliss, 1883 Previously recorded. 15, 16, 18, 19, 22, 27, 29 Plectroglyphidodon dickii (Liénard, 1839) P. aquilus Allen & Randall, 1980 P.atriaxillaris Allen, 2002 3, 14, 17 P. caeruleopunctatus Allen, 2002 1-9, 11, 12, 14-21, 23, 26-29 1-4, 8, 9, 11, 26, 28-30 P. caeruleus Quoy & Gaimard, 1825 1-12, 15-24, 26-30 P. pavo (Bloch, 1787) 1-7, 14-24, 29, 30 P. baenschi Allen, 1991* P. sulfureus Klunzinger, 1871 P. trichorus (Günther, 1867) 1 3-7, 11, 12, 15-21, 23, 24, 26-30 1-12, 14-19, 21-24, 26-30 P. trilineatus Cuvier, 1830 4-8, 14, 15, 17, 19-22, 24, 27, 29, 30 Stegastes fasciolatus (Ogilby, 1889) 1, 2, 4-7, 18, 20, 25, 26 S. limbatus (Cuvier, 1830) Previously recorded. S. lividus (Bloch & Schneider, 1801) 20 S. nigricans (Lacepède, 1802) 4, 11, 18, 20 Teixeirichthys jordani (Rutter, 1897) LABRIDAE Anampses caeruleopunctatus Rüppell, 1828 Previously recorded. A. twistii Bleeker, 1856 2, 4, 11, 26 3, 4, 8-11, 19, 23, 26, 29 1, 3-6, 8, 10-12, 16, 18, 19, 21, 23, 26, 28, 29 1-13, 17, 18, 23, 26-28 Bodianus anthioides (Bennett, 1831) 11-13, 17, 28 A. lineatus Randall, 1972* A. meleagrides Valenciennes, 1840 B. axillaris (Bennett, 1831) B. bilunulatus Lacepède, 1801) 1, 3-8, 10-19, 21, 23, 27-29 3, 6, 10, 12, 26 ABUNDANCE Moderately common, but in low numbers at each site. Photographed. Moderately common, but locally abundant. 1-12 Common to locally abundant 5-18 Only seen at site 24 over thick silt bottom with poor visibility. Moderately common, except at sheltered inshore habitats. Occasional. Abundant and one of most common damselﬁshes. Occasional, but surge environment not regularly surveyed. D E P T H (m) 1-12 10-25 1-12 1-18 2-12 0-3 Occasional. 1-15 About 12 seen (one collected) at a single site. An undescribed species. 5-15 Common. 1-20 Occasional, but locally common. Abundant, especially in areas exposed to currents. Abundant, especially at sheltered inshore reefs. 2-15 Common in live coral areas. 1-12 Common in a variety of habitats. 1-43 Moderately common and locally abundant on sheltered inshore reefs. Occasional in surge areas, but habitat not regularly surveyed. Rare, except at a single site where numerous individuals seen. Generally rare, but locally common. Rare, less than 10 seen. Occasional, usually small groups of juveniles seen. Moderately common in small groups. Moderately common. Occasional, usually on steep outer slopes. Common and most abundant member of genus. Rare, less than 10 seen. 1-20 1-20 0-6 0-5 1-5 1-12 2-30 12-42 4-60 2-30 6-60 2-40 8-108 Coral Reefs of Northwest Madagascar 115 Appendix 4 SPECIES SITE RECORDS B. diana (Lacepède, 1802) 1, 5, 6, 9-14, 17, 28 Bodianus macrourus (Lacepède, 1801) B. perdito (Quoy & Gaimard, 1834) B. trilineatus (Fowler, 1934) Cheilinus chlorurus (Bloch, 1791) Previously recorded. Previously recorded. Previously recorded. 9, 11, 12 2, 4, 5, 7, 12, 13, 15, 17-19, 21-24, 26-30 1, 5-8, 11-13, 15, 26, 28, 29 C. fasciatus (Bloch, 1791) C. oxycephalus (Bleeker, 1853) C. trilobatus Lacepède, 1802 1-7, 9, 11-13, 15-30 C. undulatus Rüppell, 1835 12, 20, 29 Cheilio inermis Forsskål, 1775 3, 25, 30 Cirrhilabrus exquisitus Smith, 1957* 10-12, 26 Coris aygula (Lacepède, 1801) 3, 12, 23, 29 C. caudimacula (Quoy & Gaimard, 1834) 1, 3, 4, 8-13, 16, 18, 26, 28, 30 C. cuvieri (Bennett, 1831) 5, 12, 26, 28, 29 C. formosa (Bennett, 1830) Cymolutes praetextatus (Quoy & Gaimard, 1834) C. torquatus (Valenciennes, 1839) Gomphosus caeruleus Lacepède, 1801 Previously recorded. Previously recorded. Previously recorded. 1-9, 11-13, 16-24, 26-30 1-30 Halichoeres cosmetus Randall & Smith, 1982* 12, 13 Epibulus insidiator (Pallas, 1770) H. dussumieri (Valenciennes,1839) H. hortulanus (Lacepède, 1802) H. iridis Randall & Smith, 1982 H. marginatus (Rüppell, 1835) H. nebulosus (Valenciennes, 1839) H. scapularis (Bennett, 1832) 10, 28 2, 7-9, 13-15, 18, 19, 25, 29 1, 3, 5, 6, 9, 25 3, 5, 11, 14-16, 19, 20, 25, 29 H. trispilus Randall, 1982* 11 H. zeylonicus (Bennett, 1833)* 8, 10 Hemigymnus fasciatus (Bloch, 1792) 1, 2, 4-9, 12-24, 27, 29 H. melapterus (Bloch, 1791) Hologymnosus annulatus (Lacepède, 1801) H. doliatus (Lacepède, 1801)* Iniistius pavo Valenciennes, 1840 I. pentadactylus (Linnaeus, 1758) Labrichthys unilineatus (Guichenot, 1847) Labroides bicolor Fowler & Bean, 1928 116 3, 5, 9, 15, 16, 22, 23, 24, 27 1-10, 12-20, 22, 23, 25-30 Rapid Assessment Program 1-10, 13, 16-19, 22, 23, 27, 29 1, 6, 12, 26, 28, 29 2, 28, 29 Previously recorded. Previously recorded. 1-7, 12, 13, 16-20, 23, 27, 29 5, 8, 11-13, 18-21, 23, 24, 26, 29 ABUNDANCE Occasional, most common on outer reef slopes. D E P T H (m) 6-25 Rare, only 3 seen. Common, several adults seen on most dives. 2-30 Moderately common. 