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Too precious to drill: the marine biodiversity of Belize Palomares, Maria Lourdes D. 2012

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  ISSN 1198-6727  Fisheries Centre Research Reports  2011 Volume 19 Number 6      TOO PRECIOUS TO DRILL: THE MARINE BIODIVERSITY OF BELIZE        Fisheries Centre, University of British Columbia, Canada  TOO PRECIOUS TO DRILL: THE MARINE BIODIVERSITY OF BELIZE     edited by  Maria Lourdes D. Palomares and Daniel Pauly                   Fisheries Centre Research Reports 19(6) 175 pages © published 2011 by  The Fisheries Centre, University of British Columbia  2202 Main Mall Vancouver, B.C., Canada, V6T 1Z4       ISSN 1198-6727  Fisheries Centre Research Reports 19(6) 2011  TOO PRECIOUS TO DRILL: THE MARINE BIODIVERSITY OF BELIZE  edited by Maria Lourdes D. Palomares and Daniel Pauly  CONTENTS  PAGE DIRECTOR‘S FOREWORD 1 EDITOR‘S PREFACE 2 INTRODUCTION 3 Offshore oil vs 3E‘s (Environment, Economy and Employment) 3 Frank Gordon Kirkwood and Audrey Matura-Shepherd The Belize Barrier Reef: a World Heritage Site 8 Janet Gibson BIODIVERSITY 14 Threats to coastal dolphins from oil exploration, drilling and spills off the coast of Belize 14 Ellen Hines The fate of manatees in Belize 19 Nicole Auil Gomez Status and distribution of seabirds in Belize: threats and conservation opportunities 25 H. Lee Jones and Philip Balderamos Potential threats of marine oil drilling for the seabirds of Belize 34 Michelle Paleczny The elasmobranchs of Glover‘s Reef Marine Reserve and other sites in northern and central Belize 38 Demian Chapman, Elizabeth Babcock, Debra Abercrombie, Mark Bond and Ellen Pikitch Snapper and grouper assemblages of Belize: potential impacts from oil drilling 43 William Heyman Endemic marine fishes of Belize: evidence of isolation in a unique ecological region 48 Phillip Lobel and Lisa K. Lobel Functional importance of biodiversity for coral reefs of Belize 52 Janie Wulff Biodiversity of sponges: Belize and beyond, to the greater Caribbean 57 Maria Cristina Diaz and Klaus Ruetzler Biodiversity, ecology and biogeography of hydroids (Cnidaria: Hydrozoa) from Belize 66 Lea-Anne Henry Documenting the marine biodiversity of Belize through FishBase and SeaLifeBase 78 Maria Lourdes D. Palomares and Daniel Pauly HABITATS 107 Evaluating potential impacts of offshore oil drilling on the ecosystem services of mangroves in Belize 107 Timothy Brook Smith and Nadia Bood Bacalar Chico Marine Reserve: Ecological status of Belize Barrier Reef's northernmost reserve 112 Mebrahtu Ateweberhan, Jennifer Chapman, Frances Humber, Alasdair Harris and Nick Jones Preparing for potential impacts of offshore petroleum exploration and development on the marine communities in the Belize Barrier Reef and lagoonal ecosystems 119 Robert Ginsburg A deep-sea coral ‗gateway‘ in the northwestern Caribbean 120 Lea-Anne Henry Natural and anthropogenic catastrophe on the Belizean Barrier Reef 125 Richard B. Aronson, Ian G. Macintyre and William F. Precht Declining reef health calls for stronger protection not additional pollution from offshore oil development 129 Melanie McField   CONTENTS (CONTINUED)  PAGE FISHERIES AND TOURISM 135 Fisheries based on Belizean biodiversity: why they're so vulnerable to offshore oil exploration 135 Eli Romero and Les Kaufman Reconstruction of total marine fisheries catches for Belize, 1950-2008 142 Dirk Zeller, Rachel Graham and Sarah Harper Under the threat of oil: assessing the value and contribution of Belizean fisheries 152 Sarah Harper, Dirk Zeller and U. Rashid Sumaila The economic value and potential threats to marine ecotourism in Belize 161 Andres M. Cisneros-Montemayor and U. Rashid Sumaila APPENDICES 167 Conference agenda 167 Letter of scientists to Belizeans 171 Conference participants 172                                       A Research Report from the Fisheries Centre at UBC 175 pages © Fisheries Centre, University of British Columbia, 2011 Fisheries Centre Research Reports are abstracted in the FAO Aquatic Sciences and Fisheries Abstracts (ASFA) ISSN 1198-6727 FISHERIES CENTRE RESEARCH REPORTS ARE FUNDED IN PART BY GRANT FUNDS FROM THE PROVINCE OF BRITISH COLUMBIA MINISTRY OF ENVIRONMENT. A LIST OF ALL FCRRS TO DATE APPEARS AS THE FINAL PAGES OF EACH REPORT. Too Precious to Drill: The Marine Biodiversity of Belize, Palomares and Pauly  1 DIRECTOR‘S FOREWORD The April 2010 Deepwater Horizon oil rig blowout in the Gulf of Mexico has sharpened attention on the oil spills occurring in many parts of the world ocean, and their potential damaging effects on marine ecosystems and the living organisms they sustain. This report focuses on the sustainability of marine fisheries of Belize in the face of potential impacts of ocean threats – in particular, oil spills. The report is timely and important in at least two ways. First, it addresses oil spills in the ocean, which occur frequently worldwide and can have significant effects on life in the ocean and the wellbeing of the people dependent on it. Second, the report focuses on a small developing country, Belize – an example of a country that does not usually receive the attention it deserves by researchers, even though the ocean and the resources it contains is the main source of existence for its citizens. Thirdly, this work is a collaboration between academic researchers, NGOs and management partners, thereby making the research output more relevant to real life problems. This report consists of several chapters that tackle issues ranging from the ecology of the marine ecosystem of Belize right through to the economic benefits currently derived from activities dependent on the ecosystem. These include fishing, angling and whale(shark) watching. A crucial point made in the report is that while oil is a non-renewable resource, fish is renewable. This means that in comparing the benefits from drilling the marine ecosystem of Belize, it is important that in the short term, possibly larger benefits from oil drilling should not be allowed to trump benefits that, if well-managed and protected, are capable of continuing to flow through time, benefiting all generations. The result of the work reported in this contribution, which is based on a broad collaboration between scientists, civil society members and managers, serves as a good example of how to produce policy relevant research that serves societal goals and objectives. I commend the authors of the report for producing a significant piece of research that has a strong potential to contribute positively to policy making in Belize. U. RASHID SUMAILA Director and Professor The Fisheries Centre, UBC  Too Precious to Drill: The Marine Biodiversity of Belize, Palomares and Pauly  2 EDITORS‘ PREFACE There is a huge amount of zoological and botanical publications on the marine biodiversity of Belize, notably because the American Museum of Natural History in New York and the Smithsonian Institution in Washington, D.C., established marine stations many years ago in Belize and used these for continuous monitoring, and for generations of graduate students to complete their theses. All these and similar materials were, however, published mainly in US and British scientific journals, with only sporadic efforts to make it accessible to the Belizean students and members of the public. Thus, those Belizeans who live with their back to the sea do not get the information that they need to turn around, and fully appreciate the beauty and wealth of the biodiversity along their shores, and its role in attracting tourists and producing seafood. This also leads to the Belizean public not fully appreciating the risk to marine biodiversity of an oil spill and the potential cost to their economy. In view of the debate and the possibility of a national referendum on offshore oil drilling in Belize, a conference entitled ‗Too Precious to Drill: the Marine Biodiversity of Belize‘ was organized jointly by Oceana Belize and the Sea Around Us project, with major funding from the Oak Foundation. This report assembles the contributions presented at this conference, and is complemented by a conference website (‗Too Precious to Drill: the Marine Biodiversity of Belize‘ at www.seaaroundus.org, under ‗Hot Topic‘) which assembles all the published material that was used in enhancing the content of SeaLifeBase (www.sealifebase.org) and FishBase (www.fishbase.org) for Belize, two global information systems documenting nomenclature, geography, ecology and biology of marine organisms of the world, and which hopefully will become tools for familiarizing Belizean students with their marine biodiversity. Also, we hope that this report and the conference website will contribute to informing the national debate on oil drilling in Belizean waters. We thank Ms Audrey Matura-Shepherd and her staff at Oceana Belize for their enthusiastic assistance with the preparation of this material and the event at which it was released, and the Oak Foundation for funding the event and the preparation of this report. The Sea Around Us project, of which this report is a product, is a scientific collaboration between the University of British Columbia and the Pew Environment Group.  Too Precious to Drill: the Marine Biodiversity of Belize, Palomares and Pauly  3 INTRODUCTION OFFSHORE OIL VS 3E‘S (ENVIRONMENT, ECONOMY AND EMPLOYMENT)1 Frank Gordon Kirkwood Independent Petroleum Engineering and Economics Consultant, Belize City; kirkwoodg@ymail.com Audrey Matura-Shepherd Vice President, Oceana in Belize, 33 Cor. Regent and Dean Sts., P.O. box 1500, Belize City Belize; amatura-shepherd@oceana.org ABSTRACT Belize has a natural resource based economy and its marine resources, particularly the Belize Barrier Reef System and its accompanying Atolls are critical to tourism, Belize‘s number one foreign exchange earner, act as a natural disaster shield and provide food security, thus being a major source of jobs. Oil concessions have been granted by the Government over most of the offshore waters of Belize, including the Princess acreage with an average water depth of 4,000 ft (1,219 m), but there has been little activity to date. However, as plans move ahead to allow offshore oil exploration and drilling in the precious Belizean waters, it is important to consider the negative impact this will have on the 3E‘s: Environment, Economy and Employment. Offshore oil is being promoted as an abundant source of revenues and jobs with minimum environmental damage, yet the oil industry experience in other areas of the world and the facts and figures about Belize are saying otherwise. While the onshore oil industry (outside of the national parks) can be beneficial to Belize, the proposed offshore oil industry activity will be potentially damaging to the 3Es and thus, should not be pursued. This applies even if the additional, non-calculable, value that the reefs and atolls provide to the welfare of Belize and that no oil industry can replace, is not taken into account. INTRODUCTION Belize has a natural resource based economy and its marine resources, particularly the Belize Barrier Reef System and its accompanying atolls are critical to tourism, are Belize‘s number one foreign exchange earner, act as a natural disaster shield and provide food security, thus being a major source of jobs. Oil concessions have been granted by the Government over most of the offshore waters of Belize, including the Princess acreage with an average water depth of 4000ft, but there has been little activity to date. However, as plans move ahead to allow offshore oil exploration and drilling in the precious Belizean waters, it is important to consider the negative impact this will have on the 3E‘s: Environment, Economy and Employment. Offshore oil is being promoted as an abundant source of revenues and jobs with minimum environmental damage, yet the oil industry‘s experience in other areas of the world and the facts and figures about Belize are saying otherwise. History of the oil industry in Belize The first exploration well in Belize was drilled in 1956 by Gulf Oil in the Yalbac area in Cayo District. Between 1956 and 1982, 41 exploration wells were drilled by major oil companies such as Gulf, Philips, Anschutz, Chevron, Esso and Placid. From 1982 to 1997, only nine further exploration wells were drilled by small or independent companies, i.e., Spartan, Central Resources, Lucky Goldstar, Dover and Bright Hawk (Belize Audubon Society, 2008). Onshore and offshore seismic data was acquired during this period over a large area of the country (see Figure 1). Exploration wells drilled in Belize before 1997 found some  1 Cite as: Kirkwood, F.G., Matura-Shepherd, A., 2011. Offshore oil vs. 3E‘s (Environment, Economy & Employment). In: Palomares, M.L.D., Pauly, D. (eds.), Too Precious to Drill: the Marine Biodiversity of Belize, pp. 3-7. Fisheries Centre Research Reports 19(6). Fisheries Centre, University of British Columbia [ISSN 1198-6727]. Offshore oil vs. 3Es, Gordon Kirkwood and Matura-Shepherd  4 oil, but there were no commercial discoveries, with majority of exploration in shallow waters, except the Gladden #1 well drilled in 1997 at 1,000 ft (304.8 m) water depth (see Figure 2). In 2000, Belize passed the Petroleum Act into law, which established the framework for opening up the Belize oil industry to new concession holders. Since 2004, 19 new oil concessions have been awarded, mainly to small, newly formed oil companies; 12 concessions are for onshore and 7 are for offshore. Current onshore oil production Belize Natural Energy Ltd. made the first commercial oil discovery In the Mike Usher #1 well that was drilled in 2005 in the farming community of Spanish Lookout, between Belmopan and San Ignacio in the Cayo district. This field, for many years the only oilfield producing in Belize, was brought onto production in 2005 and reached a peak production level of 4,500 barrels per day (bpd). All oil produced onshore is exported by road tanker from the field to Big Creek port and then by sea to its point of sale, as there are no oil refining facilities in Belize. Belize Natural Energy ships oil to buyers in Costa Rica, Panama and Corpus Christi, Texas. Some crude oil is also trucked over land to El Salvador. In addition to Spanish Lookout field, the Never Delay field, which extends under Belmopan, was discovered in 2007 and is now under development with a current production rate of about 500 bpd. Figure 3 shows the locations of these onshore oil production sites. Offshore oil concessions The oil concession map, as of October 2010, is shown in Figure 4. Offshore concessions are held by 6 companies, these being: Island Oil Belize (since May 25 2004); Miles Tropical Energy Ltd. (12 Oct 2007); PetroBelize Co. Ltd. (12 Oct 2007); Princess Petroleum Ltd. (12 Oct 2007); Providence Energy Ltd. (12 Oct 2007); Sol Oil Belize Ltd. (12 Oct 2007). OPIC Resources Corp., whose concession granted in Jan 2009, withdrew in October 2010. Offshore exploration was limited with: (i) no additional seismic being acquired since 2004 despite commitments to 550 km2 by October 2011; (ii) minor relinquishments of acreage by the concessionaires despite 50% relinquishments being due by October 2011; and (iii) 2 offshore wells (one incomplete) being drilled by Island Oil in the south of Belize off Monkey River in 2007. IMPACT OF THE OIL INDUSTRY ON THE ENVIRONMENT There is a potential conflict between the oil industry and the environment both onshore and offshore as the concessions make no special recognition of national parks, marine reserves and other conservation areas as shown in Figure 5. The risks to the environment of offshore oil exploration and development are further increased by a range of factors. The award of deepwater offshore acreage to Princess Petroleum has attracted some criticism. The average water depth in the offshore part of their concession is 4,000 ft (1219 m), with depths ranging from 0 (on Lighthouse Reef Atoll) to 12,000 ft (3658 m) further out to sea. Princess Petroleum Ltd., is a hotel company and had no oil industry experience prior to being awarded this concession, which puts in question their ability to lead successful  Figure 1. Seismic surveys done in Belize during the period 1955-1997 courtesy of Geology and Petroleum Department, Government of Belize.  Figure 2. Exploration and production wells in Belize from 1955-1997 courtesy of Geology and Petroleum Department, Government of Belize. Too Precious to Drill: the Marine Biodiversity of Belize, Palomares and Pauly  5 and accident-free operations. Moreover, the Belize government lacks the offshore oil industry resources, which, in the event of accidents, prevents immediate intervention. The petroleum Industry in Belize is controlled by the Department of Geology and Petroleum (GPD) within the Ministry of Natural Resources and the Environment (Minister Hon. Gaspar Vega) within the Government of Belize (Prime Minister Hon. Dean Barrow). The GPD department is small, consisting of a Director (Andre Cho) and 6 staff, who not only deal with the oil industry, but with all mineral extraction activities in Belize as well. Belize benefits greatly from its environment. The tourism industry, based on its marine environment, is the country‘s primary money earner. In addition, the marine environment provides a significant food source for the Belizean people, i.e., fish and crustaceans. And, the barrier reef provides large scale coastal protection for Belize against tropical storms and hurricanes. The current good health of Belize‘s marine environment is already under threat from a number of sources, and, if offshore oil exploration and development goes ahead, there will be further threats to the environment in terms of: (i) impacts of seismic surveys on fish, mammals and divers; (ii) risk of oil spills, industrial discharges, drilling mud and cuttings discharges from exploration drilling; (iii) dredging, pipelaying, platform and facilities building and installation, large scale well drilling; and (iv) impact of long term industrial discharges into the marine environment. THE IMPACT OF THE OIL INDUSTRY ON THE BELIZE ECONOMY Economic data is scarce for Belize, but according to the CIA World Factbook Data as of June 2011, Belize‘s GDP for 2010 was 2.651 B USD, which grew by a dismal 2% from the 2009 record, but which saw no growth compared to 2008 with a 3.8 % growth. This is an economy in which the service sector accounts for 54% of GDP, tourism accounting for the largest portion of this sector. With a population of just over 320,000 and a labour force of 130,000, the unemployment rate is a very high at 23% (up from 13.1% in 2009 and 8.2% in 2008), with 43% of Belizeans living below the poverty line. This can easily be appreciated by the fact that exports for 2010 were reported as 404 M USD, while imports were 740 M USD. With this big gap in the balance of payments, external debt is at 1.01 B USD (2009 estimate), and growing. With conflicting reports in the media and some inconsistent data in the CIA World Facts book, we decided to have a look at the economic facts ourselves based on available raw data. Balance of payments 2011 We looked at the estimated balance of payments for 2011, i.e., a comparison of the amount of money that flows out from a country with the money that flows into a country (see Table 1). This analysis in Belize currency of the expected balance of payments in 2011 shows that there is more money flowing out (1336.3 M BzD) of the economy than there is coming in (1149.6 M BzD). The main foreign currency earners are tourism, onshore oil production and citrus and fruit production. Tourism is Belize‘s largest economic sector, which for 2011 is expected to account for 565 M BzD.  Figure 3. Onshore oil production in Belize as obtained from Belize Natural Energy Ltd.  Figure 4. Map of current petroleum contracts in Belize showing large offshore concessions held by Princess Petroleum Ltd. (in dark violet), which include most of the eastern side of Turneffe Island and the famous dive spot, the Blue Hole (a larger version of this map is reproduced in McField, p. 133, this volume) Offshore oil vs. 3Es, Gordon Kirkwood and Matura-Shepherd  6 Table 1. Belize economy, annual gains (In) and expenses (Out) in Million BzD, from major industries based on 2009 and 2011 data projections. This analysis excludes currency movements, including debt repayments and profit repatriation by foreign companies (1 BzD=0.5 USD). Industry In Out Source Tourism 565.3 – World Travel and Tourism Economic Impact Belize (2011) Petroleum (BNE) 203.8 – Based on expected BNE production level and average oil price of $87/bbl (2011) Citrus and fruit 187.3 – External Trade Bull. (December, 2009), Statist. Inst. Belize Sugar 89.1 – Same as above Fisheries 49.4 – Same as above Other exports 54.7 – Same as above Machinery and Transportation Equipment – 266.9 External Trade Bull. (December, 2009), Statist. Inst. Belize Manufactured goods – 273.0 Same as above Mineral fuel and lubes – 209.5 Same as above Commercial free zone  – 156.5 Same as above Food and live animals  – 156.5 Same as above Chemical products – 125.2 Same as above Export processing zone  – 104.9 Same as above Other imports – 43.8 Same as above  Total 1149.6 1336.3  Impact of onshore oil Without the impact of offshore oil on the 3Es, i.e., just relying on onshore oil, it is expected that in the next decade (starting in 2021), the amount of money flowing into Belize will drastically increase to $2370 M BzD with tourism accounting for almost 50% of this, and the onshore oil industry contributing 13%, with the declining contribution on BNE being replaced by revenue from oil fields resulting from the treaty energy work for Princess Petroleum Company (under their option agreement), assuming they make the finds that treaty are targeting in the Princess onshore acreage in the south of Belize. Impact of offshore oil But if offshore oil exploration goes ahead and results in discoveries in line with government expectations, production could be started by 2018, and by 2021 offshore oil could account for 70% of revenues to economy (not accounting for the portion of the oil revenues flowing out of the country as investors transfer profits to their foreign accounts). However, due to the risk of offshore oil damaging the marine environment, offshore oil would most likely result in a decline in the tourism and fisheries.   Figure 5. Concession areas in Belize include 53 defined terrestrial and marine protected areas. Adapted from the newspaper Amandala, 26/1/2011.  Too Precious to Drill: the Marine Biodiversity of Belize, Palomares and Pauly  7 THE IMPACT OF THE OIL INDUSTRY ON EMPLOYMENT Jobs picture in 2011 Employment data are even scarcer for Belize than economic data. However, the Figure 6A shows our best estimate of the jobs picture in 2011, by industry. Forecasted jobs picture in 2021 with onshore oil only With onshore oil only, by 2021 the employment picture in Belize would be much better than today (see Figure 6B). With growth in tourism and fisheries (and other economic sectors) and some additional jobs in the onshore oil industry that could be filled by Belizean workers, as Belize Natural Energy Ltd have already proved. For example, in the tourism industry, jobs are expected to grow from 39,000 in 2011 to 61,000 jobs by 2021 (i.e., almost 40% of the workforce). Forecasted jobs picture in 2021 with onshore and offshore oil The concern is that while it appears there would be more money in the economy with offshore oil, the impact on jobs will not be good, because (unlike the onshore oil industry) the environmental risks from the offshore oil industry in Belize would make a significant percentage of a large number of people who work in the tourism and fisheries industries unemployed (see Figure 6C). While the jobs created by the offshore oil industry will not be sufficient to replace these jobs. In addition because of the expertise, skills and experience these jobs require they will largely have to be filled by foreign experts who have the qualifications and offshore experience in other locations. CONCLUSIONS Thus, it can be concluded that while the onshore oil industry (outside of the national parks) can be beneficial to Belize, the proposed offshore oil industry activity will be potentially damaging to the 3Es, ―Environment, the Economy and Employment‖, and so should not go ahead. There is an additional non-calculable value that the reefs and atolls provide to the economy and welfare of Belize that no oil industry can replace. It should be noted that while petroleum is finite, the benefits of the reef should be infinite if used sustainably. REFERENCES Belize Audubon Society, 2008. An environmental agenda for Belize 2008-2013. Belize Audubon Society, Belize City. Statistical Institute of Belize, 2009. External Trade Bulletin (December). Statistical Institute of Belize, 2011. World Travel and Tourism Economic Impact Belize. Treaty Energy, 2011. Preliminary Evaluation of the Potential for Oil within the Treaty Energy / Princess Petroleum Blocks in Belize, www.treatyenergy.com accessed June 2011. World Travel and Tourism Council, 2011. Travel and Tourism Economic Impacts.      Figure 6. Employment picture in Belize estimated by economic sector, but excluding the 2011 unemployment level of 23% (i.e., 29,900 unemployed workers). A: Employment in 2011. B: Predicted employment levels in 2021 without employment by the offshore oil industry. C: Predicted employment levels in 2021 including employment by the offshore oil industry. Tourism 38% Agriculture 28% Other 27% Fisheries 6% Petroleum 1% n=101,000 jobs A Tourism 43% Agriculture 26% Other 25% Fisheries 5% Petroleum 1% n=140,000 jobs B Tourism 27% Agriculture 34% Other 32% Fisheries 4% Petroleum 3% n=108,000 jobs C The Belize Barrier Reef, Gibson  8 THE BELIZE BARRIER REEF: A WORLD HERITAGE SITE1 Janet Gibson Wildlife Conservation Society, P.O. Box 768, 1755 Coney Drive, Belize City, Belize; jgibson@wcs.org ABSTRACT The Belize Barrier Reef Reserve System (BBRRS) World Heritage Site was declared by UNESCO in 1996 as a serial nomination composed of seven marine protected areas that represent the largest barrier reef system in the Western Hemisphere. The criteria for the listing of the BBRRS were its superlative natural phenomena and natural beauty, ongoing biological and ecological processes, and biological diversity, including several threatened species. The BBRRS has one of the highest levels of marine diversity in the Atlantic. In 2009, the Site was inscribed on the List of World Heritage in Danger for several reasons: the sale and lease of public lands within the property, destruction of fragile ecosystems due to resort and housing development, and the impact of introduced species. An additional conservation issue of concern noted was the granting of offshore oil concessions. With the prospect of offshore oil exploration and drilling added to the existing threats to the Site, particularly to its coral reefs in this era of climate change, its future integrity is at risk. In addition to the value of the BBRRS in terms of tourism and recreation, fisheries, shoreline protection and other potential economic benefits, the World Heritage Site is a source of immense national pride, as the Belize Barrier Reef is emblematic of Belize‘s outstanding heritage of marine biodiversity. INTRODUCTION Prior to the inscription of the Belize Barrier Reef Reserve System as a natural World Heritage Site in November 1996 (UNESCO, 1996), the Belize barrier reef was under consideration for this designation for many years. At a workshop held in 1993, Belize decided to submit a serial nomination. Under the leadership of the GEF/UNDP project on ‗Sustainable Development and Management of Biological Diverse Coastal Resources‘, the nomination document was prepared in 1995 and submitted formally to the World Heritage Centre by the government of Belize. In January 1996, IUCN conducted a site visit of the protected areas proposed in the nomination. At the time, three of the marine reserves to be included had not yet been established, namely the Sapodilla Cayes and South Water Caye Marine Reserves, and Bacalar Chico National Park and Marine Reserve. These three protected areas, however, were then legally declared in mid 1996. The Hol Chan Marine Reserve was also included in the original proposal, but IUCN felt that it was too small an area and did not add significantly to the nomination and recommended in particular that the Blue Hole on Lighthouse Reef should be included in the nomination, due to its unique geological formation. The review (IUCN, 1996) also mentioned that the serial nomination did not include a complete cross-section of all the elements of the system (for example the Turneffe Islands), but noted these could be added at a later phase. In view of this, the Blue Hole Natural Monument was also declared a protected area in 1996, and replaced the Hol Chan Marine Reserve in the final nomination. Another concern was in relation to the need for a wider management regime to ensure the integrity of the proposed Site. This was addressed by the explanation that Belize was committed to establishing a Coastal Zone Management Authority, which would prepare a National Coastal Zone Management Plan that would provide the necessary management controls. The World Heritage Centre had a few additional queries on the submission, including a concern about future offshore oil exploration. The government provided a statement of explanation on the nature, extent and controls applying to exploratory oil drilling offshore, such as the need to go through an Environmental Impact Assessment process. With these concerns addressed, the inscription of the Belize Barrier Reef Reserve System as a World Heritage Site proceeded at the 20th ordinary session of the World Heritage Committee held in Merida, Mexico on the 2 to 7 December  1 Cite as: Gibson, J., 2011. The Belize Barrier Reef: a World Heritage Site. In: Palomares, M.L.D., Pauly, D. (eds.), Too Precious to Drill: the Marine Biodiversity of Belize, pp. 8-13. Fisheries Centre Research Reports 19(6). Fisheries Centre, University of British Columbia [ISSN 1198-6727]. Too Precious to Drill: the Marine Biodiversity of Belize, Palomares and Pauly  9 1996. It was inscribed under the natural criteria (ii) superlative natural phenomena and natural beauty, (iii) ongoing biological and ecological processes, and (iv) biological diversity, including several threatened species, as the largest reef in the Northern Hemisphere, and as a serial nomination consisting of seven protected areas (UNESCO, 1996) covering an area of 96,300 ha (IUCN, 1996). The Belize Barrier Reef system is unique for its size and array of reef types within one relatively self- contained area. It encompasses a 220 km long barrier reef, three offshore atolls, numerous patch reefs, complex mazes of faro reefs, fringing reefs, and large offshore mangrove cayes (IUCN/UNEP, 1988) all of which are represented within the World Heritage Site. In 1842, Charles Darwin referred to it as the ‗the most remarkable reef in the West Indies‘ in his book entitled The Structure and Distribution of Coral Reefs. This highly diverse system includes at least 61 coral species (Fenner, 1999), with at least 343 additional marine invertebrate species (Jacobs and Castaneda, 1998), over 500 species of fish, 45 hydroids, 350 molluscs (IUCN, 1996), and at least 70 species of ascidians, including an endemic species (Goodbody, 2000). Threatened or endangered species include staghorn coral Acropora cervicornis and elkhorn coral Acropora palmata, three species of marine turtles (hawksbill Eretmochelys imbricata, loggerhead Caretta caretta and green turtles Chelonia mydas), the American crocodile Crocodylus acutus, the great hammerhead shark Sphyrna mokarran, goliath grouper Epinephelus itajara, Nassau grouper Epinephelus striatus, and the West Indian manatee Trichechus manatus manatus. The Belize National Biodiversity Strategy states that Belizeans, along with their global partners, are dependent on biodiversity and have a responsibility to contribute towards its conservation (Jacobs and Castaneda, 1998). Indeed, IUCN noted that the history of the Belize Barrier Reef Complex illustrates the major role that reefs have played in the history of humankind, as in Belize today a large part of the economy is dependent on the reef through fisheries and tourism (IUCN, 1996). RESULTS AND DISCUSSION The seven protected areas that comprise the World Heritage Site are: Bacalar Chico National Park and Marine Reserve, Blue Hole Natural Monument, Half Moon Caye Natural Monument, Glover‘s Reef Marine Reserve, South Water Caye Marine Reserve, Laughing Bird Caye National Park, and the Sapodilla Cayes Marine Reserve (see Figure 1). The marine reserves were established under the Fisheries Act and the natural monuments and national parks were declared under the National Parks Act. Four of the protected areas are presently managed under co-management agreements between national conservation non- government organizations and the Fisheries or Forest Departments. Bacalar Chico National Park and Marine Reserve: This protected area is located on the northern end of Ambergris Caye, on the border with Mexico. The reef is representative of the northern province (Burke, 1982), and is characterized by the unusual formation of a double reef crest. A multi-species fish spawning ground is located at the reef promontory of Rocky Point, where a queen conch Strombus gigas spawning area is also located. The barrier reef also touches the shore at Rocky Point, where outcrops of Pleistocene fossilized reefs are exposed. The terrestrial component includes lagoons, salt marsh, mangroves, unique vegetation types (e.g., the kuka palm, Pseudophoenix sargentii) and some of the best littoral forest in Belize, recognized as the most threatened and under-represented ecosystem in the country (Wildtracks, 2010). The eastern beach is a nesting site for loggerhead, green and hawksbill sea turtles and the forests and wetlands support a diverse wildlife of waterbirds, a number of Yucatan endemic birds such as the Yucatan jay Cyanocorax yucatanicus and the orange oriole Icterus auratus, 36 species of reptiles including the American crocodile, and at least 31 mammals, including several species of wild cat, such as the jaguar Panthera onca. Manatees inhabit the lagoon west of the caye. Blue Hole Natural Monument: This site is a hallmark of Belize and is famous for its unique formation and geological history. Located on Lighthouse Reef Atoll, it is a circular submerged collapsed cave or sinkhole. Such cave systems formed on the offshore limestone platforms during the Pleistocene lowering of sea levels. The rim of the hole has lush coral growth, and 24 species of coral have been noted (Graham et al., 2005). The 125 m deep hole has many large stalactites (Dill, 1971) The Blue Hole is visited by great hammerhead sharks, an endangered species, as well as lemon, bull and black tip sharks (Graham et al., 2005). Kramer and Kramer (2000) highlighted the potential of unique assemblages of cryptic and endemic species occurring in this underwater cave system.  