Coral Reef Resilience Assessment of the National Marine ,

Surveys from 31 to 7 June, 2009

IUCN Climate Change and Coral Reefs Working Group About IUCN

,8&1,QWHUQDWLRQDO8QLRQIRU&RQVHUYDWLRQRI1DWXUHKHOSVWKHZRUOG¿QGSUDJPDWLFVROXWLRQVWRRXUPRVWSUHVVLQJ environment and development challenges.

IUCN works on biodiversity, climate change, energy, human livelihoods and greening the world economy by supporting VFLHQWL¿FUHVHDUFKPDQDJLQJ¿HOGSURMHFWVDOORYHUWKHZRUOGDQGEULQJLQJJRYHUQPHQWV1*2VWKH81DQGFRPSD- nies together to develop policy, laws and best practice.

,8&1LVWKHZRUOG¶VROGHVWDQGODUJHVWJOREDOHQYLURQPHQWDORUJDQL]DWLRQZLWKPRUHWKDQJRYHUQPHQWDQG1*2 members and almost 11,000 volunteer experts in some 160 . IUCN’s work is supported by over 1,000 staff in RI¿FHVDQGKXQGUHGVRISDUWQHUVLQSXEOLF1*2DQGSULYDWHVHFWRUVDURXQGWKHZRUOG www.iucn.org

IUCN Global Marine and Polar Programme

The IUCN Global Marine and Polar Programme (GMPP) provides vital linkages for the Union and its members to all the ,8&1DFWLYLWLHVWKDWGHDOZLWKPDULQHDQGSRODULVVXHVLQFOXGLQJSURMHFWVDQGLQLWLDWLYHVRIWKH5HJLRQDORI¿FHVDQGWKH ,8&1&RPPLVVLRQV*033ZRUNVRQLVVXHVVXFKDVLQWHJUDWHGFRDVWDODQGPDULQHPDQDJHPHQW¿VKHULHVPDULQH protected areas, large marine ecosystems, coral reefs, marine invasives and the protection of high and deep seas.

The Nature Conservancy

The mission of The Nature Conservancy is to preserve the plants, animals and natural communities that represent the diversity of life on Earth by protecting the lands and waters they need to survive. The Conservancy launched the Global Marine Initiative in 2002 to protect and restore the most resilient examples of ocean and coastal ecosystems in ways WKDWEHQH¿WPDULQHOLIHORFDOFRPPXQLWLHVDQGHFRQRPLHV7KH&RQVHUYDQF\RSHUDWHVRYHUPDULQHFRQVHUYDWLRQ SURMHFWVLQPRUHWKDQFRXQWULHVDQG86VWDWHVZHZRUNZLWKSDUWQHUVDFURVVVHDVFDSHVDQGODQGVFDSHVWKURXJK transformative strategies and integrated planning and action. The focus is on: (1) Setting priorities for marine conservation XVLQJHFRUHJLRQDODVVHVVPHQWVDQGWRROVIRUHFRV\VWHPEDVHGPDQDJHPHQW  (QVXULQJFRUDOUHHIVXUYLYDOE\FUHDWLQJ UHVLOLHQWQHWZRUNVRIPDULQHSURWHFWHGDUHDV  5HVWRULQJDQGFRQVHUYLQJFRDVWDOKDELWDWVE\XWLOL]LQJLQQRYDWLYHQHZ PHWKRGV  %XLOGLQJVXSSRUWIRUPDULQHFRQVHUYDWLRQWKURXJKVWUDWHJLFSDUWQHUVKLSVDQGZRUNLQJWRVKDSHJOREDODQG national policies. Marine conservation in The Nature Conservancy builds upon the organization’s core strengths: achieving GHPRQVWUDEOHUHVXOWVZRUNLQJZLWKDZLGHUDQJHRISDUWQHUVLQFOXGLQJQRQWUDGLWLRQDOSDUWQHUVVFLHQFHEDVHGUREXVW FRQVHUYDWLRQSODQQLQJPHWKRGRORJLHVRXUH[SHULHQFHZLWKWUDQVDFWLRQVDQGSHUKDSVPRVWLPSRUWDQWO\RXUDELOLW\DQG FRPPLWPHQWWREDFNXSRXUVWUDWHJLHVZLWKKXPDQ¿QDQFLDODQGSROLWLFDOFDSLWDO)RUPRUHLQIRUPDWLRQHPDLOPDULQH# tnc.org or go to www.nature.org/marine.

Acknowledgements

This survey was conducted as part of the IUCN Climate Change and Coral Reefs Working Group global UHVLOLHQFHDVVHVVPHQWVDQGZDVPDGHSRVVLEOHE\WKHJHQHURXVVXSSRUWRIWKH1DWLRQDO)LVKDQG:LOGOLIH)RXQGDWLRQWKH Stichting Nationale Parken Bonaire and the Bonaire National manager, Ramon de Leon. Partners included The Nature Conservancy (TNC), Research and Management of Biodiversity (CARMABI), the University of Maine and Yale University.

Cover Photography

)URQWFRYHUµ%RQDLUH1DWLRQDO0DULQH3DUNPDQDJHU5DPRQGH/HRQVXUYH\LQJFRUDOUHHIV¶&UHGLW*DEULHO*ULPVGLWFK IUCN

%DFNFRYHUµ7KHVDOWSLHULQ%RQDLUH1HWKHUODQGV$QWLOOHVDWVXQVHW¶&UHGLW*HULFN%HUJVPD0DULQH3KRWREDQN Coral Reef Resilience Assessment of the Bonaire National Marine Park,

Surveys from 31 May to 7 June, 2009 The designation of geographical entities in this book, and the presentation of the material, do not imply the expression of any opinion whatsoever on the part of IUCN or The Nature Conservancy concerning the legal status of any , territory, or area, or of its authorities, or concerning the delimitation of its frontiers or boundaries. The views expressed LQWKLVSXEOLFDWLRQGRQRWQHFHVVDULO\UHÀHFWWKRVHRI,8&1RU7KH1DWXUH&RQVHUYDQF\QRUGRHVFLWLQJRIWUDGHQDPHV or commercial processes constitute endorsement.

Published by: IUCN, Gland, Switzerland Copyright: © 2011 International Union for Conservation of Nature and Natural Resources and The Nature Conservancy

Reproduction of this publication for educational or other non-commercial purposes is authorized without prior written permission from the copyright holders provided the source is fully acknowledged.

Reproduction of this publication for resale or other commercial purposes is prohibited without prior written permission of the copyright holders.

Citation: IUCN (2011). Coral Reef Resilience Assessment of the Bonaire National Marine Park, Netherlands Antilles. Gland, Switzerland: IUCN. 51pp.

ISBN: 978-2-8317-1352-6

Available from:

IUCN Global Marine and Polar Programme, IUCN, Rue Mauverney 28, 1196 Gland, Switzerland. 7HO)D[

E-mail: PDULQH#LXFQRUJ

This publication is available as a download from the IUCN Global Marine and Polar Programme website at the following address: http://www.iucn.org/dbtw-wpd/edocs/2011-008.pdf

Coral Reef Resilience Assessment of the Bonaire National Marine Park, Netherlands Antilles

Surveys from 31 May to 7 June, 2009

Gabriel Grimsditch, International Union for the Conservation of Nature Suzanne Arnold, The University of Maine Henry de Bey, Yale University Jeanne Brown, The Nature Conservancy Sabine Engel, Stichting Nationale Parken Bonaire Ramon de Leon, Stichting Nationale Parken Bonaire Mark Vermeij, Caribbean Research and Management of Biodiversity

About the Working Group

About the IUCN Climate Change and Coral Reefs Marine Working Group

The IUCN Climate Change and Coral Reefs Marine Working Group (formerly the IUCN Resilience Science Work- ing Group), focused on coral bleaching, resilience, and climate change, was established in 2006 by the Global 0DULQHDQG3RODU3URJUDPPHRI,8&1RQD\HDUJUDQWIURPWKH-RKQ'DQG&DWKHULQH70DF$UWKXU)RXQGD- tion. The goal of the working group is to draw on leading practitioners in coral reef science and management to VWUHDPOLQHWKHLGHQWL¿FDWLRQDQGWHVWLQJRIPDQDJHPHQWLQWHUYHQWLRQVWRPLWLJDWHWKHLPSDFWVRIFOLPDWHFKDQJHRQ coral reefs. The working group consults and engages with experts in three key areas: climate change and coral EOHDFKLQJUHVHDUFKWRLQFRUSRUDWHWKHODWHVWNQRZOHGJHPDQDJHPHQWWRLGHQWLI\NH\QHHGVDQGFDSDELOLWLHVRQWKH JURXQGDQGHFRORJLFDOUHVLOLHQFHWRSURPRWHDQGGHYHORSWKHIUDPHZRUNSURYLGHGE\UHVLOLHQFHWKHRU\DVDEULGJH between bleaching research and management implementation. 2QHRIWKHRXWSXWVRIWKLVJURXSZDVWKHVHWWLQJXSRIDZHEVLWHWKDWSURYLGHVOLQNVWRSURMHFWVHYHQWVSDUWQHUV and publications.

)RUPRUHLQIRUPDWLRQVHHhttp://www.iucn.org/cccr/index.cfm

This publication is part of a series of publications on management tools to promote resilience in marine ecosystems. Selected titles, also available from IUCN’s Global Marine and Polar Programme, are listed below:

Coral Reef Resilience and Resilience Assessment of Coral Reefs: Resistance to Bleaching Rapid assessment protocol for coral reefs, focusing on coral bleaching and thermal IUCN Resilience Science Group Working Paper Series – No 1 stress Gabriel D. Grimsditch and Rodney V. Salm IUCN Resilience Science Group Working Paper Series – No 5 © IUCN/TNC, October 2006 David Obura and Gabriel Grimsditch © IUCN / TNC, April 2009

Managing Mangroves for Coral Reefs, Climate Change and Resilience

Coral Reefs, Climate Change Resilience to Climate Change and Resilience An Agenda for Action from the IUCN World An Agenda for Action from the IUCN World Conservation Congress in Barcelona, David Obura & Gabriel Grimsditch Conservation Congress in Barcelona, Spain IUCN Resilience Science Group Working Paper Series – No 2 IUCN Resilience Science Group Working Paper Series – No 6 Elizabeth Mcleod and Rodney V. Salm David Obura & Gabriel Grimsditch © IUCN/TNC, October 2006 © 2009 IUCN/TNC

IUCN RESILIENCE SCIENCE WORKING GROUP PAPER SERIES ! NO 6

Managing Seagrasses for A Framework for Social Adaptation to

$)UDPHZRUNIRU6RFLDO$GDSWDWLRQ Resilience to Climate Change to Climate Change Climate Change - Sustaining Tropical Sustaining Tropical Coastal Communitites and Industries

1$0DUVKDOO3$0DUVKDOO-7DPHODQGHU'2EXUD'0DOOHUHW.LQJDQG-(&LQQHU Coastal Communities & Industries IUCN Resilience Science Group Working Paper June 2010 Series – No 3 NA Marshall, PA Marshall, J Tamelander, D Obura, D Malleret- Mats Björk, Fred Short, Elizabeth Mcleod and Sven King and JE Cinner Beer © IUCN December 2009 © IUCN/TNC, 2008

IUCN Climate Change and Coral Reefs Working Group

6 Table of Contents

Table of Contents

Executive Summary ...... 9

1. Introduction ...... 11

1.1 The Study ...... 11 1.2 Reef resilience ...... 12 1.3 Bonaire ...... 12 3XUSRVHRIWKHVWXG\ ......  2YHUYLHZRIPHWKRGV ...... 15

 0DMRU¿QGLQJV ...... 17

2YHUYLHZ ...... 17 2.2 Threats to Bonaire’s coral reef communities ...... 22 2.3 Summary ...... 26

3. Detailed Methodology ...... 27

3.1 Survey methods ...... 27 3.2 Analysis ...... 30

4. Detailed Results ...... 32

5HVLOHQFHLQGLFDWRUV ...... 32 %HQWKLFFRYHU ...... 37 &RUDOSRSXODWLRQVWUXFWXUH ......  $OJDH3RSXODWLRQV ......  )LVKFRPPXQLW\VWUXFWXUH ...... 