1-20 Moderately common, several adults seen on most dives. Rare, only 3 seen, including 2 juveniles. Rare, but weed habitat not adequately sampled. Rarely seen, but locally common, mainly on rubble slopes. Generally rare, but several seen at each of 4 sites. Photographed. Moderately common on sandrubble bottoms. Occasional on sand-rubble bottoms. 4-40 1-20 2-60 0-3 6-32 3-25 2-25 5-50 Common. 1-40 Common. Rare, except several seen on steep slopes at 2 sites. Occasional, mainly on shallow, sheltered reefs. Common, several adults seen at most sites. Rare, several seen adjacent to steep outer slopes. 1-30 Occasional. 1-30 Occasional. Occasional, but locally common in some sandy areas. Rare, only 3 seen on steep dropoff. Rare, only 5 seen. Common, but in low numbers at each site. Common, but in low numbers at each site. Occasional. Rare, only 4 seen. 1-40 Moderately common, especially in rich coral areas. Occasional, generally in smaller numbers than Labroides dimidatus. 2-30 3-25 1-30 6-43 0-15 20-56 10-35 1-20 2-30 3-35 4-35 1-20 2-40 List of the reef ﬁshes of Madagascar SPECIES SITE RECORDS L. dimidiatus (Valenciennes, 1839) 1-30 Labropsis xanthonota Randall, 1981 12 Macropharyngodon bipartitus Smith, 1957 1, 3, 5, 10, 12, 26, 28 M. cyanoguttatus Randall, 1978* 6 M, vivianae Randall, 1978 Novaculichthys macrolepidotus (Bloch, 1791) N. taeniourus (Lacepède, 1802) 3 Previously recorded. 3, 10, 12, 18, 26, 28 Oxycheilinus bimaculatus (Valenciennes, 1840) 8-10, 18, 21, 28 O. diagrammus (Lacepède, 1802) O. mentalis (Rüppell, 1828) Paracheilinus hemitaeniatus Randall & HarmelinVivien, 1977 Pseudocheilinus evanidus Jordan & Evermann, 1902 P. hexataenia (Bleeker, 1857) P. octotaenia Jenkins, 1900 Pseudocoris sp.* 1-3, 5-8, 10, 12, 13, 1523, 26-30 2-5 10-13, 28 2, 4-8, 12, 17, 21, 23 Previously recorded. 28 11-13, 26, 28 Pseudojuloides ceracinus (Snyder, 1904) 10, 14 Pteragogus cryptus Randall, 1981 P. ﬂagellifer (Valenciennes, 1839) P. pelycus Randall, 1981 P. taeniops (Peters, 1855) Stethojulis albovittata (Bonnaterre, 1785) S. interrupta (Bleeker, 1851) S. strigiventer (Bennett, 1832) Thalassoma amblycephalum (Bleeker, 1856) T. genivittatum (Valenciennes, 1839) T. hardwicke (Bennett, 1828) 5, 11-15, 23, 28 Previously recorded. Previously recorded. 8, 10 1-5, 7-13, 16, 19, 21-30 Previously recorded. 5, 15, 20, 29 1, 17, 21, 23, 25, 26 10-13 1-3, 11-14, 16-27, 29 T. hebraicum (Lacepède, 1801) 1-20, 22, 23, 25-27, 29 T. lunare (Linnaeus, 1758) T. purpureum (Forsskål, 1775) Wetmorella nigropinnata (Seale, 1901) SCARIDAE Calotomus carolinus (Valenciennes, 1839) Calotomus spinidens (Quoy & Gaimard, 1824) 1-30 Previously recorded. Previously recorded. Chlorurus atrilunula Randall & Bruce, 1983* C. cyanescens (Valenciennes, 1840) C. sordidus (Forsskål, 1775) C. strongylocephalus (Bleeker, 1862) D E P T H (m) 1-40 4-52 10-30 10-40 10-30 Occasional in rubble areas. Occasional, around rock and coral outcrops, mainly on sandy or rubble bottoms. 1-14 Common. 3-120 Rare, only 7 seen. 1-20 2-110 Previously recorded. Pseudodax moluccanus (Valenciennes, 1840) Cetoscarus bicolor (Rüppell, 1828) ABUNDANCE Moderately common; present at every site. Rare, only one seen. Occasional, and always in small numbers at each site. Rare, only one seen (and collected). Rare, only one seen. 28 Previously recorded. 1, 4-7, 13, 16-18, 2123, 26-29 2, 3, 5, 12, 16-19, 23, 26, 28, 30 16 1, 2, 4-9, 11-16, 18-24, 26-30 1, 3-8, 12, 15-19, 21, 23, 24, 27-29 Hipposcarus harid (Forsskål, 1775) 22, 23, 26, 28 Leptoscarus vaigiensis (Quoy & Gaimard, 1824) Previously recorded. Occasional, but has relatively cryptic habits. Occasional, but has cryptic habits. 6-40 2-35 Rare, only one adult male seen. Occasional, always in low numbers. Rare, only 5 seen, including an adult male. Occasional, but has cryptic habits. 10-30 Rare, but has cryptic habits. Common in variety of habitats. 4-25 0-30 Rare, only 4 seen. Occasional; usually juveniles seen. Rare, about 8 seen. Common. Common; adults and juveniles seen at most sites. Abundant and seen at every site. 0-6 1-15 4-30 0-15 Rare, only one female seen. 4-30 Moderately common, but usually in small numbers. 1-30 Occasional. 1-15 Rare, only one male seen. Common. 5-30 Common. 2-30 Occasional at sites adjacent to sandy bottoms. 5-40 3-40 2-60 4-65 1-30 1-30 1-25 Coral Reefs of Northwest Madagascar 117 Appendix 4 SPECIES SITE RECORDS S. caudofasciatus (Günther, 1862) 28 S. falcipinnis (Playfair, 1867) 3-9, 11-13, 15, 18 S. festivus Valenciennes, 1840* 28 S, frenatus Lacepède, 1802 S. ghobban Forsskål, 1775 S. globiceps Valenciennes, 1840 S. niger Forsskål, 1775 S. psittacus Forsskål, 1775 S. rubroviolaceus Bleeker, 1849 S. russelli Valenciennes, 1840* S. scaber Valenciennes, 1840 S. tricolor Bleeker, 1849 S. virdifurcatus (Smith, 1956)* TRICHONOTIDAE Trichonotus marleyi (Smith, 1961) CREEDIIDAE Limnichthys nitidus Smith, 1958 PINGUIPEDIDAE Parapercis hexophthalma (Cuvier, 1829) 118 1-8, 11-16, 18-20, 2224, 26-30 1-9, 11, 13-19, 22-24, 26-30 5, 6, 16 1, 3-8, 11, 15, 16, 18, 22, 23, 26-30 3, 5, 18, 23, 28, 29 1-3, 5, 7-11, 17, 26, 28 12, 16, 29 2, 4-7, 12-14, 18, 20, 29 1, 5, 11-13, 17, 21, 23, 26-29 2, 5, 7, 11, 13 D E P T H (m) 8-40 4-25 Moderately common. 