The Belize Barrier Reef, Gibson  10  Figure 1. Map showing the seven protected areas comprising the World Heritage Site  Half Moon Caye Natural Monument: This protected area is also located on Lighthouse Reef Atoll and includes Half Moon Caye and the surrounding atoll fringing reef and lagoon. The caye supports climax littoral forest that provides one of only two nesting sites in the Caribbean for the white color phase of the red-footed booby, Sula sula. The island is an important nesting site for frigate birds, and loggerhead, green and hawksbill sea turtles. The endemic island leaf-toed gecko Phyllodactylus insularis, and Allison‘s anole Anolis allisoni are also found on the caye. The natural monument is noted for its steep fore-reef wall dropping to over 3,000 feet (>914 m) where the greatest diversity of reef fish occurs (Graham et al., 2005), and it includes a reef fish spawning site. Forty-five species of coral have been documented in the protected area (Graham et al., 2005). Glover‘s Reef Marine Reserve: This reserve encompasses the entire Glover‘s Reef Atoll, which is the southernmost of Belize‘s three offshore atolls. Glover‘s Reef is considered the prototypic atoll of the Caribbean; it is not only the best developed biologically, but also possesses the greatest diversity of reef types (Dahl et al., 1974). Its deep lagoon is studded with over 800 patch reefs and pinnacles. Forty-five species of corals have been documented for the atoll (Bright and Lang, 2011). The northeastern corner of Too Precious to Drill: the Marine Biodiversity of Belize, Palomares and Pauly  11 the atoll is the site of one of the largest and last remaining Nassau grouper aggregations, which is also an aggregating site for more than 20 other reef fish species (Sala et al., 2001). All three species of marine turtles—loggerhead, green and hawksbill—occur on the atoll, which is an important foraging area for these reptiles, particularly the hawksbill turtle (Coleman, 2010). The endemic island leaf-toed gecko also occurs at Glover‘s (Wildtracks and Wildlife Conservation Society, 2007). South Water Caye Marine Reserve: The largest of the protected areas in the World Heritage Site, this reserve includes a portion of the barrier reef that is representative of the central province (Burke, 1982), characterized by well-developed reefs, such as the 9 km unbroken well-developed reef tract of Tobacco Reef with its extensive spur-and-groove system. It also includes several unique rhomboid or faro reefs, such as the Pelican Cayes, which are atolls situated on the continental shelf. The Pelican Cayes depict an unusual juxtaposition of fragile reef and mangrove communities and are considered a marine biodiversity hotspot, due to the extraordinary high diversity of sponges and tunicates on the mangrove roots and in the lagoons of the faro that is unparalleled in the Caribbean (Goodbody, 1995). For example, 70 species of ascidians or tunicates have been recorded from the area, including an endemic species (Goodbody, 2000), 31 species of bryozoans (Winston, 2007) 52 species of echinoderms, (Hendler and Pawson, 2000), 7 species of Foraminifera that include two new species (Richardson, 2000), 147 species of sponges, 45% of which are new species or variants (Rützler et al., 2000) and 148 species of algae (Littler et al., 1985). The reserve also provides habitat for American crocodiles, manatees, and sea turtles. South Water Caye Marine Reserve has many large offshore mangrove cayes or ranges that provide nesting habitat for brown boobies Sula leucogaster and frigate birds Fregata magnificens. The sand cayes are nesting sites for several tern species, including bridled terns Sterna anaethetus, least terns S. antillarum and roseate terns S. dougalli (Wildtracks, 2010). Laughing Bird Caye National Park: The National Park encompasses the entire Laughing Bird Caye faro reef, which encloses a spectacularly pinnacled lagoon and is considered one of the best examples of faro formation in the Caribbean (Wildtracks, 2010). The outer sides of the faro drop steeply to about 100 feet (30 m) to the deep channels surrounding the shelf atoll, including the Victoria Channel. The faro lagoon is noted as an important habitat for adult conch Strombus gigas, a species that is listed under Appendix II of CITES. The long narrow sand and shingle caye lies on the steep side of the faro, and is named for the laughing gull Larus artricilla that used to nest on the island. It is an important nesting ground for the hawksbill turtle. Sapodilla Cayes Marine Reserve: The reefs of the Sapodilla Cayes Marine Reserve are located at the southern end of the barrier reef, forming a unique hook-shaped structure (Kramer and Kramer, 2000). They are representative of the discontinuous reefs of the southern province of the barrier reef (Burke 1982) and have extensive spur-and-groove formations extending eastward. The reserve has the highest coral diversity in Belize (Wildtracks, 2010) and includes three fish aggregating sites, at Nicholas Caye, Rise and Fall Bank, and Seal Caye, all important for the endangered Nassau grouper and other reef fish. The reserve encompasses 14 sand and mangrove cayes. Hunting Caye is a nesting site for the highly endangered hawksbill turtle. As the Belize Barrier Reef Reserve System is a serial nomination, other protected areas can be added to the World Heritage Site, and recommendations have been made to include the Gladden Spit and Silk Cayes Marine Reserve and the Port Honduras Marine Reserve. Progress on nominating these additional areas, however, has been delayed due to the recent inscription of the Site on the List of World Heritage in Danger in June 2009 (UNESCO World Heritage Committee - Decision - 33 COM 7B.33). In 2008, concerns were reported to the World Heritage Centre regarding extensive mangrove cutting, dredging and infilling in the Pelican Cayes region of the South Water Caye Marine Reserve. In addition, news of an impending sale of 3,000 ha of land in the Bacalar Chico National Park also raised concerns, although plans for the sale were later cancelled (UNESCO World Heritage Committee - 08/32.COM/7B). As a result of these concerns, a mission from the World Heritage Centre visited Belize in March 2009 to assess the status of the Site. The report on the mission noted (UNESCO WHC - 09/33.COM/7B.Add), inter alia, that several dozen transfers of public lands were made for development purposes since the original inscription of the Site in 1996 and as a result the Outstanding Universal Value had been affected by this ongoing development on the cayes. It also noted that the Coastal Zone Management Authority and Institute were not able to carry out their mandate and that there was poor coordination between the government agencies responsible for overall management of the World Heritage Site. Other concerns The Belize Barrier Reef, Gibson  12 included illegal fishing, particularly within the no-take zones, and potential impacts by introduced invasive species. The report highlighted the corrective measures that Belize needed to implement. In August 2010, the World Heritage Committee decided to retain the Belize Barrier Reef Reserve System on the List of World Heritage in Danger (UNESCO World Heritage Committee - Decision - 34 COM 7A.13). The main concern noted as part of this decision was in relation to oil concessions reportedly granted within the marine area of the property, as oil exploration is considered incompatible with World Heritage status, and Belize was urged to enact legislation to prohibit oil exploration within the Site. More than 75% of coral reefs in the Caribbean are considered threatened (Burke et al., 2011). Climate- related threats are expected to increase the proportion of reefs at risk to 90% in 2030 and up to 100% by 2050 (Burke et al., 2011). The recent report card on the health of Belize‘s reefs showed that 65% of the reef is rated as being in poor or critical health, with only 1% considered in very good health (Healthy Reefs, 2010). It is clear that the threats to Belize‘s reefs need to be reduced in order to promote their recovery. Offshore exploration for oil, however, will increase the threats to the reef system. Shallow coral reefs, seagrass beds and mangroves, which characterize the Belize Barrier Reef Reserve System, are among the most sensitive environments to oil, with mangroves being the most susceptible (Guzman et al., 1991). Furthermore, it is difficult to carry out any oil spill mitigation measures for these habitats. Finally, the Belize Barrier Reef Reserve System is representative of Belize‘s marine system, which has been valued at 231-347 M USD year-1 in terms of the contribution of the coral reefs and mangroves to shoreline protection, tourism and fisheries (Cooper et al., 2009). Loss or damage to these critical ecosystems will result in a decline in the valuable services they provide to the country. Importantly, a recent study has shown that biodiversity losses due to human disturbance are raising concerns about the future functioning of ecosystems and their ability to deliver goods and services to humanity. For example, reef fish systems function better in terms of standing biomass with the addition of species or increased diversity (Mora et al., 2011). Thus the consequences of losing biodiversity are even greater than previously thought and could be devastating for coral reef fisheries. All efforts should be made to protect the biodiversity of the Belize Barrier Reef Reserve System World Heritage Site, as it is integrally connected to the human development and national heritage of Belize. ACKNOWLEDGEMENTS I wish to thank Claire Gibson for reviewing the manuscript and Virginia Burns for preparing the map. REFERENCES Bright, T., Lang, J., 2011. Picture guide to stony corals of Glover‘s Reef Atoll. Created for the Wildlife Conservation Society, Glover‘s Reef Research Station, Belize. (www.gloversreef.org) Burke, L., Reytar, K., Spalding, M., Perry, A., 2011. Reefs at Risk Revisited. World Resources Institute, Washington DC. Burke, R.B., 1982. Reconnaissance study of the geomorphology and benthic communities of the outer barrier reef platform, Belize. In: K. Rützler and I.G. Macintyre (eds.), The Atlantic Barrier Reef Ecosystem at Carrie Bow Cay, Belize I. Structure and Communities, p. 509-526. Smithsonian Inst. Press, Washington, D.C. Coleman, R., 2010. In-water surveys of marine turtles at Glover‘s Reef Marine Reserve. April 2010. Report to Wildlife Conservation Society. Cooper, E., Burke, L., Bood, N., 2009. Coastal Capital: Belize. The Economic Contribution of Belize‘s Coral Reefs and Mangroves. WRI Working Paper. World Resources Institute, Washington DC. 53 pp. Dahl, A.L., Macintyre, I.G., Antonius, A., 1974. A comparative study of coral reef research sites. Atoll Research Bulletin 172, 37-120. Dill, R.F., 1971. The Blue Hole: a structurally significant sink hole in the atoll of British Honduras. Geol. Soc. Amer., Abstracts with Programs 3, 544-545. Fenner, D., 1999. New observations on the stony coral (Scleractinia, Milleporidae, and Stylasteridae) species of Belize (Central America) and Cozumel (Mexico). Bulletin of Marine Science 64, 143-154. Goodbody, I.G., 1995. Ascidian communities in Southern Belize - a problem in diversity and conservation. Aquatic Conservation: Marine and Freshwater Ecosystems 5, 355-358. Goodbody, I.G., 2000. Diversity and distribution of ascidians (Tunicata) in the Pelican Cays, Belize. Atoll Research Bulletin 480. Graham, R.T., Hickerson, E., Barker, N., Gall, A., 2005 Rapid Marine Assessment Half Moon Caye and Blue Hole Natural Monuments Lighthouse Reef Atoll, Belize, 7-18 December 2004. Prepared for the Belize Audubon Society. Guzman, H.M., Jackson J.B.C., Weil, E., 1991. Short-term ecological consequences of a major oil spill on Panamanian subtidal reef corals. Coral Reefs 10, 1-12. Too Precious to Drill: the Marine Biodiversity of Belize, Palomares and Pauly  13 Healthy Reefs for Healthy People, 2010. Report Card for the Mesoamerican Reef. An evaluation of ecosystem health 2010. Hendler, G., Pawson, D.L., 2000. Echinoderms of the Rhomboidal Cays, Belize: Biodiversity, Distribution and Ecology. Atoll Research Bulletin 479, 275-299. IUCN, 1996. World Heritage Nomination – IUCN Summary, Belize Barrier Reef Reserve System (Belize). May 1996. IUCN, 1996. World Heritage Nomination IUCN Technical Evaluation. Belize Barrier Reef Reserve System (Belize). October 1996. IUCN/UNEP, 1988. Coral Reefs of the World. Volume 1: Atlantic and Eastern Pacific. UNEP Regional Seas Directories and Bibliographies. IUCN, Gland, Switzerland and Cambridge, U.K./UNEP Nairobi, Kenya xivii + 373 pp., 38 maps. Jacobs, N.D., Castaneda, A. (Editors), 1998. Belize National Biodiversity Strategy. National Biodiversity Committee, Ministry of Natural Resources and the Environment, Belmopan, Belize, Central America. Kramer, P.A., Kramer, P.R., 2000. Ecological status of the Mesoamerican Barrier Reef System: impacts of hurricane Mitch and 1998 coral bleaching. Final Report to the World Bank. Littler, D.S., Littler, M.M., Brooks, B.L., 2000. Checklist of Marine Algae and Seagrasses from the Ponds of the Pelican Cays, Belize. Atoll Research Bulletin 474, 153-208. Mora, C., Aburto-Oropeza, O., Ayala Bocos, A., Ayotte, P.M., Banks, S. et al., 2011. Global human footprint on the linkage between biodiversity and ecosystem functioning in reef fishes. PLoS Biol 9(4), e1000606. Richardson, S.L., 2000. Epiphytic Foraminifera of the Pelican Cays, Belize: Diversity and Distribution. Atoll Research Bulletin 475, 209-230 Rützler, K., Diaz, M. C. , van Soest, R.M.W., Zea, S., Smith, K.P., Alvarez, B., Wulff, J., 2000. Diversity of sponge fauna in mangrove ponds, Pelican Cayes, Belize. Atoll Research Bulletin 467, 230-250 Sala, E., Ballesteros, E., Starr, R.M., 2001 Rapid decline of Nassau grouper spawning aggregations in Belize: fishery management and conservation needs. Fisheries 26(10), 23-30. UNESCO, 1996. World Heritage Committee Report on Twentieth Session, Merida, Yucatan, Mexico, 2 – 7 December 1996. Wildtracks and Wildlife Conservation Society, 2007. Management Plan Glover‘s Reef Marine Reserve and World Heritage Site 2008- 2013. Wildtracks, 2009 Management Plan South Water Caye Marine Reserve World Heritage Site 2010-2015. Wildtracks 2010. Directory of Protected Areas of Belize. Winston, J.E., 2007. Diversity and distribution of bryozoans in the Pelican Cays, Belize, Central America. Atoll Research Bulletin 546, 1-26.  Threats to bottlenose dolphins from oil exploration, Hines  14 BIODIVERSITY THREATS TO COASTAL DOLPHINS FROM OIL EXPLORATION, DRILLING AND SPILLS OFF THE COAST OF BELIZE1 Ellen Hines Marine and Coastal Conservation and Spatial Planning Center, Department of Geography and Human Environmental Studies, San Francisco State University 1600 Holloway Ave., San Francisco, CA 94132 USA, ehines@sfsu.edu ABSTRACT Protected from import/export, wildlife trade, and hunting by Belize‘s 1981 Wildlife Protection Act, threats from human induced mortality to coastal dolphins are currently minimal. Dolphins along the coast, islands and offshore areas of Belize are distributed in small, thus vulnerable population groups. Currently, unsustainable fishing (overfishing and illegal fishing) causing prey depletion and indirect capture, rapid coastal development (mangrove clearing, dredging, and coastal development), increasing vessel traffic and pollution have been identified as growing threats as human populations and coastal use grow. However, oil exploration, drilling and the possibility of spills off the coast of Belize are additional threats. The increases in shipping due to oil and gas exploration are likely to lead to acoustic disturbances and ship strikes for Belizean dolphins. Seafloor exploration for oil resources involves seismic testing. The loud, broad band sounds produced by seismic air guns have been shown to cause avoidance and other behavioural responses in beluga whales and other odontocete (toothed) species, which could lead to long- term adverse effects on populations. Seismic impulses can travel for long distances, and in some cases have been detected over 3000 km from their source. While the effects of air gun noise vary, other observed effects include auditory damage and decompression sickness. Seismic airguns may also affect prey including fish and squid. During drilling and production, populations of dolphins would be vulnerable to the cumulative effects of chronic oil pollution from small tanker spills, pipeline leaks and other accidents. A catastrophic oil spill would be extremely harmful. While cetaceans are less vulnerable to oiling than many other marine species such as otters and seabirds, oil may damage the eyes, and inhalation of surface vapours can damage their lungs. Also, oil spills may have long-term impacts on prey populations such as fish and benthic invertebrates. INTRODUCTION Protected from import/export, wildlife trade, and hunting by Belize‘s 1981 Wildlife Protection Act, threats from human induced mortality to coastal dolphins are currently minimal. Dolphins along the coast, islands and offshore areas of Belize are distributed in small, thus vulnerable population groups. Currently, unsustainable fishing (overfishing and illegal fishing) causing prey depletion and indirect capture, rapid coastal development (mangrove clearing, dredging, and overdevelopment), increasing vessel traffic and pollution have been identified as growing threats as human populations and coastal development increase. As the possibility of off-shore oil drilling in Belize is explored, research results and recommendations as to the effects of petroleum development need to be heeded. Threats to Belize‘s dolphins can be identified in all stages of offshore petroleum industry activities: exploration, production, transportation and accidents. There are two species of dolphins found along the coast of Belize: bottlenose (Tursiops truncatus) and rough-toothed (Steno bredanensis), bottlenose being the most predominant. The IUCN does not currently list a population estimation for bottlenose dolphins in the Caribbean (Hammond et al., 2008a). Although considered a species of ‗least concern‘ internationally, specific demographic information about bottlenose  1 Cite as: Hines, E., 2011. Threats to coastal dolphins from oil exploration, drilling and spills off the coast of Belize. In: Palomares, M.L.D., Pauly, D. (eds.), Too Precious to Drill: the Marine Biodiversity of Belize, pp. 14-18. Fisheries Centre Research Reports 19(6). Fisheries Centre, University of British Columbia [ISSN 1198-6727]. Too Precious to Drill: the Marine Biodiversity of Belize, Palomares and Pauly  15 dolphins in Belize has not been established. While projects have been conducted since 1992 on various topics, research has not been country-wide. We cannot estimate the number of animals within Belize as a whole, or clarify dolphin movements and home ranges. Research on bottlenose dolphins has led scientists to consider them as inhabitants of distinct long-term stocks. For instance, there are 32 provisionally identified stocks or communities identified in the Gulf of Mexico, based on geographic, genetic and social relationships (Waring et al., 2009; Wells et al., 1987). While these communities are not closed demographic populations, as there is known interbreeding with adjacent communities, residents interact with each other, share habitat areas, and have similar distinct genetic profiles. These communities have multi-year, multi-generational patterns within a geographic area such that the dolphins are considered integral units within the ecosystems they inhabit. Thus, if the home range of such a community were eradicated or severely disturbed, it would take a long time to repopulate the dolphin population and resulting effects of this on local ecology is unknown. Also, bottlenose dolphins are known to be predominantly coastal. They have been seen in pelagic areas, and classified as either coastal or offshore, with varying foraging behavior. Offshore animals have been seen as residents on offshore islands, which is relevant to the outer atolls of Belize (Wells and Scott, 1999). Without concrete data to support large-scale management, it is more appropriate to consider a precautionary approach to conservation-oriented management at the community or stock level. Rough-toothed dolphins are found globally in tropical to subtropical oceans in both oceanic and deeper continental shelf waters (Hammond et al., 2008b). Limited survey efforts have been conducted in the U.S. Gulf of Mexico (although recent status reports cautiously estimate 2,653 animals). No formal surveys have been conducted in the Caribbean which leaves no information or current population estimate for Belizean waters. As there is no recorded major human disturbance currently, the dolphin is not on the U.S. endangered species list and is considered a species of least concern globally on the IUCN Red List (IUCN, 2011). However, there are insufficient data from which to determine population trends for this species (Waring, 2009). While rough-toothed dolphins are not seen in the same structured communities as bottlenose dolphins, lack of knowledge of their regional numbers, distribution and habitat use should be considered cause for risk-averse management. Dolphins in Belize Bottlenose dolphins in Belize have primarily been studied in the Drowned Cayes and Turneffe Atoll (Figure 1). Both sites have dolphins present year-round within the unique and highly productive combination of mangrove, seagrass and coral habitats. Drowned Cayes The latest information on bottlenose dolphin research in the Drowned Cayes is from Kerr et al., (2005). This paper documents photo-identification research between 1997-1999. The authors estimated the 122 (95% CI = 114 -140) animals there as a closed population. When comparing this population with research at Turneffe Atoll, they noted similar small group sizes, variable levels of site fidelity and low abundance, but did not find any overlap in sightings. Kerr et al., (2005) noted the proximity of the Drowned Cayes to mainland Belize, exposing the dolphins to increased risks of pollution, boat traffic, resource extraction and  Figure 1. The Drowned Cayes and Turneffe Atoll (adapted from belizetravelmall.com).  Drowned Cayes Threats to bottlenose dolphins from oil exploration, Hines  16 increasing levels of fishing extraction. They mentioned that research has noted that long-term overfishing of Caribbean reefs has already affected fish community structures and coral ecosystems (Sedberry et al., 1999; Jackson et al., 2001). Turneffe Atoll Turneffe Atoll is relatively pristine, further from the coastal development of mainland Belize, with few year-round inhabitants. However, increasing development due to ease of access since an airstrip was built in 2004, and the expansion of tourism and cruise ship visits are unaddressed threats. Various researchers, since 1992, have studied the Atoll‘s dolphins (a summary of Turneffe Atoll dolphin research can be found in Dick, 2008). Relevant here are results that showed a combination of continuous and seasonal residents with an estimated population of 216 dolphins (CV = 27.7%), with most sightings in channels between mangroves and reefs, and a relatively large seasonal population of mothers and calves (Grigg and Markowitz, 1997; Grigg, 1998; Dick and Hines, 2011). Threats to dolphins noted here include unsustainable fishing and illegal gillnetting by Guatemalan and Honduran fishers. Dredging of mangrove and seagrass for development can impact local fish and trophic levels (Dick and Hines, 2011). Threats Offshore oil activities can be threatening to marine mammals in various ways. Habitats can be altered and behavior disturbed by noise from seismic surveys, shipping and drilling. Related pollutants can be chronically released, and there is a real risk of an accidental oil spill from platforms and tankers. As oil activities grow, they can create cumulative impacts on near and offshore ecosystems. As oil tanker traffic increases, the chances for shipboard spills and collisions increase as well (Huntington, 2009). As shown in Figure 2, each phase of the offshore oil activity listed has its own potential threats. Note that noise as a potential hazard is associated with each listed activity: seismic surveying, drilling, air and ship support, construction and operation (Geraci and St. Aubin, 1980). Exploration Seafloor exploration for oil resources involves increases in shipping, which generate their own noise and dangers of collision with dolphins. However, seismic testing associated with offshore oil exploration is one of the most intense anthropogenic noises in the ocean and often are implemented over large areas for extended periods (Gordon et al., 2003). The loud, broad-band sounds produced by seismic air guns have been shown to cause avoidance and other behavioral responses in beluga whales and other odontocete (toothed) species, which could lead to long-term adverse effects on populations. Seismic impulses can travel for long distances, and in some cases have been detected over 3000 km from their source. While the effects of air gun noise vary, other observed effects include auditory damage and decompression sickness. Seismic airguns may also affect prey including fish and squid. And while these are problematic, seismic surveys are only one element of the noise contribution of exploration. Field development and construction,  Figure 2. Proximate factors associated with offshore oil exploration and their effects on marine mammals (Geraci and St.Aubin, 1980). Too Precious to Drill: the Marine Biodiversity of Belize, Palomares and Pauly  17 exploratory drilling, underwater acoustic communication, equipment placement, and sea-floor processing all generate their own particular noises (Fernández et al., 2004; Gordon et al., 2003; Geraci and St. Aubin, 1980; M. Stocker, 2011, pers.comm.). While most of the sound of the seismic air guns is around 100 Hz, there is leakage of sounds at higher frequencies (around 200-500 Hz), closer to the hearing range of odontocetes (between 200 Hz and 100 kHz; Ketten, 1998), that are audible between 10 to 100km. Dolphins have been observed both less frequently and not vocalizing as usual during seismic surveys (Harwood and Wilson, 2001). The potential effects of low-frequency sounds on marine mammals as assessed by the U.S. Marine Mammal Commission (Anon., 1998) includes, in decreasing order of severity:  Death from lung hemorrhage or other tissue trauma;  Permanent or temporary hearing loss or impairment;  Disruption of feeding, breeding, nursing, acoustic communication and sensing, or other vital behavior. If severe, frequent or long lasting this could lead to a decrease in individual survival and productivity and a corresponding decrease in population size and productivity;  Abandonment or avoidance of traditional feeding, breeding or other biologically important habitats, again with possible effects on survival, productivity and population size;  Psychological and physiological stress making animals more vulnerable to disease, parasites and predators;  Changes in the distribution, abundance or productivity of important marine mammal prey species with consequent effects on individual survival, productivity and population size. Production and transportation During drilling and production, populations of dolphins would be vulnerable to the cumulative effects of chronic oil pollution from small tanker spills, pipeline leaks and other accidents. There are increased risks of ship collisions. As seen in Figure 2, noise from drilling is also present. Ships employed in transportation of oil products and movement of materials and personnel are of varied sizes, but can be responsible for underwater noise that can spread from dozens to hundreds of kilometers. Larger ships generate more low- frequency sounds that are less dangerous to small cetaceans such as dolphins, however, Würsig and Greene (2002) studied the effects of small and medium tankers in Hong Kong on small local cetaceans and found that noise from these vessels could interfere with passive listening for prey, and possibly communication. Results also suggested possibilities for increased stress or permanent shifts in hearing levels. Accidents A catastrophic oil spill would be extremely harmful. Ninety-two percent of dolphins found in the oiled areas within a year of the Deepwater Horizon BP oil spill were dead (NOAA, 2011). Marine mammals such as dolphins and whales must come to the surface to breathe; inhaling spilled petroleum products can expose them to floating oil. Ingested oil can kill animals immediately; more often it results in lung hemorrhaging, liver, and kidney damage which can lead to death. Oil accumulated on the skin of animals can make it difficult to breathe and move in the water. While cetaceans, with their smoother skin, are less vulnerable to oiling than many other marine species such as otters and seabirds, oil may damage the eyes. Oil spills may also have long-term impacts on prey populations such as fish and benthic invertebrates (Geraci and St. Aubin, 1980; IWC, 2010). DISCUSSION Dolphins in Belize are already exposed to the cumulative effect of unsustainable fishing (over fishing and illegal fishing) causing prey depletion and indirect capture, rapid coastal development (mangrove clearing, dredging, and overdevelopment), increasing vessel traffic and pollution have been identified as growing threats as human populations and coastal use grow. While there is more information about bottlenose than rough-toothed dolphins, both species are found in small population groups, susceptible to loss of animals. Mother and calf pairs of bottlenose dolphins found seasonally (see above) in Turneffe Atoll could be especially vulnerable. Habitat destruction would damage the structure of bottlenose dolphin population communities which would increase recovery time and amplify disturbance to local trophic systems. Oil exploration, drilling and the possibility of spills off the coast of Belize are a quite severe danger to small Threats to bottlenose dolphins from oil exploration, Hines  18 cetaceans. Toxicity as a result of oil spill accidents, long term seepages, and leaks from vessels can add to a synergistic mixture of threats whose exact effects are still largely unknown. It is clear however, that a decision to reject offshore oil is a precautionary measure that will go a long way towards protecting Belize‘s dolphins. ACKNOWLEDGEMENTS I would like to thank Sea Around Us Project, in particular Daniel Pauly and Deng Palomares for organizing the workshop at which this was presented, and for their invitation to participate. My research in Belize was funded by the Oceanic Society and Birgit Winning, and assisted by my graduate students, Suzanne Holguin, Stefanie Egan, Dori Dick and Sadie Waddington. The Oceanic Society and associated researchers have laid the foundation for what knowledge we have about dolphins in Belize. REFERENCES Anon, 1998. Annual Report to Congress 1997. Marine Mammal Commission, Bethesda, Maryland. Belize Wildlife Protection Act, Chapter 220, Revised Edition 2000. Dick, D.M., 2oo8. Abundance and spatial analysis of bottlenose dolphins at Turneffe Atoll, Belize. Masters Thesis, Department of Geography and Human Environmental Planning, San Francisco State University, San Francisco. Dick, D.M., Hines, E.M., 2011. Development and implementation of distance sampling techniques to determine bottlenose dolphin abundance at Turneffe Atoll, Belize. Marine Mammal Science 27(3), 606-621. Fernández, A., Arbelo, M., Deaville, R., Patterson, I.A.P., Castro, P., Baker, J.R., Degollada, E., Ross, H.M., Herráez, P., Pocknell, A.M., Rodríguez, F., Howie, F.E., Espinosa, A., Reid, R.J., Jaber, J.R., Martin, V., Cunningham, A.A. and Jepson, P. 2004. Whales, sonar and decompression sickness. Nature 428, 1-2. Gordon, J., Gillespie, D., Potter, J., Frantzis, A., Simmons, M.P., Swift, R., Thompson, D., 2003. A review of the effects of seismic surveys on marine mammals. Marine Technology Society Journal 37, 16-34. Hammond, P.S., Bearzi, G., Bjørge, A., Forney, K., Karczmarski, L., Kasuya, T., Perrin, W.F., Scott, M.D., Wang, J.Y., Wells, R.S., Wilson, B., 2008a. Tursiops truncatus. In: IUCN 2011. IUCN Red List of Threatened Species. Version 2011.1. <www.iucnredlist.org>. Hammond, P.S., Bearzi, G., Bjørge, A., Forney, K., Karczmarski, L., Kasuya, T., Perrin, W.F., Scott, M.D., Wang, J.Y., Wells, R.S. and Wilson, B. 2008b. Steno bredanensis. In: IUCN 2011. IUCN Red List of Threatened Species. Version 2011.1. <www.iucnredlist.org>. Harwood, J., Wilson, B., 2001. The implications of developments on the Atlantic Frontier for marine mammals. Continental Shelf Research 21, 1073-1093 IWC (International Whaling Commission), 2010. Opening statement of the animal welfare institute to the 62nd meeting of the International Whaling Commission. http://iwcoffice.org/_documents/commission/IWC62docs/62-OS%20NGO.pdf Jackson, J.B.C., et al., 2001. Historical overfishing and the recent collapse of coastal ecosystems. Science 293:629-638. Kerr, K.A., R.H. Defran, Campbell, G.S., 2005. Bottlenose dolphins (Tursips truncates) in the Drowned Cayes, Belize: Group size, site fidelity and abundance. Caribbean Journal of Science 41, 172-177. Ketten, D.R., 1998. Marine mammal auditory systems: a summary of audiometric and anatomical data and its implications for underwater acoustic impacts. NOAA Technical Memorandum NMFS. NOAA Office of Protected Resources, 2011. http://www.nmfs.noaa.gov/pr/pdfs/oilspill/species_data.pdf Sedberry, G.R., Carter, H.J., Barrick, P.A., 1999. A comparison of fish communities between protected and unprotected areas of the Belize reef ecosystem: implications for conservation and management. Proceedings of the Gulf and Caribbean Fisheries Institute 45:95-127. Waring G.T., Josephson E., Maze-Foley K., Rosel, P.E., Editors, 2009. U.S. Atlantic and Gulf of Mexico Marine Mammal Stock Assessments -- 2011. NOAA Tech Memo NMFS NE 219; 598 p. Available from: National Marine Fisheries Service, 166 Water Street, Woods Hole, MA 02543-1026, or online at http://www.nefsc.noaa.gov/nefsc/publications/ Wells, R.S., Scott, M.D., Irvine, A.B., 1987. The social structure of free-ranging bottlenose dolphins. In Current Mammology Vol. I. ed. H.H. Genoways, 247-305. Plenum Press, New York and London. Wells, R.S., Scott, M.D., 1999. Bottlenose dolphin Tursiops truncatus. In Ridgeway, S.H., Harrison, R.J. (eds.), Handbook of Marine Mammals:Volume VI, The Second Book of Dolphins and Porpoises, pp. 137-182. Academic Press, San Diego, CA . Würsig, B., Greene, Jr., C.R., 2002 Underwater sounds near a fuel receiving facility in western Hong Kong: relevance to dolphins. Marine Environmental Research 54, 129-145.  