References ...... 51

7

Executive Summary

Executive Summary

Although Bonaire’s coral reefs remain among the healthiest and most resilient in the Caribbean, this IUCN report based on the IUCN Resilience Assessment of Coral Reefs highlights some of the threats that exist to Bonaire’s coral reefs, and which could have serious implications for resilience to future FOLPDWHFKDQJHDQGRWKHUWKUHDWV7KHUHSRUWLGHQWL¿HGUHFRPPHQGDWLRQVIRUDGGUHVVLQJWKHFXUUHQW threats, as well as high and low resilience sites.

7KHWKUHDWVDQGUHFRPPHQGDWLRQVLGHQWL¿HGLQFOXGH

&RDVWDOGHYHORSPHQWDQGDUWL¿FLDOEHDFKHV.Recommendation: All coastal construction on Bonaire should be strictly regulated and follow the construction guidelines. The guidelines should become law in order to be enforced appropriately.

Leaching from septic tanks. Recommendation: It is strongly recommended that Bonaire invest in appropriate sewage treatment facilities to improve water quality and increase the resilience of its valuable coral reefs. It is also recommended that a water quality monitoring program be set up and sustained.

,QFUHDVLQJGDPVHO¿VKSRSXODWLRQV5HFRPPHQGDWLRQ,WLVUHFRPPHQGHGWKDWWKH¿VKLQJRISUHGD- WRU\¿VKVSHFLHVRQ%RQDLUH¶VFRUDOUHHIVEHFRQWUROOHGDQGPDQDJHGWRDVXVWDLQDEOHOHYHOWRSUHYHQW SRSXODWLRQH[SORVLRQVRISUH\¿VKFDSDEOHRIPRGLI\LQJWKHUHHIKDELWDW

Trididemnum and Lobophora. Recommendation: It is recommended that the populations of Tri- didemnum and Lobophora are closely monitored and the factors contributing to the unnatural abun- dance of these coral-overgrowing organisms should be studied and then eliminated.

Due to a variety of factors affecting resilience which were assessed using the IUCN methodology, sites were also ranked according to their overall resilience:

Resilience rating Sites Management High North NDA High 3OD\D)UDQV NDA High .DUSDWD MPA High Margate Bay MPA High Vista Blue MPA High South Bay MPA Medium 3OD\D)XQFKL MPA Medium Wayaka II MPA Medium 2LO6OLFN/HDS MPA Medium Cliff )3$

9 Executive Summary

Resilience rating Sites Management Medium Bari )3$ Medium Something Special )3$ Medium 18th Palm )3$ Medium Bachelor’s Beach MPA Medium Angel City MPA Medium City MPA Medium Tori’s Reef MPA Medium Carl’s Hill MPA Medium Mi Dushi MPA Medium .HHSVDNH MPA Low Chachacha )3$

It is noteworthy that sites with lowest resilience ratings (e.g. Chachacha) are those most impacted by FRDVWDOGHYHORSPHQWZKLOHVLWHVZLWKKLJKHVWUHVLOLHQFHUDWLQJV HJ0DULQH5HVHUYH1RUWK3OD\D)UDQV .DUSDWD0DUJDWH%D\9LVWD%OXHDQG6RXWK%D\ DUHWKRVHIXUWKHVWDZD\DQGOHDVWLPSDFWHGE\FRDVWDO development.

10 Introduction

1. Introduction

2 To survey the resistance and resilience of 1.1 The Study coral reefs in Bonaire to coral bleaching and climate change; This survey was conducted as part of the IUCN Climate Change and Coral Reefs Working Group 3 To train regional global coral reef resilience assessments, and managers and other partners in implemen- was made possible by the generous support tation of the aforementioned surveys; and RI WKH 1DWLRQDO )LVK DQG :LOGOLIH )RXQGDWLRQ the Stichting Nationale Parken Bonaire and the 4 To make recommendations on manage- Bonaire National Marine Park manager, Ramon de ment of coral reefs within the Marine Pro- Leon. Partners included The Nature Conservancy WHFWHG$UHDEDVHGRQWKHVXUYH\¿QGLQJV (TNC), Caribbean Research and Management of Biodiversity (CARMABI), the University of Maine and Yale University. This document aims to provide 7KLVUHSRUWFRQWDLQVLQIRUPDWLRQRQREMHFWLYHV information on how to incorporate resilience DQGZKLOHREMHFWLYHZDVPHWWKURXJKDWUDLQ- information and climate change responses into LQJ ZRUNVKRS RUJDQLVHG RQ %RQDLUH LQ FRQMXQF- Marine Protected Area design and management. tion with The Nature Conservancy and the Na- 6SHFL¿FDOO\WKHVWXG\REMHFWLYHVDUH WLRQDO 2FHDQLF DQG$WPRVSKHULF$GPLQLVWUDWLRQ Members of the survey team incorporated staff 1 To design and implement a rapid assess- from the following partner organisations: ment protocol to monitor and quantify bleaching and reef resilience tailored to the needs the Bonaire Marine Park;

Table 1

Name Organisation Data collected Suzanne Arnold The University of Maine Coral recruits, algae Henry de Bey Yale University 3UHGDWRU\¿VK Jeanne Brown The Nature Conservancy +HUELYRURXV¿VK Sabine Engel Stichting Nationale Parken Bonaire Coral size class Gabriel Grimsditch IUCN Benthic cover Ramon de Leon Stichting Nationale Parken Bonaire Coral recruits, algae 0DUN9HUPHLM Caribbean Research and Management Coral size class of Biodiversity

11 Introduction

1.2 Reef resilience The reefs considered in this study were on the lee- ward western shore of the island. In general, such Reef health is largely determined by a reef’s “re- reefs consist of a shallow terrace extending from silience”, i.e. its ability to resist threats and to re- the shore to a drop-off at a depth of approximately cover to its former state after a disturbance has PHWHUV2QWKHZLQGZDUGHDVWHUQVKRUHRI occurred. Bonaire, the terrace generally extends 100-200 meters offshore to a depth of 12 meters and is The natural resilience of reefs is being under- covered primarily with crustose coralline algae and mined by stresses associated with human activi- Sargassum or gorgonians. The shallow terrace ties. These local pressures reduce the resilience on the leeward shore was characterized by coral of the system by undermining its ability to cope communities dominated by Acropora palmata and with additional stresses, such as those associated A. cervicornis mixed with Montastrea annularis and with climate change. Increasingly, policy-makers, gorgonians. Unfortunately, which conservationists, scientists and the broader com- passed through in November 1999 and Hurricane munity are calling for management actions to re- 2PDU LQ 2FWREHU  FDXVHG VLJQL¿FDQW GDP- store and maintain the resilience of the coral reefs age to these communities and wiped out nearly all in order to minimize the negative impacts of cli- $FURSRUDFRORQLHVOHDYLQJDUXEEOHWHUUDFH)URP mate change. the drop-off the fore reef slopes with an inclination of between 30° and 60° to a sediment-covered The approach used in this study was developed platform at a depth of around 50-80 meters. About by the IUCN Climate Change and Coral Reefs 65 species of scleractinian coral are found on Bo- working group (http://cms.iucn.org/cccr), led by naire’s coral reefs. &25',2(DVW$IULFDZKLFKKDVRXWOLQHGDVHULHV of protocols that include basic resistance and resil- The Bonaire National Marine Park (BNMP) was ience indicators in coral reef assessments. These created in 1979, and has had consistent manage- methods are designed to assist management au- ment since 1991. It includes all the waters sur- thorities in focusing management effort to priority URXQGLQJ%RQDLUHDQG.OHLQ%RQDLUH VLQFH  areas. from the high-tide mark to 60 meters of depth. It comprises 2,700 hectares of coral reef, seagrass and mangrove ecosystems and provides habitat for 1.3 Bonaire a diverse range of marine species including about VSHFLHVRIVWRQ\FRUDODQGPRUHWKDQVSH- Bonaire lies in the approxi- FLHVRIUHHI¿VK)XUWKHUPRUHWKHUHDUH¿YHLQWHU- mately 100km (60 miles) north of and nationally-recognized Ramsar sites on the islands 12’ north of the equator, separated from the South ± /DF .OHLQ %RQDLUH 6DOLxD 6ODJEDDL *RWRPHHU American mainland by a deep water trench. Bo- and Pekelmeer. QDLUH LV SDUW RI WKH .LQJGRP RI WKH 1HWKHUODQGV and is regarded by the as an The park is managed by a local nongovernmental, 2YHUVHDV7HUULWRU\ QRWIRUSUR¿WRUJDQL]DWLRQ67,1$3$%RQDLUHZKLFK has a co-management structure with stakeholders, 7KHUHHIVDURXQG%RQDLUHDQG.OHLQ%RQDLUHIRUP conservationists and local interest groups repre- a narrow fringing reef, which starts at the shoreline sented on the Board. The day to day management and extends to a maximum of 300 meters offshore. is carried out under the supervision of a Director by The entire reef system is protected as part of the the Marine Park manager, Chief Ranger and Rang- Bonaire National Marine Park (BNMP). ers which are all employed by STINAPA Bonaire.