3-25 Common. 3-30 Rare, only 4 males seen. 2-15 Moderately common. 2-20 Occasional. Moderately common. Occasional. 4-25 1-30 1-20 Moderately common. 5-35 Moderately common. 8-40 2-25 Previously recorded. Previously recorded. 1-3, 5-8, 12-15, 17, 18, 21-23, 26-30 10, 16, 17, 28 4, 11, 24 P. punctulata (Cuvier, 1829) P. sp. TRIPTERYGIIDAE Enneapterygius elegans Peters, 1876 Previously recorded. Enneapterygius tutuilae Jordan & Seale, 1906 6 E. sp. 27b Helcogramma fuscopinna Holleman, 1982 H. obtusirostre (Klunzinger, 1871) BLENNIIDAE Alloblennius parvus Smith-Vaniz & Springer, 1978 Alticus kirki (Günther, 1868) Previously recorded. Previously recorded. Moderately common. 5-25 Occasional. Rare, only 5 seen. 5-25 4-20 One specimen collected with rotenone. Three specimens collected with rotenone from intertidal zone. 0-32 0-5 Previously recorded. Previously recorded. Previously recorded. Antennablennius biﬁlum (Günther, 1861) Aspidontus taeniatus tractus Fowler, 1903 Atrosalarias fuscus (Rüppell, 1835) Blenniella chrysospilos (Bleeker, 1857) Previously recorded. Previously recorded. Previously recorded. B. cyanostigma (Bleeker, 1849) 27b B. gibbifrons (Quoy & Gaimard, 1824) B. periophthalmus (Valenciennes, 1836) Previously recorded. Previously recorded. Cirripectes castaneus Valenciennes, 1836 1, 5 C. ﬁlamentosus (Alleyne & Macleay, 1877) 13, 19, 22 C. perustus Smith, 1959 C. stigmaticus Strasburg & Schultz, 1953 Previously recorded. Previously recorded. Rapid Assessment Program ABUNDANCE Rare, several ﬁsh seen on relatively steep outer slope. Occasional. Rare, only one male seen. Photographed. One specimen collected with rotenone from intertidal zone. Rarely observed, but easily overlooked. Rarely observed, but easily overlooked. 0-2 1-5 1-20 List of the reef ﬁshes of Madagascar SPECIES SITE RECORDS Ecsenius nalolo Smith, 1959* 3, 20, 21, 27 Entomacrodus lemuria Springer & Fricke, 2000 E. striatus (Quoy & Gaimard, 1836) E. thalassinus (Jordan & Seale, 1906) E. vermiculatus (Valenciennes, 1836) Exallias brevis (Kner, 1868) Glyptoparus delicatulus Smith, 1959 Hirculops cornifer (Rüppell, 1830) Istiblennius bellus (Günther, 1861) Previously recorded. Previously recorded. USNM collection. Previously recorded. Previously recorded. Previously recorded. Previously recorded. Previously recorded. Istiblennius dussumieri (Valenciennes, 1836) 27b I. edentulus Bloch & Schneider, 1801 27b I. spilotus (Springer & Williams, 1994) I. steindachneri (Pfeffer, 1893) Meiacanthus mossambicus Smith, 1959 Previously recorded. Previously recorded. 1-24, 26-30 Omobranchus elongatus (Peters, 1855) 27b O. fasciolatus (Valenciennes, 1836) Plagiotremus rhinorhynchus (Bleeker, 1852) P. tapeinosoma (Bleeker, 1857) Petroscirtes breviceps (Valenciennes, 1836) P. mitratus Rüppell, 1830 Salarias fasciatus (Bloch, 1786) Xiphasia setifer Swainson, 1839 CALLIONYMIDAE Previously recorded. 9, 22 2, 3, 8, 9, 15, 17, 22 Previously recorded. Previously recorded. Previously recorded. Previously recorded. Callionymus delicatulus Smith, 1963 30 C. marleyi Regan, 1919 Diplogrammus infulatus Smith, 1963 Synchiropus stellatus Smith, 1963 ELEOTRIDAE Calumia godeffroyi (Günther, 1877) GOBIIDAE Amblyeleotris diagonalis Polunin & Lubbock, 1979 Previously recorded. Previously recorded. Previously recorded. A. periophthalma (Bleeker, 1853) 17, 20, 26, 27 A. steinitzi (Klausewitz, 1974) 1-6, 7, 15, 23, 27-29 A. wheeleri (Polunin & Lubbock, 1977) 3-5, 8, 12, 15, 18, 26 A. sp. 14 Several specimens collected with rotenone from intertidal zone. Two specimens collected with rotenone from intertidal zone. D E P T H (m) 2-15 0-3 0-2 Common. Several specimens collected with rotenone from intertidal zone. 1-20 Rare, only 2 seen. Occasional. 1-40 1-25 One specimen collected with dipnet. 0-20 Previously recorded. 4-6, 15, 17, 18, 23, 27 A. sphynx (Valenciennes, 1837) A. tekomaji Smith, 1959 Asterropteryx semipunctatus Rüppell, 1830 1, 15, 17, 21, 22, 24, 26, 29 Previously recorded. 20 Previously recorded. Bathygobius albopunctatus (Valenciennes, 1837) 27b B. cocosensis (Bleeker, 1854) B. cyclopterus (Valenciennes, 1837) B. fuscus (Rüppell, 1830) Previously recorded. Previously recorded. Previously recorded. Amblygobius albimaculatus (Rüppell, 1830) ABUNDANCE Occasional, usually on silty inshore reefs on or near Porites coral. Occasional, locally common in sandy areas. Occasional, locally common in sandy areas. Occasional, locally common in sandy areas. Occasional, locally common in sandy areas. One specimen collected with spear. 