Too Precious to Drill: the Marine Biodiversity of Belize, Palomares and Pauly  19 THE FATE OF MANATEES IN BELIZE1 Nicole Auil Gomez #14 Princess Margaret Drive, Belize City, Belize; nauilgomez@gmail.com ABSTRACT Sirenians (manatees and dugongs) are the only fully aquatic, herbivorous marine mammals existing today and Belize boasts the largest number of Antillean manatees in the world. Yet, the country‘s manatee population is considered threatened and may be declining. Manatees have to contend with high-speed watercraft that account for over 20% of their mortality. Also, intentional habitat alteration and industrial practices fragment and destroy the ecosystem they depend upon. Land-based effluent has decimated sub- aquatic vegetation and has likely compromised individual manatee health in areas such as Placencia Lagoon. High levels of toxic trace elements, including lead, were also found in manatees captured there. With limited data on the threats of contaminants to manatees, a pilot study showed that organic contaminants (polychlorinated biphenyls - PCBs) in manatees from Chetumal Bay may currently present a threat to their immune function and reproduction. Marine currents may allow PCBs to be present at a regional level. Also, as radio-tracked manatees have been documented to travel between Belize and Chetumal Bay, they are further exposed to organic compounds through inadvertent consumption of sediment during grazing. Added petrochemicals would further contaminate and destroy manatee feeding areas as the toxic components of oil are thought to accumulate in seagrass leaves, making vegetation vulnerable to these stressors. After an oil spill, manatees, dolphins and turtles are exposed to volatile hydrocarbons while traveling and feeding, as shown from surveys following the 2010 BP Deepwater Horizon catastrophe in the Gulf of Mexico. While experiments on captive marine mammals indicate that manatees can withstand small amounts of exposure to, or ingestion of, oil, it is not certain if these animals can detect, avoid, or leave a contaminated area before experiencing significant harmful effects. With very limited data on the effect of oil-related stressors to sirenians, we know that the threats they face today, compounded with the incalculable environmental damage of an oil-related disaster, would certainly affect the chances for survival of the endangered manatees in Belize. INTRODUCTION Three species of manatees are included within the Mammalian Order Sirenia: the Amazonian manatee (Trichechus inunguis), the West African manatee (T. senegalensis), and the West Indian manatee (T. manatus). Dugongs (Dugong dugon) are also in this Order, and live exclusively in the Pacific Ocean. The West Indian manatee is divided into two subspecies: Florida (T. manatus latirostris) and Antillean (T. m. manatus). This species inhabits fresh, brackish and marine waters in the Wider Caribbean, from Florida to the northeastern coast of South America. They need access to freshwater, and thus remain in relatively shallow coastal waters (within 3 m depth; Hartman, 1979). They are the only fully aquatic extant herbivorous marine mammal. With a low reproductive rate and historically hunted throughout its range, manatees are considered vulnerable to extinction by the IUCN (Thornback and Jenkins 1982). Given the small population size and past exploitation, today Belize‘s manatee population has a low genetic diversity (low levels of haplotype diversity, microsatellite heterozygosity and allelic variation), as compared with other marine mammals, and endangered or bottlenecked populations (Hunter et al., 2010). Belize harbors the highest known density of Antillean manatees in the Caribbean (O‘Shea and Salisbury, 1991; Auil, 1998). Status surveys were first conducted 32 years ago by Charnock-Wilson (1968) and Charnock-Wilson, Bertram and Bertram (1974), and later by Bengtson and Magor (1979) and O‘Shea and Salisbury (1991). Further countrywide surveys have shown that while manatees are present along most of Belize‘s coastline from the Rio Hondo to Sarstoon, areas of consistently high presence include the  1 Cite as: Auil Gomez, N., 2011. The fate of manatees in Belize. In: Palomares, M.L.D., Pauly, D. (eds.), Too Precious to Drill: the Marine Biodiversity of Belize, pp. 19-24. Fisheries Centre Research Reports 19(6). Fisheries Centre, University of British Columbia [ISSN 1198-6727]. The fate of manatees of Belize, Auil Gomez  20 Southern Lagoon, the Belize River and Belize City Caye area, Placencia Lagoon, Port Honduras, Corozal Bay, and Indian Hill Lagoon (O‘Shea and Salisbury, 1991; Gibson, 1995; Auil, 1998; Morales-Vela et al., 2000). Additionally, while there are higher numbers of manatees in the larger caye and coast habitats, the lagoon and river systems have a higher probability of manatee occurrences (Auil, 2004). The highest count during a survey was 338 manatees, so today, population estimates are approximated at about 1000 manatees in Belizean waters (O‘Shea and Salisbury, 1991; Auil, 1998; Auil, 2004). Although a developing country with limited available resources, Belize has proven to be a successful manatee conservation site. A rapidly growing and lucrative ecotourism industry is attractive to economically depressed communities, offering excellent prospects for touristic development that values wildlife and their habitats. There are several communities along the coast of Belize with facilities or activities for receiving visitors, and the clear coastal waters provide good conditions for viewing manatees for both scientific study and tourism. Fortunately, the country has taken the lead in manatee conservation and coastal conservation efforts in Central America, where Belize‘s efforts provide a model that can be used by other countries. Belize has an interagency management working group (Belize Manatee Working Group) that comprises government representatives, NGOs and scientists that carry out research and conservation initiatives throughout the country. Additionally, there are three sites declared as wildlife sanctuaries specifically for manatees: Corozal Bay, Swallow Cay, and Southern Lagoon. Unfortunately, very little data exists on chemical contaminants on manatees, including heavy metals, trace elements and organic compounds. Virtually no data exists on the effect of oil on these coastal marine mammals. When indentified in samples, the biological response of an individual manatee to a contaminant is not always determined and cumulative response to toxins is unknown. In this report, I outline some research to describe our current knowledge of threats faced by manatees in Belize. THREATS Consistent assessment of manatee status began in August 1996 under the Coastal Zone Management Project at the commencement of the National Manatee Project. Threats to manatees, particularly in Belize, are predominantly anthropogenic. From January 2005 to December 2010, 76 reports of manatee strandings were received (Figure 1; Galves, 2011). Twenty-nine percent of these were unverified. For the carcasses that were located, examiners could not determine the cause of death for 38%, primarily as 43% of this ‗undetermined‘ category was in an advance stage of decomposition or was not recorded. The number of strandings per year ranged from six in 2008 to 18 in 2010. There has been a recent increase in the number of strandings over the years that have researchers concerned. Watercraft collision has been the primary cause of identified death for each year (range 14% - 27%). Habitat alteration While we are unable to directly quantify manatee loss based on habitat destruction, we know that a reduction of the quality or quantity of the submerged, natant or overhanging vegetation they rely upon impacts manatee health. This is particularly so as their habitat, primarily seagrass beds, become fragmented. The health of seagrass meadows in shallow aquatic ecosystems varies with sediment and nutrient loading. Excessive nutrient loading can smother seagrass by stimulating the production of algae that block out light. Sediment re-suspension due to wind, riverine transport or boat traffic can also lower water clarity and produce the same result. Human activities that accelerate contaminant loading or over- exploit sensitive species can seriously impair and threaten these ecosystem services. These changes may persist long after disturbances have ceased, rendering habitats unsuitable. One case of significant sub- aquatic vegetation loss in a primary manatee habitat is in Placencia Lagoon, where formerly viable seagrass meadows have deteriorated rapidly due to contaminated run-off.   Figure 1. Cause of manatee strandings 2005 - 2010. Undetermined 38% Unverified 29% Watercraft 17% Live 7% Poached 5% Perinatal 3% Other 1% Too Precious to Drill: the Marine Biodiversity of Belize, Palomares and Pauly  21 The seagrass Halophila baillonii provided the preferred forage for manatees in Placencia Lagoon (Short et al., 2006; Auil Gomez, personal observation). A baseline study revealed that many areas in the lagoon retained critical ecosystem functions, populations of ecologically unique species, and seagrasses, and had little effluent influence prior to 2003 (Smith and Mackie, 2005). Recent data indicate those seagrass meadows were lost in the three years following, coinciding with aquaculture effluent (Gallego, 2004; Ledwin, 2010). The far northern basin of the lagoon has demonstrated low water clarity and sparse populations of submersed aquatic plants since sampling began in 2003 (Smith and Mackie, 2005). In 2006, in preparation for the first manatee capture event, the predominant vegetation recorded in the northern to central part of the lagoon was Halophila, along with some patches of Halodule and Chara. The leaves of the vegetation in the northern part of the lagoon were covered with strands of filamentous green algae, likely an effect of over-enrichment nutrient loading from shrimp farm and community septic systems (Auil et al., 2007). The lagoon today has low water clarity and sparse submersed aquatic vegetation, with approximately 7% vegetation coverage within the system, compared with 83% reported back in 2003 (Ledwin, 2010). Parasites In 2007, an adult captive manatee from the Corozal Bay died of what appeared to be a verminous pneumonia during medical treatment. That case was very similar to a couple of cases reported by researchers in the southeastern US. In one of those cases a verminous pneumonia involving the fluke (Pulmonicola cochleotrema, formally referred to as Cochleotrema cochleotrema) was suspected as the proximate cause of death in an adult wild manatee from Georgia in the 1980s where a total count of 490 flukes resulted in interstitial pneumonia (Buergelt et al., 1984). In another case over 250 flukes in the manatee‘s respiratory system resulted in severe rhinitis and pulmonary edema. Occlusion of the airways by Pulmonicola, especially if the animal is manipulated, could cause airway blockage and result in asphyxiation. Generally, with Pulmonicola infections there are only a few parasites present in the airways, but due to some underlying factors contributing to accelerated growth in the local manatee population, this condition could lead to vulnerability in the population and even death. It is suspected that another such case was encountered in a manatee captured in Placencia Lagoon in April 2007 (Auil et al., 2007). Other manatees during that capture experienced mucoid discharge from the nasal passages that could have been due to the presence of an irritating parasite. More detailed studies are necessary in order to determine the etiology and significance of this debilitating parasitic condition in manatees in Belize. We hypothesize that effluent from shrimp farms has contributed to a bloom in algae providing food for a small snail which is likely the intermediate host of this parasite. These snails in turn attach to the vegetation consumed by the manatees. Contaminants Trace elements are introduced into coastal systems by industrial activities, including agriculture runoff, and they can reach potentially toxic levels. While many are essential to biological function, some, such as lead, are not. Trace elements can be directly and indirectly harmful to specific organs, or cause immune, neurological, or reproductive problems (Ramey, 2010). As manatees use nearshore habitats, in particular mangrove areas for grazing, they are at great risk as the substrate has a greater load of trace metals than other shoreline sediments (Ramey, 2010). Tonya Ramey analyzed red blood cell samples of 95 manatees captured between 1998 and 2009 to establish a baseline for Belize of eleven trace elements (silver, arsenic, cadmium, cobalt, chromium, copper, iron, nickel, selenium, lead and zinc). There were measureable levels of each trace element in the manatee population, except for cadmium, which was found in only one individual (Ramey, 2010). While some manatees had high concentrations of certain trace elements, they were not thought to be at toxic levels (Ramey, 2010). The author determined that while the sex of the manatee did not have a significant effect on mean trace element level, age did, as younger (juvenile) manatees exhibited significantly higher concentrations of copper. Season and location also affected trace element concentration; for example, lead concentrations were higher in wet season than dry and were higher in the Southern Lagoon manatees than the Drowned Caye animals. One case of interest was a juvenile male caught in Placencia Lagoon that had 10 times the colbalt concentration and three times the lead and zinc concentration than the population average (Ramey, 2010). It is surmised that this animal could have such high concentrations due to foraging in areas of high agricultural contamination risk; and while it cannot be confirmed, immunosuppression may have occurred. These factors could certainly reduce not only the fitness of individuals, but the population on a whole. The fate of manatees of Belize, Auil Gomez  22  It is assumed that because manatees are near the bottom of the food chain, they are not likely to bioaccumulate organic contaminants. A study by Mote Marine Lab and ECOSUR was carried out in the Chetumal Bay, Mexico, and in Florida, USA to examine a variety of organic compounds such as oil-related toxicants called polycyclic aromatic hydrocarbons (PAHs) and other persistent organic pollutants (POPs) including organochlorine pesticides (OCPs), brominated flame retardants (polybrominated diphenyl ethers—PBDEs) and polychlorinated biphenyls (PCBs). PAHs, which attach to suspended particles then settle into the sediment, were undetectable in both sample sets (Wetzel et al., 2008); PAHs are mutagenic and carcinogenic. Additionally, the exposure subsequent to the spilled oil in Charlotte Harbor following Hurricane Charley did not appear to be a problem (Wetzel et al., 2008). POPs were higher in Mexican samples than the Florida samples, but this was not significant; PCBs and OCPs affect the immune system and reproductive success (Wetzel et al., 2008). While Chetumal Bay and the corresponding Belize portion of Corozal Bay constitute good manatee habitat, the land-based pollution is a concern and the area is considered high risk (Wetzel et al., 2008). The PCB levels in Chetumal Bay in particular were high and are of concern; although the threshold for manatees is unknown, it could be affecting reproduction and immune function. Additionally, the contaminant levels in the environment are a concern as PCBs in sediments and seagrasses exceed sediment values alone (Wetzel et al., 2008). As marine currents likely carry these PCBs into Belize‘s waters, and manatee tracking data confirm movement of Belize‘s manatees to Chetumal waters (Auil et al., 2007), it is very likely that the contamination exposure in the north of the country affect manatees that may be coming from the central or southern parts of the country. Examining the complex nature of a manatee‘s surroundings, we see that even if there are areas of available vegetation, manatees are susceptible to encounter toxins that they cannot detect or likely avoid in order to survive. Pockets of habitat have been negatively impacted, and when looked at cumulatively, the risk is high; therefore informed consideration needs to be given when dredging, clearing or filling projects are proposed. Manatee samples should be routinely processed to detect contaminants. Organic compounds, as well as trace elements in substrate and aquatic vegetation should also be examined in Belize to better understand what the manatees truly face. CASE STUDY: DEEPWATER HORIZON OIL SPILL Approximately 185 million gallons of oil was released into the Gulf of Mexico after the Deepwater Horizon oil rig exploded on April 20, 2010, impacting five states in the US. The Gulf is not only an area for several fisheries, but is also used by threatened megafauna such as turtles, dolphins and manatees. As a response to the spill a study was initiated to determine the estimated abundance and distribution of manatees within and adjacent to the spill site, and to document locations of fouled areas or animals. A reconnaissance aerial survey was carried out prior to the spill and aerial surveys were carried out weekly from June 2010 to November 2010 (n = 76). A total of 21 manatees, 1,409 dolphins (97% Tursiops truncatus) and 9 turtles (Caretta caretta) was identified; visible oil was seen during 13 surveys (Ross et al., 2011). Peak manatee sightings (July) were made after oil observations, but some occurrences overlapped with manatee sightings, however, none were observed in actively oiled surface waters (Ross et al., 2011). The team determined that the distribution of manatees and other species was within the oiled region, including feeding areas, an area of approximately 7.4 km2 of patchy seagrass habitat along the survey route (Ross et al., 2011). Additionally, while manatee carcasses were examined after the spill, none in the Gulf of Mexico were determined to have had petrochemical contamination (A. Aven and A. Garrett, personal communication). CONCLUSIONS Marine mammals are exposed to a wide range of toxins in the oceans and coasts where they feed, breed and travel. It is unclear in most species how chemicals are absorbed, distributed, metabolized and excreted and what are their effects (O‘Hara and O‘Shea, 2001). As manatees are the primary consumers of vegetation in the marine mammal world, they would show rather different effects from exposure than most other marine mammals that are carnivorous. Oil, with its varied compositions (crude to mixtures of compounds), can come in direct contact with a manatee‘s external membranes or orifices including eyes, or can be inhaled, and ingested. While it is likely that they can withstand some level of ingested oil by metabolizing and removing it, as demonstrated in captive experiments (O‘Hara and O‘Shea, 2001), it is not clear how this would translate to wild animals in a complex system. Furthermore, it is not known if Too Precious to Drill: the Marine Biodiversity of Belize, Palomares and Pauly  23 individuals would be able to remove themselves from a contaminated area after receiving an exposure level that could be fatal. The manatees‘ food base is certainly vulnerable to the effects of oil by direct mortality, reducing their ability to withstand additional stress, reducing flowering and leaf chlorophyll content, and oil penetration into sediment preventing new shoot growth. Given the complex nature of the aquatic system, especially within the coastal zone which is heavily impacted by land based activities, manatees would have another obstacle to contend with for survival if the petrochemical industry takes root in Belize‘s waters. In contrast, if added protection and management are afforded the species and the coastal habitat, it would benefit not only the status of manatees in Belize, but neighboring countries as well, since Belize is thought to be a manatee source population for Central America. ACKNOWLEDGEMENTS My thanks go to the organizers of this conference and to Janet Gibson (WCS) for advising on the inclusion of manatees in the discussion. Thanks to the manatee capture and tracking project team, Dr. James Powell (Sea to Shore Alliance), Dr. Robert Bonde (USGS) and Jamal Galves (Sea to Shore Alliance and CZMAI) who provide the unique opportunity for the collection of a wealth of data that help us to learn new things about the manatee. Thanks also to Monica Ross (Sea to Shore Alliance) for providing information on the manatee surveys following the BP oil spill, and Dr. Benjamin Morales (ECOSUR) for the valuable information from the Chetumal Bay studies. REFERENCES Auil, N., 1998. Belize Manatee Recovery Plan. Sustainable development and management of biologically diverse coastal resources – Belize project no. BZE/92/G31. UNEP. 67pp. Auil, N., 2004. Abundance and distribution trends of the West Indian manatee in the coastal zone of Belize: implications for conservation. Masters Thesis. Department of Wildlife and Fisheries Sciences. Texas A&M University. College Station, USA. Auil, N., Powell, J., Bonde, R., Andrewin, K., Galves, J., 2007. Belize conservation program, ten year summary. Report to Liz Claiborne Art Ortenberg Foundation. Wildlife Trust, USA. Bengtson, J.L., Magor, D., 1979. A survey of manatees in Belize. Journal of Mammalogy 60, 230-232. Buergelt, C.D., Bonde, R.K., Beck, C.A., O'Shea, T.J., 1984. Pathologic findings in manatees in Florida. Journal of the American Veterinary Medical Association 185(11), 1331-1334. Charnock-Wilson, J., 1968. The manatee in British Honduras. Oryx 9, 293-294. Charnock-Wilson, J., Bertram, K., Bertram, C., 1974. The manatee in Belize. Belize Audubon Society Bull. 6, 1-4. Galves, J., 2011. Manatee strandings along the coastal zone of Belize, 2005-2010. Report to Sea to Shore Alliance and Coastal Zone Management Authority and Institute. Belize. Gibson, J., 1995. Managing manatees in Belize. MS Thesis. Univ. of Newcastle-Upon-Tyne. Gallego, O., 2004. Mapeo del Fondo Lagunar, Laguna de Placencia. Final Report for Friends of Nature. Hartman, D.S., 1979. Ecology and behavior of the manatee (Trichechus manatus) in Florida. The American Society of Mammalogists, Lawrence, Kansas. Hunter, M.E., Auil Gomez, N.E., Tucker, K.P., Bonde, R.K., Powell, J., McGuire, P.M., 2010. Low genetic variation and evidence of limited dispersal in the regionally important Belize manatee. Animal Conservation 13(6), 592-602. Ledwin, S., 2010. Assessment of the ecological impacts of two shrimp farms in Southern Belize. Masters Thesis. School of Natural Resources and Environment. University of Michigan. Ann Arbor, USA. Morales-Vela, B., Olivera-Gomez, D., Reynolds, J.E., Rathbun, G.B., 2000. Distribution and habitat use by manatees (Trichechus manatus manatus) in Belize and Chetumal Bay, Mexico. Biological Conservation 95, 67-75. O‘Hara, T.M., O‘Shea, T.J., 2001. Toxicology. In: Diefrauf, L.A., Gulland, M.D. (eds.), CRC Handbook of Marine Mammal Medicine, pp. 471-500. Second Edition. New York. O‘Shea, T., Salisbury, C.A., 1991. Belize: A last stronghold for manatees in the Caribbean. Oryx 25(3), 154-164. Ramey, T.L., 2010. Trace element concentrations in red blood cells of Antillean manatees (Trichechus manatus manatus) in Belize. Masters Thesis. Department of Ecology, Evolution and Environmental Biology. Columbia University. New York, USA. Ross, M., Carmichael, R., Aven, A., Powell, J., 2011. Manatee Aerial Surveys in Mississippi, Alabama and Louisiana under the NRDA Program: Results of the proposed data collection plan to assess injury to West Indian manatees outside of Florida from the Deepwater Horizon Oil Spill. Report to NRDA marine mammal and sea turtle Technical Working Group. The fate of manatees of Belize, Auil Gomez  24 Short, F.T., Fernandez, E., Vernon, A., Gaeckle, J.L., 2006. Occurrence of Halophila baillonii meadows in Belize, Central America. Aquatic Botany 85, 249-251. Smith, T., Mackie, R., 2005. Use of stable isotopes to detect anthropogenic nutrients in Placencia Lagoon. Final Report for Friends of Nature. Thornback, J., Jenkins, M., 1982. The IUCN Mammal Red Data Book. Part I . Threatened Mammalian Taxa of the Americas and the Australasian Zoogeographic Region. IUCN, Gland, Switzerland. Wetzel, D.L., Pulster, E., Reynolds, J.E. III, Morales, B. Gelsleichter, J., Oliaei, F., Padilla, J., 2008. Organic contaminants in West Indian manatees from Florida to Mexico: A Pilot Study. 42 p.  