12 Introduction

7KH PLVVLRQ RI WKH %103 LV µSURWHFW DQG PDQ- such as groupers and barracudas as current and age the island’s natural, cultural and historical re- growing threats to the reefs. Increasing macroal- sources, while allowing ecologically sustainable gal cover means that corals are being outcom- XVHIRUWKHEHQH¿WRIIXWXUHJHQHUDWLRQV¶,WKRVWV peted for space and light, with a subsequent loss approximately 60,000 visitors a year of which of diversity and habitat as well as a decrease in  DUH 6&8%$ GLYHUV 2QH RI WKH SULPDU\ reef accretion and growth. Increased macroalgae challenges of managing the BNMP is dealing with FDQDOVRUHÀHFWORVVRIKHUELYRUHVRUSRRUHUZDWHU the various stakeholders who use the park and TXDOLW\'HFOLQLQJKHUELYRU\E\SDUURW¿VKDQGRWK- enabling sustainable use of natural resources. er reef herbivores causes increased macroagal Stakeholders are varied and include the gov- FRYHU DV KHUELYRUHV HJ SDUURW¿VK VXUJHRQ- ernment, tour operators, hotel owners, schools, ¿VK VHD XUFKLQV  DUH D FUXFLDO IXQFWLRQDO JURXS building and zoning departments, environment in maintaining the competitive advantage of hard and nature management department, legal de- corals over macroalgae on a reef. They scrape SDUWPHQWKDUERURI¿FHDJULFXOWXUDOGHSDUWPHQW algae off the reef substrate to prepare it for coral dive operators and other water sport activity recruitment and are therefore a crucial driver of providers, non-governmental organizations, law coral reef resilience. Declining herbivore popula- enforcement, maintenance, research and moni- WLRQVFDQEHFDXVHGE\LQFUHDVHG¿VKLQJHIIRUWV toring, education, advisory and volunteer groups ,Q DGGLWLRQ WKH QXPEHU RI GDPVHO¿VK KDV LQ- among others. Some of the main challenges to FUHDVHGDVWKHVHVPDOO¿VKLQFUHDVHLQQXPEHUV PDQDJHPHQW LQFOXGH RYHU¿VKLQJ QXWULHQW HQ- now that their predators (e.g. groupers or bar- richment, land-use change, poaching, heavy UDFXGDV KDYHEHHQUHGXFHGGXHWRRYHU¿VKLQJ recreational use, sedimentation, terrestrial run- 'DPVHO¿VK FDXVH DGGLWLRQDO FRUDO PRUWDOLW\ E\ RIILOOHJDOVDQGPLQLQJDUWL¿FLDOEHDFKFUHDWLRQ µFXOWLYDWLQJ¶WHUULWRULDOWXUIDOJDOSDWFKHVRIIZKLFK LQYDVLYHOLRQ¿VK$OOWKHVHWKUHDWVFRXOGEHFRP- they feed, on large coral colonies. They maintain pounded in the near future by climate change- and defend these turf algae, which then overgrow related threats such as coral bleaching or ocean the coral leading to mortality and reduced repro- DFLGL¿FDWLRQ duction.

Despite the many threats, the successful man- $VZHOODVGHFOLQLQJKHUELYRU\RYHU¿VKLQJDQGLQ- agement of BNMP means that Bonaire’s coral FUHDVLQJGDPVHO¿VKWKHHIIHFWVRIFOLPDWHFKDQJH reefs remain among the healthiest in the Carib- DUHDOVRDSRVVLEOHPDMRUWKUHDWWR%RQDLUH¶VUHHIV bean. The islands lie outside the path of most Sustained above-average water temperature can KXUULFDQHV +XUULFDQHV 2PDU DQG /HQQ\ ZHUH OHDG WR µFRUDO EOHDFKLQJ¶ D SKHQRPHQRQ ZKHUH H[FHSWLRQV 7KH\DUHDOVRVXEMHFWWRWKHVWURQJ the symbiotic relationship between the coral host &DULEEHDQ &XUUHQW ZKLFK FRQVWDQWO\ ÀXVKHV WKH and the unicellular micro-algae that live in its tis- reefs with fresh oceanic waters. Therefore, coral sue is disrupted. The micro-algae living in the mortality seems to have been delayed compared coral tissue photosynthesize and are thus crucial to most other Caribbean sites. for the coral’s energy needs, as well as providing the coral with pigments. However, with sustained However, a study by Steneck et al. (2007) shows above-average temperatures and high UV-ra- some disturbing trends that are affecting the diation from the sun, the symbiosis between the health and resilience of Bonaire’s coral reefs. coral and the micro-algae is damaged and the mi- 7KH\LGHQWL¿HGLQFUHDVLQJPDFURDOJDHGHFOLQLQJ cro-algae are expelled from the coral. This leaves KHUELYRU\IURPSDUURW¿VKLQFUHDVHVLQGDPVHO¿VK the coral weak and in a vulnerable state to over- populations and loss of large-bodied predators growth by seaweed, infection by disease or mor-

13 Introduction

WDOLW\7KHSKHQRPHQRQLVFDOOHGµEOHDFKLQJ¶EH- 1.4 Purpose of the study cause as the coral loses its pigments, the calcium carbonate (aragonite) skeleton becomes visible Different coral reefs react differently to bleaching and the corals appear white. With increasing sea events and other threats because of various factors surface temperatures around the world due to WKDW LQÀXHQFH WKHLU UHVLOLHQFH WR GLVWXUEDQFHV climate change, bleaching events are becoming Because of their various biological compositions more frequent and more intense. In 1998-1999 a DQG SK\VLFDO FRQGLWLRQV VHH 2YHUYLHZ RI global bleaching event caused an estimated 16% Methods) we can expect different coral reefs, even mortality of corals worldwide (Willkinson, 2002). RQWKHVDPHLVODQGWRIROORZGLIIHUHQWWUDMHFWRULHV More recently in 2005, a mass bleaching event in with rising sea temperatures and increased the Caribbean caused mortality from 0-27% in dif- bleaching events. Therefore it is important for a ferent Caribbean countries. Bleaching events are manager to understand which resilience factors predicted to increase in frequency and intensity are characteristic of the coral reefs in their as seawater warms up and the climate changes. management zone. This study aims to assess the Bonaire is also at risk from the bleaching threat, various resilience characteristics of the coral reefs and it is thus important for the management of in the Bonaire National Marine Park, therefore the park to take this into account in management giving the manager a better understanding of the schemes. potential responses of the reefs to future climate change and bleaching events.

Map of Bonaire and survey sites 14 Introduction

Table 2 - Sites surveyed in Bonaire in June 2009. Geographic coordinates, management regime and depth of sampling are shown.

Date 2009 Site Depth (m) Lat (N) Long (W) Management 31 May 3OD\D)XQFKL 10 12°16’56.18” ƒ¶´ MPA

31 May Wayaka II 10 ƒ¶´ ƒ¶´ MPA -XQH 3OD\D)UDQV 10 ƒ¶´ ƒ¶´ NDA -XQH Marine Reserve 10 12°13’12.60” 68°22’26.76” NDA North 31 May .DUSDWD 10 12°13’8.21” 68°21’7.83” MPA 3 June 2LO6OLFN/HDS 10 12°11’59.69” 68°18’31.86” MPA 3 June Cliff 10 12°10’25.23” 68°17’25.72” )3$ 2 June Bari 10 12°10’3.37” ƒ¶´ )3$ 1 June Something 10 ƒ¶´ 68°17’7.22” )3$ Special 30 May 18th Palm 10 12°8’16.30” 68°16’37.79” )3$ 5 June Chachacha 10 ƒ¶´ ƒ¶´ )3$ Beach -XQH Bachelor’s 10 12° 7’32.31” ƒ¶´ MPA Beach -XQH Angel City 10 ƒ¶´ ƒ¶´ MPA 5 June Salt City 10 ƒ¶´ 68°16’56.76” MPA 5 June Tori’s Reef 10 ƒ¶´ 68°16’55.16” MPA 1 June Margate Bay 10 12°2’50.93” 68°16’18.02” MPA 1 June Vista Blue 10 12°1’57.39” 68°15’55.06” MPA 2 June Carl’s Hill 10 12°9’51.79” 68°19’23.01” MPA 7 June Mi Dushi 10 12°9’32.99” ƒ¶´ MPA 2 June South Bay 10 ƒ¶´ 68°19’21.56” MPA 7 June .HHSVDNH 10 12°8’57.19” ƒ¶´ MPA

MPA = Marine Protected Area; NDA = No Diving Area; FPA = Fish Protected Area

1.5 Overview of methods were later analyzed using Coral Point Count soft- ware. The methods applied in this study were developed by the IUCN working group on Climate Change 2) Fleshy algae – provides information on the DQG&RUDO5HHIVVSHFL¿FDOO\WRH[DPLQHWKHUHVLO- main competitors to corals on degrading reefs. ience of coral reefs to climate change (i.e., future )OHVK\DOJDHFRYHU  DQGKHLJKW FP ZDVHVWL- increases in seawater temperature). Several com- mated in 1m2 quadrats. ponents of the reef ecosystem were measured at varying levels of detail, as follows: 3) Coral size class distribution – provides de- tailed information on the demography and sizes of 1) Benthic cover – provides the main overall in- coral colonies, and can show indications of past dicators of reef state, and particularly the balance impacts by the presence or not of large colonies. It between corals and algae. Benthic photographs includes sampling of recruitment and small corals were used to assess benthic cover. Photos were in 1 m2 quadrats, and larger corals in 25x1 m belt taken from about 1 meter above the substrate and transects.

15 Introduction

 Fish herbivores and other functional groups ±¿VKH[HUWSULPDU\FRQWURORQWKHUHHIFRPPXQLW\ and on algae through herbivory, thus controlling competition between algae and corals. The num- EHUVRI¿VKLQGLIIHUHQWIXQFWLRQDOJURXSVLQFOXG- ing herbivore functional groups, was measured in 50x5 m belt transects.

5) Resilience indicators – these are factors that affect the resistance of corals to bleaching and the resilience or recovery potential of the reef commu- nity. A broad range of indicators in different classes is measured, including those overviewed in 1-6 above and some at qualitative levels. The main 2QH RI WKH VXUYH\RUV HQMR\LQJ WKH ¿VK FRXQWV RQ classes of indicators are listed below: Bonaire’s reefs.

Table 3

Indicator Type Description

Benthic Cover Cover Primary indicators of reef health, particularly of coral and algal dominance and competition.

Coral community Current Indicators of the current condition of the coral community, including recruitment, aspects of size class structure, condition, etc. Historic Indicators of the historic condition of the coral community, including past impacts and recovery to date.

Ecological – reef community Positive Abundance of organisms that are positive indicators of coral health – HJVHDXUFKLQVSUHGDWRU\¿VKHV Negative Abundance of organisms that are negative indicators of coral health – HJERULQJRUJDQLVPVHQFUXVWLQJVSRQJHVGDPVHO¿VKHWF Herbivory +HDOWKRIWKH¿VKKHUELYRUHFRPPXQLW\

Physical Substrate Substrate health, critical for settlement and survival of young corals. &RROLQJ ÀXVKLQJ )DFWRUVWKDWFDXVHPL[LQJDQGFRROLQJRIZDWHUZKLFKFDQUHGXFHWKH high temperatures experienced by a reef. Shading & screening )DFWRUV WKDW UHGXFH OLJKW SHQHWUDWLRQ LQ WKH ZDWHU WKXV UHGXFLQJ synergistic stress to corals from temperature and light. Acclimatization )DFWRUVWKDWFDXVHKLJKYDULDELOLW\LQHQYLURQPHQWDOFRQGLWLRQVWKDW promote acclimatization of corals to stress, for example exposure of corals at low tide or ponding and pooling of water leading to high temperature variability.