8-35 8-15 6-30 5-20 5-20 Occasional, in sand-rubble areas. 1-5 Rare, only one pair seen. 3-20 Several specimens collected with rotenone from intertidal zone. 0-2 Coral Reefs of Northwest Madagascar 119 Appendix 4 SPECIES SITE RECORDS Bryaninops amplus Larson, 1985* 29 B. natans Larson, 1986* 5, 6 B. yongei (Davis & Cohen, 1968)* 1 Callogobius ﬂavobrunneus (Smith, 1958) C. irasus (Smith, 1959) Previously recorded. Previously recorded. C. plumatus (Smith, 1959) 12 C. sclateri (Steindachner, 1880) 13, 29 C. sp. 12 Coryogalops anomolus Smith, 1958 C. sordida (Smith, 1959) Cryptocentrus fasciatus (Playfair & Günther, 1867) Previously recorded. Previously recorded. C. strigilliceps (Jordan & Seale, 1906) 4-6, 15, 20, 33, 24 Eviota distigma (Jordan & Seale, 1906) E. nebulosa Smith, 1958 3, 5, 6, 15, 17, 21, 22, 27, 29, 30 Previously recorded. Previously recorded. E. prasina (Klunzinger, 1871) 13 E. sebreei Jordan & Seale, 1906* 3, 21 E. sigillata Jewett & Lachner, 1983* 6, 29 E. sp. 1 12 E. sp. 2 12 Exyrias bellisimus (Smith, 1959)* Flabellogobius fourmanoiri Smith, 1956 22 Previously recorded. Fusigobius duospilus Hoese & Reader, 1985* 12 F. longispinis (Goren, 1978)* 6, 13, 29 F. neophytes (Günther, 1877) Gnatholepis anjerensis (Bleeker, 1851) Gobiodon citrinus (Rüppell, 1838) Previously recorded. Previously recorded. 1, 5, 12, 22, 24, 26, 29, 30 Previously recorded. G. rivulatus (Rüppel, 1830) 30 G. unicolor (Castelnau, 1873)* 23 Hetereleotris tentaculatus Smith, 1958 H. vulgaris (Klunzinger, 1871) H. zanzibarensis (Smith, 1958) Previously recorded. Previously recorded. Previously recorded. Ctenogobiops crocineus Smith, 1959 G. cauerensis Bleeker, 1853 120 15, 20 Rapid Assessment Program ABUNDANCE One specimen collected with rotenone, but not doubt common where seawhips are abundant. Rare, but relatively inconspicuous due to tiny size. Only one seen, but difﬁcult to detect. No doubt common wherever seawhips are abundant. One specimen collected with rotenone. Several specimens collected with rotenone. One specimen collected with rotenone. Rare, but sand habitat not adequately surveyed. Occasional, but locally common on silt-sand fringe of shallow reefs. Occasional, but locally common on sandy fringe of reefs. One specimen collected with rotenone. Rarely observed, but locally common on live corals. Several specimens collected with rotenone. Two specimens collected with rotenone. One specimen collected with rotenone. Rare; several large adults seen. One specimen collected with rotenone. Three specimens collected with rotenone. Rarely noticed, but locally common on sand-rubble bottoms. Only one collected, but probably common among tabular Acropora coral. Only one collected, but probably common among tabular Acropora coral. D E P T H (m) 10-40 6-27 5-40 3-25 3-25 5-30 2-15 1-6 2-15 8-10 5-20 8-10 4-12 4-12 1-25 5-25 10-30 1-45 3-10 2-12 List of the reef ﬁshes of Madagascar SPECIES Istigobius decoratus (Herre, 1927) I. ornatus (Rüppell, 1830) SITE RECORDS 1, 2, 4-7, 9, 14, 15, 22, 24, 27, 29 27b Macrodontogobius wilburi Herre, 1936* 1, 2, 4-7, 15, 17-24, 27-29 20 Mahidolia mystacina (Valenciennes, 1837) 20 Oplopomus oplopomus (Valenciennes, 1837) 15, 20, 21 Paragobiodon echinocephalus (Rüppell, 1830) Previously recorded. Koumansetta hectori (Smith, 1957) Phyllogobius platycephalops (Smith, 1964)* 10 Pleurosicya elongata Larson, 1990* 29 P. fringilla Larson, 1990 Previously recorded. P. mossambica Smith, 1959 1 Priolepis cincta (Regan, 1908) P. inhaca (Smith, 1949) - USNM P. kappa Winterbottom & Burridge, 1993 P. nocturna (Smith, 1956) Trimma corallinus (Smith, 1959) Previously recorded. Previously recorded. Previously recorded. Previously recorded. Previously recorded. T. macrophthalma (Tomiyama, 1936)* 6, 11, 12, 13 T. naudei Smith, 1957 6, 29 T. haima Winterbottom, 1984* 6, 13, 29 T. sheppardi Winterbottom, 1984* 13 T. sp. 1 13 T. sp. 2 29 Valenciennea helsdingeni (Bleeker, 1858) Previously recorded. V. parva Hoese & Larson, 1994* 27 V. puellaris (Tomiyama, 1936) 3-5, 15, 24, 26, 29 V. sexguttata (Valenciennes, 1837) 4, 24 V. strigata (Broussonet, 1782) 3, 4, 18, 19, 26, 28 Vanderhorstia ambanoro (Fourmanoir, 1957)* 15, 20 V. delgoae (Barnard, 1937) Vanderhorstia ornatissima Smith, 1959 KRAEMERIIDAE Kraemeria samoensis Steindachner, 1906 MICRODESMIDAE Gunnelichthys copleyi (Smith, 1951) Gunnelichthys curiosus Dawson, 1968* Previously recorded. Previously recorded. ABUNDANCE Moderately common on sandy fringe of reefs. Several specimens collected with rotenone from intertidal zone. D E P T H (m) 1-18 0-5 Common. 0-5 Rare, only one seen. Rare, but sand habitat not adequately surveyed. Rarely seen, but locally common on ﬁne sand bottoms away from reefs. 