Too Precious to Drill: the Marine Biodiversity of Belize, Palomares and Pauly  25 STATUS AND DISTRIBUTION OF SEABIRDS IN BELIZE: THREATS AND CONSERVATION OPPORTUNITIES1 H. Lee Jones 7 West Street, Punta Gorda, Belize; lee_jones@att.net Philip Balderamos 19/21 Turneffe Avenue, Belmopan, Belize; pbala@btl.net ABSTRACT The Belize cays and atolls offer a wealth of opportunities for seabirds. Indeed, seabird breeding colonies once proliferated off the coast of Belize. These colonies have been under near constant threat from a variety of sources as far back as the mid-Nineteenth Century, and some have long since vanished. But, how many remain? Where are they located? What are the current threats to their survival? Can some of the extirpated colonies be re-established on cays that are now protected? The answers to these questions are largely unknown. Before the nature and severity of continuing threats and potential future threats, such as those emanating from oil development and transport in Belize, can be adequately assessed, a comprehensive baseline inventory of existing colonies must be established. Only then can we determine the most appropriate measures necessary to preserve and expand these colonies and perhaps to re- establish some of the colonies that have been lost over the years. Does oil development loom as the next significant threat to what remains of the seabird populations in Belize? If so, what measures can be taken to minimize or compensate for this threat? INTRODUCTION Seabirds play an important role in maintaining a healthy marine ecosystem. Most of the seabird species in Belize prey on small to medium-sized fish, and to a lesser extent on arthropods, mollusks, and other invertebrates. As important components of the marine ecosystem, seabirds are efficient tools for monitoring ocean conditions and, at least in some cases, as predictors of stocks of important fisheries (Cairns, 1992; Roth et al., 2007). Because seabirds congregate in large flocks around schools of fish, they have been revealing optimal fishing locales ever since man took to the sea in his quest for food (Au and Pitman 1986, Erdman 1967, Johannes 1981). Additionally, tropical seabirds, especially those that nest in mangroves, enrich shallow-water fish nurseries with their nitrogen-rich excrement, or guano. Although it may seem counterintuitive, seabirds cull smaller and younger fish from schools, thereby reducing competition for food and allowing more fish to attain larger size–a benefit to sport fisheries that many modern-day fishers fail to recognize. The collapse of seabird colonies around the world has had many causes, typically working in synergy. And while diminished seabird populations have frequently been concomitant with diminished or failed fisheries, it is often difficult to pin fisheries collapses directly on the collapse in seabird populations. Overfishing often goes hand in hand with over-harvesting of seabirds or their eggs, habitat conversion, and introduction of non-native predators as human populations expand beyond the capacity of the local resources to support them. For example, Christmas Island, part of the Republic of Kiribati in the western Pacific Ocean, had one of the largest seabird colonies in the world, with several million sooty terns and tens of thousands of 17 other species (Jones, 2000). Now, the numbers of seabirds there have been reduced by more than 90 percent, the result of rat infestations in their nesting colonies, poaching of their eggs for food, and in some cases the massacre of birds for both food and sport (Jones, 2000). At the same  1 Cite as: Jones, H.L., Balderamos, P., 2011. Status and distribution of seabirds in Belize: threats and conservation opportunities. In: Palomares, M.L.D., Pauly, D. (eds.), Too Precious to Drill: the Marine Biodiversity of Belize, pp. 25-33. Fisheries Centre Research Reports 19(6). Fisheries Centre, University of British Columbia [ISSN 1198-6727]. Status and distribution of seabirds in Belize, Jones and Balderamos  26 time, the human population of Christmas Island expanded from a few hundred people in the mid-20th Century to more than 5,000 people at the beginning of the 21st Century. Christmas Island‘s fisheries industry is now on the verge of collapse, but for reasons that that are only partially related to the collapse of its seabird colonies. With increasing pressure on fish populations from overfishing and the introduction of dynamite, cyanide, and modern longline and gillnet fishing techniques, the latter of which have been stretched across narrow openings in the lagoon to catch bonefish as they head out to sea to spawn, the rapid demise of the industry was a foregone conclusion. This scenario in Belize, while not as severe, has several parallels. Belize‘s human population has nearly doubled in the past three decades. Rats have been inadvertently introduced to nesting islands. Seabird eggs have been collected for food. Fish populations upon which the seabirds, as well as humans, depend have been decimated by insufficient or inadequately enforced regulations, especially with respect to the inappropriate use of gill nets and a steep increase in the number of commercial boats, both domestic and foreign, fishing Belize‘s waters. The result: fish populations in Belize are in serious decline and no longer sustainable at present levels of harvesting (see also Zeller et al., this volume). While the general decline in fish populations has certainly had its impact on seabirds, the main direct cause of seabird declines in Belize has been the conversion of habitat for resorts, private residences, and seasonal fishing camps and the associated impacts caused by dogs, cats, and rats. Habitat conversion primarily involves mangrove cutting, removal of littoral forest and dredging of the seabed. These problems are ongoing. They have not been resolved. With new threats on the horizon, including the cumulative effects of climate change and, possibly, contamination of the marine ecosystem from offshore oil extraction and transport, seabirds in the waters off Belize could soon be a thing of the past–unless the ongoing threats are diminished and the potential new threats are addressed proactively. MATERIALS AND METHODS For the purposes of this paper, a seabird is any bird that nests on marine islands and forages in the marine environment. In Belize, that includes members of the Fregatidae, Sulidae, Phalacrocoracidae, Pelecanidae, Ardeidae, Threskiornithidae, Pandionidae, and Laridae. This paper provides a literature review of the past and current status of seabirds in Belize, along with an analysis of past, present, and perceived future threats to their continued presence in Belize. It also includes management and recovery recommendations designed to assure their survival, and in some cases, the re-establishment of populations that have been extirpated. RESULTS AND DISCUSSION Credible information on seabird populations in Belize is sparse. Other than a few key publications that include brief synopses of seabirds (Oates, 1901; Russell, 1964; Jones, 2003) or largely anecdotal accounts (Salvin, 1864; Sclater and Salvin, 1869), most available information comes from the field notes and verbal accounts of biologists who have visited the cays, often only briefly. The one exception is Jared Verner‘s Master‘s thesis (1959) and subsequent publication (Verner, 1961) on the Red-footed Booby (Sula sula) colony on Half Moon Caye. Gaps in our knowledge in some cases span several decades, thus making it all, but impossible to determine any meaningful population trends over time. Almost nothing in the literature, or otherwise, documents anthropogenic threats to seabirds or the consequences of these threats. In short, we know very little about the past or current status of seabirds in Belize. We know that a few species that once nested in Belize have been extirpated or nearly so. We also suspect that a few species now nest in Belize that did not occur historically. A summary of seabirds found historically in Belize is presented in an Appendix at the end of this contribution (see also Paleczny, this volume). Past and current status The first records of seabirds in Belize come from Osbert Salvin (1864) who spent two weeks in May 1862 on several of the Belize cays collecting seabirds and their eggs. We have very little information on Belize seabirds after 1862, until nearly a hundred years later when two ornithologists from Louisiana State University independently visited Belize: Jared Verner, whose studies pertained specifically to one species, the Red-footed Booby, and Stephen M. Russell who, from 1955 to 1961, conducted an inventory and literature review of all birds then known to occur in Belize (Russell, 1964). The Belize Audubon Society sponsored field trips to several of the cays, primarily in the 1980s and early 1990s, led mostly by W. Ford Young, Dora Weyer, Meg Craig, and Martin Meadows. Meadows and Lee Jones (unpublished notes) Too Precious to Drill: the Marine Biodiversity of Belize, Palomares and Pauly  27 visited most of the southern cays in late May 1998. Luz Hunter, Philip Balderamos, Erneldo Bustamante, and Tony Rath documented a significant mixed-species tern colony on Tobacco Caye in July 2002, but it has since vanished. In late February and early March 2007, Betty Ann Schreiber (unpublished notes) and Robert Fleischer visited many cays where seabirds were known to have nested in the past, but their visit was too early in the season to capture the breeding season of terns and a few other species. Thus, to date, the only efforts that even approach a comprehensive inventory of Belize seabirds was Salvin‘s two-week visit to the northern cays in 1862 and Jones‘ and Meadows‘ brief visit to the southern cays in 1998. Salvin was the first to characterize the Red-footed Booby colony on Half Moon Caye, Magnificent Frigatebird (Fregata magnificens) colonies on Half Moon Caye and Man-O‘-War Caye, and significant colonies of Brown Noddy (Anous stolidus) and Black Noddy (Anous minutus) at Glovers Reef. He found Double-crested Cormorants (Phalacrocorax auritus), but not Brown Pelicans (Pelecanus occidentalis) nesting on Man-O‘-War Caye; whereas, when visited 94 years later Russell found the latter nesting, but not the former. Salvin also found Snowy Egrets (Egretta thula) nesting there, but no nesting of this species in Belize has been documented since. Small colonies of cormorants and pelicans have recently been found on several other cays. Salvin estimated the presence of several thousand Red-footed Boobies on Half Moon Caye. The colony was already well known to Belizeans at that time, but the literature contains no specific references to this colony prior to the publication of Salvin‘s 1862 expedition in 1864. When next reported in the literature 96 years later, Verner (1959) counted 1,389 nests, but did not estimate total number of birds present. Belize Audubon Society (1992) similarly estimated 1,325 nests in late 1991. In 2007, however, Schreiber counted only 157 occupied nests, a number that is consistent with Jones‘ impressions from visits to the cay in 1999, 2004, and 2010. While numbers of Red-footed Boobies at Half Moon Caye appear to have decreased dramatically in the last two decades, Magnificent Frigatebird numbers appear to have remained relatively constant at around 60 to 80 pairs, although precise numbers are not available for any period. The same appears to be true for frigatebirds on Man-O‘-War Caye, where reported numbers have ranged from 60 to 100–110 occupied nests. Although apparently known for a number of years previously, in early 1984 the Belize Audubon Society reported on a small Tricolored Heron (Egretta tricolor) and Reddish Egret (Egretta rufescens) colony on two small mangrove cays, Little Guana Caye and Cayo Pajaros, on the Chetumal Bay side of Ambergris Caye (Belize Audubon Society, 1984a). That year, Belize Audubon Society (1984a) also documented Great Egret (Ardea alba) nesting well to the south on Little Monkey Caye near the mouth of Monkey River. In 1990 and 1994, Meadows (Belize Audubon Society, 1991; and unpublished notes) found Tricolored and Reddish egrets, as well as Double-crested Cormorants, Great Blue Herons (Ardea herodias), and Roseate Spoonbills (Platalea ajaja) nesting on Cayo Rosario not far from Little Guana Caye and Cayo Pajaros, and found White Ibises (Eudocimus albus) nesting there in 1994. Estimates of the number of breeding birds or nests were not given in these brief accounts. Several species of terns have been found nesting from time to time on various cays. In 1862, Salvin found ―many thousands‖ of Brown Noddies nesting at Southwest Caye on Glovers Reef and others nesting at Ellen (now known as Carrie Bow), Curlew, and South Water cayes. Although not recorded on later surveys at these three cays, they persisted on Southwest Caye in numbers exceeding 100 pairs at least through 1956 (Russell, 1964), and five birds (nesting status not mentioned) were seen there as late as 1986 (Triggs, unpublished notes). It apparently has not nested there in recent decades, and a resort now occupies most of the cay. Nearby Middle Caye was restored in the 1990s by the Wildlife Conservation Society and is uninhabited except for research facilities and a small staff at the north end, but no noddies or other seabirds currently nest there. In 2002, ten adults on Tobacco Caye were behaving as if they were nesting, but no direct nesting evidence was obtained. There are no other recent records of Brown Noddy nesting in Belize. In addition to Brown Noddy, Salvin also found Black Noddy nesting on Southwest Caye in 1862 (Table 1), but he gave no estimate of its numbers. Berry (cited in Russell, 1964) also found it there and on Morgan Caye (now known as Northeast Caye, also at Glovers Reef) in 1907. We have not been able to find any definitive records of Black Noddy nesting in Belize since 1907; in fact, there are only a handful of credible reports of the species at all in Belize since then. Two other congeners, Sooty Tern (Onychoprion fuscatus) and Bridled Tern (Onychoprion anaethetus), have also nested in Belize. Salvin (1866) ―only met with a few solitary‖ Sooty Terns in 1862, and it was not found nesting in Belize until 1958 when Verner (cited in Status and distribution of seabirds in Belize, Jones and Balderamos  28 Russell, 1964) found a colony with nests containing eggs on Round Caye. In 1971, Henry Pelzel (unpublished ms) had 200-400 pairs on the Silk Cayes. Sometime later, a colony of similar size was discovered on Middle Snake Caye (first mentioned in the literature in 1990; Belize Audubon Society, 1991), but there have been no more confirmed reports from the Silk Cayes. The colony on Middle Snake Caye persisted until around 2008, but was recently abandoned. It is reported to now be on Tom Owens Caye, but this has not been confirmed. Table 1. Nesting history of Brown and Black noddies in Belize. Cay 1861–1910 1911–1960 1961–2010 Morgan (=Northeast) Black nested in considerable numbers (Salvin, 1864) – – Southwest 1,000s of Brown, unknown number of Black Brown nested in the hundreds 5 Brown ―present‖ in 1986 Ellen (=Carrie Bow) A few Brown nested – – Curlew A few Brown nested – – Pompion Not visited Brown nested Tom Owens Black may have nested – – Tobacco – – 10 Brown behaving as if nesting in 2002  Salvin found nesting colonies of Bridled Tern on Saddle, Ellen, and Curlew cays, and possibly South Water Cay in 1862 (Salvin, 1864; Russell, 1964), but it was not reported again from Belize until April 1994 when Meadows (personal communication) observed six pairs attempting to nest on a small artificial cay between Caye Caulker and Ambergris Caye. Four years later, Jones (unpublished notes) and Meadows found a few pairs nesting on several cays along the reef off southern Belize. Lastly, 12 adults were observed by Luz Hunter and her colleagues behaving as if they had nests on Tobacco Caye in July 2002 (Jones, 2002). Laughing Gull (Leucophaeus atricilla) and Sandwich Tern (Thalasseus sandvicensis) are both common along the coast and cays of Belize, but there are few confirmed records of either species breeding in the country. Although Laughing Gull was rumored to nest in Belize for many years, no direct evidence was obtained until May 1998 when Jones (unpublished notes) and Meadows found about 20 nests with eggs on Lawrence Rock at Seal Caye and one nest with eggs on Black Rock. Although never documented, Laughing Gulls almost certainly nested on Laughing Bird Caye before the island was decimated by Hurricane Greta in 1978 and ultimately driven away, presumably by egg collectors, fishers, and tourists, about ten years later (Malcolm Young, personal communication to Lee Jones). It has not nested there since the island, associated reefs, and surrounding waters were designated a national park in 1991. Although Sandwich Tern eggs were collected by Salvin on Northern Two Cayes presumably in 1862 (Oates, 1901), the species was not recorded in Belize again until the early 1960s when a few were seen in Chetumal Bay and Belize Harbor (Russell, 1964). The species has increased dramatically in number since then, but primarily as a non-breeding visitor. Jones and Meadows found about 100 pairs nesting in a dense colony on a small sandbar near North Spot (coordinates 16°15' N, 88°12' W) in 1998. The only other record of nesting in Belize comes from Tobacco Caye where Luz Hunter and her colleagues found 50 birds with large chicks and fledglings in July 2002 (Jones, 2002). Roseate Tern (Sterna dougallii) has also nested in Belize, although little information on this species is available. In 1862, Salvin collected a male from three to four birds present on Grassy Caye where he thought they were ―preparing to breed‖. Luz Hunter and her colleagues counted roughly 200 chicks on Tobacco Caye 140 years later (Jones, 2002), and L. Cottle (fide Betty Ann Schreiber) found Roseate Terns breeding at two sites on the Grassy Caye Range in June 2006. They do not currently breed on Tobacco Caye as confirmed by Philip Balderamos, and their current status on the Grassy Cayes is not known. Roseate Tern is a threatened species in the Caribbean (USFWS, 1987). Belize could play a significant role in its recovery based on the fact that it is occasionally seen in Belizean waters in numbers that exceed 100 birds, has bred as recently as 2006, and may currently be breeding, but undetected. Although seldom documented, Least Tern (Sternula antillarum) is known to nest at various locations along the mainland coast, as well as on a few cays. Salvin found a few pairs ready to lay on Long Caye and ―above a hundred pairs‖ nesting on Grassy Caye in 1862 (Salvin, 1864). According to Belize Audubon Society (1984b), it nests (or nested) on one of the Drowned Cayes near Gallows Pt. Reef 11-12 miles (18- 19 kilometers) east of Belize City. Meadows (unpublished notes) found 70 birds and 12 nests with eggs and Too Precious to Drill: the Marine Biodiversity of Belize, Palomares and Pauly  29 chicks in Bella Vista outside Belize City in May 1988 where they are now reported to nest annually. He also found about ten grown juveniles on a small sandbar near Cayo Rosario in July 1994. Hunter et al. found 20 large chicks on Tobacco Caye in July 2002 (Jones, 2002). Schreiber (unpublished notes) reported that L. Cottle found Least Terns nesting at two sites on the Grassy Caye Range in June 2006, and Jim and Dorothy Beveridge (personal communication) believe that it nests each summer north of the airstrip on the lagoon side of Caye Caulker, although they have not been able to access the site and have not observed eggs or chicks. According to the definition used in this paper, Ospreys of the subspecies Pandion haliaetus ridgwayi that is endemic to the Caribbean are seabirds. In Belize and elsewhere in the Caribbean, they nest exclusively on cays and feed on fish that they catch in nearshore waters. These Ospreys have shown a remarkable ability to adapt to human activities, and one or more pairs nest on most of the cayes, even those that have long been inhabited. We could find no evidence that numbers of this species have declined in Belize, although, as with other seabird species in Belize, specific nesting information is scant and no definitive conclusions can be drawn. Historical and ongoing impacts Anthropogenic impacts on seabirds in Belize have included deliberate intervention in the form of egg collecting, shooting, and vandalism, along with unintentional impacts resulting from tourists, fishers, and others repeatedly entering breeding colonies and causing abandonment. Less direct, but equally destructive, and often much longer lasting impacts have included dredge-and-fill operations, along with replacement of mangroves and littoral forest, for coconut plantations, fishing camps, private homes, and resorts. An inevitable result of repeated human visitation and habitation has been the introduction of non- native predators such as cats, dogs, and rats. While there is no evidence that the limited amount of specimen and egg collecting in the past has resulted in colony failure, persistent shooting, vandalism, and egg harvesting by local fishers, recreational boaters, and others have certainly played a major role in the demise of seabird colonies in Belize. Although potentially severe, these impacts usually do not result in permanent abandonment. Elimination of breeding habitat, on the other hand, does result in permanent loss of breeding colonies. An example of this may be Middle Caye on Glovers Reef. When Salvin visited Glovers Reef in 1862, he found terns nesting on all the cays except Middle Caye, which must have had nesting seabirds in the past, but was already inhabited by the mid-1800s. The native vegetation had been cleared to make way for a coconut plantation, undoubtedly the reason seabirds were no longer breeding there in the 1860s. Now, 150 years later, the coconut plantation is gone and the native vegetation has been restored. The island has a small marine station at its northern end and is fully protected. Yet, there are still no seabirds breeding on the island. Permanent developments and associated habitat conversion have replaced seabird colonies on the other three cays at Glovers Reef and at South Water Caye, Round Caye, Pompion Caye, and perhaps a few others where seabird colonies were never documented prior to their development. Associated with human habitation on many islands are domestic dogs and cats and, unintentionally, rats of the genus Rattus. Introduced non-native species are a leading cause of extinctions in island communities (Atkinson, 1985). Rats, alone, are responsible for 40 to 60 percent of all recorded bird and reptile extinctions worldwide. Although rats have not been implicated in the loss of any seabird colonies in Belize, they have surely played a role, along with other, more direct, human intervention. Black Rats (Rattus rattus) are a suspected culprit in the decimation of the Red-footed Booby colony on Half Moon Caye. Although booby colonies worldwide have tended to survive rat infestations, rat depredation has been mentioned as a possible cause of depletion of all three booby species that occur in the Caribbean (Nelson, 1978; del Hoyo et al., 1992; Priddel et al., 2005). Lastly, climate change is likely to have impacts of uncertain magnitude on seabird colonies in Belize and worldwide in coming decades. The warming of the oceans has already been demonstrated to have had a profound effect on both the intensity and frequency of tropical storms, including hurricanes, and prolonged droughts in many regions of the world. Recent studies have also demonstrated that the oceans have become more acidic as they absorb human-generated carbon dioxide from the atmosphere, and more oxygen-deprived as they absorb agricultural runoff, factors that in turn will further accelerate climate change (Rogers and Laffoley, 2011). Status and distribution of seabirds in Belize, Jones and Balderamos  30 Potential impacts of oil extraction and transport If we are to be in a position to assess the potential impacts from oil development on seabirds we need to first know what species still breed in Belize, where they breed, and how large their colonies are. This will require a comprehensive survey of all known sites, past and present, and perhaps other sites where seabirds may be breeding, but as yet undetected. We also must be able to document the nature and extent of existing threats and the degree to which these threats can be rectified or managed. Only then will we have the tools necessary to evaluate the nature and extent of future impacts and to devise effective measures to avoid, eliminate, reduce, or compensate for those impacts. More specifically, we need to determine what the threat of oil development is, relative to existing threats, and design management and conservation programs that place these ongoing and perceived future threats in perspective. If oil development in Belize poses potentially catastrophic threats to the marine ecosystem in the Gulf of Honduras, as some have asserted, then every effort should be directed toward rethinking the extraction and transport process. The relative benefits and costs of oil development should be carefully weighed against the potential costs to Belize‘s precious marine resources and the economic and cultural benefits that derive from their protection. If, on the other hand, the amount of oil ultimately extracted from and transported through Belizean waters is relatively small and can be extracted and handled safely with proper precautionary measures in place and being enforced, then conservation efforts should perhaps be focused elsewhere where they can be of greater benefit. Different groups of birds, depending on their specific foraging behavior, nesting substrates, and other factors, have differing degrees of vulnerability to oil contamination. Of the seabirds that breed or otherwise reside in Belize, Double-crested Cormorant, Brown Pelican, and Laughing Gull are most vulnerable to offshore oil contamination, as these species spend much of their time in the water. They are all locally abundant near the mainland coast and around the cays and essentially absent beyond the reef and atolls. Boobies spend less time on the water, and terns spend essentially no time on the water, but both groups feed by plunging into the water from the air. Boobies detect fish by sight and, as fish cannot be seen through oily waters, they generally avoid foraging in or landing on oil slicks (del Hoyo et al., 1992). Terns, like boobies, are plunge divers, but unlike boobies they do not rest on the water. Whether or not terns will forage in an oil slick is not known to us, but because most species nest on the ground often just above the high tide line, they could be vulnerable to contamination from oil that washes up on beaches, especially during spring tides. The most common tern species in Belize are the Sandwich Tern and Royal Tern, although only Sandwich has bred in Belize and documented instances are few. Both are common in nearshore waters, including near and at the cayes, where they would be most vulnerable. Least tern is seasonally common along the mainland coast from March to October and breeds (or has bred) locally on several cays. It does not typically venture far from shore, however, and would be most vulnerable to spills near land. Several other species of terns breed or formerly bred on the outer cays, but most of these are now rare or absent or their current status is not known. Sooty Tern can be seasonally abundant near its breeding colonies, but its current status in Belize is unclear. Outside the breeding season (roughly September to March) it is found far offshore over deep waters in the Caribbean. Very little is known about the current breeding status of three other species: Brown Noddy, Bridled Tern, and Roseate Tern. Black Noddy is no longer part of the regularly occurring Belize avifauna. It is unknown if the long gaps between breeding or suspected breeding of many of the terns in Belize are due to their absence or near absence in the western Caribbean during these periods or if they have simply been overlooked. With the paucity of visits to many of the small outer cays where most species are most likely to breed, the latter is certainly feasible. Because so little is known about these species, it would be impractical to assess their vulnerability to oil spills in Belize waters at this time. In the overall scheme of things, however, their vulnerability must be small because they are so rare and/or local in the country and only seasonally present, not year-round inhabitants. Ospreys typically grab fish at the surface with their talons, but occasionally plunge into the water to catch their prey. Like terns, they do not rest on the water, and like boobies they are not likely to forage over oil slicks; thus, their vulnerability to oil contamination must be minimal. Too Precious to Drill: the Marine Biodiversity of Belize, Palomares and Pauly  31 Magnificent Frigatebirds generally are not susceptible to oiling, although they may ingest some oil with their prey. They do not land on the water and catch their prey either by pirating it from other birds in flight or picking it off the water‘s surface with their bill while in flight. Herons, ibises, and spoonbills are long-legged wading birds that feed in shallow water. Only those species that nest in colonies on the cays are considered in this paper. Members of this group rarely if ever swim or float in water. They are most vulnerable to oil contamination along inshore waters where feeding groups congregate, and near their rookeries. While they are not as likely to have their plumage saturated with oil from direct contact, they are vulnerable to the toxic effects of ingesting oil that may be present in or on their prey. They may also transfer small amounts of oil from their beaks and feet to their feathers when preening or scratching. Conserving what we have and restoring what we have lost As discussed above, many of the cays that supported seabird colonies in the past are now developed and have few or no remaining seabirds. Others like Middle Caye (Glovers Reef), Laughing Bird Caye, and Middle Snake Caye are now ostensibly protected, but have no nesting seabirds, although Sooty Terns may return to the latter as they have in the past. Some, like Tom Owens Caye, are either developed or support fishing camps, but still have small numbers of breeding seabirds. For many others, we have no recent information or seabirds tend to nest on them only sporadically, perhaps due to periodic disturbance by fishers, tourists, and vandals. Very few cays with seabird colonies are both protected and patrolled regularly. Half Moon Caye may be the only example. But, being protected and patrolled is often insufficient. On Half Moon Caye, Black Rats are abundant. They readily climb trees and are well known predators on the eggs and young of unattended nests of many species, although little information has been published on their effect on boobies. Regular patrols, coupled with increased enforcement of existing laws will, however, help in reducing poaching, vandalism, wanton habitat destruction, and unauthorized access to sensitive seabird areas. But, patrolling an area as vast as the Belize cays necessitates a considerable increase in personnel, patrol boats, equipment, and training and a considerable expenditure of money. Educational programs in the schools and community centers of Belize would also go a long way toward altering the mindset of those who may not otherwise appreciate the economic value and benefits that accrue from responsible management and conservation of Belize‘s seabirds and other natural resources. Such benefits include an increase in ecotourism, a cleaner, healthier marine environment, and improved commercial and recreational fisheries. In the last few decades, rats have been successfully eradicated from several hundred islands around the globe (Taylor and Thomas, 1993; Howland et al., 2007; Fischer and Dunlevy, 2010), including some much larger than Half Moon Caye. In case after case, seabirds that had been eradicated or nearly eradicated from these islands by rats (and sometimes cats) have returned and are now flourishing (Seniloli, 2008). The same could be accomplished on Half Moon Caye at modest expense. Recent successes in attracting seabirds back to islands where they once bred have also met with success (Kress, 1983, 1998; Kress and Nettleship, 1988; Parker et al., 2007). Typically, decoys and broadcast calls of the target species are set up on the desired island at the onset of the breeding season, and if birds are in the area, they may settle in and form the nucleus of a new colony. But, beforehand, all rats, cats, and other non-native predators must be removed if any new colony is to have a chance of succeeding. Middle Caye on Glovers Reef is ideally suited for this purpose. Suitable habitat for both Brown Noddy and Black Noddy is present, and they both formerly nested in large numbers at Glovers Reef. Economic incentives abound for re-establishing seabird colonies in Belize. Ecotourism is an obvious one. The oil industry can play an important role in assuring that these once flourishing colonies return. With the implementation of proven measures designed to prevent oil leakage and spills during the processes of extraction, handling, and transport, the threat of further damage to the already decimated seabird populations in Belize can be all but eliminated. ACKNOWLEDGEMENTS Betty Ann Schreiber kindly furnished her unpublished notes from a trip she and Robert Fleischer made to the Belize cays in 2007. Status and distribution of seabirds in Belize, Jones and Balderamos  32 REFERENCES Au, D.W.K., Pitman, R.L., 1986. Seabird interactions with dolphins and tuna in the eastern tropical Pacific. Condor 88: 304-317. Belize Audubon Society. 1984a. Belize Audubon Society Newsletter Vol. 15 No. 11. Belize Audubon Society. 1984b. Belize Audubon Society Newsletter Vol. 16 No. 3. Belize Audubon Society. 1991. Belize Audubon Society Newsletter Vol. 23 No. 1. Belize Audubon Society. 1992. Belize Audubon Society Newsletter Vol. 24 No. 3. Cairns, D.K., 1992. 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A fortnight amongst the seabirds of British Honduras. Ibis 1864: 372-387. Salvin, O., 1866. A further contribution to the ornithology of Guatemala. Ibis 1866: 188-206. Sclater, P.L., Salvin, O., 1859. On the ornithology of Central America. Ibis 1859: 1-22, 117-138, 213-234. Seniloli, E., 2008. Restoring the seabird colony on Mabualau Island, Bau waters, Fiji. BirdLife International. Taylor, R.H., Thomas, B.W., 1993. Rats eradicated from rugged Breaksea Island (170 ha), Fiordland, New Zealand. Biological Conservation 65: 191-198. U.S. Fish and Wildlife Service. 1987. Endangered and threatened wildlife and plants; determination: two populations of the Roseate Tern and Bonamia grandiflora (Florida bonamia), Final Rules. U.S. Federal Register 52:211. Verner, J. 1959. Nesting activities of the Red-footed Booby in British Honduras. M.S. thesis. Louisiana State University. Verner, J. 1961. Nesting activities of the red-footed booby in British Honduras. Auk 78: 573-594.  Too Precious to Drill: the Marine Biodiversity of Belize, Palomares and Pauly  33 APPENDIX: GAZETTEER OF HISTORICAL SEABIRD COLONY SITES IN BELIZE Chetumal Bay  Shipstern Caye:  White Ibis Cayo Rosario:  Double-crested Cormorant, Great Blue Heron, Tricolored Heron, Reddish Egret, Roseate Spoonbill, Least Tern (nearby) , Wood Stork(?), Brown Pelican(?) Little Guana Caye: Tricolored Heron, Reddish Egret, White Ibis Cayo Pajaros:  Tricolored Heron, Reddish Egret, White Ibis Unspecified cayes: Wood Stork, Roseate Spoonbill, Bridled Tern(?)  