Anthropogenic Water Human impacts to water quality, that reduce the recovery ability of reefs and increase stress to corals. Substrate Human impacts to the reef substrate, that reduce the recovery ability of reefs and increase stress to corals. )LVKLQJ 'HJUHHRI¿VKLQJDQGLWVLPSDFWRQUHFRYHU\DELOLW\RIUHHIV

16 0DMRU¿QGLQJV

0DMRU¿QGLQJV

7KLV VHFWLRQ VXPPDUL]HV WKH PDLQ ¿QGLQJV IURP 3OD\D)UDQV6RXWK%D\.DUSDWDDQG0DULQH5H- WKH'HWDLOHGUHVXOWV VHFWLRQ ZKLFKFDQEHFRQ- serve North stand out for their healthy, resilient sulted for additional details and background infor- FRUDO SRSXODWLRQV JUDSK   7KH\ KDYH WKH PDWLRQRIWKH¿QGLQJVRYHUYLHZHGKHUH highest live coral cover of all sites surveyed with FRYHUVEHWZHHQ JUDSKVDQG  2.1 Overview and low macroalgal cover. It appears that these sites have recovered best from the devastation 7KH UHVLOLHQFH GDWD VHFWLRQ   KLJKOLJKWV WKDW FDXVHGE\+XUULFDQHV2PDUDQG/HQQ\)XUWKHU- certain sites on Bonaire are expected to have more, these sites, as well as Margate Bay in the higher overall resilience to stressors, including south, provide suitable habitat conditions favoring climate change. It also indicates sites that are bleaching-susceptible coral species (for the pur- predicted to be more susceptible to future stress. SRVH RI WKLV VWXG\ GH¿QHG DV Acropora palmata, 7DEOH  EHORZ VKRZV WKH FODVVL¿FDWLRQ RI VLWHV Acropora cervicornis, Eusmilia fastigata, Meandri- according to their resilience category (high, na meandrites, Montastrea annularis, Montastrae PHGLXPDQGORZ EDVHGRQGDWDLQJUDSKV faveolata, Montastrea franksii) that occur in great DQG7KHVHJUDSKVFOHDUO\LGHQWLI\GLVWLQFW DEXQGDQFHDWWKHVHVLWHV JUDSK 2YHUDOO groups of which those characterized by “high 3OD\D)UDQV6RXWK%D\.DUSDWDDQG0DULQH5H- resilience” can be considered to have greater reef serve North score well on all resilience indicators “health” of all sites surveyed: UHODWHG WR ZDWHU TXDOLW\ JUDSK   VXEVWUDWH FRQGLWLRQV  DQGFRUDOSRSXODWLRQV   High resilience sites: 3OD\D )UDQV 0DULQH 5H- VHUYH1RUWK.DUSDWD0DUJDWH%D\9LVWD%OXHDQG However, coral recruitment in some of the high South Bay. coral cover and high environmental quality sites is relatively low compared to other sites surveyed Medium resilience sites3OD\D)XQFKL:D\DND JUDSK   )RU H[DPSOH 3OD\D )UDQV KDV  ,, 2LO 6OLFN /HDS &OLII %DUL 6RPHWKLQJ 6SHFLDO recruits (colonies sized 0-2.5cm) per 100 m2.DU- 18th Palm, Bachelor’s Beach, Angel City, Salt City, pata has 83 recruits per 100 m2, and South Bay 7RUL¶V5HHI&DUO¶V+LOO0L'XVKLDQG.HHSVDNH has 102 recruits per m2, all lower than the overall average of 152 recruits per 100 m2. This is often Low resilience site: Chachacha. the case in healthier coral communities with higher coral cover where there is less space available for Unsurprisingly, the highly resilient sites are located recruitment so this is not necessary a cause for DZD\IURP.UDOHQGLMNWKHPDLQXUEDQFHQWUHDQG worry, however it is important to ensure that coral where the main coastal development impacts are. mortality in these sites remains low by ensuring 3OD\D)UDQV.DUSDWDDQG0DULQH5HVHUYH1RUWK that the quality of the local environment remains DUH DZD\ WR WKH QRUWK IXUWKHUPRUH 3OD\D )UDQV high, i.e., unchanged. and Marine Reserve North are No Diving Areas, meaning that there is even less human impact. A worrying trend is the increasing cover of Tri- Margate Bay and Vista Blue are to the southern didemnum, a tunicate that overgrows living coral, HQGRIWKHLVODQG6RXWK%D\LVRQ.OHLQ%RQDLUH LQQRUWKHUQVLWHV JUDSKVDQG VXFKDV but on the opposite side of the island away from 0DULQH5HVHUYH1RUWK.DUSDWDDQG3OD\D)UDQV .UDOHQGLMN Although the abundance of this tunicate is low

17 0DMRU¿QGLQJV

DW WKH PRPHQW PD[LPXP  FRYHU DW P the proportion of local coral colonies that showed depth), there is anecdotal evidence that cover is some sign of disease. These results show that in increasing at deeper depths and this trend should Bonaire predators control the population of dam- be monitored given this species’ ability to rapidly VHO¿VKUDWKHUWKDQKDELWDWDQGWKDWWKHPRUHGDP- overgrow large patches of living coral. VHO¿VKWKHUHDUHWKHPRUHWXUIDOJDHDQGGLVHDVHG corals there are too. A further worrying trend is the increasing popula- WLRQRIGDPVHO¿VKDVQRWHGLQ6WHQHFNHWDO $V SUHGDWRU ¿VK DUH OLNHO\ LPSRUWDQW IRU UHJXODW- DQG WKLV ,8&1 VWXG\ FRQ¿UPV WKH WUHQG 7KHVH LQJ WKH QXPEHU RI GHVWUXFWLYH GDPVHO¿VK WKHLU VPDOOWHUULWRULDO¿VKJDUGHQWXUIDOJDOµ\DUGV¶ZKLFK populations are also important for coral reef re- they use as food sources and zealously protect VLOLHQFH7KHPDMRULW\RISUHGDWRUELRPDVVRQ%R- them against invaders. However, they kill live coral naire is made up of grunts and snappers, although to maintain and expand their yards, and are caus- Margate Bay also has high biomass of groupers LQJDORWRIGDPDJHLQ%RQDLUH¶VUHHIV*UDSK JUDSK ,WVHHPVDVLIWKHKLJKUHVLOLHQFHRI VKRZV0DULQH5HVHUYH1RUWK6RXWK%D\2LO6OLFN sites such as Margate Bay and Vista Blue are in /HDS DQG .HHSVDNH VKRZ WKH KLJKHVW UHODWLYH WKLVFDVHLVGULYHQE\JRRG¿VKSRSXODWLRQV JUDSK DEXQGDQFH RI FRUDOGHVWUR\LQJ GDPVHO¿VK ZKLOH  WKDWFDQKHOSPHGLDWHFRUDODOJDOLQWHUDFWLRQ :D\DND ,, DQG 3OD\D )XQFKL H[KLELW WKH ORZHVW and competition. Both these sites are also next relative abundances. The high resilience of sites to a Ramsar site (i.e., salt plains), but it is pos- such as Marine Reserve North and South Bay is sible that the salt discharge affects reef resilience thus potentially at risk if the trend of increasing as there is a wide layer of anoxic sediment at the GDPVHO¿VKSRSXODWLRQVFRQWLQXHV GLVFKDUJHSRLQWV*UDSKVKRZVWKDW0DUJDWH Bay and Vista Blue don’t have the highest live cor- 'DPVHO¿VKSRSXODWLRQVRQ%RQDLUHDUHOLNHO\FRQ- DOFRYHU DQGUHVSHFWLYHO\ KRZHYHU WUROOHG E\ SUHGDWRU SRSXODWLRQV 9HUPHLM HW DO LQ they have among the highest coral recruitment SUHSDUDWLRQ 2IDOOEHQWKLFDQG¿VKJXLOGVFRQVLG- (215 and 188 recruits per 100 m2 respectively) in- HUHGRQO\WKHDEXQGDQFHRISUHGDWRU\¿VKHVFRXOG dicating high potential for recovery. EH UHODWHG WR WKH ORFDO DEXQGDQFH RI GDPVHO¿VK *UDSK ZLWKDQHJDWLYHFRUUHODWLRQEHWZHHQ GDPVHO¿VK DQG SUHGDWRU ELRPDVV 'DPVHO¿VK Stegastes planifrons were observed on colonies of 12 different coral species, and only on four massive coral species (Montastraea annularis, M. faveolata, M. franksi and Colpophylia natans) did they occur in greater densities than expected. &OHDUO\WKHGDPVHO¿VKSUHIHUODUJHPDVVLYHFRUDO species over smaller branching, i.e., structurally more complex species (Eusmilia fastigiata, Mil- lepora complenata, Madracis decactis and M. mi- rabilis +RZHYHUWKHDEXQGDQFHRIGDPVHO¿VK¶V preferred habitat (Montastraea coral species) had no positive effect on the local abundance of GDPVHO¿VK *UDSK   7KH DEXQGDQFH RI S. 3DUURW¿VKVFUDSLQJDOJDHRIIWKHVXEVWUDWH7KLVDFWLRQ planifrons was positively correlated with the lo- cleans the substrate and prepares it for coral colonization, cal cover of turf- and crustose coralline algae and and thus increasing reef resilience.