2-15 Only a few seen, but easily escapes notice due to small size. Commensal with sponges (Phyllospongia). Only a few noticed, but easily overlooked. Commensal with barrel sponges. A new record for the W. Indian Ocean. Two collected. Photographed. Only a few seen on Porites coral, but easily escapes notice due to small size. Several specimens collected with rotenone. Two specimens collected with rotenone. Two specimens collected with rotenone. One specimen collected with rotenone. Several specimens collected with rotenone. Two specimens collected with rotenone. Rare, one specimen speared in 12 m. Occasional in sandy areas. Rare, only 4 pairs seen in sandy areas. Rare, less than 10 seen. Rarely seen, but locally common on ﬁne sand bottoms. 1-20 2-25 3-20 1-25 1-35 5-30 3-25 0-36 20-48 15-30 10-20 2-30 8-30 1-10 1-25 4-20 Previously recorded. Previously recorded. 26 Rare, only 3 seen. 4-25 Coral Reefs of Northwest Madagascar 121 Appendix 4 SPECIES G. monostigma Smith, 1958 Paragunnellichthys seychellensis Dawson, 1967 PTERELEOTRIDAE Nemateleotris magniﬁca Fowler, 1938 Parioglossus philippinus (Herre, 1945) Ptereleotris evides (Jordan & Hubbs, 1925) SITE RECORDS 4 Previously recorded. ABUNDANCE Rare, only one seen. D E P T H (m) 2-30 11, 12 Previously recorded. 3, 5, 8, 13, 17, 28 Rare, only 4 pairs seen. 6-61 2-15 P. microlepis (Bleeker, 1856) 4, 22, 24 Occasional. Rare, except about 20 seen at site 24. Rare, less than 10 seen. Rare, only one juvenile and one adult seen. Rare, only one large adult seen. 1-20 XENISTHMIDAE Xenisthmus africanus (Smith, 1958) EPHIPPIDAE Platax boersi Bleeker, 1852* 3, 12, 21, 22, 26 P. orbicularis (Forsskål, 1775) 27b, 29 P. teira (Forsskål, 1775) SIGANIDAE 23 Siganus argenteus (Quoy & Gaimard, 1824) S. luridus (Rüppell, 1829) S. stellatus (Forsskål, 1775) S. sutor (Valenciennes, 1835) ZANCLIDAE Zanclus cornutus Linnaeus, 1758 1-12, 14, 16, 18, 19, 21-24, 26-30 1, 3, 4, 6, 8-10, 14, 1719, 21 1-7, 9-13, 15-19, 2124, 26-30 20, 21 1-4, 7-13, 15-19, 2124, 26-30 ACANTHURIDAE Acanthurus blochii Valenciennes, 1835 A. dussumieri Valenciennes, 1835 A. leucosternon Bennett, 1832 8, 11, 15, 24, 29 6, 8, 9, 12-16 5, 12, 16, 18, 19 A. lineatus (Linnaeus, 1758) 2, 8, 9, 14, 18, 19, 26 A. mata (Cuvier, 1829) 1, 3, 7, 9, 14, 15, 17, 24 A. nigricaudus Duncker & Mohr, 1929 A. nigrofuscus (Forsskål, 1775) 3, 4, 8, 10-13, 22, 24, 26, 28, 30 2, 4-7, 11, 12, 16, 21, 25, 26, 28 A. tennenti Günther, 1861 11, 12 A. thompsoni (Fowler, 1923)* 13 A. triostegus (Linnaeus, 1758) 2, 5, 13, 20, 25, 27b Ctenochaetus binotatus Randall, 1955* 1, 5, 8, 9, 15, 17, 20, 22, 23, 25-27, 29, 30 7, 11-13, 28, 29 C. striatus (Quoy & Gaimard, 1824) 1-5, 7-21, 23, 24, 26-29 C. truncatus Randall & Clements, 2001* 11-13, 17-19, 21 Naso annulatus (Quoy & Gaimard, 1825) N. brachycentron (Valenciennes, 1835) 2, 3, 9 1, 3, 11 1-3, 9-13, 15, 17, 23, 26, 30 9 1, 2, 6, 7, 11-13, 17-19, 23, 26-28 A. xanthopterus Valenciennes, 1835 N. brevirostris (Valenciennes, 1835) N. caeruleacauda Randall, 1994* N. elegans Rüppell, 1829 122 Previously recorded. Rapid Assessment Program Common, often in large schools. Moderately common. Common, either solitary or in pairs. Occasional in seagrass beds. 1-30 0-2 1-30 1-12 1-27 0-5 Common. 1-180 Occasional. Occasional. Rare, less than 10 seen. Occasional, usually in shallow surge-affected areas. Occasional, frequently on dropoffs in turbid water. 3-20 4-130 1-25 Occasional. 3-30 Occasional, but easily overlooked. 2-20 Generally rare, but locally common at two sites. Generally rare, but common on edge of dropoff at site 13. Occasional, usually in shallow wave-affected areas. Moderately common, usually over sandy bottoms adjacent to reefs. Occasional. Common, usually in depths less than 10 m. Only a few noticed, but hard to differentiate from C. striatus at a distance. Rare, less than 10 seen. Rare, less than 10 seen. 1-15 5-30 1-20 4-75 0-90 3-90 10-55 2-30 3-25 15-40 15-50 Moderately common. 4-50 One school seen. 5-40 Moderately common. 1-30 List of the reef ﬁshes of Madagascar SPECIES N. hexacanthus (Bleeker, 1855)* N. tuberosus Lacepède, 1802 N. unicornis (Forsskål, 1775) SITE RECORDS 4-7, 11-13, 15-17, 22, 23, 29, 30 Previously recorded. 4-7, 11-13, 15-17, 22, 23, 29, 30 N. vlamingii (Valenciennes, 1835) 1, 2, 5, 12, 26, 28 Paracanthurus hepatus (Linnaeus, 1758) 8, 10 1, 5, 10, 13, 14, 15, 17, 19, 20, 22-24 Previously recorded. 1-8, 11-24, 26-30 Zebrasoma desjardinii (Bennett, 1836) Z. gemmatum (Valenciennes, 1835) Z. scopas (Cuvier, 1829) SPHYRAENIDAE Sphyraena acutipinnis Day, 1876 S. barracuda (Walbaum, 1792) S. ﬂavicauda Rüppell, 1838 Previously recorded. 