Inner Cayes  Northern Inner Cayes  Hick‘s Cayes: Brown Pelican Drowned Cayes Least Tern  Southern Inner Cayes  Laughing Bird Caye Laughing Gull (never confirmed, but almost certainly nested there) Little Monkey Caye Great Egret Middle Snake Caye Sooty Tern, Bridled Tern(?) East Snake Caye Brown Pelican Mangrove Cayes Brown Pelican, Great Blue Heron  Outer Cayes Caye Caulker Least Tern(?) Sergeant‘s Caye Brown Noddy specimen taken here Man-O‘-War Caye Magnificent Frigatebird, Double-crested Cormorant (1862), Brown Pelican, Snowy Egret (?), Brown Booby allegedly Tobacco Caye Least Tern, Roseate Tern, Sandwich Tern, Brown Noddy(?), Bridled Tern(?) South Water Caye Brown Noddy, Bridled Tern (?) Carrie Bow Caye Brown Noddy, Bridled Tern Curlew Caye Brown Noddy, Bridled Tern Tarpum Caye Important roosts of Magnificent Frigatebird and Brown Pelican Silk Cayes Sooty Tern Round Caye Brown Noddy, Sooty Tern, Bridled Tern Pompion Caye Brown Noddy; Bridled Tern North Spot Sandwich Tern Red Rock and Black Rock Laughing Gull, Bridled Tern Tom Owen‘s Caye Bridled Tern, Sooty Tern(?), Black Noddy(?) Lawrence Rock Laughing Gull, Bridled Tern  Atolls  Lighthouse Reef Northern Two Cayes Sandwich Tern Saddle Caye Bridled Tern Half Moon Caye Magnificent Frigatebird, Red-footed Booby  Turneffe Islands Mauger Caye Brown Booby allegedly Grassy Caye Least Tern, Roseate Tern, Great Egret(?), White Ibis(?) Unspecified caye Great Blue Heron  Glovers Reef Northeast Caye Black Noddy Long Caye Least Tern Middle Caye Apparently there are no historical records of seabirds breeding on this now protected and restored caye Southwest Caye Brown Noddy, Black Noddy  Seabirds of Belize and oil drilling, Paleczny  34 POTENTIAL THREATS OF MARINE OIL DRILLING FOR THE SEABIRDS OF BELIZE1 Michelle Paleczny Sea Around Us Project, Fisheries Centre, University of British Columbia 2202 Main Mall, Vancouver BC V6T 1Z4 Canada; m.paleczny@fisheries.ubc.ca ABSTRACT In their 2011 report, the Belize Audubon Society conlcudes that seabirds are an important component of the marine ecosystem and internationally renowned ecotourism industry in Belize. This paper intends to inform decision makers about the potential threats that marine oil drilling could have on this important component. Included is a brief review of the literature on the interactions between seabirds and marine oil drilling and a summary of the status and distribution of seabirds of Belize. This is followed by an assessment of probability of negative impacts to seabirds caused by marine oil drilling in Belize, based on the knowledge and experience described in the literature. Results indicate that marine oil drilling would negatively impact the seabirds of Belize. INTRODUCTION: INTERACTIONS BETWEEN SEABIRDS AND MARINE OIL DRILLING Marine oil drilling affects seabirds in three key ways, as: Attractants: Marine oil platforms attract seabirds because of increased prey concentration, roosting refuge and interest in lights and flares (Wiese et al., 2001). Seabirds have been observed at concentrations up to 38 times higher surrounding marine oil platforms than in adjacent waters (Wiese et al., 2001). Obstacles: Marine oil platforms can be an obstacle to seabird flight, causing collisions either by accident during low-visibility conditions or because of attraction to lights and flares (Wiese et al., 2001). These collisions cause episodic mortality that is poorly documented by independent scientists, but can cause mortality of up to tens of thousands of seabirds per collision event (Montevecchi, 2006). Pollution: Marine oil platforms release oil into the water, via accidental spills and intentional discharge. Accidental spills from marine oil platforms can be very large (e.g., Ixtoc 476,000 tonnes, Nowruz 272,000 tonnes, Deepwater Horizon 700,000 tonnes), although intentional release during normal operation probably amounts to greater volume than accidental spills (GESAMP, 2007; US Gov, 2010). In total (i.e., accidental and intentional), an average of 16,400 tonnes of oil are reported spilled into the world‘s oceans from marine oil platforms every year (GESAMP, 2007). Oil pollution can cause seabird mortality in two notable ways. First, oil in water can be ingested during feeding or preening, causing digestive and osmoregulatory disorders, reproductive failure, reduced immunity, and mutations in seabirds (Burger and Fry, 1993). Second, oil in water can foul seabird feathers, reducing insulation and buoyancy, which then causes hypothermia, exhaustion and starvation (O‘Hara and Morandin, 2010). Throughout history, seabird die-offs have been documented after large oil spills. For example, the Exxon Valdez 1989 and Gulf of Mexico 2010 spills killed 250,000 and several thousand seabirds respectively (Piatt and Ford, 1996; Safina, 2011). It is important to note that seabird mortality caused by small oil spills is not typically reported, yet the cumulative impact on seabird populations may be greater than that of large spills (Wiese and Robertson, 2004). Recent research has revealed that even the smallest of oil spills, sheens invisible to the naked eye, can be lethal to seabirds (O‘Hara and Morandin, 2010).  1 Cite as: Paleczny, M., 2011. Potential threats of marine oil drilling for the seabirds of Belize. In: Palomares, M.L.D., Pauly, D. (eds.), Too Precious to Drill: the Marine Biodiversity of Belize, pp. 34-37. Fisheries Centre Research Reports 19(6). Fisheries Centre, University of British Columbia [ISSN 1198-6727]. Too Precious to Drill: the Marine Biodiversity of Belize, Palomares and Pauly  35 STATUS AND DISTRIBUTION OF SEABIRDS IN BELIZE A review of the status and distribution of seabirds in Belize is available in this volume (Jones and Balderamos, this volume). In general, breeding colonies are small and scattered, with the exception of two important sites, Half Moon Caye and Man-O-War Caye (Miller and Miller, 2006). Here I also present a map of all documented seabird colonies in Belize (Figure 1), which demonstrates widespread use of islands and cays; and a table describing the approximate abundance and habitat use for all species (Table 1), which concisely describes the seabird community of Belize and its use of both coastal and pelagic habitat. Several species have been affected by threats such as introduced predators, habitat destruction, poaching, persecution, pollution and unsustainable fishing (for more details, see: Jones and Balderamos, this volume; Miller and Miller, 2006). MARINE OIL DRILLING AND SEABIRDS OF BELIZE Should Belize choose to go forward with marine oil drilling, the potential for marine oil platforms overlapping with seabird habitat is high, since seabirds are widely dispersed throughout islands, cays, and the pelagic environment. Furthermore, we can expect spatial overlap to be increased by seabird behaviour (i.e., concentrating around platforms). The probability of collisions between sea birds and oil platforms is also high, since seabird collisions have been documented in all marine oil drilling regions. The frequency and overall mortality caused by collisions is impossible to predict because of the abovementioned lack of independent research on this topic, particularly for the Belize seabird fauna. In the nearby Gulf of Mexico, collisions with marine oil platforms have been estimated to cause 200,000 deaths of migrating birds per year, an unspecified fraction of which are seabirds (Russell, 2005). The probability of pollution having negative effects on seabirds is also very high, given the guaranteed operational discharge plus the chance of accidental spills. Rate of mortality is impossible to predict, given the unpredictable nature of accidental spills and the lack of data on non-trivial seabird mortality caused by operational discharge. Although it is impossible to predict the amount of mortality that will be caused by marine oil drilling, it is possible to predict based on experience that mortality will occur. Given that many seabird populations in Belize are small and/or already threatened, it is unlikely that they can withstand additional threats without facing population declines. Thus, it is very probable that this mortality will have population-level impacts. CONCLUSIONS Based on the status and distribution of seabirds in Belize, and the threats associated with marine oil drilling, it is apparent that marine oil drilling is an activity that will have negative effects on the seabirds of Belize. A precautionary approach of banning marine oil drilling would benefit seabirds and the related ecotourism economy. ACKNOWLEDGEMENTS This is a contribution from the Sea Around Us project a collaboration between the University of British Columbia and the Pew Environment Group.  Figure 1. Map of all documented seabird breeding sites (black dots) within Belize (Miller and Miller, 2006; Jones, 2003; Jones and Balderamos, this volume). Seabirds of Belize and oil drilling, Paleczny  36 Table 1. Classification (Peters, 1979), habitat (Jones 2003; Sea Around Us Project Database, 2011) and abundance estimate (Miller and Miller, 2006; updated by Jones, personal communication) for all seabird species that occur (breeding and non-breeding; but not vagrant) in Belize. Common name Order Family Genus Species Habitat (c=coastal, p=pelagic) Abundance (# individuals) Black Noddy Charadriiformes Laridae Anous minutus P 0-50 Black Tern Charadriiformes Laridae Chlidonias niger C 500-1000 Bridled Tern Charadriiformes Laridae Sterna anaethetus C 0-50 Brown Noddy Charadriiformes Laridae Anous stolidus P 0-50 Common Tern Charadriiformes Laridae Sterna hirundo C 50-100 Forster's Tern Charadriiformes Laridae Sterna forsteri C 0-50 Herring Gull Charadriiformes Laridae Larus argentatus C 50-100 Caspian Tern Charadriiformes Laridae Sterna caspia C 50-100 Laughing Gull Charadriiformes Laridae Larus atricilla CP 500-1000 Least Tern Charadriiformes Laridae Sterna antillarum C 50-1000 Ring-billed Gull Charadriiformes Laridae Larus delawarensis C 0-50 Roseate Tern Charadriiformes Laridae Sterna dougallii CP 0-50 Royal Tern Charadriiformes Laridae Sterna maxima C 500-1000 Sandwich Tern Charadriiformes Laridae Sterna sandvicensis C 1000-5000 Sooty Tern Charadriiformes Laridae Sterna fuscata P 1000-5000 Bonaparte's Gull Charadriiformes Laridae Larus philadelphia C 0-50 Pomarine Jaeger Charadriiformes Stercorariidae Catharacta pomarinus P 0-50 Magnificent Frigatebird Pelecaniformes Fregatidae Fregata magnificens CP 1000-5000 Brown Pelican Pelecaniformes Pelecanidae Pelecanus occidentalis CP 1000-5000 Double-crested Cormorant Pelecaniformes Phalacrocoracidae Hypoleucos auritus C 500-1000 Brown Booby Pelecaniformes Sulidae Sula leucogaster P 500-1000 Masked Booby Pelecaniformes Sulidae Sula dactylatra P 0-50 Red-footed Booby Pelecaniformes Sulidae Sula sula P 1000-5000 Audubon's Shearwater Procellariiformes Procellariidae Puffinus lherminieri P 0-50 Too Precious to Drill: the Marine Biodiversity of Belize, Palomares and Pauly  37 REFERENCES Belize Audubon Socity, 2011. Belize Audubon Society‘s position on offshore oil exploration, extraction and production. Available at: http://belizeaudubon.org/news/2011/01/12/bas-position-on-offshore-oil-exploration-extraction-and-production/ Burger, A.E., Fry, D.M., 1993. Effects of oil pollution on seabirds in the northeast Pacific. Pacific Seabird Group Publication. GESAMP, 2007. Estimates of oil entering the marine environment from sea-based activities. GESAMP (IMO/FAO/UNESCO- IOC/UNIDO/WMO/IAEA/UN/UNEP Joint Group of Experts on the Scientific Aspects of Marine Environmental Protection) Report and Studies No. 75. Jones, H.L., 2003. Birds of Belize. University of Texas Press, Austin Texas. Miller, B.W., Miller, C.W., 2006. Waterbirds in Belize. Final Report for the Belize Audubon Society – Wildlife Conservation Society. Montevecchi, W.A., 2006. Chapter 5: Influences of Articial Light on Marine Birds. In: Rich, C., Longcore, T. (eds.), Ecological Consequences of Artificial Night Lighting. Island Press, 112 p. O‘Hara, P.D., Morandin, L.A., 2010. Effects of sheens associated with offshore oil and gas development on the feather microstructure of pelagic seabirds. Marine Pollution Bulletin 60, 672-678. Peters, J.L., 1979. Checklist of Birds of the World. Vol. 1. Second Edition. Harvard University Press, Cambridge, MA, USA. Piatt, J.F., Ford, R.G., 1996. How many seabirds were killed by the Exxon Valdez Oil Spill? American Fisheries Society Symposium 18, 712-719. Russell, R.W., 2005. Interactions between migrating birds and offshore oil and gas platforms in the northern Gulf of Mexico: Final Report. U.S. Dept. of Interior, Minerals and Management Service, Gulf of Mexico OCS Region, New Orleans, LA. OCS Study MMS 2005-009. 348 p. Safina, C., 2011. The 2010 Gulf of Mexico oil well blowout: A little hindsight. PloS Biol. 9(4), e10111049. US Gov., 2010. Unites States Government Press Release: U.S. Scientific Teams Refine Estimates of Oil Flow from BPs Well Prior to Capping. United States Government Press Release, available at: http://www.restorethegulf.gov/release/2010/08/02/us- scientific-teams-refine-estimates-oil-flow-bps-well-prior-capping Wiese, F.K., Montevecchi, W.A., Davoren, G.K., Huettmann, F., Diamond, A.W., Linke, J., 2001. Seabirds at risk around offshore oil platforms in the Northwest Atlantic. Marine Polluation Bulletin 42(12), 1285-1290. Wiese, F.K., Robertson, G.J., 2004. Assessing seabird mortality from chronic oil discharges at sea. Journal of Wildlife Management 68(3), 627-638.  Elasmobranchs of Glovers Reef, Chapman  38 THE ELASMOBRANCHS OF GLOVER‘S REEF MARINE RESERVE AND OTHER SITES IN NORTHERN AND CENTRAL BELIZE1 Demian Chapman Institute for Ocean Conservation Science and School of Marine and Atmospheric Science, Stony Brook University, Stony Brook, NY 11794, USA; demian.chapman@stonybrook.edu Elizabeth Babcock Rosenstiel School of Marine and Atmospheric Science, 4600 Rickenbacker Cswy, Miami, FL 33149, USA Debra Abercrombie Abercrombie and Fish Consulting, Port Jefferson Station NY 11776, USA Mark Bond and Ellen Pikitch Institute for Ocean Conservation Science and School of Marine and Atmospheric Science, Stony Brook University, Stony Brook, NY 11794, USA ABSTRACT Glover‘s Reef Marine Reserve (GRMR) is one of the largest marine reserves in Belize. In 2000, our group initiated a study of the sharks and rays (elasmobranchs) at this site in order to (1) characterize local biodiversity, (2) determine the significance of GRMR as an elasmobranch nursery area and (3) broadly assess the potential of marine reserves for the conservation of sharks. Our surveys encompass the lagoon, the forereef and the deep benthic habitat (~400 m) off the edge of the reef slope. We documented the presence of at least 15 elasmobranch species at GRMR. Two species recorded in our survey, Galapagos sharks (Carcharhinus galapagensis) and night sharks (Carcharhinus signatus), had never been recorded in Belize before. We found evidence of local breeding in at least 7 elasmobranch species at GRMR and have maintained a standard time series of shark abundance since 2001. This survey indicates that the abundance of at least some species have remained stable at this site, suggesting that marine reserves can help protect certain shark species. Automated acoustic telemetry of Caribbean reef (Carcharhinus perezi, N = 34) and nurse sharks (Ginglymostoma cirratum, N = 25) showed that both species exhibit a high degree of fidelity to GRMR, which helps to explain the observed stable abundance trends. Since 2005, we have started surveying other areas in northern and central Belize, including Caye Caulker Marine Reserve (CCMR), Turneffe atoll (TU) and Southwater Caye (SW). Fished reefs (TU, SW) exhibit depressed populations of sharks relative to reserve reefs (GRMR, CCMR). However, both sites provide important nursery habitat for a variety of sharks, including scalloped and great hammerheads (Sphyrna lewini, S. mokarran), lemons (Negaprion brevirostris) and blacktips (Carcharhinus limbatus). Population genetic studies by us and others indicate that Mesoamerica harbors differentiated stocks of certain shark species that are not regularly replenished by immigration. Overall, we conclude that Belize has a diverse, largely self-sustaining elasmobranch fauna that is under serious threat from overexploitation and habitat loss.   1 Cite as: Chapman, D., Babcock, E., Abercrombie, D., Bond, M., Pikitch, E., 2011. The elasmobranchs of Glover‘s Reef Marine Reserve and other sites in northern and central Belize. In: Palomares, M.L.D., Pauly, D. (eds.), Too Precious to Drill: the Marine Biodiversity of Belize, pp. 38-42. Fisheries Centre Research Reports 19(6). Fisheries Centre, University of British Columbia [ISSN 1198-6727]. Too Precious to Drill: the Marine Biodiversity of Belize, Palomares and Pauly  39 INTRODUCTION Global exploitation of sharks is increasing and expanding, yet basic research and management continues to lag far behind (FAO, 2000). This trend is evident in the Belize, a country experiencing a level of population growth and increasing demand for its natural resources, which threatens the health of its marine ecosystems. Sharks and rays are some of the largest marine predators in Belize and some, such as the Caribbean reef shark, Carcharhinus perezi, southern stingray, Dasyatis americana, and nurse shark, Ginglymostoma cirratum, form the basis of a lucrative dive tourism industry. Indeed, a diver survey by the Coral Reef Alliance found that ‗seeing sharks‘ was the primary attraction to Belize for the majority of respondents. Sharks are also fished for local consumption and for export, especially to the Asian dried fin market (Gibson et al., 2005; Pikitch et al., 2005). Beyond these direct commercial uses, sharks may perform critical ecosystem services that may even exceed their direct economic value (Heithaus et al., 2008). Directed shark fisheries have already drastically reduced shark populations in many parts of the world (Musick et al., 2000). Sharks are extremely vulnerable to overexploitation because they exhibit a K- selected life-history strategy, reproducing more like mammals than teleost fishes (FAO, 2000; Musick et al., 2000). Despite this, management and conservation of sharks has been largely reactive, proceeding only after marked declines in abundance and diversity have already occurred (FAO, 2000; Musick et al., 2000). Such a situation is primed to occur in the Belize. Not only is shark exploitation largely unregulated and unmanaged, but only the whale shark, Rhincodon typus, is legally protected, a species that was never even part of the commercial catch. There is no National Plan of Action for sharks in Belize, as called for by the United Nations Food and Agriculture Organization, and there are no restrictions on the landing of sharks. The main fishing gear used for shark fishing is monofilament gillnetting, which is indiscriminant with regard to size and species and is also rapidly lethal to any captured shark. The reliance on this gear type currently makes it impossible to develop species specific shark legislation, as it would not be possible to catch and release alive any species that were prohibited from the fishery. There are, however, some parts of Belize where the shark populations may be less impacted by fishing. Together with a few remote, lightly fished locations, the Belize Marine Protected Area Network may be among these because marine reserves within the network provide a spatial respite from fishing pressure. Glover‘s Reef Marine Reserve (GRMR) is one such location. In this paper, we will review our studies of the elasmobranchs of GRMR and other sites in central and northern Belize. The objective of this extended abstract is to describe the remaining elasmobranch biodiversity in this region. MATERIALS AND METHODS Glover‘s reef atoll (16o 44‘ N, 87o 48‘W) lies approximately 25 km to the east of the Mesoamerican Barrier Reef and 45 km east of the Belizean mainland. The atoll is ~30 km from north to south and ~10 km at its widest. The reef crest partially separates the narrow fore-reef (<500 m in most areas) from the lagoon, which is basin shaped and as deep as 18 m. The forereef drops off nearly vertically at the reef slope, to depths of >400 m on the west side of the atoll and >1000 m on the east side of the atoll. Glover‘s Reef Marine Reserve (GRMR) was established in 1993 and is zoned for multiple uses. The southern third of the atoll is designated as a no-take zone, called the ‗conservation zone‘, where no fishing is permitted. The rest of the atoll interior of the reef slope is zoned for restricted fishing where, among other regulations, commercial-scale shark fishing has been eliminated by a moratorium on the use of longlines and gillnets. Between July 2000 and July 2011, we deployed several types of elasmobranch surveying gear at GRMR and several other sites in northern and central Belize (Turneffe atoll, Caye Caulker Marine Reserve and Southwater Caye Marine Reserve). Standard longlines and methods associated with handling, tagging, sampling and releasing elasmobranchs caught on this and the other gear are described in Pikitch et al., (2005) and Chapman et al., (2005). Since October 2007, we have deployed deep water longlines at depths of 250-400 m. Deep lines consist of a ¼ inch‘ nylon rope mainline that is set on the seafloor by a cement block tied to one end and suspended near vertically by a set of large floats tied to the other end. Five gangions were placed at 15 m intervals starting from the bottom, each one consisting of a tuna clip attached to 3.5 m of nylon line, a swivel and 3.5 m of stainless steel aircraft cable terminating in a baited 16 o/o circle hook. Since 2009, we have deployed baited remote underwater video (BRUV) units to quantify the relative abundance of sharks between sites. BRUVs consist of a video camera (Sony Handycam DCR-HC52) inside an underwater housing that is mounted on a metal frame that has a small, pre-weighed bait source (1 kg of crushed baitfish) mounted on a pole in the camera‘s field of view. Sharks Elasmobranchs of Glovers Reef, Chapman  40 are identified and counted as they swim into the field of view over the standard 70 minute deployment. Our final method of documenting elasmobranchs is opportunistic surveys of fishers and their catches in Belize City, Dangriga, Turneffe atoll, Glover‘s Reef atoll and Southwater Caye. One fisherman from Turneffe atoll provided us with a tissue sample from every shark he landed from June 2008-June 2011. We identified these samples to species using DNA barcoding (Wong et al., 2009). RESULTS AND DISCUSSION We documented 15 species of elasmobranchs at GRMR from 2000-2011. The dominant species are the nurse shark, Ginglymostoma cirratum, a demersal mesopredator, and the Caribbean reef shark, Carcharhinus perezi, an active top predator. Both species are common in the lagoon and fore-reef habitats. Nurse sharks also frequent shallow seagrass flats, while Caribbean reef sharks were recorded diving down the reef slope to depths of up to 352 m (Chapman et al., 2007). Acoustic telemetry shows that both nurse and Caribbean reef sharks are mostly year round residents of GRMR (Chapman et al., 2005, Bond et al., submitted, Pikitch et al., submitted). Caribbean sharpnose sharks, Rhizoprionodon porosus, are small demersal mesopredators that are also quite common in the lagoon. In contrast, lemon sharks, Negaprion brevirostris, an active top predator species, are now uncommon at GRMR. In the early years of the study we frequently observed and captured neonate lemon sharks on the seagrass flats around Middle Caye and Southwest Caye. Genetic studies showed that many of these were siblings, indicating that a relatively small number of adult females were using GRMR for parturition. From 2006 through to this year, we have not observed any neonates in these areas. However, in 2011 a group of at least 3 subadult lemon sharks has been frequently sighted at Middle and Northeast Cayes scavenging at fish cleaning stations. We have tagged and measured 2 of these at 189 and 193 cm TL (one male and one female). Southern stingrays (Dasyatis americana) and spotted eagle rays (Aetobatus narinari) are the most common rays at GRMR and can be found from shallow seagrass flats all the way to the edge of the reef slope. Both are mesopredators that feed on benthic invertebrates and small fish. All 6 of these shark and ray species are known to breed at GRMR, as we have captured specimens of every age class (neonate to adults of each sex). We have also observed a few specimens of yellow stingray (Urobatis jamaicensis) and captured 2 Caribbean whiprays (Himantura schmarde). These rays are also likely to be residents of GRMR, given its isolation. GRMR also provides temporary habitat for several other migratory or highly mobile elasmobranchs. Over the course of the study we have captured 6 tiger sharks (Galeocerdo cuvier) ranging from 220-260 cm TL. We fitted 2 of these with coded acoustic transmitters. Both were detected at the capture site on the day they were released, but never after that. One was later detected sporadically on the other side of the atoll after a hiatus of ~150 days. Tiger sharks therefore appear to be transient at GRMR, although it is notable that we have only recently (from 2008) started regularly captured them on our standard longline sets. Great hammerhead sharks (Sphyrna mokarran) of ~200-400 cm TL are also occasionally observed at GRMR, but to date we have not captured or tagged any of them. On June 8, 2009, we captured and tagged the first blacktip shark (Carcharhinus limbatus) recorded at GRMR. The individual was a 184 cm TL female and therefore probably mature. Another notable capture in our survey was a juvenile Galapagos shark (Carcharhinus galapagensis). This was the first documented capture of this species in the western Caribbean and only the second verified capture in the whole Caribbean since 1963 (Pikitch et al., 2005). Lastly, we are aware of at least one whale shark (Rhincodon typus) sighting at GRMR (Pikitch et al., 2005). Our deep water longline survey of the reef slope and pelagic habitat around Glover‘s Reef has revealed several additional species using the atoll. The most notable captures have been night sharks (Carcharhinus signatus), a relatively large mesopelagic species that had never before been recorded in Mesoamerica. We discovered that adults of this species form large aggregations at certain locations around Glover‘s Reef in 200-400 m depth. Twelve individuals have been captured, ranging in size from 197 to 249 cm TL (6 females, 6 males). Deep lines have also captured 3 silky sharks, Carcharhinus falciformis, a large epipelagic species, ranging in length from 221 to 286 cm TL (2 males, 1 female). Lastly, we have recorded 5 adult specimens of the smooth dogfish (Mustelus canis insularis) at depths of 200-400 m off the reef slope from fisheries catches. We hypothesize that there is a rich demersal elasmobranch fauna at these depths, but we have not yet set our lines in such a way to capture them over concerns about causing mortality and/or losing gear. Too Precious to Drill: the Marine Biodiversity of Belize, Palomares and Pauly  41 The elasmobranch fauna of Turneffe atoll is generally similar to that of GRMR in terms of species composition and diversity (16 species). Nurse and Caribbean reef sharks dominated our standard longline surveys in 2005 and 2006, where we also captured Caribbean sharpnose sharks, southern stingray, blacktip sharks, lemon sharks and Caribbean whiprays. Our survey of the catches of one net fisher at Turneffe from 2008-2011 indicate that Caribbean sharpnose and Caribbean reef sharks are the main commercial species, followed by blacktip, lemon and great hammerhead sharks. These three last species all appear to be more common at Turneffe atoll than Glover‘s, which probably reflects habitat differences between these sites. Turneffe has much more extensive mangrove forest than Glover‘s Reef, which means that Turneffe is more likely to serve as a nursery for these shark species. We also observed scalloped hammerhead (Sphyrna lewini) and bull sharks (Carcharhinus leucas) in the net catches, although much less frequently than the other species. We have been shown numerous underwater photographs of aggregations of large scalloped hammerheads taken off the southern end of Turneffe. All net-captured scalloped hammerheads were small, which indicates that this area serves as both juvenile habitat and an aggregation area for adults. We have also surveyed fisheries catches in Belize City, Dangriga and Southwater Caye Marine Reserve opportunistically since 2000. In Pikitch et al. (2005), we reported the following composition from Belize City and Dangriga fishmarket collections: ‗Shark collections at the 2 coastal fish markets yielded a total of 57 intact specimens, consisting of 30 blacktips Carcharhinus limbatus (18 neonates, 12 juveniles <90 cm TL), 2 Negaprion brevirostris (1 neonate, 1 juvenile 156 cm TL), 22 bonnetheads Sphyrna tiburo (all juveniles <60 cm TL), 1 scalloped hammerhead S. lewini (neonate), 1 great hammerhead S. mokkaran (juvenile <90 cm TL), and 1 Rhizoprionodon porosus (neonate). Discussions with fishers indicated these were all captured inshore on the coastal side of the barrier reef between Dangriga and Belize City. Since that report we have observed that juvenile blacktips, adult Caribbean sharpnose are still observed in these two markets. We have observed more Caribbean reef and nurse sharks in Belize City from 2005 to the present and have observed no bonnetheads at all. On January 25, 2011 in Southwater Caye Marine Reserve, we sampled recent catches of a gillnetter that fishes the area. We found 5 fin sets from large great hammerhead sharks, as well as ~ 20 fin sets from juvenile Caribbean reef sharks. Our surveys reveal that the parts of Belize we sampled retain a reasonably diverse elasmobranch fauna (at least 18 species) and possess large tracts of important habitat for them. However, there is evidence that fishing is severely impacting shark populations. Our BRUV deployments show that there are significantly fewer Caribbean reef shark observed on fished reefs (Turneffe, Southwater Caye prior to reserve establishment) than the marine reserves GRMR and Caye Caulker (Bond et al., submitted). Many of these species rely on the Barrier Reef and associated seagrass and mangrove forests as juvenile habitat. At least one of the offshore atolls, Glover‘s Reef, is an important refuge from fishing for several shark species. Given the potentially precarious state of shark populations in Belize, we suggest that additional anthropogenic stressors affecting these habitats could greatly accelerate fisheries-induced population declines. Oil exploration and potential spills from drilling are two potential stressors that could seriously impact these habitats and the shark populations that rely on them. REFERENCES Bond, M.E., Babcock, E.A., Pikitch E.K., Abercrombie, D.L., Lamb, N., Chapman, D.D., (submitted). Reef sharks exhibit site-fidelity and higher relative abundance in marine reserves on the Mesoamerican Barrier Reef. Chapman, D.D., Pikitch E.K., Babcock, E.A., Shivji, M.S., 2005. Marine reserve design and evaluation using automated acoustic telemetry: a case-study involving coral reef-associated sharks in the Mesoamerican Caribbean. Marine Technology Society Journal 39(1), 40-53. Chapman, D.D., Pikitch, E.K., Babcock, E.A., Shivji, M.S., 2007. Deep-diving and diel changes in vertical habitat use by Caribbean reef sharks, Carcharhinus perezi. Marine Ecology Progress Series 344, 271–275. FAO, 2000. Conservation and Management of sharks. FAO Technical Guidelines for Responsible Fisheries 4, Suppl. 1. Rome. 37 pp. Gibson, J., McField, M., Heyman, W., Wells, S., Carter, J., Sedberry, G., 2004. Belize‘s Evolving System of Marine Reserves. In: Sobel, J., Dahlgren, C. (eds.), Marine Reserves: A Guide to Science, Design and Use, pp. 287-316. Island Press, Washington, U.S.A. Heithaus, M.R., Frid, A., Wirsing, A. J., Worm, B., 2008. Predicting ecological consequences of marine top predator declines. Trends in Ecology and Evolution 23, 202-210. Musick, J.A., Burgess, G., Caillet, G., Camhi, M., Fordham, S., 2000. Management of sharks and their relatives (Elasmobranchii). Fisheries 25(3), 9-13. Elasmobranchs of Glovers Reef, Chapman  42 Pikitch, E.K., Chapman, D.D., Babcock, E.A., Shivji, M.S., 2005. Habitat use and demographic population structure of elasmobranchs at a Caribbean atoll (Glover‘s Reef, Belize). Marine Ecology Progress Series 302, 187-197. Pikitch, E.K., Chapman, D.D., Babcock, E.A., Karnaukus, M., Submitted. Movements and residency of nurse sharks, Ginglymostoma cirratum, within an insular Marine Protected Area in Belize. Wong, E.H.-K, Shivji, M.S., Hanner, R.H., 2009. Identifying sharks with DNA barcodes: assessing the utility of a nucleotide diagnostic approach. Molecular Ecology Resources. 9, 243-256.  