18 0DMRU¿QGLQJV

7KLVJURXSRIµPHGLXP¶UHVLOLHQFHVLWHVLOOXVWUDWHV the complex nature of coral reef resilience. The JURXS FDQ DFWXDOO\ EH GLYLGHG LQWR µKLJKHU¶ DQG µORZHU¶ UHVLOLHQFH FDWHJRULHV DV ZHOO :LWKLQ WKH medium resilience group, there is a gradient of UHVLOLHQFHDQGVLWHVVXFKDV:D\DND,,3OD\D)XQ- chi (to the north) and Angel City (to the south) that DUHIXUWKHUDZD\IURP.UDOHQGLMNKDYHKLJKHUUHVLO- LHQFHVFRUHVWKDQVLWHVFORVHUWR.UDOHQGLMNVXFK as Bari, Something Special, Bachelor’s Beach and &OLII JUDSK :D\DND,,DQG3OD\D)XQFKLDUH located next to the protected Washington Slagbaai National Park, and there is little to no coastal de- 'HQXGLQJ VXUJHRQ¿VK IHHGLQJ RQ DOJDH 6FKRROV RI YHORSPHQWLQWKHDUHD6LWHVRQ.OHLQ%RQDLUH D WKHVH¿VKFUXLVHDORQJWKHUHHIHDWLQJDOJDHDQGDUHYLWDO for keeping algal growth in check thereby increasing reef RAMSAR area with no coastal development at all) resilience. VXFKDV.HHSVDNH0L'XVKLDQG&DUO¶V+LOODOVR suffer less land-based stress and fall in the higher High resilience sites are characterized by high resilience category. abundances (i.e, biomass) of important reef herbi- vores. South Bay has the highest biomass of par- 2QWKHRWKHUKDQG6RPHWKLQJ6SHFLDOLVORFDWHG URW¿VKFDSDEOHRIVFUDSLQJDOJDHRIIWKHVXEVWUDWH next to a marina with boats mooring directly on the and making it suitable for coral recruitment (graph UHHIDQGVXEMHFWHGWRSK\VLFDOGXPSLQJRIZDVWH   0DUJDWH %D\ KDV WKH KLJKHVW ELRPDVV RI and affected by runoff from a large catchment EURZVHUV ¿VKWKDWIHHGRQPDFURDOJDOIURQGVHJ area. Bari is located next to a pier, a residential FKDHWRGRQVDQGSRPDFDQWKLGVJUDSK DQG DUHD DQG DQ DUWL¿FLDO EHDFK &OLII LV ORFDWHG QH[W GHQXGHUV ¿VKWKDWUHPRYHHSLOLWKLFDOJDOWXUIIURP the reef substratum, but do not scrape the sur- face, e.g. acanthurids and microspathodon, graph  ERWKFUXFLDOIRUNHHSLQJDOJDOJURZWKXQ- der control and thus preventing overgrowth and/or shading of neighbouring corals.

)LQDOO\KLJKUHVLOLHQFHVLWHVVXFKDV3OD\D)UDQV DQG6RXWK%D\KDYHWKHPRVWIDYRXUDEOHFDOFL¿FD- WLRQ UDWHV$ FDOFL¿FDWLRQ UDWH KLJKHU WKDQ  LQGL- cates a growing reef, i.e., there are more calcifying organisms (live coral) than non-calcifyers (other VXEVWUDWHW\SHV $FDOFL¿FDWLRQUDWH LQGLFDWHV a reef that is not growing due to the dominance RIQRQFDOFLI\HUVRYHUFDOFLI\HUV3OD\D)UDQVDQG 6RXWK %D\ KDYH WKH KLJKHVW FDOFL¿FDWLRQ UDWHV DQGUHVSHFWLYHO\JUDSK EXWWKH\ Salt plains to the south of the island. High salinity are still lower than 1 meaning that they may be discharges from these plains could affect development of corals that are close to the discharge points such as susceptible to erosion. Salt City or Tori’s Reef.

19 0DMRU¿QGLQJV

WRDZDWHUIDFWRU\DQGLVVXEMHFWWRUXQRIIIURPD It has the lowest live coral cover (9.2%, graphs large catchment area as well as leaching from DQG DQGVHQVLWLYHFRUDOVSHFLHVDUHQR septic tanks. Bachelor’s Beach is located next to a ORQJHUSUHVHQW JUDSK DQGKDVRQHRIWKH UHVLGHQWLDODUHDDQGDOVRVXEMHFWHGWRQXWULHQWUXQ ORZHVWFDOFL¿FDWLRQUDWHVRIDOOVLWHV JUDSK RII7KHRQO\VLWHVWKDWDUHIDUDZD\IURP.UDOHQGLMN  PHDQLQJWKDWLWLVKLJKO\VXVFHSWLEOHWRHUR- and that score poorly are Salt City and Tori’s Reef sion. This degraded and impacted site serves ZKLFKDUHERWKVXEMHFWWRODQGEDVHGVWUHVVIURP as a stark reminder of what can happen to coral the very high salt discharges directly on to them reefs if they are not protected from the effects from the salt plains. In summary, land based hu- of land-based human stressors. man activities had great negative impacts on near- by reef communities and likely reduced their ability +RZHYHULWLVD)LVK3URWHFWHG$UHDVRLWLVKRSHG to recover from future (natural) disturbances. WKDW¿VKSRSXODWLRQVZLOOEHDEOHWRUHFRYHURYHU time and add a possible layer of resilience, at least 7KHUH LV RQH FOHDU RXWOLHU LGHQWL¿HG E\ WKH GDWD to the food security provided by Bonaire’s reefs. In ZKLFKIDOOVLQWRLWVRZQµORZ¶UHVLOLHQFHFDWHJRU\GXH IDFWWKH)LVK3URWHFWHG$UHDV &OLII%DUL6RPH- to the degradation and impacts it suffers: Playa thing Special, 18th Palm, Something Special and &KDFKDFKD7KLVVLWHLVULJKWQH[WWR.UDOHQGLMNDQG Chachacha) are all to the lower end of the resil- suffers impacts of coastal development and run-off LHQFH VSHFWUXP JUDSK   EHFDXVH WKH\ DUH from land-based sources of pollution and sewage. ORFDWHGDURXQG.UDOHQGLMN*UDSKLOOXVWUDWHV This site scored poorly in resilience indicators re- WKLVZHOO)LVK3URWHFWHG$UHDVDOOVFRUHEDGO\RQ ODWLQJWRDOJDH JUDSK FRUDOV JUDSK  resilience factors relating to coral populations, sub- ¿VK JUDSK  VXEVWUDWH TXDOLW\ JUDSK   strate conditions, water quality, and anthropogenic DQGZDWHUTXDOLW\ JUDSK ZKLFKFRXOGDOOEH stressors, yet they are positively correlated with UHODWHGWRWKHODQGEDVHGLPSDFWVIURP.UDOHQGLMN ¿VKSRSXODWLRQV7KLVVKRZVWKDWDOWKRXJKWKH)LVK

Dead Acropora stands. Acropora beds in the shallows were completely wiped out during Hurricane Lenny in 1999 and in 2008. Their recovery will depend on recruitment and environmental quality of the affected sites.

20 0DMRU¿QGLQJV

7DEOH

Resilience rating Sites Management High Marine Reserve North NDA High 3OD\D)UDQV NDA High .DUSDWD MPA High Margate Bay MPA High Vista Blue MPA High South Bay MPA Medium 3OD\D)XQFKL MPA Medium Wayaka II MPA Medium 2LO6OLFN/HDS MPA Medium Cliff )3$

Medium Bari )3$ Medium Something Special )3$ Medium 18th Palm )3$ Medium Bachelor’s Beach MPA Medium Angel City MPA Medium Salt City MPA Medium Tori’s Reef MPA Medium Carl’s Hill MPA Medium Mi Dushi MPA Medium .HHSVDNH MPA Low Chachacha )3$

Protected Area management seems to be working 6LWHV ZLWK KLJK FRUDO FRYHU 3OD\D )UDQV 0D- IRU¿VKRWKHUODQGEDVHGVRXUFHVRIVWUHVVRYHU- ULQH5HVHUYH1RUWK.DUSDWDDQG6RXWK%D\ DUH ride this positive effect and lead to loss of resil- marked with a coral symbol. ience. Degradation of water quality due to land- based activities thus appears to an acute threat to 6LWHV ZLWK KLJK GDPVHO¿VK GHQVLW\ 0DULQH 5H- Bonaire’s reefs that needs to be resolved. VHUYH1RUWK2LO6OLFN/HDS6RXWK%D\DQG.HHS- VDNH DUHPDUNHGZLWKDGDPVHO¿VKV\PERO The map on the following page shows the resil- ience ratings and factors by site. High resilience Sites with high herbivory levels (Bari, Mi Dushi, VLWHV 0DULQH 5HVHUYH 1RUWK 3OD\D )UDQV .DU- South Bay and Salt City) are marked with a par- pata, Margate Bay, South Bay and Vista Blue) are URW¿VKV\PERO marked by green circles. Medium resilience sites 3OD\D )XQFKL :D\DND ,, 2LO 6OLFN /HDS &OLII Sites with high predator biomass (Margate Bay Bari, Something Special, 18th Palm, Bachelor’s and Vista Blue) are marked with a snapper sym- Beach, Angel City, Salt City, Tori’s Reef, Carl’s Hill, bol. 0L'XVKL.HHSVDNH DUHPDUNHGE\\HOORZFLUFOHV Low resilience sites (Chachacha) are marked by a red circle.

21 0DMRU¿QGLQJV

2.4 DISTRIBUTION

A vegetation buffer around your entire building site

2.2 Threats to Bonaire’s coral reef small area of damage on a coral sets of a reaction communities that affects the whole coral colony. Clearing native vegetation in order to construct buildings on the &RDVWDOGHYHORSPHQWDQGDUWL¿FLDOEHDFKHV coast destabilizes the topsoil and high winds or rainfall then blow or washes the soil into the sea. Coastal development and construction is one of &UHDWLQJDUWL¿FLDOEHDFKHVOLNHWKHRQHVFORVHWR WKHPDMRUWKUHDWVWRWKHFRUDOUHHIVRI%RQDLUH7KH\ 18th Palm and Bari, also has a negative effect are very vulnerable to pollution and sedimentation. RQ FRUDOV $UWL¿FLDO EHDFKHV UHTXLUH VHD VDQG Debris, sand, cement, stones and other runoff of not the crushed sand you can buy, and Bonaire coastal development and erosion that are washed does not have a supply of natural sea sand that in the sea can cause serious damage or mortality can be used for this purpose. What sea sand the to corals by smothering them and blocking their LVODQGGRHVKDYHLVDVFDUFHFRPPRGLW\GLI¿FXOW access to the sunlight they need for energy. As to extract or occurs within protected areas, such corals use energy to clean themselves, even a as Lac Bay and the Wasington Slagbaai National