11 16, 18, 21, 22, 26, S. jello Cuvier, 1829 23 S. qenie Klunzinger, 1870 23 SCOMBRIDAE Gymnosarda unicolor (Rüppell, 1836) Rastrelliger kanagurta (Cuvier, 1816) Scomberomorus commerson (Lacepède, 1800) BOTHIDAE 12 Previously recorded. 26 Arnogolossus sp. 6 Bothus mancus (Broussonet, 1782) B. myriaster (Temminck & Schlegel, 1846) B. pantherinus (Rüppell, 1830) SAMARIDAE Previously recorded. Previously recorded. Previously recorded. Samariscus triocellatus Woods, 1966* 12 SOLEIDAE Pardachirus marmoratus (Lacepède, 1802) BALISTIDAE Abalistes stellatus (Lacepède, 1798) Balistapus undulatus (Park, 1797) ABUNDANCE D E P T H (m) Moderately common. 6-140 Moderately common. 4-80 Occasional, but locally common adjacent to dropoffs. Rare, less than 10 seen. Occasional, but locally common at site 22. 4-50 2-40 4-30 Common. 1-60 Rare, only one seen. Occasional, usually in schools. Rare, one mixed school (with S. qenie) of 50 ﬁsh seen. Rare, one mixed school (with S. jello) of 50 ﬁsh seen. 0-20 1-20 Rare, only one seen. 0-100 Rare, only one seen. 0-30 Only one seen, but difﬁcult to detect due to cryptic coloration. 5-30 One specimen collected with rotenone. 10-30 Occasional. 3-50 1-20 5-40 Previously recorded. Previously recorded. 6, 7, 11-13, 16, 18, 23, 24, 26, 27, 29 Balistoides conspicillum (Bloch & Schneider, 1801) B. viridescens (Bloch & Schneider, 1801) Canthidermis maculatus (Bleeker, 1865) 10, 12 Rare, only 3 seen. 10-50 9, 21, 26, 28 Previously recorded. Rare, only 4 seen. 5-45 Melichthys indicus Randall & Klausewitz, 1973 12 Generally rare, but about 10 seen at site 12. 2-30 Odonus niger (Rüppell, 1836) Pseudobalistes ﬂavimarginatus (Rüppell, 1829) Previously recorded. Previously recorded. P. fuscus (Bloch & Schneider, 1801) 1, 3, 10, 14, 15, 26 Rhinecanthus aculeatus (Linnaeus, 1758) 15, 25 R. rectangulus (Bloch & Schneider, 1801) Sufﬂamen bursa (Bloch & Schneider, 1801) S. chrysoptera (Bloch & Schneider, 1801) S. fraenatus (Latreille, 1804) MONACANTHIDAE Aluterus scriptus (Osbeck, 1765) Amanses scopas (Cuvier, 1829) Cantherines dumerilii (Hollard, 1854)* Previously recorded. 12, 13, 28 1-19, 21-30 10, 26 Rare, less than 10. Common. Rare, only 5 seen. 3-90 1-35 8-185 Previously recorded. 1, 5, 18, 23 6 Rare, only 6 seen. Rare, only one seen. 3-20 1-35 Occasional on sand-rubble fringe of reefs. Generally rare, except about 15 seen at site 15. 1-50 0-3 Coral Reefs of Northwest Madagascar 123 Appendix 4 SPECIES C. pardalis (Rüppell, 1837) Oxymonacanthus longirostris (Bloch & Schneider, 1801) Paraluteres prionurus (Bleeker, 1851)* SITE RECORDS 2, 3, 8, 12, 18, 23, 26, 27 Previously recorded. 1, 4-8, 11, 14-16, 1820, 23, 26, 29 3, 5, 6, 13, 18, 27 Pervagor janthinosoma (Bleeker, 1854) 30 C. fronticinctus (Günther, 1866) Pseudalutarius nasicornis (Temminck & Schlegel, 1850) OSTRACIIDAE Lactoria cornuta (Linnaeus, 1758) Lactoria fornasini (Bianconi, 1846) Ostracion cubicus Linnaeus, 1758 O. meleagris Shaw, 1796 Tetrasomus gibbosus (Linnaeus, 1758) TETRAODONTIDAE Arothron hispidus (Linnaeus, 1758) A. immaculatus (Bloch & Schneider, 1801) A. mappa (Lesson, 1830)* A. nigropunctatus (Bloch & Schneider, 1801) A. stellatus (Schneider, 1801) Canthigaster bennetti (Bleeker, 1854) C. janthinoptera (Bleeker, 1855) C. solandri (Richardson, 1844) C. valentini (Bleeker, 1853) DIODONTIDAE Diodon hystrix Linnaeus, 1758 Lophodiodon calori (Bianconi, 1855) 124 Rapid Assessment Program ABUNDANCE D E P T H (m) Occasional. 2-40 Moderately common in rich coral areas. Occasional. Rare, only one seen. Photographed. 1-30 2-25 2-18 Previously recorded. Previously recorded. 26 3, 14-16, 18, 22, 23, 27, 29, 30 5, 6, 12, 28 Previously recorded. 14 Previously recorded. 11, 12 1-3, 12, 15, 26, 28 29 2, 3, 8, 10, 11, 14, 20, 26 Previously recorded. 1-9, 11, 13-15, 17-21, 23, 24, 26-30 1-24, 26-30 6, 14, 26, 29 Previously recorded. Rare, only one seen. 1-30 Occasional. 1-40 Rare, less than 10 seen. 2-30 Rare, only one seen. 1-50 Rare, only 2 seen. Occasional. Rare, only one seen. 4-40 2-35 3-58 Occasional in sand-rubble areas. 1-10 Common, usually in pairs. 3-36 Common. 3-55 Rare, only 4 seen. 1-30 Additional Published Reports of the Rapid Assessment Program South America * Bolivia: Alto Madidi Region. Parker, T.A. III and B. Bailey (eds.). 1991. A Biological Assessment of the Alto Madidi Region and Adjacent Areas of Northwest Bolivia May 18 - June 15, 1990. RAP Working Papers 1. Conservation International, Washington, DC. * Bolivia: Lowland Dry Forests of Santa Cruz. Parker, T.A. III, R.B. Foster, L.H. Emmons and B. Bailey (eds.). 1993. The Lowland Dry Forests of Santa Cruz, Bolivia: A Global Conservation Priority. RAP Working Papers 4. Conservation International, Washington, DC. † Bolivia/Perú: Pando, Alto Madidi/Pampas del Heath. Montambault, J.R. (ed.). 