Too Precious to Drill: the Marine Biodiversity of Belize, Palomares and Pauly  43 SNAPPER AND GROUPER ASSEMBLAGES OF BELIZE: POTENTIAL IMPACTS FROM OIL DRILLING1 William Heyman and Shinichi Kobara Department of Geography, Texas A&M University, TX 77843-3147 USA; wheyman@tamu.edu; shinichi@tamu.edu ABSTRACT The grouper/snapper species complex is made up of top predators within coral reef ecosystems and together is the most commercially important group. As top predators, they are also indicative of the health of coral reef systems. This paper summarizes nearly 20 years of research on the life history of snappers and groupers in Belize, with a focus on spawning aggregations, larval transport, and juvenile settlement and development in nursery areas. The contribution then addresses the possible impacts of oil on the various life stages and habitats utilized by snappers and groupers. Though the likelihood of spill or contamination from oil exploration and development may be very small, the potential effects of such events are so catastrophic that the risk may outweigh the potential benefits. INTRODUCTION Coral reef ecosystems, along with tropical rain forests are the most biologically diverse ecosystems on earth. These systems contribute to regional economic and ecological health in many ways including fisheries productivity, marine tourism, and coastal protection from storms. The snapper/grouper complex includes many of the reef‘s top predators and serves as indicators of ecosystem health. The Belize Barrier Reef System is the largest in the hemisphere, the second largest in the world, and is locally and internationally recognized for its cultural, ecological, and economic values (IUCN World Heritage Site Designation). Snappers and groupers share similar life history characteristics—starting their juvenile period in seagrass and mangrove habitats, migrating into shallow patch reefs, and after 3-6 years reach reproductive maturity (see contributions in Arreguín-Sanchez et al., 1996). They then spawn in aggregations at predictable times and places, releasing pelagic eggs that are fertilized in the water column. After 12-24 hours, the eggs metamorphose into larvae, which swim in the plankton for 1 to three weeks and then settle as juveniles into protected nursery habitats, generally mangroves and seagrass. While there are many variations on this general pattern, most  1 Cite as: Heyman, W., Kobara, S., 2011. Snapper and grouper assemblages of Belize: potential impacts from oil drilling. In: Palomares, M.L.D., Pauly, D. (eds.), Too Precious to Drill: the Marine Biodiversity of Belize, pp. 43-47. Fisheries Centre Research Reports 19(6). Fisheries Centre, University of British Columbia [ISSN 1198-6727].   Figure 1. The locations of 14 multi-species reef fish spawning aggregation sites in Belize. The sites all occur at sharp bends in reefs otherwise referred to as reef promontories (after Figure 1 in Kobara and Heyman, 2010). Snapper and grouper assemblages of Belize, Heyman and Kobara  44 commercially important groupers and snappers in Belize (e.g., mutton snapper, cubera snapper, yellow tail snapper, Nassau grouper, black grouper, yellowfin grouper, tiger grouper) can be described this way. The goals of this paper are to: 1) summarize recent research and provide further details on the timing and location of spawning aggregations; 2) discuss the value of these fishes and the ecosystems on which they rely to the local and national economy; and 3) discuss the possible effects on these systems from oil exploration and development. RESULTS AND DISCUSSION With the support and collaboration of the Belize National Spawning Aggregations Working Committee, we have participated in research that characterizes multi-species reef fish spawning aggregations in Belize. In short, aggregations of multiple reef fish species aggregated to spawn at specific times and places that can be explained by the ‗multi-species promontory‘ hypothesis which states: Large and commercially important reef fishes in the Caribbean typically spawn in large aggregations at shelf edges (25-40 m water depth) and reef promontories (relatively sharp bends in the shelf) adjacent to deep water (>200 m) (Figures 1 and 2). These aggregations occur at predictable times and places for each species according to seasonal, lunar, tidal, and diel cycles—with grouper spawning in Belize occurring around sunset, 4-12 days after full moon from December-March and most snapper spawning occurring around sunset, 2-8 days after full moon from March-June and a smaller peak August-September (Figure 3). These data are detailed and summarized in a series of recently published scientific papers (Heyman et al., 2001; 2005; Heyman and Kjerfve, 2008; Kobara and Heyman, 2008; 2010).  Figure 2. Geomorphology of Gladden Spit, perhaps the best studied multi-species reef fish spawning aggregation site in the world, and representative of other sites in terms of geomorphology and species composition. The figure shows the location of the aggregations (fish symbol) at the promontory tip, along the shelf edge (30 m water depth) adjacent to deep water (>200 m; from Heyman and Kobara, 2011). From 1998 to the present, the Belize National Spawning Aggregations Working Committee has described and monitored between 9 and 12 multi-species reef fish spawning aggregations throughout Belize. They also worked as a coalition to protect these sites, which were declared as no-take marine reserves in 2002, along with legislation to protect the endangered Nassau Grouper (Gibson et al., 2007). Importantly, the legislation to protect these sites was fully supported by the great majority of commercial fishers, including those who have traditionally fished the spawning sites. Their support came from their understanding of Too Precious to Drill: the Marine Biodiversity of Belize, Palomares and Pauly  45 the value of the spawning sites to the long-term health of the stocks and the reef. They gained this understanding in part, through their intensive participation in the research program that described the sites. Their support was likely increased by their access to training in economic alternatives to fishing, such as SCUBA and snorkel ecotourism guiding, provided by members of the Spawning Aggregations Working Committee (especially TIDE, SEA, and Green Reef; Heyman, 2011).   Figure 3. Gladden Spit has at least 17 species of fish from nine families that aggregate to spawn including: (A) Dog snapper, Lutjanus jocu spawning event in late afternoon (B) Mutton snapper, Lutjanus analis releasing gametes in a group spawning event in mid afternoon, (C) Black grouper, Mycteroperca bonaci, shown here in courtship coloration, (D) Smooth trunkfish, Lactophrys triqueter showing courtship behaviour and coloration, (E) Horse-eye jack, Caranx latus, in courtship coloration, (F) Jack crevalle, Caranx hippos showing courtship coloration, (G) White margate, Haemulon album aggregation in mid afternoon (adapted from Figure 3 in Heyman and Kjerfve, 2008). Figure 4. Life cycle of Cubera snappers (representative of the grouper snapper complex) include various stages and habitats (shown clockwise from upper left): A) adult spawning aggregations which occur at reef promontories (photo by Douglas David Seifert), B) fertilized eggs which float in the plankton for 18 hours (photo by Carol Farnetti Foster), C) planktonic larvae which start feeding after four days and often drift or swim inshore, and D) juveniles that settle and live in seagrass and amongst the shallow prop roots of red mangroves. The most vulnerable period in the life history of most groupers and snappers occur during the egg and larval phases (Figure 4). These delicate stages are far more susceptible to changes in water quality and/or toxicity. Studies of the impacts from oil drilling, exploration or spillage should be conducted on these stages, rather than on the adults, which may have the ability to simply swim away. The effects of oil, drilling muds, and dispersants have been evaluated for many marine and coastal fish species and the results are uniformly negative and significant. The larval and early juvenile phases of most organisms, including fishes, are highly vulnerable to even the slightest concentrations of oil-derived pollutants and dispersants (Wilson, 1977; NRC, 2005; Tunnel, 2011). The value of reef fish spawning aggregations cannot be overstated. They include the most important and most vulnerable stages in the life cycle of most commercially important reef fishes. The adults are most vulnerable to fishing at these predictable times and locations and the eggs and juveniles are highly susceptible to contamination. The value of these aggregations therefore includes the value of the fish in a healthy fishing industry. The value also includes the existing and potential ecotourism on these dramatic events. Recent work has illustrated that the value of spawning aggregations for ecotourism far exceeds Snapper and grouper assemblages of Belize, Heyman and Kobara  46 their value in commercial fisheries (Sala et al., 2001). Further, dive ecotourism does not appear to negatively affect the fishes‘ courtship or spawning behaviour (Heyman et al., 2010). As Belize has the best well documented and dramatic reef fish spawning aggregations in the world, ecotourism on spawning aggregations represents significant and relatively untapped, sustainable tourism development potential. CONCLUSION Are the risks of oil drilling worth taking? If well managed, there is a very low probability of large spills and accidents in oil drilling, exploration and development. But, in the unlikely event that such a spill does occur, there will likely be catastrophic effects on coral reef ecosystems. The long term economic and employment benefits from fisheries and tourism industries that can persist with a healthy coral reef system will be significantly degraded by an oil spill. However, small the risk, and however large the economic gains from oil extraction, the risk of the catastrophic damages that will ensue are probably not worth taking. ACKNOWLEDGEMENTS First and foremost, we wish to acknowledge the collaboration of the Belize National Spawning Aggregations Working Committee and all its local and international individuals and entities including the Belize Fisheries Department, Belize Fisheries Cooperative Association, University of Belize, Belize Audubon Society, Toledo Institute for Development and Environment, Southern Environmental Association, Wildlife Conservation Society, World Wildlife Fund for Nature, the Nature Conservancy, and local fishers. Most importantly, we wish to acknowledge the support and collaboration of fishers throughout Belize including Eloy Cuevas, Eloy Cuevas, Jr., Sonny Garbutt, Jason Williams, Anselmo J. Nunez, K.K. Nunez, Jack Young, Brian Young, Matteus (Tamba) Nicholas, and Victor Jacobs, Jr., Victor Jacobs, Sr. (deceased), and Jack Cabral (deceased). We acknowledge the extensive scientific and logistical contributions of Nicanor Requena, Chris Houser, and Tal Ezer. The Nature Conservancy, the Summit and Oak Foundations, Conservation International, and Texas A&M University have supported this research over the years. Thanks to Oceana and The Sea Around Us Project for hosting this meeting. All of this research has been conducted under various permits from the Fisheries Department of Belize. REFERENCES Arreguín-Sanchez, F., Munro, J.L., Balgos, M.C., Pauly, D. (Editors), 1996. Biology, fisheries and culture of tropical groupers and snappers. ICLARM Conference Proceedings 48. Manila, Philippines. 449 p. Cooper, E., Burke, L., Bood, N. 2008. Coastal Capital: Economic Contribution of Coral Reefs and Mangroves to Belize. Washington DC: World Resources Institute. Ezer, T., Heyman, W.D., Houser, C., Kjerfve, B. 2010. Modeling and observations of high frequency flow variability and internal waves at a Caribbean reef spawning aggregation site. Ocean Dynamics 60(5), 1307-1318. Ezer, T., Thattai, D.V., Kjerfve, B., Heyman, W.D., 2005. On the variability of the flow along the Meso-American Barrier Reef System: A numerical model study of the influence of the Caribbean Current and eddies. Ocean Dynamics 55, 458-475. Gibson, J., Pott, R.F., Paz, G., Majil, I., Requena, N., 2007. Experiences of the Belize Spawning Aggregation Working Group. Proceedings of the Gulf and Caribbean Fisheries Institute 59, 389-396. Heyman, W.D., 2011. Elements for building a participatory ecosystem-based marine reserve network. The Professional Geographer, 63(4), DOI:10.1080/00330124.2011.585078. Heyman, W.D., Carr, L., Lobel, P., 2010. Diver disturbance to reef fish spawning aggregations: It is better to be disturbed than to be dead. Marine Ecology Progress Series 419, 201-210. Heyman, W.D., Kjerfve, B., 2008. Multi-species reef fish spawning aggregations at Gladden Spit, Belize. Bulletin of Marine Science 83(3), 531-551. Heyman, W.D., Kjerfve, B., Ezer, T., 2008. Mesoamerican reef spawning aggregations help maintain fish populations: a review of connectivity research and priorities for science and management. NOAA Marine Sanctuaries Conservation Series NMSP- 08-07, 150-169. Heyman, W.D., Kjerfve, B., Johannes, R.E., Graham, R., 2001. Whale sharks, Rhincodon typus, aggregate to feed on fish spawn in Belize. Marine Ecology Progress Series 215, 275-282. Heyman, W.D., Kjerfve, B., Rhodes, K.L., Graham, R.T., Garbutt, L., 2005. Cubera snapper, Lutjanus cyanopterus, spawning aggregations on the Belize Barrier Reef over a six year period. Journal of Fish Biology 67(1), 83-101. Heyman, W.D., Kobara, S., 2011. Reef geomorphology influences sites for reef fish spawning aggregations in the Caribbean. Ch. 26 In: Harris, P.T., Baker, E.K. (eds.), Seafloor Geomorphology as Benthic Habitat: Geohab Atlas of Seafloor Geomorphic Features and Benthic Habitats. Elsevier, Amsterdam. Too Precious to Drill: the Marine Biodiversity of Belize, Palomares and Pauly  47 Kobara, S., Heyman, W.D., 2010. Sea bottom geomorphology of multi-species spawning aggregation sites in Belize. Marine Ecology Progress Series 405, 243-254. Kobara, S., Heyman, W.D., 2008. Geomorphometric patterns of Nassau grouper (Epinephelus striatus) spawning aggregation sites in the Cayman Islands. Marine Geodesy 31(4), 231-245. National Research Council (NRC), 2005. Understanding Oil Dispersants: Efficacy and Effects. National Academies Press, Washington, DC. 396 p. Sala, E., Ballesteros, E., Starr, R.M., 2001. Rapid decline of Nassau grouper spawning aggregations in Belize: fishery management and conservation needs. Fisheries 26, 23-30. Tunnel, J.W., Jr., 2011. An expert opinion of when the Gulf of Mexico will return to pre-spill harvest status following the BP Deepwater Horizon MC 252 oil spill. Harte Research Institute for Gulf of Mexico Studies, Texas A&M Corpus Christi. 52 pp. Wilson, K.W., 1977. Acute toxicity of oil dispersants to marine fish larvae. Marine Biology 40, 65-64.  Discovering new fish species in Belize, Lobel and Lobel  48 ENDEMIC MARINE FISHES OF BELIZE: EVIDENCE OF ISOLATION IN A UNIQUE ECOLOGICAL REGION1 Phillip S. Lobel and Lisa K. Lobel Department of Biology, Boston University, 5 Cummington Street, Boston, MA 2215 USA; plobel@bu.edu ABSTRACT The Meso-Amercian Barrier Reef (MABR) forms a physical boundary enclosing a large coastal lagoon that runs the length of the country of Belize. This creates a semi-enclosed body of water that is a mix of oceanic water and freshwater river input. Thus, the Belize lagoon ecosystem is unique in the western hemisphere and represents a distinct biogeographical province with special water quality. The majority of coral reef fishes have pelagic larvae that spend weeks in the plankton during their early development. This is a highly dispersive phase of the life history of marine fishes. As a result, most Caribbean species range throughout the Caribbean Sea. Even so, some taxa show strong local selection and restricted biogeographic distributions. In Belize, this is especially evident and probably due in large part to the physical barrier of the reef and the special quality of the marine water in the lagoon system. Recent studies of the fishes inside the MABR have discovered several new species of fishes and many of these are endemic to Belize. A preliminary estimate of endemic fishes in Belize yields a count of 12 species found only in the lagoonal area and another 8 species found on the outer barrier reef and the atolls. INTRODUCTION Belize is in a corner of the Caribbean Sea that is bounded by unique oceanographic conditions and the distinctive barrier reef. The majority of fishes that occur in Belize are the same species that are found everywhere else in the Caribbean. But, there are exceptions; there are some species found only in Belize and nowhere else. These are scientifically designated as ‗endemics‘. The endemic marine fishes of Belize are evidence of biological isolation in a unique ecological region. The biological questions are ‗what local ecology are these endemic fishes adapted to?‘ and ‗how do these species maintain their genetic uniqueness?‘ Such questions are complicated and require learning the details about the fish‘s natural history. Many marine animals have early life histories as embryos and small larvae which drift in the plankton. These animals spawn free floating eggs and their offspring drift away. As such, these propagules are easily dispersed over vast distances creating large populations of single species. Such is the case for many of the most familiar fishes such as groupers, snappers, surgeonfishes and many other reef fishes. But, some other fishes have evolved different reproductive tactics that result in their planktonic larvae staying close to natal reefs. Some of these fishes have developed non-dispersing embryos and larvae and others have adapted their reproduction and larval dispersal to occur during times when oceanographic conditions entrain and retain the larvae near home reefs. The Belize Barrier Reef System is a semi- enclosed marine habitat. Even though it does pulse with tides and there is an influx of oceanic waters through myriad channels. The bulk of the water inside the Barrier Reef has a higher retention or residency time than it would have otherwise if that barrier reef was not present. It is important to describe biogeographically restricted species for purposes of conservation and as key indicator species used in ecological monitoring for local effects. To date, there have been about 500 fish species found in Belize waters (see Palomares and Pauly, this volume). David Greenfield pioneered ichthyological surveys in Belize in the 1970s. Subsequently scientists from the Smithsonian museum have been the leading force in documenting Belize reef communities and describing new species (e.g. Baldwin, Faust, Lavett-Smith, Ruetzler, Tyler and others).  1 Cite as: Lobel, P.S., Lobel, L.K., 2011. Endemic marine fishes of Belize: evidence of isolation in a unique ecological region. In: Palomares, M.L.D., Pauly, D. (eds.), Too Precious to Drill: the Marine Biodiversity of Belize, pp. 48-51. Fisheries Centre Research Reports 19(6). Fisheries Centre, University of British Columbia [ISSN 1198-6727]. Too Precious to Drill: the Marine Biodiversity of Belize, Palomares and Pauly  49 This report highlights the several new fish species that we have found in Belize with help and collaboration of colleagues. Big fishes are easier to find than little fishes. The new discoveries that we are making of previously unknown species in Belize waters have one key common trait. The new fishes are small and easily overlooked or mistakenly misidentified. In some cases, it required DNA data combined with color photographs of live fish to make the case distinguishing the Belize population as a distinct species.   Cleaner Goby, Elacatinus lobeli (Randall and Colon, 2009). This species is similar in color to the neon goby found in Florida, but it is genetically and morphologically distinct. It is a cleanerfish and removes ectoparasites from host fishes. It is endemic to the MABR, and is found broadly inside the MABR and on the atolls. It was named for Phillip S. Lobel who photographed and collected type specimens. This probable new species of Banner Goby, Microgobius is found only in deep sand flats areas (>15 m) in the inter-reefal channels between mangrove islands in Belize MABR. It is very hard to see as it prefers murky water, is very skittish and avoids divers. This species is being diagnosed for description by the authors with J. Randall.   Sponge Goby, Elacatinus coloni (Randall and Lobel, 2009). This species is a sponge dweller and feeds on a polychaete sponge parasite. It is endemic and found only inside the MABR usually in tube sponges. Atoll Goby, Elacatinus nov. sp. (Lobel and Kaufman, in prep.). On their first scuba dive at Lighthouse Atoll, Lobel and Kaufman discovered this new species! This was the weekend before the February 2010 MMAS meetings (Belize City). We returned to the atoll after the meetings and collected the type specimens. It has blue stripes similar to E. lobeli but this new fish is distinct by having a white-ish nose spot and is only found at deeper depths (>30 m).  Discovering new fish species in Belize, Lobel and Lobel  50  The ‘Maya hamlet’ is a new species of Hypoplectrus found only in Belize. The manuscript describing this species is in press (Lobel, 2011). The Social wrasse, Halichoeres socialis Randall and Lobel, 2003. Male and female (in back). This new Belize fish species was discovered during the Lobels‘ first field trip to Belize in 1993. Continued study of its biology and biogeographic distribution is an annual project conducted by students in Professor P.S. Lobel‘s coral reef ecology field course, Boston University Marine Program.  A preliminary listing of marine fishes with distributions mainly within the MABR system including the outer reef and atolls is presented in Table 1. A few of these species (e.g., E. lori, T. clarkii, T. briggsi) possibly range as far as the Bay Islands, Honduras, which are nearby the southern margin of the MABR. More research is needed to better define the biogeography of these species. ACKNOWLEDGEMENTS This report summarizes research accomplished under annual permits from the Department of Fisheries, Belize, 1997-2010. Research was supported by Conservation International Marine Management Area Science program, Boston University, The Legacy Program USA, The Ross and Edwards families of Lighthouse Atoll, The former Friends of Nature, and the Southern Environmental Association. Much of the field work was based from the Wee Wee Cay Marine Laboratory. We are appreciative that this work needed the help of friends and colleagues to succeed: Horace and Sharon Andrews, Mary and Paul Shave, Shelly and Clifford Robinson, Udell Foreman, David Greenfield, Jack Randall, Pat Colin, Will Heyman, Eli Romero, Les Kaufman, Margo Stiles, Lindsay Garbut and many others, many thanks. All photographs by Phillip S. Lobel (copyrighted, all rights retained). Table 1. Preliminary listing of marine fishes with distributions mainly within the MABR system including the outer reef and atolls. Species whose distribution is so far known only from the lagoons inside of the MABR are noted with an asterisk (*). Family Genus and species Reference Batrachoididae Sanopus greenfieldorum * Collette 1983  Triathalassothia gloverensis Greenfield and Greenfield 1973  Opsanus dichrostomus * Collette 2001  Sanopus astrifer Robins and Starck 1965 Chaenopsidae Acanthemblemaria paula * Johnson and Brothers 1989  Emblemariopsis ruetzleri * Tyler and Tyler 1997  Emblemariopsis dianae * Tyler and Hastings 2004 Gobiesocidae Tomicodon lavettsmithi * Williams and Tyler 2003  Tomicodon clarkei Williams and Tyler 2003  Tomicodon briggsi Williams and Tyler 2003 Gobidae Elacatinus coloni * Randall and Lobel 2009  Elacatinus lobeli Randall and Colin 2009  Elacatinus lori Colin 2002  Elacatinus sp nov Lobel and Kaufman ms  Microgobius sp nov * Lobel, Lobel and Randall ms  Psilotris amblyrhynchus Smith and Baldwin 1999 Blennidae Starksia weigti * Baldwin et al. 2011  Starksia sangreyae * Baldwin et al. 2011 Serrranidae Hypoplectrus sp nov * Lobel et al. 2009, Lobel in press Labridae Halichoeres socialis * Randall and Lobel 2003   New species of invertebrate found in Belize, December 2010. P. Lobel found this undescribed phoronid worm in the muddy bottom habitat in deep channels between cays in the lagoon. In February 2011, Lobel returned with Prof. G. Giribet (Harvard) and collected specimens. The description of this new species is in preparation. The worm has a burrow in the sand and it will extend itself about 5 cm above the surface, but retracts when disturbed. It is one of only about 12 species of this kind of phoronid worldwide and one of the few tropical ones.     Too Precious to Drill: the Marine Biodiversity of Belize, Palomares and Pauly  51 REFERENCES Baldwin, C.C., Castillo, C.I., Weigt, L.A., Benjamin, V.C., 2011. Seven new species within western Atlantic Starksia atlantica, S. lepicoelia, and S. sluiteri (Teleostei, Labrisomidae), with comments on congruence of DNA barcodes and species. Zookeys 79, 21- 72. Colin, P.L., 2002. A new species of sponge-dwelling Elacatinus (Pisces: Gobiidae) from the western Caribbean. Zootaxa 106, 1-7. Collette, B.B., 1983. Two new species of coral toadfishes, family Batrachoididae, genus Sanopus, from Yucatan, Mexico, and Belize. Proceedings of the Biological Society of Washington 96, 719-724. Collette, B.B., 2001. Opsanus dichrostomus, a new toadfish (Teleostei: Batrachoididae) from the Western Caribbean Sea and Southern Gulf of Mexico. Ocassional Papers of the Museum of Zoology University of Michigan 731, 1-16. Greenfield, D.W., Greenfield, T., 1973. Triathalassothia gloverensis, a new species of toadfish from Belize (=British Honduras) with remarks on the genus. Copeia, 1973(3), 560-565. Johnson, G.D., Brothers, E.B., 1989. Acanthemblemaria paula, a new diminutive chaenopsid (Pisces: Blennioidei) from Belize, with comments on life history. Proceedings of the Biological Society of Washington 102, 1018-1030. Lobel, P.S., 2011. A review of the hamletfishes (Serranidae, Hypoplectrus) with description of two new species. Zootaxa in press. Lobel, P.S., Rocha, L., Randall, J.E., 2009. The color phases and distribution of the western Atlantic labrid fish, Halichoeres socialis. Copeia 2009(1), 171-174. Randall, J.E., Colin, P.L., 2009. Elacatinus lobeli, a new cleaning goby from Belize and Honduras. Zootaxa 2173, 31-40 Randall, J.E., Lobel, P.S., 2003. Halichoeres socialis, a new labrid fish from Belize, Caribbean. Copeia 2003(1), 124-130 Randall, J.E., Lobel, P.S., 2009. A literature review of the sponge-dwelling gobiid fishes of the genus Elacatinus from the western Atlantic, with description of two new Caribbean species. Zootaxa 2133, 1-19 Robins, C.R., Starck, W.A. II, 1965. Opsanus astrifer, a new toadfish from British Honduras. Proceedings of the Biological Society of Washington 78, 247-250. Smith, S.G., Baldwin, C.C., 1999. Psilotris amblyrhynchus, a new seven-spined goby (Teleostei: Gobiidae) from Belize, with notes on settlement-stage larvae. Proceedings of the Biological Society of Washington 112, 433-442. Tyler, J.C., Hastings, P.A., 2004. Emblemariopsis dianae, a new species of chaenopsid fish from the western Caribbean off Belize (Blennioidei). Aqua, Journal of Ichthyology and Aquatic Biology 8(4), 9-60. Tyler, D.M., Tyler, J.C., 1997. A new species of chaenopsid fish, Emblemariopsis ruetzleri, from the western Caribbean off Belize (Blennioidei), with notes on its life history. Proceedings of the Biological Society of Washington 110, 24-38. Williams, J.T., Tyler, J.C., 2003. Revision of the western Atlantic clingfishes of the genus Tomicodon (Gobiesocidae), with descriptions of five new species. Smithsonsonian Contribributions to Zoolology 621, 1-26.  Funcitonal importance of Belize coral reefs, Wulff  52 FUNCTIONAL IMPORTANCE OF BIODIVERSITY FOR CORAL REEFS OF BELIZE1 Janie Wulff Department of Biological Science, Florida State University, Tallahassee, FL, USA; wulff@bio.fsu.edu ABSTRACT A thriving coral reef results from an intricate collaboration among many different kinds of animals, plants, and micro-organisms. Some of the key collaborators include nearby seagrasses and mangroves that capture and control sediments and transform dissolved nutrients into plant biomass, and herbivorous fishes and sea urchins that prevent quickly growing algae from overwhelming reefs. But most central to the building and maintenance of the reefs are corals and sponges, and the microbial collaborators that live within their bodies. Reef-building corals deposit solid carbonate skeletons as they grow, building a sturdy 3-dimensional framework within which fishes, crustaceans, and other animals shelter and find food, while sponges glue living corals onto the reef frame and protect them from excavators, facilitate regeneration of damaged reefs, and keep the water clear by efficiently filtering bacteria and phytoplankton. All of these functional roles must be played for a reef to remain healthy and capable of recovering from damage. Coral reefs, as shallow-water tropical ecosystems, have always been challenged by physical damage due to hurricane-charged water movement, and more recently, by pulses of freshwater and sediment due to heavy coastal rains, and temporarily extreme temperatures. Recovery from effects of these challenges is a normal part of the dynamics of healthy coral reefs. High species diversity of corals and sponges is essential to successful recovery because species differ in their ability to: a) resist challenges (physical disturbance, disease, high or low temperatures, sediment, etc.), b) recover from challenges (by regeneration, regaining symbionts after bleaching, halting the advance of disease, etc.), c) recover in the sense of recolonization by the next generation, and d) host symbionts and engage in other interactions that increase survival of participants. As well, individuals within a species vary in their ability to resist or recover from challenges and to interact positively with other organisms. When high biodiversity is protected, there are always at least some species capable of performing each of the roles essential to the functioning of the reef - even when other species are temporarily diminished by their vulnerability to a particular environmental challenge. However, when multiple challenges occur together, or when the challenges are novel (i.e., exposure to substances that humans have manufactured or released from inside the earth), too many species may be diminished or deleted simultaneously, impairing the natural growth and recovery processes. INTRODUCTION Coral reefs, as shallow-water tropical ecosystems, have always been challenged by physical damage due to hurricane-charged water movement, and more recently by pulses of fresh-water and sediment due to heavy rains on deforested coasts, and temporarily extreme temperatures. Recovery from effects of environmental challenges is a normal part of the dynamics of healthy coral reefs. High species diversity of corals and sponges is essential to successful recovery because species differ in their ability to: a) resist challenges (physical disturbance, disease, high or low temperatures, sediment, etc.); b) recover from challenges (by regeneration, regaining symbionts after bleaching, halting the advance of disease, etc.); c) recover in the sense of recolonization by the next generation; and d) host symbionts and engage in other interactions that increase survival of participants. As well, individuals within a species vary in their ability to resist or recover from challenges and to interact positively with other organisms. When high biodiversity is protected, there are always at least some species capable of performing each of the roles essential to the functioning of the reef—even when other species are temporarily diminished by their vulnerability to a particular environmental challenge. However, when multiple challenges occur together,  1 Cite as: Wulff, J., 2011. Functional importance of biodiversity for coral reefs of Belize. In: Palomares, M.L.D., Pauly, D. (eds.), Too Precious to Drill: the Marine Biodiversity of Belize, pp. 52-56. Fisheries Centre Research Reports 19(6). Fisheries Centre, University of British Columbia [ISSN 1198-6727]. Too Precious to Drill: the Marine Biodiversity of Belize, Palomares and Pauly  53 or when the challenges are novel (i.e., exposure to substances that humans have manufactured or released from inside the Earth, such as oil), too many species may be diminished or deleted simultaneously, impairing the natural growth and recovery processes. CORALS AND SPONGES Corals and sponges spend their adult lives attached to the substratum on which they settle as