22 0DMRU¿QGLQJV

3DUN$UWL¿FLDOEHDFKHVZDVKDZD\JUDGXDOO\ZLWK ticides, soil and debris from your yard and off the time through natural processes and are removed streets into the sea. Therefore, by preserving as immediately during conditions of wind reversal or much existing vegetation as possible and choos- high waves. Because of the highly negative im- ing native and other drought resistant plants and pacts of sedimentation on coral reefs, the sites in trees this pollution can be reduced. Native plants, Bonaire closest to highly developed coastal areas once established, do not require much, if any, wa- HJ.UDOHQGLMNDQGWKHSRUW KDYHORZHUUHVLOLHQFH ter, fertilizer or pesticides. This will save you time ratings and overall health (e.g. Chachacha Beach) and money. It also reduces pollution from excess than those that are removed from coastal devel- fertilizer and pesticides that could wash off your RSPHQW 0DULQH5HVHUYH1RUWK3OD\D)UDQV.DU- yard. This is especially important on properties pata, Margate Bay, Vista Blue, South Bay). near the sea. Any irrigation with grey or black wa- ter, deposits extremely high quantities of nutrients The following excerpt is from the construction (fertilizers) onto the coral reefs. These nutrients guidelines for Bonaire: promote algal growth which kills the coral. If you have plants, like palm trees, that require irriga- µ,Q DQ HIIRUW WR PLQLPL]H HURVLRQ DQG UXQ RII WKH tion, locate them as far away from the shore as period of time spent in construction must be as possible…. It can take at least six weeks to get a short as possible. That is why you should not clear building permit and numerous months to select a your location until construction begins. Especially builder. Waiting until the last possible moment will during construction near the shore, erosion control help hold your topsoil in place, prevent erosion, WHFKQLTXHV OLNH VLOW VFUHHQV DQG ¿OWHU FORWK PXVW and if you are building near the sea, it will keep the EHHPSOR\HGLI¿OOPDWHULDOLVOHIWH[SRVHG)ROORZ- sediment off the reef. ing construction, the shoreline must be immedi- ately stabilized with native vegetation…. Leave as The vegetation you must remove can be re-used. much natural vegetation on the site as possible. Wood can be used for wood chips and to produce 7KHOLPLWHGUDLQIDOOPHDQVLWLVGLI¿FXOWDQGH[SHQ- charcoal. Trees can be transplanted. Topsoil that sive to grow and maintain plants. However, the becomes available after removing the vegetation native vegetation already on your property is very can be re-used in the landscaping. When you start well adapted to the environment of Bonaire and construction, it is most important that you set up requires no watering, so it is worth looking after. the building site in such manner that prevents pol- 2QRXUDULGLVODQGLWLVZLVHWROHDYHDVPXFKH[- lution, erosion and damage to terrestrial and ma- isting vegetation as possible, using it as the basis ULQHHQYLURQPHQW.HHSLQJWKHEXLOGLQJVLWHFOHDQ of your garden. This saves you money by reducing during construction is the best way to prevent this. costs of clearing the site and re-landscaping your It also saves you time and energy at the end of the garden as well as reducing costs for irrigation. This construction because you do not have to clean as will also help conserve the islands vegetation and much and scrape off materials from the building preserve your valuable topsoil. If you remove too site. much vegetation you will loose your topsoil. It will blow away, creating a dusty, barren yard, or will If you are not able to screen the whole site, you run off after rainfall. Reducing pollution from runoff should at least screen the leeward side of the site is another way you can help the environment and to catch waste that can blow away or runoff. If your marine life. Plants that are not native to the island SODQVFDOOIRUDIHQFHRUZDOODURXQG\RXU¿QDOFRQ- require a lot of water, fertilization and pest control, VWUXFWLRQ FRQVLGHU EXLOGLQJ LW ¿UVW 7KLV ZD\ \RX non-native plants can also bring diseases. Water will not have the added expense of a temporary from rain and irrigation washes fertilizers and pes- screen.

23 0DMRU¿QGLQJV

is a very good start for your future garden. It also species composition and abundance. The 3 com- prevents soil run off and keeps waste materials SRQHQWV RI VHZDJH HIÀXHQW PRVW GHWULPHQWDO WR from littering the streets, nature and the sea. coral communities are nutrients, sediments, and toxic substances. Nutrient enrichment by sewage During construction be careful not to unnecessar- HIÀXHQWPD\HQKDQFHEHQWKLFDOJDOELRPDVVDQG ily damage the existing vegetation when stacking primary production in the water column. Increased materials or placing containers or equipment. Ce- primary production in the water column favours ment and concrete are especially harmful to plants EHQWKLF¿OWHUIHHGLQJLQYHUWHEUDWHVZKLFKZLWKWKH and marine life. benthic algae, may out-compete corals and other reef-building organisms. Anthropogenic inputs of ,WLVSURKLELWHGWRFUHDWHDUWL¿FLDOEHDFKHVRQ%R- dissolved nutrients and organic particulate mat- naire and a permit is required to replenish natural- ter may also depress oxygen levels. While heavy ly occurring beaches. Sand can wash or be blown sediment loads on corals may be lethal, lesser into the sea and kill our coral reefs. quantities may inhibit growth, causing changes in the growth forms of colonies, decreasing coral Bonaire has no fresh water supply. Therefore, all cover, altering species composition of reef-build- fresh water needs to be distilled from seawater ing organisms, and inhibiting coral recruitment. through the process of reverse osmoses, which Toxic substances may induce metabolic changes uses an enormous amount of energy, thus making in corals, decrease rates of growth and reproduc- ZDWHU H[SHQVLYH )LOOLQJ DQG UH¿OOLQJ \RXU VZLP- tion, or reduce viability of corals. ming pool with fresh water will cost you a lot of money. To keep the pool clean and bacteria free, Recommendation: It is strongly recommended chloral and other chemicals need to be added. that Bonaire invest in appropriate sewage treat- However, you are not allowed to dispose of chlo- ment facilities to improve water quality and in- rinated water into the sea or on land. You are not crease the resilience of its valuable coral reefs. DOORZHGWR¿OOWKHSRROZLWKVHDZDWHU7KLVLVEH- It is also recommended that a water quality FDXVHDSRROZLWKVHDZDWHUQHHGVDWKURXJKÀRZ monitoring program be set up and sustained. to the sea, which may result in chemical or organic (algae) pollution.’ 'DPVHO¿VK

Recommendation: All coastal construction on Bonaire should be strictly regulated and follow the construction guidelines. The guidelines should become law in order to be enforced ap- propriately.

Leaching from septic tanks

A wide range of sewage impacts on coral reef com- munities has been reported. Little or no impact has EHHQREVHUYHGRQVRPHUHHIVLQZHOOÀXVKHGZD- WHUVWKDWUHFHLYHVPDOOTXDQWLWLHVRIHIÀXHQWZKHUH- DVODUJHGLVFKDUJHVRIHIÀXHQWLQWRSRRUO\ÀXVKHG ODJRRQVDQGED\VKDYHFDXVHGPDMRUFKDQJHVLQ 'DPVHO¿VKSURWHFWLQJLWVDOJDO\DUG7KHVHWHUULWRULDO¿VK kill live coral to maintain and expand their algal yards

24 0DMRU¿QGLQJV

7HUULWRULDO GDPVHO¿VK DUH D ¿VK VSHFLHV WKDW µJDUGHQ¶ DOJDO WXUI µ\DUGV¶ ZKLFK WKH\ GHIHQG IRU food. Unfortunately, they maintain and expand their yards by killing corals. Populations of WHUULWRULDOGDPVHO¿VKKDYHEHHQREVHUYHGWRKDYH increased recently (Steneck et al, 2008) and are causing coral mortality in many areas of Bonaire. This population increase hampers coral reef resilience and can stop reefs from recovering from VWUHVVDQGPRUWDOLW\HYHQWV,QFUHDVLQJGDPVHO¿VK SRSXODWLRQV LV OLNHO\ OLQNHG WR RYHU¿VKLQJ RI predators that feed on as there is a negative FRUUHODWLRQ EHWZHHQ GDPVHO¿VK Stegastes planifrons) and predator biomass, as shown by *UDSK

Recommendation: It is recommended that the Trididemnum (left) overgrowing live coral (right) ¿VKLQJRISUHGDWRU\¿VKVSHFLHVRQ%RQDLUH¶V coral reefs be controlled and managed to a sustainable level to prevent population H[SORVLRQV RI SUH\ ¿VK FDSDEOH RI PRGLI\LQJ Trididemnum and Lobophora the reef habitat. Trididemnum solidum cover, a tunicate that competes with and overgrows coral, has increased LQQRUWKHUQVLWHV JUDSKVDQG DQGLQ deeper sections of the reef. Lobophora variegata cover, a brown macroalgae that also smothers and overgrows coral, has also been observed to have increased in sites around the island (graphs DQG Lobophora population increases could be natural population cycles, but could also be linked to decreasing water quality and QXWULHQWLQÀX[IURPODQG HJVHSWLFWDQNOHDFKLQJ fertilization by erosion of iron rich geological structures). Trididemnum, on the other hand, is possibly an invasive species.

Recommendation: It is recommended that the populations of Trididemnum and Lobophora are closely monitored and the factors contributing to the unnatural abundance of these coral-

School of predatory snappers. It is important to maintain overgrowing organisms should be studied and SRSXODWLRQVRISUHGDWRU\¿VKVXFKJURXSHUVRUVQDSSHUV then eliminated. in order to maintain a balanced population on Bonaire’s reefs.

25 0DMRU¿QGLQJV

Therefore, protecting the resilience and health of Bonaire’s coral reefs from the threats described in this report is of paramount importance to the island’s welfare. The recommendations outlined above should be implemented in order to continue PRQLWRULQJ DQG ¿QGLQJ VROXWLRQV WR SURWHFW WKH resilience of this valuable ecosystem. Bonaire should invest in appropriate sewage treatment facilities to improve water quality and increase the resilience of its valuable coral reefs. It is also recommended that a water quality monitoring SURJUDP EH VHW XS DQG VXVWDLQHG )LVKLQJ RI SUHGDWRU\ ¿VK VSHFLHV RQ %RQDLUH¶V FRUDO UHHIV should also be controlled and managed to a sustainable level to prevent population explosions RI SUH\ ¿VK VXFK DV GDPVHO¿VK FDSDEOH RI Brown algae Lobophora overgrowing live coral modifying the reef habitat. Trididemnum and Lobophora should also be closely monitored and the factors contributing to the unnatural 2.3 Summary abundance of these coral-overgrowing organisms should be studied and then eliminated. Some of Bonaire’s economy is dependent on , with WKHVH WKUHDWV DUH %RQDLUHVSHFL¿F GDPVHO¿VK between 50 and 70% of the island’s economy Trididemnum, Lobophora), so even though they DWWULEXWHG WR WKLV VHFWRU  WRXULVWV YLVLWHG may not be recognized as threats to coral reefs the island in 2008, and of these approximately at a global scale it is crucial for managers to 52,000 used the BNMP and 36,000 were SUBA understand and take into account these local divers. Bonaire is known as a diver’s paradise and IDFWRUV WKDW LQÀXHQFH FRUDO UHHI UHVLOLHQFH ZKHQ LVDPDMRUGHVWLQDWLRQIRU1RUWK$PHULFDQGLYHUV designing and implementing management plans ZKR PDNH XS  RI DOO DUULYDOV '(=$   and monitoring programmes. %HFDXVH FRUDO UHHIV DUH WKH PDMRU DWWUDFWLRQ IRU most tourists, their health and attractiveness to 6&8%$ GLYHUV FRXOG GLUHFWO\ LQÀXHQFH %RQDLUH¶V economy. Therefore, protecting this valuable natural resource is crucial for the livelihoods of the island’s inhabitants and there is much to lose economically (as well as in terms of food security, biodiversity and other ecosystem services) and socially if the coral reefs become too degraded. Already some of Bonaire’s coral reefs are at the tipping point of becoming functionally extinct, DQG ZH KDYH LGHQWL¿HG &KDFKDFKD ERUGHULQJ .UDOHQGLMNDVWKHPRVWYXOQHUDEOHVLWHWREHFRPH irreversibly damaged in the near future.