2002. Informes de las evaluaciones biológicas de Pampas del Heath, Perú, Alto Madidi, Bolivia, y Pando, Bolivia. RAP Bulletin of Biological Assessment 24. Conservation International, Washington, DC. * Bolivia: South Central Chuquisaca Schulenberg, T.S. and K. Awbrey (eds.). 1997. A Rapid Assessment of the Humid Forests of South Central Chuquisaca, Bolivia. RAP Working Papers 8. Conservation International, Washington, DC. * Bolivia: Noel Kempff Mercado National Park. Killeen, T.J. and T.S. Schulenberg (eds.). 1998. A biological assessment of Parque Nacional Noel Kempff Mercado, Bolivia. RAP Working Papers 10. Conservation International, Washington, DC. * Bolivia: Río Orthon Basin, Pando. Chernoff, B. and P.W. Willink (eds.). 1999. A Biological Assessment of Aquatic Ecosystems of the Upper Río Orthon Basin, Pando, Bolivia. Bulletin of Biological Assessment 15. Conservation International, Washington, DC. § Brazil: Rio Negro and Headwaters. Willink, P.W., B. Chernoff, L.E. Alonso, J.R. Montambault and R. Lourival (eds.). 2000. A Biological Assessment of the Aquatic Ecosystems of the Pantanal, Mato Grosso do Sul, Brasil. RAP Bulletin of Biological Assessment 18. Conservation International, Washington, DC. § Ecuador: Cordillera de la Costa. Parker, T.A. III and J.L. Carr (eds.). 1992. Status of Forest Remnants in the Cordillera de la Costa and Adjacent Areas of Southwestern Ecuador. RAP Working Papers 2. Conservation International, Washington, DC. * Ecuador/Perú: Cordillera del Condor. Schulenberg, T.S. and K. Awbrey (eds.). 1997. The Cordillera del Condor of Ecuador and Peru: A Biological Assessment. RAP Working Papers 7. Conservation International, Washington, DC. § Guyana: Kanuku Mountain Region. Parker, T.A. III and A.B. Forsyth (eds.). 1993. A Biological Assessment of the Kanuku Mountain Region of Southwestern Guyana. RAP Working Papers 5. Conservation International, Washington, DC. * Guyana: Eastern Kanuku Mountains. Montambault, J.R. and O. Missa (eds.). 2002. A Biodiversity Assessment of the Eastern Kanuku Mountains, Lower Kwitaro River, Guyana. RAP Bulletin of Biological Assessment 26. Conservation International, Washington, DC. * Paraguay: Río Paraguay Basin. Chernoff, B., P.W. Willink and J. R. Montambault (eds.). 2001. A biological assessment of the Río Paraguay Basin, Alto Paraguay, Paraguay. RAP Bulletin of Biological Assessment 19. Conservation International, Washington, DC. Cover photos Top: Nosy Kivinjy Photo by G. Allen Center: New species of damselﬁsh (Pomacentrus caeruleopunctatus) Photos by G. Allen Bottom: Coral reef Photo by J. Veron Central America § Belize: Columbia River Forest Reserve. Parker, T.A. III. (ed.). 1993. A Biological Assessment of the Columbia River Forest Reserve, Toledo District, Belize. RAP Working Papers 3. Conservation International, Washington, DC. * Guatemala: Laguna del Tigre National Park. Bestelmeyer, B. and L.E. Alonso (eds.). 2000. A Biological Assessment of Laguna del Tigre National Park, Petén, Guatemala. RAP Bulletin of Biological Assessment 16. Conservation International, Washington, DC. Asia-Pacific * Indonesia: Wapoga River Area. Mack, A.L. and L.E. Alonso (eds.). 2000. A Biological Assessment of the Wapoga River Area of Northwestern Irian Jaya, Indonesia. RAP Bulletin of Biological Assessment 14. Conservation International, Washington, DC. * Indonesia: Togean and Banggai Islands. Allen, G.R., and S.A. McKenna (eds.). 2001. A Marine Rapid Assessment of the Togean and Banggai Islands, Sulawesi, Indonesia. RAP Bulletin of Biological Assessment 20. Conservation International, Washington, DC. * Indonesia: Raja Ampat Islands. McKenna, S.A., G.R. Allen and S. Suryadi (eds.). 2002. A Marine Rapid Assessment of the Raja Ampat Islands, Papua Province, Indonesia. RAP Bulletin of Biological Assessment 22. Conservation International, Washington, DC. * Indonesia: Yongsu - Cyclops Mountains and the Southern Mamberamo Basin. Richards, S.J. and S. Suryadi (eds.). 2002. A Biodiversity Assessment of Yongsu - Cyclops Mountains and the Southern Mamberamo Basin, Papua, Indonesia. RAP Bulletin of Biological Assessment 25. Conservation International, Washington, DC. * Papua New Guinea: Lakekamu Basin. Mack, A.L. (ed.). 1998. A Biological Assessment of the Lakekamu Basin, Papua New Guinea. RAP Working Papers 9. Conservation International, Washington, DC. † Papua New Guinea: Milne Bay Province. Werner, T.B. and G. Allen (eds.). 1998. A Rapid Biodiversity Assessment of the Coral Reefs of Milne Bay Province, Papua New Guinea. RAP Working Papers 11. Conservation International, Washington, DC. * Papua New Guinea: Southern New Ireland. Beehler, B.M. and L.E. Alonso (eds.). 2001. Southern New Ireland, Papua New Guinea: A Biodiversity Assessment. RAP Bulletin of Biological Assessment 21. Conservation International, Washington, DC. * Papua New Guinea: Milne Bay Province. Allen, G.R., J.P. Kinch, S.A. McKenna and P. Seeto (eds.). 