waterborne larvae, and they illustrate great variety in shape and size, facility at asexual propagation and regeneration, and tendency to host microbes within their bodies. Corals and sponges differ from each other in important ways that underlie the compatible roles they play in building, maintaining, and repairing coral reefs. Corals Corals deposit rock-like calcium carbonate skeletons as they grow, creating the basic building blocks of the reef structure. Among the 45 reef-building corals inhabiting the Belize Barrier Reef (Bright and Lang, 2011) are a great variety of shapes, including branching, plate-shaped, pillar, massive mounds, and encrusting forms. Whatever the overall shape of a colony, the living tissue is always a very thin layer over the surface. Thus even shallow wounds expose skeleton, making it vulnerable to colonization by quickly growing algae that can inhibit regeneration, and also by excavating organisms which can bore into the solid carbonate of the coral skeleton, weakening its attachment to the reef frame. Although coral polyps can capture plankton with their tentacles, they acquire most of their food from the single celled algae, called zooxanthellae, that live at high densities within their tissue. Like all plants, zooxanthellae convert sunlight energy into food energy. Their position within the corals enhances their access to nutrients due to recycling of metabolic wastes. Although this collaboration is unquestionably beneficial to the corals, as their chief food source, dependence on zooxanthellae makes corals vulnerable to the possibility that the association may break down under stressful environmental conditions. In particular, abnormally high sea surface temperatures cause zooxanthellae to be expelled by their coral hosts. Moderation of temperatures can allow recolonization of corals, but bleaching can weaken corals, making them more vulnerable to other threats such as diseases. If zooxanthellae are unable to recolonize quickly, the corals die. Sponges Most of the over 800 species of sponges (Diaz and Rützler, this volume) that inhabit Caribbean coral reefs and associated habitats have soft bodies with living tissue throughout. Their skeletons, which homogeneously pervade the living tissue, are made of fine meshworks of protein fibers, generally augmented by silica spicules. Sponges are also pervaded by a system of canals through which they pump water, from which they filter bacteria and other very small particles extremely efficiently. The extraordinarily simple internal structure of sponges bestows on them great flexibility in growing around obstacles, adjusting to changes in orientation, and accommodating close associations with other organisms. Because sponges are living tissue throughout, wound healing can be achieved quickly, by simply reconstituting the layer of specialized cells that cover the surface; thus sponges are masters of regeneration after damage or fragmentation (Wulff, 2011). ROLES OF CORALS AND SPONGES IN BUILDING, MAINTAINING, AND REPAIRING CORAL REEFS Growth of corals is required for generating the solid carbonate building blocks of reef framework. But, even as they accrete, coral skeletons are also eroded by grazing fishes and sea urchins, and by a handful of bivalve and sponge species that transform solid carbonate to fine sediment, as they excavate burrows for themselves. Excavations can erode coral basal attachments to the point that corals relinquish their grip on the reef frame, often perishing in the surrounding sediments or cascading into deeper water. Fortunately, sponges associated with corals can increase coral survival by gluing them to the reef frame. Experimental removal of sponges from fore-reef patches resulted in 40% of the corals becoming disengaged from the reef frame; while on similar patch reefs with intact sponges coral mortality was only 4% (Wulff and Buss, 1979). This collaboration of solid rock-generating corals with sponges capable of adhering corals to the reef frame is further enhanced as the sponges filter the entire water column above the reef each day, maintaining water clarity that allows corals to receive adequate sunlight for their zooxanthellae. Physical damage to coral reefs, on scales ranging from small patches to many square kilometers, is inevitable given the coincident geographic distribution of coral reefs and tropical storms. The ability to recover is a normal part of the life histories of coral and sponge species, and repair and regeneration is a Funcitonal importance of Belize coral reefs, Wulff  54 normal part of coral reef growth. At any moment portions of a reef system have been recently damaged by a storm, so the process of regeneration of rubble mounds into solid reef frame onto which living corals can flourish once again is required for continued growth of coral reefs to keep pace with rising sea level. Large pieces of damaged or dead coral may remain stable where they fall at the end of a storm, but smaller rubble pieces can continue to be churned by foraging fishes or water motion, impeding their incorporation into a stable structure. Coral larvae that settle on loose rubble tend to be smashed, as rubble pieces are moved against each other. Because sponges can adhere quickly to solid carbonate with any part of themselves, the same gluing capability that allows them to bind living corals to the reef frame also allows them to bind piles of loose rubble into continuous structures. Once loose rubble pieces are stabilized, crustose coralline algae can grow from one piece of rubble to the next, cementing them together, rendering them more hospitable to small corals (Wulff, 1984). Sediment generated by grazing and excavating organisms fills in the holes in the frame, increasing solidity. Growth of corals continues the cycle. Tropical storms have challenged coral reefs as long as they have existed, but additional challenges have been increasing in importance: pulses of freshwater and sediment running off of deforested land, bleaching due to increased sea surface temperatures, coral predators that are no longer kept in check by their larger predators that have been overfished, and diseases that are poorly fended off by animals that are stressed by other challenges. Each of the many species of corals and sponges that participate in reef building and re-building is characterized by a unique set of strengths and vulnerabilities, and no single species is the best at coping with all environmental challenges. Species that rebound gracefully after a storm may succumb to disease, while species that resist bleaching may be overwhelmed by uninhibited predators, and those most resistant to predators may be devastated by storms. In the following section, examples illustrating the wide range of variation in resistance to and recovery from a few of the challenges faced by sessile animals on reefs are drawn from the diverse species inhabiting the Belize Barrier Reef. VARIATION AMONG SPECIES IN RESISTANCE TO, AND RECOVERY FROM, CHALLENGES Physical damage by storms Massive corals, such as Montastraea species, are champion survivors of hurricanes, remaining standing amidst a litter of fragments of branching species and broken off corals with small basal attachments. Branching species of both corals and sponges, although readily broken tend to be especially adept at recovering from breakage, as fragments can reattach to the substratum, and branching patterns adjust to their new orientation as fragments continue to grow. Thus moderate storms can result in propagation, but the violent water motion of major hurricanes can overdo breakage to the point of destruction (e.g., Woodley et al., 1981). Corals with smaller forms and shorter life spans may be readily damaged by storms, but tend to be successful at replenishing their populations by efficient settlement of larvae (e.g., Bruckner and Hill, 2009). Sponge species also balance resistance to damage with recovery in a variety of ways. After Hurricane Allen in Jamaica in 1980, monitoring of nearly six hundred sponges over 5 weeks for recovery revealed an inverse relationship between ability to resist damage and ability to recover from damage (Woodley et al., 1981; Wulff, 2006b). Erect branching species suffered the most damage, but they were also most adept at recovering; while at the opposite extreme, many sponges of species that live confined to cryptic spaces within the reef frame eluded damage altogether in their protected microhabitat; however, those that were exposed as the framework was ripped apart did not recover at all. Massive sponges with tough skeletons were highly resistant to being damaged, but when they were damaged, recovery was ilusive. These massive, tough species were able to recoup their substantial losses, however, by recolonizing the battered reefs with their next generation (Wilkinson and Cheshire, 1988). Bleaching Variation in susceptibility to bleaching varies with the coral species, clade of zooxanthellae hosted, and habitat details (e.g., Baker, 2003). Variation among species can be extreme, as in a 2005 bleaching event during which 85% of colonies of the relatively small massive coral Porites astreoides were resistant, but fewer than 5% of the colonies of the large massive corals in the Montastraea annularis species complex remained unbleached (Bruckner and Hill, 2009). Closely related coral species can differ in vulnerability, for example the plate-shaped Agaricia agaricites tends to be able to cope with higher temperatures better than closely related Agaricia tenuifolia (Robbart et al., 2004). The net result of bleaching is a combination Too Precious to Drill: the Marine Biodiversity of Belize, Palomares and Pauly  55 of susceptibility to bleaching and ability to recover. Ultimate results of very similar rates of severe bleaching in the brain coral Colpophyllia natans and the short thickly branched Porites porites (92% and 97% of colonies, respectively) were very different, with 88% of completely bleached Colpophyllia recovering, but only 28% of completely bleached Porites recovering (Whelan et al., 2007). Individuals within a species also vary in their ability to cope with environmental challenges. In the case of species that are capable of propagation by fragmentation, it is possible that relatively resistant genotypes will be able to quickly increase in abundance. Genotypes of staghorn and elkhorn coral that have demonstrated particular resistance are currently being propagated in nurseries in Laughing Bird Caye National Park, Belize, in order to bolster natural replenishment of reefs (Carne, in press). Disease Coral diseases are not generally specific to a single species, but there are patterns in the tendency of a particular disease to affect certain corals (Bruckner, 2009), complicated by the recent history of bleaching and other weakening circumstances. For example, the ultimate fates of the Porites porites and Colpophyllia natans colonies in the bleaching recovery study mentioned in the previous paragraph were high mortality all around, because the Colpophyllia colonies that recovered from bleaching succumbed to White-plague type II disease (Whelan et al., 2007). Diseases have disproportionately influenced populations of some of the most important and abundant Caribbean reef coral species. The near demise of the Acropora species, staghorn and elkhorn corals, that contributed rapid growth and facile recovery from damage to shallow reef zones, has been attributed to white band disease; and populations of the large, long-lived massive Montastraea species, so highly resistant to physical damage, have been heavily influenced by yellow band and white plague diseases. Short-lived, smaller-colony species have been less affected (e.g., Bruckner et al., 2009). Sponge diseases by contrast tend to be quite specific to particular species. Disease may be having a profound effect on sponge species diversity. By the end of a 14 year study on a shallow reef at a remote site, 20 of the 39 sponge species present at the start had vanished, with disease the most likely culprit (Wulff, 2001, 2006a). CONCLUSIONS Complementary roles played by corals and sponges in reef building, maintenance, and repair are all required to the point that if any are not performed, the entire enterprise can fail. But, why do we need to be concerned about keeping more than a few species of each alive and well? Species of corals and sponges that build, maintain, and repair coral reefs have evolved in a context that has provided the selective impetus for an effective balance between resistance to, and recovery from, physical disturbance by tropical storms. Species less resistant to damage make up for that by effective individual recovery by regeneration or by population level recovery by recolonization. When threats are relatively novel, as are bleaching and disease, strategies that compensate for lack of resistance are much less evident, perhaps reflecting the lack of time for evolution in response to these threats. Species that appear especially vulnerable are failing to exhibit effective recovery. Oil is not a substance to which corals and sponges have had a chance to evolve strategies for either resistance or recovery. In 1986, an oil spill in Bahia las Minas, near the Caribbean terminus of the Panama canal, killed many corals outright, resulting in an immediate decrease in coral cover by 76% at 3 m depth or less, and 45% at 9 to 12 m depth (Jackson et al., 1989). After 5 years, recovery was still not apparent. Corals on oiled reefs had slower growth and higher injury rates, and there was practically no recruitment of the next generation (Guzmán et al., 1994). Effects of oil on sponges are much less understood, in large part because sponges vanish so quickly after they are killed that they are invisible to any monitoring that is not immediate. Highly efficient filtering of large volumes of water may render sponges especially vulnerable to oil that has been broken into fine suspended droplets with chemical dispersants. Lingering effects of the Panama oil spill were in part due to continual re-oiling, every time sediments in which oil had become buried were resuspended by water movement (Levings et al., 1994). High biodiversity ensures functional redundancy of species that differ in how gracefully they cope with temperature extremes, disease, and physical damage so that there are always at least some species capable of performing each of the roles essential to the functioning of the reef - even when other species are temporarily diminished by their vulnerability to a particular environmental challenge. However, when multiple challenges occur together, or when the challenges are novel, as oil is, too many species may be diminished or deleted simultaneously, impairing the natural growth and recovery processes. Given the Funcitonal importance of Belize coral reefs, Wulff  56 inability of slow-recovering species to resist novel threats, it seems rash to risk the addition of oil to the many other threats currently facing corals and sponges of the Belize Barrier Reef. ACKNOWLEDGEMENTS My funding sources for research in Belize are the Caribbean Coral Reef Ecosystems Program CCRE) of the US National Museum of Natural History, the Marine Science Network of the Smithsonian Institution, funded in part by the Hunterdon Oceanographic Research Endowment, and the National Science Foundation (Grant number 0550599). I am grateful to Deng Palomares and Daniel Pauly for organizing the symposium on the Biodiversity of the Belize Barrier Reef of which this paper is a part. This is CCRE Contribution #907. REFERENCES Baker, A.C., 2003. Flexibility and specificity in coral-algal symbiosis: Diversity, ecology, and biogeography of Symbiodinium. Annual Review of Ecology and Systematics 34, 661-689. Bright, T., Lang, J.C., 2011. Picture Guide to Stony Corals of Glover‘s Reef Atoll. Wildlife Conservation Society http://www.gloversreef.org/grc/pdf/stony_corals_picture_guide_1-30-11.pdf. Bruckner A.W., 2009. Field Guide to Western Atlantic Coral Diseases. USDC National Oceanic and Atmospheric Administration, Silver Spring, MD. http://cdhc.noaa.gov/disease/default.aspx>http://cdhc.noaa.gov/disease/default.aspx. Bruckner, A.W., Hill, R.L., 2009. Ten years of change to coral communities off Mona and Desecheo Islands, Puerto Rico, from disease and bleaching. Diseases of Aquatic Organisms 87, 19-31. Carne, L., in press. Strengthening coral reef resilience to climate change impacts: A case study of reef restoration at Laughing Bird Caye National Park, southern Belize. World Wildlife Fund. Guzmán, H.M., Burns, K.A., Jackson, J.B.C., 1994. Injury, regeneration and growth of Caribbean reef corals after a major oil spill in Panama. Marine Ecology Progress Series 105, 231-241. Jackson, J.B.C., Cubit, J.D., Keller, B.D., Batista, V., Burns, K., Caffey, H.M., Caldwell, R.L., Garrity, S.D., Getter, C.D., Gonzalez, C., Guzmán, H.M., Kaufmann, K.W., Knap, A.H., Levings, S.C., Marshall, M.J., Steger, R., Thompson, R.C., Weil, E., 1989. Ecological effects of a major oil spill on Panamanian coastal marine communities. Science 243, 37-44. Levings, S.C., Garrity, S.D., Burns, K.A., 1994. The Galeta oil spill 3. Chronic re-oiling, long-term toxicity of hydrocarbon residues on epibiota in the mangrove fringe. Estuarine, Coastal and Shelf Science 38, 365-395. Robbart, M.L., Peckol, P., Scordilis, S.P. Curran, H.A., Brown-Saracino, J., 2004. Population recovery and differential heat shock protein expression for the corals Agaricia agaricites and A. tenuifolia in Belize. Marine Ecology Progress Series 283, 151-160. Whelan, K.R.T., Miller, J., Sanchez, O., Patterson, M., 2007. Impact of the 2005 coral bleaching event on Porites porites and Colpophyllia natans at Tektite Reef, US Virgin Islands. Coral Reefs 26, 689-693. Wilkinson, C.R., Cheshire, A.C., 1988. Growth rate of Jamaican coral reef sponges after Hurricane Allen. Biological Bulletin 175, 175- 179. Woodley, J.D., Chornesky, E.A., Clifford, P.A., Jackson, J.B.C., Kaufman, L.S., Lang, J.C., Pearson, M.P., Porter, J.W., Rooney, M.C., Rylaarsdam, K.W., Tunnicliffe, V.J., Wahle, C.W., Wulff, J.L., Curtis, A.S.G., Dallmeyer, M.D., Jupp, B.P., Koehl, M.A.R., Neigel, J., Sides, E.M., 1981. Hurricane Allen‘s impact on Jamaican coral reefs. Science 214, 749-755. Wulff, J.L., Buss, L.W., 1979. Do sponges help hold coral reefs together? Nature 281, 474-475. Wulff, J.L., 1984. Sponge-mediated coral reef growth and rejuvenation. Coral Reefs 3, 157-163. Wulff, J.L., 2001. Assessing and monitoring coral reef sponges: Why and how? Bulletin of Marine Science 69, 831-846. Wulff, J.L., 2006a. Rapid diversity and abundance decline in a Caribbean coral reef sponge community. Biological Conservation 127, 167-176. Wulff, J.L., 2006b. Resistance vs. recovery: morphological strategies of coral reef sponges. Functional Ecology 20, 699-708. Wulff, J.L., 2006c. Ecological interactions of marine sponges. Canadian Journal of Zoology Special Series 84, 146-166. Wulff, J.L., 2011. Sponges. In: Hopley, D. (ed.), Encyclopedia of Modern Coral Reefs: Structure, Form and Process. Springer, Heidelberg.  Too Precious to Drill: the Marine Biodiversity of Belize, Palomares and Pauly  57 BIODIVERSITY OF SPONGES: BELIZE AND BEYOND, TO THE GREATER CARIBBEAN1 Maria Cristina Diaz Museo Marino de Margarita, Blvd. El Paseo, Boca del Río, Margarita, Edo. Nueva Esparta, Venezuela; taxochica@gmail.com Klaus Rützler Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560-0163, USA; ruetzler@si.edu ABSTRACT Sponges represent one of the most diverse benthic faunal groups in subtidal habitats of Caribbean coral reefs and mangroves. On coral reefs, sponges (100-261 species) surpass the species richness of other conspicuous reef organisms, such as octocorals (60-80 species) and scleractinian corals (50-60 species). In the past 35 years, researchers supported by the Caribbean Coral Reef Ecosystems program (Smithsonian Institution) have produced more than 125 publications about marine sponges. These studies have covered many disciplines, including traditional morphological descriptions of new species, but also developmental biology, ecology, symbioses, disease, and evolutionary analyses revealing population affinities throughout the Caribbean using DNA fingerprinting. Various studies have shown that the Belizean corals reefs and mangroves harbor the third richest sponge fauna in the greater Caribbean (after Cuba, and the Florida peninsula). Comparisons between reef and mangrove faunas show that, throughout the Caribbean, they are consistently distinct in their species composition. Many more species will be discovered once the less accessible habitats, such as mesophotic reefs and deeper hard bottoms, are explored. The importance of sponges as a marine resource in Belize is substantial, with respect to services relevant to both their own communities and the human domain. First, they are well known as unique biological pumps and filters, due to great living biomass combined with high water filtration capacity (up to 1 liter per cubic-centimeter sponge per hour), and to complex bacterial assemblages living symbiotically in their bodies (cyanobacteria, nitrifying bacteria, archaebacteria). Secondly, a varied morphologic diversity (shape and color), some with large sizes (up to several meters in diameter), makes them one of the most attractive and intriguing creatures to the curious sport diver visiting Belizean coral reefs. Some sponges are the main dietary component for marine turtles, and a food supplement for many reef fishes (butterfly fishes, angelfishes). Besides their nutritional benefit to sea turtles and fishes, they also provide habitats to hundreds of species of invertebrates and fishes living in cavities inside sponges. In mangrove habitats, too, we have found that sponges are diverse and abundant, particularly on stilt roots of red mangrove lining the tidal channels, and that they probably have developed a long-standing relationship with these plants, offering protection from root borers and possibly exchanging nutrients with them. Besides their attractiveness to underwater tourism, sponges, together with algae and bacteria, are among the marine organisms with highest pharmacological potential for human use, mainly from secondary metabolites produced as defensive chemicals. This well-known capacity makes them a unique resource that must be protected for the future benefit of marine as well as human communities. INTRODUCTION While oceans harbor approximately 80% of animal life on the planet, the Caribbean contains the greatest concentration of species in the Atlantic Ocean and is a global-scale hot spot for marine Biodiversity (Roberts et al., 2002). The Caribbean Sea is a semi enclosed basin of the western Atlantic Ocean, with an  1 Cite as: Diaz, M.C., Rützler, K., 2011. Biodiversity of sponges: Belize and beyond, to the greater Caribbean. In: Palomares, M.L.D., Pauly, D. (eds.), Too Precious to Drill: the Marine Biodiversity of Belize, pp. 57-65. Fisheries Centre Research Reports 19(6). Fisheries Centre, University of British Columbia [ISSN 1198-6727]. Biodiversity of sponges, Diaz and Ruetzler  58 area of about 2,754,000 km2, bathed by currents that enter through the Lesser Antilles and the Windward Passage, and leave northwesterly towards the Gulf of Mexico to form the Gulf Stream. The most prominent marine ecosystems in the Caribbean are sea grass beds (66,000 km2), coral reefs (26,000 km2), and mangroves (11,560 km2) (Miloslavich et al., 2010). Coral reef and mangrove ecosystems are among the most productive and biodiverse tropical marine communities. Coral reefs harbor 4-5% of all known species and are responsible for the highest recorded oceanic productivities (1,500-5,000 gC·m-2·year-1). Mangroves forests line as much as 60-75 % of tropical coasts and may constitute ‗biodiversity hotspots‘ themselves (Rützler et al., 2000), which have been demonstrated to increase reef fish productivity (Mumby et al., 2007). In recent decades, these ecosystems have suffered the consequences of uncontrolled human development (waste water pollution, habitat destruction, clear cutting, among others), and global warming. The area coverage of mangrove has decreased about 1% per year since 1980 (Agard et al., 2007), while live coral coverage has decreased 80% during the last two decades (Gardner et al., 2003; Wilkinson, 2004). Therefore, these ecosystems and the organisms within them are not in their prime conditions and must be studied to understand their role and function and preserved if we intend for the next human generations to continue benefitting from them. Sponges may represent the most diverse benthic faunal component on Coral Reef and mangroves in the Caribbean (Figure 1). Reef sponges may reach four times the diversity of hard and soft corals (Diaz and Rützler, 2001), and mangrove sponges may equal or surpass the richest groups of macroalgae and ascidians, representing from 10 to 70% of the total root epiphytic diversity in various Caribbean sites (Diaz and Rützler, 2009). Marine sponges are essential to the ecology of these systems, mainly owing to their high capacity of water filtration and their role in metabolic processes, including those of their microbial associates (Diaz and Rützler, 2001; Lesser, 2006; de Goeij et al., 2008). In 1972, the Smithsonian Institution‘s Caribbean Coral Reef Ecosystems Program (CCRE) established a field station on Carrie Bow Cay, a tiny sand islet off southern Belize formed by reef-crest debris, to provide year-round support for research by varied experts concerned with investigating biodiversity in the broadest sense, developmental biology, species interaction, oceanographic and carbonate-geological processes, community development over time, starting in the Pleistocene, and distributional, physiological, and chemical ecology. Early on, program participants consisted of staff of the National Museum of Natural History, but eventually, despite financial constraints, collaborators were brought in from other academic institutions worldwide. Numerous studies examined the biological and geological role of Porifera in the reef communities. At last count, 113 researchers focused on sponges of the Carrie Bow area, with 88 (78%) conducting fieldwork and the remainder coauthoring publications. Of the fieldworkers, 63 (72%) studied sponges directly, while the rest (25 or 28%) dealt with sponge associates. To date, 125 scientific papers have been published on the results of this research, while many more are in progress (Rützler, 2011). The present paper reviews our understanding of marine sponges in Belize and beyond to the greater Caribbean. We intend to reflect on the importance of these organisms to the marine communities they inhabit and to the human domain. MATERIAL AND METHODS We carried out a historical review of research in marine sponge biodiversity from Belize and the Caribbean from the early 1800s to the present using a comprehensive taxonomic list that contains classification and authorship information for all sponge species described for the Caribbean (Diaz, van Soest, Rützler and Guerra-Castro, in progress) The list can be found in the World Porifera database: http://www.marinespecies.org/porifera/ (Van Soest et al., 2010), or on the Porifera Tree of Life (PorToL) website (http://www.portol.org/resources). We compiled our own data and published data from other authors and summarized information about the ecological role and pharmacological use of tropical marine sponges, updating our previous review (Diaz and Rützler, 2001). Too Precious to Drill: the Marine Biodiversity of Belize, Palomares and Pauly  59   Figure 1. Sponges are conspicuous components of coral reef and mangrove fauna. (Left) Neofibularia nolitangere (brown mounds) and Callyspongia plicifera (light bluish-gray tube) on a patch reef near Carrie Bow Cay, Belize. (Right). Three common mangrove species, Mycale magniraphidiphera (translucent), M. microsigmatosa (orange), and Haliclona manglaris (light green) covering red mangrove (Rhizophora mangle) rootlet tips in a tidal channel at Twin Cays.  RESULTS AND DISCUSSION Porifera biodiversity in time From the earliest descriptions by P.S. Pallas and J.B. Lamarck (mid-1700s and early 1800s) to the present, approximately 100 authors have contributed taxonomic descriptions of some 800 species of sponges from the greater Caribbean (Figure 2). The earliest comprehensive study of Caribbean sponges, published in 1864 by P. Duchassaing and G. Michelotti, dealt exclusively with collections from the Lesser Antilles, and included approximately 43 species. Subsequent work by J. S. Bowerbank, H. J. Carter, A. Dendy, O. Schmidt, and E. Topsent between 1858 and 1890 covered mainly the Gulf of Mexico and the West Indies, and added more than 150 species. The most prolific authors were the Austrian naturalist Oscar Schmidt, who contributed more than 165 species in 1870-80, and the North American Max Walker de Laubenfels who contributed more than 60 species during 1932-1954 (see Wiedenmayer, 1977 for literature review). The first sponge known from Belize (then British Honduras) was a tiny (5x12 mm) Polymastia biclavata (now genus Coelosphaera), sent to England by a local collector and described by B.W. Priest before the Quekett Microscopical Club of London in 1881. This remained the only record from Belize for the next 56 years, until the British Rosaura Expedition of 1937/38 collected five species from Belize City harbor and Turneffe Island atoll; even those specimens were not described until M. Burton‘s treatise in 1954. When the participants of the CCRE program (Smithsonian Institution, National Museum of Natural History) arrived in Belize in the early 1970s, studies centered on systematics and faunistics, including the quantitative distribution of benthic organisms among the various shallow-water habitats (reachable with scuba diving). Over the next 30 years or so, taxonomy was approached by methods ranging from basic morphology to fine structure, DNA barcoding, and ecological manipulations. One highlight of these years was a workshop for six experts on Caribbean Porifera held at Carrie Bow Cay in 1997. CCRE studies have identified 30 new species, many as part of taxonomic revisions, local or Caribbean-wide, for instance of Biodiversity of sponges, Diaz and Ruetzler  60 the families Clionaidae, Mycalidae, Chalinidae and Axinellidae, and the genera Lissodendoryx (Coelosphaeridae) and Iotrochota (Iotrochotidae). Several species first described from Belizean mangroves were later found distributed on mangroves Caribbean wide. Until now, 189 sponge species have been reported from Belize reef and mangrove habitats (Diaz et al., in progress) This number represents only part of the diversity because many species that we collected remain unclassified and enigmatic and several prime sponge habitats remain unexplored for logistical reasons, such as the deep (below scuba) forereef, mesophotic bottoms, and cryptic environments. Experts estimate than once many regions, depths and habitats get explored, sponge biodiversity might nearly double, from the approximately 10,000 species recognized worldwide so far. The cumulative curve of number of species described per year in the greater Caribbean (Figure 2) shows that the sponge diversity in this region is still underestimated, and that whenever new geographic areas or different habitats are explored, undiscovered species are encountered. Such is the case of the recent description of thirteen new species from sciophilous habitats (cryptic areas of reefs, caverns, or small caves) from Curaçao and Colombia (Van Soest, 2009). Sponges are the most species-rich benthic animal group (165-265) in Cuba, Belize, and Jamaica, a higher diversity than elsewhere in the Caribbean (Miloslavich et al., 2010). Belizean sponges (189 species) represent the third most diverse fauna in the greater Caribbean after Cuba (265 species) and South Florida with (228 species; Diaz et al., in progress.). Comparing the diversities of five important marine animal groups (mollusk, crustaceans, echinoderms, corals and sponges) from 17 countries within various Caribbean marine ecoregions, Miroslavic et al. (2010) found that Belize ranked seventh in species richness. But, when they related species richness to the coastal area of each country, Belize ranked the fourth richest country, with 248 species/100 km, after Cayman islands (388 species/100 km), Costa Rica (362 species/100 km), and Puerto Rico (262 species/100 km). Porifera in the Caribbean and habitat preservation A classical approach to species conservation is to preserve the habitats where they live. This approach becomes even more critical when the species have specific habitat preferences. Scientists, park managers, or government officials might wonder how distinct mangrove and reef faunas are, and which habitat might be more important to protect. We have found that despite geographic contiguity between both habitats, their sponges present biological distinctness, which shows the importance of preserving both ecosystems. Diaz (in press) compared mangrove and coral reef sponge species composition in four distant Caribbean regions (Belize, Cuba, Panama, and Venezuela) and showed that the compositions of these faunas were statistically different. The taxonomic distinctness among faunas was observed at various supraspecific levels (genera, families). For example, major reef players such as species in the family Petrosiidae (genera Xestospongia, Neopetrosia, Petrosia), the family Agelasidae (Agelas spp.) and the order Verongida (Aplysina, Verongula), are basically absent from contiguous sponge-rich mangrove communities. On the other hand, the family Chalinidae (Haliclona, Chalinula) and the family Mycalidae (Mycale spp.) are more species-rich in mangroves than on coral reefs. It is assumed that differences might reflect distinct histories for both faunas. These results place in evidence the need to preserve both ecosystems in order to protect such distinctive faunal components.  