26 Detailed Methodology

3. Detailed Methodology

3.1 Survey methods

Benthic cover

Benthic photographs were used to assess benthic cover. Photos were taken from about 1 meter above the substrate and were later analyzed using Coral Point Count software. Categories used ZHUHWKHIROORZLQJZLWKFRUDODQGDOJDHIXUWKHULGHQWL¿HGWRWKHJHQXVOHYHO

7DEOH%HQWKLFFDWHJRULHVIRULGHQWL¿FDWLRQ

Invertebrates Algae Other Substrate Coral )OHVK\DOJDH Microbial Rock Recent Dead Coral Algal Assemblage Seagrass Rubble Soft coral Coralline Algae 8QLGHQWL¿HG Sand Invertibrates Halimeda Sponge

Coral population structure

&RUDOSRSXODWLRQVWUXFWXUHZDVTXDQWL¿HGXVLQJ¿[HGVL]HFODVVHVRIFRUDOVIURPWKHVPDOOHVWUHFUXLWVWRWKHODUJ- est adults at a site. A belt transect 25 m long and 1 m wide was used to record the number of colonies larger than FP)RUFRUDOVVPDOOHUWKDQFPVXEVDPSOLQJZDVGRQHXVLQJVL[P2 quadrats at the 0, 5, 10, 15, 20, and PDUNV2QO\FRORQLHVZKRVHFHQWHUOLHVZLWKLQWKHVDPSOHGXQLWVZHUHFRXQWHG±ODUJHFRORQLHVZLWKWKHLUFHQ- ter outside the transect were ignored. A 1 m stick was used to help guide estimation of transect width, mark the 1 m2 quadrats and help guide size estimation of coral heads (Table 6a). Genera that covered a range of bleaching susceptibility from high to low (Table 6b), and that are generally common on Caribbean reefs were selected.

Table 6a - Size classes of corals for size class measurements. Size classes Sampling method (cm) (1) 0-2.5 Recorded in six 1m2 (2) 2.5-5 quadrats per transect (3) 5-10 (4) 10-20 Recorded in 25*1 m (5) 20-40 belt transects (6) 40-80 (7) 80-160 (8) 160-320 (9) > 320

27 Detailed Methodology

Table 6b - Selected species in classes of bleaching susceptible, intermediate and resistant.

Susceptible Intermediate Resistant Acropora palmata Agaricia lamarki Colpophyllia natans Acropora cervicornis Diploria clivosa Dendrogyra Eusmilia fastigata Diploria labyrinthiformis cylindrica Meandrina meandrites Diploria strigosa Madracis decactis Montastrea annularis Millepora spp Madracis mirabilis Montastrae faveolata Mycetophyllia spp Montastrea Montastrea franksii Porites asteroides cavernosa Siderastrea siderea Porites porites Stephanocoenia Agaricia agaricites intersepta

Replication of transects depended on logistics at a site and the complexity of the coral community, YDU\LQJEHWZHHQDQG.

Fish community structure

)LVKVXUYH\VIRFXVHGRQKHUELYRUHIXQFWLRQDOJURXSVIROORZLQJ*UHHQHWDO  DQGVHH,8&1&&&5  ZLWKVL]HFODVVHVDOVRHVWLPDWHGIRURWKHU¿VK+HUELYRU\LVLPSRUWDQWDVLWOLPLWVFRPSHWLWLRQDQG REVWUXFWLRQE\DOJDH)RXUIXQFWLRQDOJURXSVRIKHUELYRURXV¿VKHVZHUHXVHGQRQGHQXGLQJGHQXGLQJ excavators/scrapers, browsers (Table 7), each playing a unique ecological role in coral reef resilience.

7DEOH)XQFWLRQDOJURXSVRIKHUELYRURXVDQGSUHGDWRU\¿VKHVUHFRUGHGLQWKLVVXUYH\

Functional group Taxonomic groups Function and notes Non-denuding Territorial Pomacentridae Habitat engineers, create algal µ\DUGV¶RQOLYHFRUDODQG protect these yards against other herbivores. Responsible for coral mortality and retarded coral recovery. Denuding Acanthurids, Microspathadon Algal control. Remove epilithic algal turf from the reef substratum, but do not scrape the surface, prevent coral overgrowth and shading by macroalgae. Excavators, scrapers Scarids Bioerosion, colonization surfaces. Remove algae, sediment and other material by closely cropping or scraping the substrate. Browsers Chaetodons, Pomacanthids $OJDOFRQWURO)HHGRQPDFURDOJDO fronds, reduce coral overgrowth and shading by macroalgae. Carnivores /XWMDQLGDH+DHPXOLGDH 3UHGDWHRQRWKHU¿VKLQFOXGLQJ Serranidae, Carangidae, herbivores. Sphyraenidae 5HVSRQVLEOHIRUNHHSLQJ¿VK populations in check and avoiding population explosions.

Sampling was done in three 50 x 5 m belt transects. The transects were separated by at least 5 to PIURPWKHHQGRIWKHSUHYLRXVWUDQVHFW$OO¿VKLQWKHDERYHFDWHJRULHVZHUHFRXQWHG

28 Detailed Methodology

Resilience Indicators

Resilience indicators were measured or estimated during each sampling dive, generally towards the second half of the dive to allow time for familiarization with the site. The indicators and their overall grouping is shown in the table below.

Table 8 - Indicators recorded.

Group Factor Variable 1-Cover Coral Hard coral Gorgonians Algae )OHVK\$OJDH Dead coral with algae Coralline algae Substrate Rubble

2-Physical Substrate Topogr. Complex. - micro Topogr. Compl. - macro Sediment texture Sediment layer &RROLQJ ÀXVKLQJ Currents Wave energy/ exposure Deep water (30-50m) Depth of reef base Shading & Aspect screening Slope (degrees) Physical shading Canopy corals Visibility (m)/ turbidity Acclimatization Exposed low tide Ponding/pooling 3-Coral community Size/age Largest corals (3) Condition Coral bleaching Mortality-new Mortality-old Recovery-old Coral disease &RUDODVVRFLDWHV Positive Branching residents Negative Competitors %LRHURGHUV XUFKLQVQRQ¿VK Bioeroders (internal, spo) Corallivores (negative) )LVKJURXSV Herbivory Browsers Denuding Excavating Non-denuding )LVKLQJ Predators 6-Anthropogenic Water Nutrient input Pollution (chemical) Substrate Pollution (solid) Turbidity/Sedimentation Physical damage Storm damage )LVKLQJ 'HVWUXFWLYH¿VKLQJ )LVKLQJSUHVVXUH Management MPA Resources ,&=0

29 Detailed Methodology

A semi-quantitative 5-point scale was used for estimation of most of the indicators, except for those (such as temperature, visibility) that could easily be measured or estimated quantitatively. &ODVVL¿FDWLRQRIWKHSRLQWVFDOHZDVGRQHXVLQJORFDODQGUHJLRQDONQRZOHGJH,QWKHSRLQWVFDOH general principles were to assign them as follows: minimum (1), maximum (5) and moderate (3) OHYHOIRUHDFKLQGLFDWRUIRUWKHUHJLRQRIDSSOLFDWLRQDQGLQWHUPHGLDWHOHYHOVRIORZ  DQGKLJK  

)RUDQDO\VLVWZRRSHUDWLRQVKDGWREHDSSOLHGWRWKHUDZGDWDFROOHFWHGLQVLWX

‡ )RUYDULDEOHVPHDVXUHGTXDQWLWDWLYHO\WUDQVIRUPDWLRQVZHUHDSSOLHGWRDVVLJQWKHPWRDSRLQW scale for consistency in multivariate analysis of the data. In general terms, the distribution of values could be even across sites (resulting in even numbers of sites assigned to levels 1 to 5), concentrated around the middle (large number of sites at moderate level 3), or strongly skewed to one side (most sits high or low for a variable).

‡ ,QVLWXHVWLPDWLRQRISRLQWVFDOHVZHUHGRQHEDVHGRQWKHSDUDPHWHULWVHOILHIURPORZWRKLJK )RUFRQVLVWHQWPXOWLYDULDWHDQDO\VLVVRPHLQGLFDWRUVKDGWREHUHYHUVHGVRWKDWDOOYDOXHVµJRRG¶ IRUFRUDOVVFRUHGDQGDOOYDOXHVEDGIRUFRUDOVVFRUHG)RUH[DPSOHDOJDOOHYHOVLQWKH¿HOG PLJKWKDYHEHHQVFRUHGµ¶IRUKLJKOHYHOVEXWLQDQDO\VLVWKLVZDVUHFRGHGDVµ¶EHLQJEDGIRU corals.

9DULDEOHVOLNHYLVLELOLW\ZHUHHVWLPDWHGGXULQJWKHGLYHKRZHYHUDUHEHVWTXDQWL¿HGXVLQJFRQWLQXRXV data recorders.

3.2 Analysis

Analysis proceeded through the following broad steps, for each dataset collected:

1) Calculation and plotting of basic distributions for each variable, across all study sites. These are GRQH¿UVWWRLOOXVWUDWHWKHEDVLFSDWWHUQVVKRZQE\LQGLYLGXDOYDULDEOHVDQGLQGLFDWRUV

2) Multi-dimensional Scaling (MDS) analysis helps to reveal patterns in datasets that include PXOWLSOH YDULDEOHV DQG SDUWLFXODUO\ XVHIXOQHVV ZKHUH SDUDPHWULF WHVWV HJ$129$  DUH QRW appropriate.

30 Detailed Methodology

%\SURMHFWLQJDOOYDULDEOHVRQWR[DQG\D[HVDQ0'6SORWKHOSVLOOXVWUDWHZKLFKSRLQWVDUHFORVHWR one another and which are distant. Thus the physical distance of points on the plot (Graph 3.2.1) illustrates their relative distance in the dataset. By superimposing a variable in the dataset on the SRLQWVZKHUHWKHVL]HRIDFLUFOHUHSUHVHQWVWKHPDJQLWXGHRIWKHYDULDEOHµEXEEOHSORWV¶ *UDSK 3.2.2) can help to illustrate which variables are most important in determining the relatedness among SRLQWVRQWKHSORW7KHFLUFOHVDURXQGFOXVWHUVRISRLQWVLOOXVWUDWHVLJQL¿FDQWJURXSLQJVRIVLWHVDQG help interpretation of the results.

Graph 3.2.1

Graph 3.2.2

31 Detailed Results

4. Detailed Results

Due to the complex datasets in this study, results and discussion will be presented together in QXPEHUHG VHFWLRQV IRU HDFK GDWDVHW ZLWK PRUH V\QWKHWLF GLVFXVVLRQ DQG ¿QGLQJV SUHVHQWHG LQ section 3.