2003. A Rapid Marine Biodiversity Assessment of Milne Bay Province, Papua New Guinea - Survey II (2000). RAP Bulletin of Biological Assessment 29. Conservation International, Washington, DC. † Philippines: Palawan Province. Werner, T.B. and G. Allen (eds.). 2000. A Rapid Marine Biodiversity Assessment of the Calaminanes Islands, Palawan Province, Philippines. RAP Bulletin of Biological Assessment 17. Conservation International, Washington, DC. Africa & Madagascar * Perú: Tambopata-Candamo Reserved Zone. Foster, R.B., J.L. Carr and A.B. Forsyth (eds.). 1994. The Tambopata-Candamo Reserved Zone of southeastern Perú: A Biological Assessment. RAP Working Papers 6. Conservation International, Washington, DC. * Botswana: Okavango Delta. Alonso, L.E. and L. Nordin (eds.). 2003. A Rapid Biological Assessment of the aquatic ecosystems of the Okavango Delta, Botswana: High Water Survey. RAP Bulletin of Biological Assessment 27. Conservation International, Washington, DC. * Perú: Cordillera de Vilcabamba. Alonso, L.E., A. Alonso, T. S. Schulenberg and F. Dallmeier (eds.). 2001. Biological and Social Assessments of the Cordillera de Vilcabamba, Peru. RAP Working Papers 12 and SI/MAB Series 6. Conservation International, Washington, DC. † Côte d’Ivoire: Marahoué National Park. Schulenberg, T.S., C.A. Short and P.J. Stephenson (eds.). 1999. A Biological Assessment of Parc National de la Marouhe, Côte d’Ivoire. RAP Working Papers 13. Conservation International, Washington, DC. * Venezuela: Caura River Basin. Chernoff, B., A. Machado-Allison, K. Riseng and J.R. Montambault (eds.). 2003. A Biological Assessment of the Aquatic Ecosystems of the Caura River Basin, Bolívar State, Venezuela. RAP Bulletin of Biological Assessment 28. Conservation International, Washington, DC. * Guinea: Pic de Fon. McCullough, J. (ed.). 2004. A Rapid Biological Assessment of the Foret Classée du Pic de Fon, Simandou Range, Southeastern Republic of Guinea. RAP Bulletin of Biological Assessment 35. Conservation International, Washington, DC. * Venezuela: Orinoco Delta and Gulf of Paria. Lasso, C.A., L.E. Alonso, A.L. Flores and G. Love (eds.). 2004. Rapid assessment of the biodiversity and social aspects of the aquatic ecosystems of the Orinoco Delta and the Gulf of Paria, Venezuela. RAP Bulletin of Biological Assessment 37. Conservation International, Washington, DC. * Madagascar: Ankarafantsika. Alonso, L.E., T.S. Schulenberg, S. Radilofe and O. Missa (eds). 2002. A Biological Assessment of the Réserve Naturelle Intégrale d’Ankarafantsika, Madagascar. RAP Bulletin of Biological Assessment 23. Conservation International, Washington, DC. * Available through the University of Chicago Press. To order call 1-800-621-2736; www.press.uchicago.edu † Available only through Conservation International. To order call 202-912-1249. § Out of Print 31 Participants ..................................................................................... 4 Organizational Profiles ................................................................. 5 Acknowledgments ......................................................................... 7 Executive Summary ....................................................................... 8 Overview ....................................................................................... 11 Map and Photos ........................................................................... 19 Technical Reports ....................................................................... 23 Appendices ................................................................................... 85 Conservation International 1919 M Street, NW Suite 600 Washington, DC 20036 USA TELEPHONE: 202-912-1000 FAX: 202-912-0772 WEB: www.conservation.org www.biodiversityscience.org A Rapid Marine Biodiversity Assessment of the Coral Reefs of Northwest Madagascar A Rapid Marine Biodiversity Assessment of the Coral Reefs of Northwest Madagascar Rapid Assessment Program A Rapid Marine Biodiversity Assessment of the Coral Reefs of Northwest Madagascar Editors Sheila A. McKenna and Gerald R. Allen RAP Bulletin Biological Assessment of 31 Protocole of Agreement CI & FAO no PO 1081138 9 0 0 0 0> 9 781881 173847 C ONSERVATION I NTERNATIONAL IS B N 1-881173-84-4 Center for Applied Biodiversity Science (CABS) Conservation International Australian Institute of Marine Science Western Australian Museum

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A rapid marine biodiversity assessment of the coral reefs of northwest Madagascar

A rapid marine biodiversity assessment of the coral reefs of northwest Madagascar

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