Figure 2. Cumulative number of sponge species described in the Caribbean from 1766 to the present. Too Precious to Drill: the Marine Biodiversity of Belize, Palomares and Pauly  61 Ecosystem services Biological pumps Vast volumes of water (up to 1 liter·cm-3 sponge tissue per hour) can be pumped and filtered by marine sponges (Reiswig, 1974). Estimating the biomass of sponge populations in three reef types in Belize, Wilkinson (1989) found the highest values (in wet weight) on inner (lagoon) reefs (1,011-2,458 g·m-2), followed by barrier back reefs (99-1354 g·m-2), and outer reefs (368-702 g·m-2). Assuming an average daily pumping activity of 12 hours (Pile et al., 1997), and a wet volume to weight ratio for sponge tissue of 0.5 (Corredor et al., 1988), we can extrapolate that sponges in Belize reefs may pump 594- 14,748 l water m-2·day-1. This large capacity of water filtration makes sponges not only filter feeders par excellence (Vacelet and Boury Esnault, 1995) but—owing to animal and microbial metabolic processes referred to below—gives these animals a unique role in water transformation with unprecedented ecological consequences. For example, sponges are well known to have high removal rates of particular organic carbon (POC; Reiswig, 1971; Richter et al., 2001; Scheffers et al., 2004) and even higher rates (up to two orders of magnitude) of removal of bulk dissolved organic carbon (DOC; Yahel et al., 2003; de Goeij and van Duyl, 2007). De Goeij et al., (2008) conclude that the three Caribbean thinly encrusting sponges Halisarca caerulea, Merlia normani, and Mycale microsigmatosa, are dissolved organic matter (DOM)- feeders and thus act as sinks of DOC on the reefs they inhabit. The microbial processes of nitrification (aerobic transformation of ammonium to nitrite and nitrate) and denitrification (anaerobic reduction of nitrate to nitrogen gas) have been shown to occur among Caribbean and Mediterranean sponges, and project the highest benthic nitrification rates in tropical waters (Diaz and Ward, 1997; Southwell et al., 2007; Schläppy et al., 2010). Therefore, sponge population size and composition could strongly influence the concentration and speciation of dissolved inorganic nitrogen (DIN) in the reef and mangrove water column, affecting the new production in the ecosystems where they abound. Other metabolic pathways must be evaluated to further predict the role of sponges in these marine systems. Space competitors Various encrusting sponges have been found to overgrow corals and other sessile invertebrates in the Caribbean (Vicente, 1978; Suchanek et al., 1983; Aerts and van Soest, 1997). Chondrilla nucula (now, C. caribensis) has been the principal aggressor at least at three Caribbean sites: Puerto Rico (Vicente, 1978), St. Croix (Suchanek et al., 1983), and Belize (Rützler, et al., 2007). Two recently discovered thinly encrusting reef sponges, Xestospongia bocatorensis and Haliclona walentinae, both containing filamentous cyanobacteria as endosymbionts, were reported to overgrow even some highly aggressive species, fire coral (Millepora sp. and the toxic sponge Neofibularia notilangere; Diaz et al., 2007). These species as well were shown to be phototrophs, acting like plants, sustaining photosynthetic rates much higher than their respiratory rates (Thacker et al., 2007). Studies in the Colombian Caribbean identified the thickly encrusting Desmapsamma anchorata and the ramose species Aplysina cauliformis and Callyspongia armigera as the most frequent overgrowers (Aerts and van Soest, 1997). Morphological plasticity of sponges, their ability to attach to one another without causing harm (Rützler, 1970; Sarà, 1970), diverse chemistry (Faulkner, 2002), and microbial associations (Taylor et al., 2007) are probably among the most important causes for their capacity to overgrow other organisms and avoid being overpowered by them. Calcium carbonate cycle in the reefs Sponges have a dual effect on reef frameworks: firstly, high levels of boring sponge activity may result in net decrease of reef accretion (Rützler, 2002) and, secondly, non-excavating demosponges are found to increase the rates of carbonate accretion by binding coral colonies, in both shallow and deep reef areas, reinforcing the reef frame and decreasing considerably the loss of coral colonies due to dislodgement by wave action, fish predation, and other forces (Wulff and Buss, 1979). Some burrowing clioanid species have become more abundant and have started to be considered as pests for Caribbean coral reefs (Williams and Bunkley-Williams, 2000). Biodiversity of sponges, Diaz and Ruetzler  62 Food and home for others Spongivory is a common life style among several endangered turtle species and among coral reef fishes and seastars (Wulff, 1994; 2005). Hawksbill turtles (Eretmochelys imbricata) are known to feed mainly on a large variety of sponges in the Caribbean, and the green turtle (Chelonia mydas) includes among its varied diet several species of marine sponges. Besides being food for other organisms, sponges may harbor hundreds of animal and algal species during their lives (Villamizar and Laughlin, 1991). Human services Traditionally, some sponge species have been appreciated for their natural softness and resistance to tearing, and their ability to hold and discharge large amounts of water. Since the Roman times, they have been used to hold water or wine, to bath, of for medicinal uses. Nowadays, they are still used for cleaning cars or boats and for cosmetic purposes. For the past 50 years, marine sponges have been considered a potential gold mine, owing to the diversity of their chemicals compounds called secondary metabolites (Sipkema et al., 2005). They produce an enormous array of antitumor, antiviral, anti-inflammatory, immunosuppressive, antibiotic, and other bioactive molecules that can affect the pathogenesis of many human diseases. Sponges, in particular, are responsible for more than 5300 different chemical products, and every year hundreds of new compounds are being discovered (Faulkner, 2002), such as Ara-C, the first marine-derived anticancer agent, and the antiviral drug Ara-A (Proksch et al., 2002). Ara-C is currently used in the routine treatment of patients with leukemia and lymphoma. Ara-A (Acyclovir) is an important antiviral agent. The marine biotech company Porifarma is developing sponge farms in western Turkey to supply sponge metabolites and act as biofilters for neighboring fish farms (de Goeij and Osinga, personal communication). Porifarma will farm two sponge species: Dysidea avara, which produces avarol that has antitumor, antibacterial, and antifungal properties, and Chondrosia reniformis, which is a good source of collagen that can be converted into nano-particles and used to deliver drugs to the target location (Duckworth, 2009). An important question for the future remains how to actually prepare the potential novel drugs on a large scale (Sipkema et al., 2005). Belize‘s sponge biodiversity represents an unexplored ‗treasure trunk‘ for metabolites with high pharmacological potential. Last but not least, sponges, together with fishes, stony corals and soft corals are one of the most attractive members of the coral-reef community, thus having commercial importance for the diving industry. Their variety in shape, size, and intensity of colors, makes them stars in professional as well as amateur photography. One of the most attractive species in Caribbean coral reefs, the giant barrel sponges (Xestospongia muta), is considered the ‗redwood‘ of the reef, for its size and presumed old age. McMurray et al. (2008) estimated that a sponge of 1 m diameter could be 100 years old, certain very large specimens as old as 2,300 years of age. Threats to marine sponges Caribbean sponges are under the same threats that menace their habitats. Among them are habitat destruction, sewage discharge, storm water run-off from polluted land, and global warming which has been increasing water temperatures and altering the ocean food chain and sea floor environment. Various diseases and mass mortalities have already been reported (Williams and Bunkley-Williams, 2000; Olsen et al., 2006; Gochfeld et al., 2007). In particular, although data are still scarce, these ancient animals should be sensitive to oil pollution as their survival depends on large volumes of water processed through their bodies. Zahn et al., (1983) demostrate irreversible DNA damage of polycyclic aromatic hydrocarbon (PAH) through the binding of these oil derived compounds to macromolecular fractions in sponges. The effect of large-scale oil spills, or long-term oil contamination has been recorded both from mangroves and coral reef ecosystems in the Caribbean. The largest oil spill in the Americas occurred in 1986 when more than 8 M liters of crude oil spilled into a complex region of mangroves, seagrasses, and coral reefs just east of the Caribbean entrance to the Panama Canal (Jackson et al., 1989). Extensive mortality of shallow subtidal reef corals, mangrove communities, and infauna of seagrass beds were reported. After 1.5 years, only some organisms in areas exposed to the open sea had recovered. The results of chronic oil pollution from a refinery in Aruba (Netherlands Antilles), including spills and clean-up efforts are, after 60 years, still clearly discernible over a distance of 10 to 15 km along the reef, and includes deteriorated Too Precious to Drill: the Marine Biodiversity of Belize, Palomares and Pauly  63 reef structure, low living-coral cover, and fewer juveniles of reef organisms in front and down current of the refinery (Bak, 1987). CONCLUSIONS Sponges are one of the most diverse marine animals in coral reefs and mangroves of the Caribbean, representing an important structural and functional component of these ecosystems. Belize, with respect to its marine habitat diversity and benthic marine fauna (echinoderms, mollusks, corals and sponges), is one of the richest countries in the Caribbean; therefore, its ecological integrity is important for the entire Caribbean ecosystem. Estimates by experts indicate that probably at least 5000 sponge species worldwide remain to be discovered. Sponges are the source for the highest benthic nitrification rates on the bottom of the oceans, the largest ‗dissolved inorganic carbon sink‘ on Caribbean coral reefs, and the most diverse source of natural products from the ocean. The protection of Belizean coral reefs and mangroves, and the waters that sustain them, is essential to the future existence of these important organisms and the potential of new discoveries and possible exploitation of their biomedical properties. ACKNOWLEDGEMENTS We would like to thank our colleagues Rob van Soest and Edlin Guerra-Castro for allowing the use of our jointly obtained data for this publication. 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Hydroids of Belize, Henry  66 BIODIVERSITY, ECOLOGY AND BIOGEOGRAPHY OF HYDROIDS (CNIDARIA: HYDROZOA) FROM BELIZE1 Lea-Anne Henry Centre for Marine Biodiversity and Biotechnology, Heriot-Watt University, Edinburgh, United Kingdom, EH14 4AS; l.henry@hw.ac.uk ABSTRACT An account of the species richness, assemblage composition, ecology and biogeography of the marine benthic hydroids (Cnidaria: Hydrozoa) across key marine habitats of Belize is provided. Patterns in species richness and composition stress the importance of local hydrography and seabed topography in controlling hydroid biodiversity across coral reef, mangal, seagrass and shallow wharf piling settings. The lack of knowledge regarding the biodiversity of hydroid species from deep-sea (>200 m water depth) settings in Belizean waters is striking, but species richness is expected to be very high given the large number of hydroid species identified from the deep-sea Caribbean in general. Hydroid species richness and composition are also closely governed by finer-scale features such as the diversity of suitable substrata, e.g., coral framework, seagrass thalli and other fauna (mainly sponges and molluscs), each of which differentially selects for species with adaptive morphologies and life history traits. The hydroid fauna of Belize show strong biogeographical affinities with that of the wider Caribbean, West Indies and the south-eastern US, demonstrating a significant influence of large-scale oceanographic features in structuring Belizean marine biodiversity. INTRODUCTION ‗Hydroids‘ are a group of benthic cnidarian invertebrates belonging to the Class Hydrozoa. Unlike other cnidarians, many hydrozoans alternate between sessile polyp and (sometimes almost entirely suppressed) medusa phases as part of their life histories including the Orders Limnomedusae, Anthoathecata, Leptothecata, but not the hydrozoan Orders Siphonophorae, Trachymedusae or Narcomedusae. The polyp phase is generally referred to as the hydroid phase, which may comprise a single individual polyp or a physiologically integrated colony of multiple and sometimes specialized polyps. While many hydroids are gelatinous, several anthoathecate families such as the Stylasteridae, Milleporidae, Hydractiniidae and Rosalindidae) possess calcified skeletons or basal encrusting mats. Hydrozoans comprise mostly marine species, but some inhabit brackish and freshwater habitats. Over 3500 species have been identified, but molecular characterization of cryptic species is likely to reveal many more. They are typically hard substrate generalists, but species exhibit a range of life history strategies that differentially assembles species across space. Bacteria, algae, plants and other animals use hydroids themselves as hard substrata, and thus the occurrence and biodiversity of hydroids further enhances marine biodiversity. Hydroids are preyed upon by a variety of animals including sea turtles, fish, echinoderms, sea spiders, crustaceans and sea slugs. Hydroids themselves are generally suspension feeders, and actively feed in moderately strong water currents by capturing food with their polyp tentacles. Hydroids also paralyze living prey such as zooplankton by discharging stinging nematocyst cells from feeding tentacles. While hydroids are typically out competed for space by sponges, ascidians and octocorals, hydroid nematocysts are sometimes clustered into larger nematophores over the whole colony itself and offer an effective defense and offense strategy. However, the full tentacular and colony nematocyst complement is also often unfortunately and rather painfully discharged onto the exposed skin of divers, snorkelers and aquarists.  1 Cite as: Henry, L.-A., 2011. Biodiversity, ecology and biogeography of hydroids (Cnidaria: Hydrozoa) from Belize. In: Palomares, M.L.D., Pauly, D. (eds.), Too Precious to Drill: the Marine Biodiversity of Belize, pp. 66-77. Fisheries Centre Research Reports 19(6). Fisheries Centre, University of British Columbia [ISSN 1198-6727]. Too Precious to Drill: the Marine Biodiversity of Belize, Palomares and Pauly  67 The wider Caribbean region has the highest number of hydroid species in the Atlantic Ocean (Medel and López-González, 1998). Although the marine environments of Belize are some of the most species-rich in the Caribbean (Miloslavich et al., 2010), study of the hydroid fauna has not been as intensive as it has been for that carried out for example on sponges or scleractinian corals. Regardless, targeted taxonomic investigation of hydroids in Belize consistently reveals new species or extends biogeographic ranges (e.g. Spracklin, 1982; Calder, 1988; Puce et al., 2005). Existing information has been driven almost entirely by activities carried out directly by or in collaboration with the Smithsonian Institution. Data collection has been guided mostly by the long-running Caribbean Coral Reef Ecosystems Program from its field station site at Carrie Bow Cay and in mangrove habitats at Twin Cays and Wee Wee Caye. More recently, the Marine Invasions Research Lab at the Smithsonian Environmental Research Center has been monitoring Belize for non-native and exotic species introductions. Others have recorded hydroids incidentally as part of marine surveys conducted by the British Coral Cay Conservation at Turneffe Atoll, independently at Bacalar Chico (Ambergris Caye) and in lagoonal habitats at Snake Cays (southern Barrier Reef). Because about 50% of hydroids also produce a well-developed pelagic medusa as part of their life cycle, research into Belizean medusae must also be considered, however, this knowledge seems to be restricted to the medusa fauna from Carrie Bow Cay. MATERIALS AND METHODS Herein, this research and the author‘s unpublished data are reviewed in their entirety to produce a contemporary species checklist of the hydroid fauna from Belize, inclusive of all Hydrozoa. These included Cairns (1982), Larson (1982), Spracklin (1982), Calder (1988, 1991), Ellison and Farnsworth (1992), Kaehler and Hughes (1992), Fenner (1999) and Puce et al. (2005). Taxonomic revision required many published lists to be updated; for consistency, synonymies and taxonomy followed that of the World Registry of Marine Species (Appeltans et al., 2011). Substrate and habitat affinities of Belizean hydroids are also reviewed to understand the role of environmental forcing across multiple spatial scales, including the importance of ocean circulation in creating the biogeographic affinities of hydroids from Belize in both shallow water and deep-sea (>200m) contexts. RESULTS AND DISCUSSIONS A total of 117 hydrozoan species were identified from Belizean waters (Appendix 1), 103 of which were hydroids (i.e., those species producing a polyp phase as part of their life cycle). Of these 103 species, 30 were species records based solely from their medusa phase; thus the polyp phases of 73 hydroid species were listed. Excluding these 30 records of medusa phases to remove any taxonomic overlap, the unidentified corymorphid species from Spracklin (1982) as well as the Amphinema sp., Leuckartiara sp., and Aequorea sp. from Calder (1991a) can now be included. Thus, the polyp phases of 76 species of hydroid species have now been recorded from Belize (Table 1). Notably, none of these 76 species have been recorded in depths deeper than about 67 m (Spracklin, 1982). However, a large hydrocoral has been observed at nearly 300m depth off Glover‘s Reef (in Lutz and Ginsberg, 2007). The leptothecate Kirchenpaueria halecioides (Figure 1) is the most commonly encountered hydroid across all habitats in Belize.   Figure 1. The most commonly encountered hydroid in Belize, Kirchenpaueria halecioides. Inset picture details the fixed sporosac gonophore of K. halecioides that contains its dispersive larval phase.  Hydroids of Belize, Henry  68 Table 1. Valid hydroid species of Belize and their common wider global distributions. Spracklin‘s (1982) unidentified species and other genera from Calder (1991a) are now included. Species Common distribution Order Anthoathecata (29 species) Suborder Capitata Family Cladonematidae Eleutheria dichotoma De Quatrefages, 1842  North Atlantic, Mediterranean Family Corymorphidae unidentified corymorphid sp. (in Spracklin, 1982) Family Corynidae Coryne sargassicola Calder, 1988  western North Atlantic Family Halocordylidae Pennaria disticha (Goldfuss, 1820)  North Atlantic, Gulf of Mexico, Mediterranean, New Zealand, Red Sea, South Africa Family Sphaerocorynidae Sphaerocoryne bedoti Pictet, 1893  North Atlantic, Mediterranean, Red Sea Family Milleporidae Millepora alcicornis Linnaeus, 1758  Gulf of Mexico, Caribbean, Mozambique Millepora complanata Lamarck, 1816  Gulf of Mexico, Caribbean Millepora squarrosa Lamarck, 1816  Gulf of Mexico, Caribbean Family Tubulariidae Ectopleura grandis Fraser, 1944  Gulf of Mexico Zyzzyzus warreni Calder, 1988  tropical circumglobal Family Zancleidae Zanclea alba (Meyen, 1834)  North Atlantic, Gulf of Mexico Zanclea costata Gegenbaur, 1857  North Atlantic, Barents Sea, Mediterranean, Gulf of Mexico Suborder Filifera Family Bougainvilliidae Bimeria vestita Wright, 1859  North Atlantic, Caribbean, Black Sea, South Africa, Indian and Pacific Oceans Millardiana longitentaculata Wedler and Larson, 1986  tropical western Atlantic Pachycordyle napolitana Weismann, 1883  North and central Atlantic, Mediterranean Family Eudendriidae Eudendrium attenuatum Allman, 1877  Gulf of Mexico, sub-tropical northwest Atlantic Eudendrium bermudense Calder, 1988  Bermuda Eudendrium eximium Allman, 1877  Gulf of Mexico, sub-tropical north-western Atlantic Eudendrium klausi Puce, Cerrano, Marques and Bavestrello, 2005  Belize Myrionema amboinense Pictet, 1893  possibly North Atlantic, mostly west Pacific, Indo-Pacific, Indian Ocean Myrionema hargitti (Congdon, 1906)  Gulf of Mexico, Caribbean Family Hydractiniidae Hydractinia arge (Clarke, 1882)  northwest Atlantic Family Oceaniidae Corydendrium parasiticum (Linnaeus, 1767)  North Atlantic, Gulf of Mexico, Caribbean, Mediterranean Turritopsis fascicularis Fraser, 1943  Gulf of Mexico Turritopsis nutricula McCrady, 1857;  North Atlantic, Gulf of Mexico, Caribbean Turritopsoides brehmeri Calder, 1988  Belize Family Pandeidae Amphinema sp. (in Calder, 1991) Leukartiara sp. (in Calder, 1991) Family Stylasteridae Stylaster roseus (Pallas, 1766)  Caribbean, Gulf of Mexico, northeast Brazil Order Leptothecata (47) species Family Aequoreidae Aequorea sp. (in Calder, 1991) Family Campanulariidae Clytia hemisphaerica (Linnaeus, 1767)  circumglobal Clytia latitheca Millard and Bouillon, 1973  South Africa, Red Sea Clytia linearis (Thorneley, 1900)  Circumglobal, but not high Arctic or Southern Ocean Clytia macrotheca (Perkins, 1908)  tropical western Atlantic, Gulf of Mexico Clytia noliformis (McCrady, 1859)  Atlantic, Gulf of Mexico, Mediterranean Clytia paulensis (Vanhöffen, 1910)  North Atlantic, Gulf of Mexico, Mediterranean Clytia tottoni (Leloup, 1935)  eastern South Pacific Obelia bidentata Clark, 1875  circumglobal Obelia dichotoma (Linnaeus, 1758)  Atlantic and Pacific Oceans, Gulf of Mexico, Mediterranean, Red Sea, New Zealand Orthopyxis sargassicola (Nutting, 1915)  western Atlantic, Brazil Family Campanulinidae Egmundella grandis Fraser, 1941  Chesapeake Bay Lafoeina tenuis Sars, 1874  North Atlantic, Gulf of Mexico, Arctic Ocean, Mediterranean, west Indian Ocean  Too Precious to Drill: the Marine Biodiversity of Belize, Palomares and Pauly  69  Table 1. Continued Species Common distribution Family Eirenidae Eutima sp. (in Calder, 1991) Family Haleciidae Halecium bermudense Congdon, 1907  Gulf of Mexico, western Atlantic Halecium nanum Alder, 1859  Atlantic and Pacific Oceans Halecium speciosum Nutting, 1901  eastern Pacific Halecium tenellum Hincks, 1861  circumglobal Hydrodendron mirabile (Hincks, 1866)  Atlantic Ocean, West Indies, Mediterranean, southwest Indian Ocean, New Zealand Nemalecium lighti (Hargitt, 1924)  western Atlantic, Indo-Pacific Family Hebellidae Hebella scandens (Bale, 1888)  North Atlantic, Gulf of Mexico, Mediterranean, South Africa Hebella venusta (Allman, 1877)  Gulf of Mexico, Caribbean, Red Sea Scandia mutabilis (Ritchie, 1907)  Gulf of Mexico Family Phialellidae Phialella sp. (in Calder, 1991) Family Sertulariidae Diphasia tropica Nutting, 1904  Caribbean, tropical Atlantic Dynamena crisioides Lamouroux, 1824  North Atlantic, Gulf of Mexico, South Africa, Mozambique, Red Sea Dynamena disticha Bosc, 1802  North Atlantic, Gulf of Mexico, Mediterranean, Red Sea Sertularella diaphana (Allman, 1885)  warm circumglobal Sertularella peculiaris Leloup, 1974  South Atlantic, Lesser Antilles Sertularia distans (Lamouroux, 1816) North Atlantic, Mediterranean Sertularia marginata (Kirchenpauer, 1864)  North Atlantic, Mediterranean, New Zealand Sertularia tumida (Allman, 1877)  Gulf of Mexico Sertularia turbinata (Lamouroux, 1816)  North Atlantic, Mediterranean, Caribbean Symmetroscyphus intermedius (Congdon, 1907)  western North Atlantic, Caribbean Thyroscyphus marginatus (Allman, 1877)  North Atlantic, Gulf of Mexico, Australia Superfamily Plumularioidea Family Agalopheniidae Aglaophenia latecarinata Allman, 1877  North Atlantic, Gulf of Mexico, Red Sea Aglaophenia pluma (Linnaeus, 1758) North Atlantic, Mediterranean, South Africa Family Halopterididae Antennella quadriaurita Ritchie, 1909  Atlantic, Gulf of Mexico, Indo-Pacific, New Zealand Antennella secundaria (Gmelin, 1791)  North Atlantic, Gulf of Mexico, Mediterranean, Red Sea, New Zealand Halopteris alternata (Nutting, 1900)  Atlantic, Caribbean, Gulf of Mexico Halopteris carinata Allman, 1877  Atlantic, Caribbean, Gulf of Mexico Halopteris diaphana (Heller, 1868)  North Atlantic, Mediterranean Monostaechas quadridens (McCrady, 1859)  North Atlantic, Gulf of Mexico Family Kirchenpaueriidae Kirchenpaueria halecioides (Alder, 1859)  North Atlantic, Caribbean, Mediterranean, Gulf of Mexico, Red Sea Family Plumulariidae Plumularia margaretta (Nutting, 1900)  North Atlantic, Gulf of Mexico, Mediterranean Plumularia setacea (Linnaeus, 1758)  North Atlantic, Gulf of Mexico, Mediterranean, Red Sea, South Africa Plumularia strictocarpa Pictet, 1893  Gulf of Mexico, South Africa  Two species are endemic to Belize, Turritopsoides brehmeri (from mangal habitat, Twin Cays) and Eudendrium klausi (from coral reef habitat, Carrie Bow Cay). However, it is likely that with targeted taxonomic studies, there is no a priori reason to expect that these species are found exclusively in Belize. This is particularly true of T. brehmeri, which colonises both Thalassia seagrass and sponge substrata and is therefore likely to colonize other substrata outside mangroves (Calder, 1988). Currently, the International Union for Conservation of Nature (IUCN) Red List of Threatened Species includes three hydroids found in Belize waters, all hydrocorals: Millepora alcircornis, M. complanata and M. squarrosa (since M. striata has been synonymized with M. squarrosa, the former species is provisionally not included here, but as of 2011 the IUCN still includes M. striata on its Red List). Only M. squarrosa is listed as decreasing in occurrence, but according to the IUCN, all three species remain categorized as being of ‗least concern‘ in terms of becoming endangered or extinct. Hydroids of Belize, Henry  70 Species diversity, composition and abundance of hydroids in Belize are all closely dependent on the occurrence, nature and variety of substrata. The richest species assemblages are found on dead or exposed tissue of scleractinians and gorgonian coral substrata including Muriceopsis flavida, Pseudopterogorgia acerosa, P. bipinnata, Gorgonia flabellum and G. ventalina (Spracklin, 1982; Puce et al., 2005). The seagrass Thalassia testudinum and algal substrata provided by Halimeda and Sargassum are also important in providing substrata for hydroids in Belize (Spracklin, 1982; Calder, 1988; Kaehler and Hughes, 1992), as are the sponges Monanchora arbuscula, Cliona caribbaea, Tedania (Tedania) ignis, (Spracklin, 1982; Calder, 1991a). Although the nature of the association is unclear, the hydrocorals Millepora alcicornis and M. squarrosa have been observed to encrust living corals as well (Fenner, 1999). Within mangrove habitats formed by Rhizopora, the most diverse assemblages are found on mangal prop roots (Calder, 1991b), particularly in areas with moderate to strong wave action or water currents (Calder, 1991a); this environmental setting helps prevent sedimentation and smothering of hydroids while delivering an adequate food supply. While largely substrate generalists, certain species are better adapted to living on for example, ephemeral substrata such as Thalassia seagrass in Belize. Their morphologies, growth patterns and distribution on these substrata are selected for maximizing ‗residence time‘ on seagrass leaves (Kaehler and Hughes, 1992), helping to ensure the hydroid survives to reproduce. Others, such as the hydrocorals Millepora alcicornis and M. squarrosa, can survive in spatially competitive environments like coral reef habitats by being able to rapidly encrust other corals. High risk of desiccation, competition, smothering and predation restrict the leptothecate hydroid Dynamena crisioides to a narrow depth range in Belize from the lower intertidal to the very shallow subtidal zone, where it exhibits significant intra- and interpopulation variation in morphology and reproduction to cope with environmental conditions (Calder, 1991c). The biodiversity of the hydroid fauna also varies across wider habitat types. Based on the data to date, species richness and composition of the hydroid fauna varies between habitat categories (seagrass, man-made fouling plates, mangrove, patch reefs, back reefs, reef crest, spur and groove habitats, sand troughs, outer ridge, fore-reef slopes and floating Sargassum) (Figure 1). This review identified the highest number of hydroid species from mangroves (53 species), followed by patch reefs (21 species). Increasing proximity of mangroves to the barrier reef itself also enhances species richness of Rhizopora epibionts including hydroids (Ellison and Farnsworth, 1992), which may help to explain the high species richness of hydroid species seen at Twin Cays (Calder, 1991a). The relationship between mangal epifauna is also mutualistic: root epibionts help prevent boring by the root-boring isopod Phycolimnoria that reduces plant growth (Ellinson and Farnsworth, 1992). Given that so few studies have targeted the hydroid fauna in general, it remains to be seen whether mangroves truly support the richest assemblages. Notwithstanding lack of knowledge, it is clear that the habitat heterogeneity conferred by lagoonal habitats such as mangroves and patch reefs is of particular importance in sustaining high levels of hydroid biodiversity in Belize. Thus, the conservation of such habitats is vital in helping to sustain the marine biodiversity of Belize. Natural groupings of all habitat types occur according to whether they are artificial, lagoonal, reef or oceanic in nature (Figure 2), with lagoonal habitats exhibiting more consistent, i.e., more homogenous species composition than those from reef habitats where species composition varies more widely between  Figure 2. Non-metric multidimensional scaling plot of species Bray-Curtis dissimilarity indices based on presence/absence data (using PRIMER v6 software), comparing hydroid assemblages across wider habitat types (lagoonal, those collected on dock settlement plates, reef habitats and oceanic Sargassum). The overall low stress (0.13) indicates that hydroid assemblages are shaped differently across wider habitat niches. back reef fouling plate reef crest seagrass Sargassum mangrovepatch reef fore-reef slope outer ridge low-relief spur & groove sand trough relief spur & groove 20 stress: 0.13 lagoon dock oceanic reef Too Precious to Drill: the Marine Biodiversity of Belize, Palomares and Pauly  71 habitats (Figure 2). Thus, despite having the highest numbers of species or ‗alpha‘ diversity, lagoonal habitats had lower species turnover, or ‗beta‘ diversity. Hydroid biodiversity in Belize, therefore, depends both on forces driving high species richness in lagoonal habitats, and those that control high species turnover between coral reef habitats: thus, the total or ‗gamma‘ diversity of hydroids in Belize will require the conservation of both habitat categories. The general affinity of the hydroid fauna from Belize is West Atlantic Tropical (Briggs, 1