4.1 Resilence indicators

Table 9 : Each factor was scaled from 1 (poor conditions for corals) to 5 (good conditions for corals), and the sites ranked from highest overall resilience to the lowest.

Group Explanation Factor Explanation Cover Benthic cover Benthic Benthic cover – combined estimates of hard and soft corals, and algae Coral Condition of coral Current Current status shown by bleaching, disease, community sexual recruitment and fragmentation of corals. Historic Past impacts to coral community as shown by evidence of past mortality, evidence of recover potential and size class distributions Ecological Broader ecological Negative Negative associates of corals – such as factors that affect predators and epiphytes on coral surfaces corals Positive Positive associates of corals, such as obligate IHHGHUV EXWWHUÀ\¿VK DQGLQYHUWHEUDWHVDQG¿VK in branching corals. Herbs +HUELYRURXV¿VKSRSXODWLRQV Physical Environmental and Acclimatization Past and present temperature dynamics that habitat features may protect corals by acclimatization/adaptive that affect corals responses Cooling & 'HJUHHRIFRROLQJÀXVKLQJRIGHHSHUDQGRU ÀXVKLQJ oceanic waters Shading & Degree of shading or screening of corals by screening turbid water, reef slope, canopy corals, etc. Substrate Substrate quality, such as sediment type and thickness, amount of rubble. Anthropogenic Human pressures )LVKLQJ 'HJUHHRI¿VKLQJVKRZQE\¿VKSRSXODWLRQV on reef sites and/or other data Substrate Anthropogenic alterations to substrate – from sediment, damage, etc. Water Anthropogenic alterations to water quality – from runoff, pollution, etc.

32 Detailed Results

70 68 66  62 60 58 56  52 50     

Cliff Bari )XQFKL .DUSDWD Salt City MarResN Carl's Hill Mi Dushi South Bay Vista Blue Wayaka IIAngel City.HHSVDNH 18th Palm Tori's Reef Chachacha Margate Bay3OD\D)UDQV 2LO6OLFN/HDS

Bachelor's Beach Something Special Graph 4.1.1

Graph 4.1.2

7KHJUDSKVKRZVWKHDJJUHJDWHVFRUHVRIUHVLOLHQFHLQGLFDWRUVIRUDOOVLWHVDQGWKHJUDSK shows sites clustered together according to their relative resilience scores. It is clear that there are 3 GH¿QHGFOXVWHUHGDFFRUGLQJWRWKHVLWHV¶UHODWLYHUHVLOLHQFH7KHµKLJK¶UHVLOLHQFHJURXSLQFOXGHV0DUJDWH %D\3OD\D)UDQV6RXWK%D\.DUSDWD9LVWD%OXHDQG0DULQH5HVHUYH1RUWK7KHµPLGGOH¶UHVLOLHQFHJURXS LQFOXGHV:D\DND,,$QJHO&LW\.HHSVDNH3OD\D)XQFKL&DUO¶V+LOO0L'XVKL&OLII%DUL(LJKWHHQWK3DOP 6DOW&LW\7RUL¶V5HHI2LO6OLFN/HDSDQG6RPHWKLQJ6SHFLDO7KHµORZ¶UHVLOLHQFHJURXSLQFOXGHVRQO\RQH site, Chachacha.

33 Detailed Results

Graph 4.1.3

7KHSORWDERYHVKRZVWKHVSUHDGRIVLWHVDFFRUGLQJWRWKHLUUHVLOLHQFHVFRUHVDQGSURWHFWLRQVWDWXV The vectors show resilience factors that are driving the differences in resilience. Sites to the left of the plot have lower resilience scores, sites to the right have higher resilience scores. It is clear to see that )LVK3URWHFWHG$UHDVIDOOWRWKHOHIWRIWKHSORWDQGKDYHORZHUUHVLOLHQFHVFRUHV7KLVPD\EHEHFDXVH WKH )3$V DUH ORFDWHG FORVH WR WKH XUEDQ FHQWUH .UDOHQGLMN DQG WKXV PRUH YXOQHUDEOH WR DGGLWLRQDO DQWKURSRJHQLF VWUHVVRUV )3$V DUH QHJDWLYHO\ FRUUHODWHG ZLWK DOO UHVLOLHQFH IDFWRUV H[FHSW IRU µ)LVK *URXSV¶LQGLFDWLQJWKDWDWOHDVW¿VKSRSXODWLRQVLQ)3$VDUHPRYLQJLQWKHULJKWGLUHFWLRQ&RQYHUVHO\1R Diving Areas (NDAs) have the highest resilience scores and correlations with resilience factors. NDAs DUHGHVLJQDWHGWRWKHQRUWKRIWKHLVODQGIDUIURP.UDOHQGLMNDQGDQWKURSRJHQLFVWUHVVRUV5HPRYLQJWKH additional stress from divers seems to be working well, and these sites appear to be among the most resilient to stress.

Graph 4.1.4

Graph 4.1.4

7KHEXEEOHSORWDERYH  VKRZVWKHUHODWLYHHIIHFWRIDOJDOSRSXODWLRQVRQWKHRYHUDOOHFRORJLFDO resilience of sites. Large bubbles mean that algal populations are favorable for resilience, while small bubbles mean they are unfavorable. It can be observed that the sites with most unfavorable algal populations for include Wayaka II, Something Special, Bari and Chachacha.

34 Detailed Results

Graph 4.1.5

7KHEXEEOHSORWVKRZVWKHUHODWLYHHIIHFWRIFRUDOSRSXODWLRQVRQWKHRYHUDOOHFRORJLFDOUHVLOLHQFH of sites. Large bubbles mean that coral populations are favourable for resilience, while small bubbles mean they are unfavourable. It can be observed that Wayaka II and Chachacha have the unhealthiest, OHDVWUHVLOLHQWFRUDOSRSXODWLRQVZKLOH0DULQH5HVHUYH1RUWKDQG3OD\D)UDQVKDYHWKHKHDOWKLHVWPRVW resilient populations.

Graph 4.1.6

7KHEXEEOHSORWVKRZVWKHUHODWLYHHIIHFWRI¿VKSRSXODWLRQVRQWKHRYHUDOOHFRORJLFDOUHVLOLHQFHRI VLWHV/DUJHEXEEOHVPHDQWKDW¿VKSRSXODWLRQVDUHIDYRXUDEOHIRUUHVLOLHQFHZKLOHVPDOOEXEEOHVPHDQ WKH\DUHXQIDYRXUDEOH,WFDQEHREVHUYHGWKDW¿VKSRSXODWLRQVDUHPRVWIDYRXUDEOHDW0DUJDWH%D\DQG :D\DND,,ZKLOHWKH\DUHOHDVWIDYRXUDEOHDW.HHSVDNH

35 Detailed Results

Graph 4.1.7

7KHEXEEOHSORWVKRZVWKHUHODWLYHHIIHFWRIVXEVWUDWHFRQGLWLRQVRQWKHRYHUDOOHFRORJLFDOUHVLOLHQFH of sites. Large bubbles mean that substrate conditions are favourable for resilience, while small bubbles mean they are unfavourable. It can be observed that Chachacha and Cliff have the most unfavourable substrate conditions, while sites to the right of the plot have the most favourable conditions.

Graph 4.1.8

7KH EXEEOH SORW  VKRZV WKH UHODWLYH HIIHFW RI ZDWHU TXDOLW\ RQ WKH RYHUDOO HFRORJLFDO UHVLOLHQFH RI sites. Large bubbles mean that water quality is favourable for resilience, while small bubbles mean that LWLVXQIDYRXUDEOH,WFDQEHREVHUYHGWKDWVLWHVFORVHWRWKHXUEDQFHQWUH.UDOHQGLMNVXFKDV&KDFKDFKD Something Special, Cliff or Bachelor’s Beach have the least favourable water quality, while sites further DZD\IURP.UDOHQGLMNKDYHPRUHIDYRXUDEOHZDWHUTXDOLW\FRQGLWLRQV

36 Detailed Results

4.2 Benthic cover

70 Benthic cover all sites 60 50  30 20 10 0

Sponges Tunicate 2WKHUOLYH =RDQWKLGV Hard CoralGorgoniansMacroalgae Sand, rubble Coralline algae Diseased corals

Dead coral with algae Graph 4.2.1

Dead coral overgrown by turf algae dominates the substrate, with an average cover of 38.2% (Graph  7KLVPD\UHÀHFWFRUDOPRUWDOLW\FDXVHGE\GDPVHO¿VKµ\DUGV¶RQODUJHFRUDOKHDGV0DFURDOJDH FRYHULVORZDYHUDJLQJLQGLFDWLQJJRRGKHUELYRU\OHYHOV+DUGFRUDOFRYHUDYHUDJHVZKLFKLV high for the Caribbean region, however the high incidence of dead coral overgrown by algae is worrying.

 Macroalgae cover 35 30 25 20 15 10 5 0 Cliff Bari )XQFKL .DUSDWD Salt City Margate Mi DushiMi Carl's Hill Carl's MarResN Wayaka II 18th Palm .HHSVDNH Angel City Vista Blue South Bay Tori's Reef Tori's Chachacha 3OD\D)UDQV 2LO6OLFN/HDS Bachelor Beach Something Special Graph 4.2.2

Live coral cover 70 60 50  30 20 10 0 Cliff Bari )XQFKL .DUSDWD Salt City Margate Mi DushiMi Carl's Hill Carl's MarResN Wayaka II 18th Palm .HHSVDNH Angel City Vista Blue South Bay Tori's Reef Tori's Chachacha 3OD\D)UDQV 2LO6OLFN/HDS Bachelor Beach Something Special

Graph 4.2.3

37 Detailed Results

Graph 4.2.4

Graph 4.2.5

7KHEDUJUDSKVKRZVVKRZWKDWKDUGFRUDOFRYHUUDQJHVIURPLQ3OD\D)UDQVWRLQ %DUL7KHEDUJUDSKVKRZVWKDWPDFURDOJDHFRYHUUDQJHVIURPLQ%DULWRLQ6DOW&LW\ Macroalgae cover is low (under 10%) in every site except for Bari which exhibits 5 times the average macroalgal cover and appears to be suffering from macroalgal overgrowth and low coral cover (8.2% - the lowest of the sites surveyed). The highest hard coral cover sites are situated either on the northern coast RI%RQDLUH 3OD\D)UDQV.DUSDWD0DULQH5HVHUYH1RUWK RU.OHLQ%RQDLUH 6RXWK%D\.HHSVDNH 7KHVH VLWHVDOVRDOOH[KLELWDPRQJWKHORZHVWPDFURDOJDHFRYHU EHORZ 7KHEXEEOHSORWV *UDSKVDQG  SURYLGHDQLFHYLVXDOUHSUHVHQWDWLRQRIWKHEDODQFHEHWZHHQKDUGFRUDODQGPDFURDOJDHFRYHU

38 Detailed Results

90 De ad cor al cove r 80 70 60 50  30 20 10 0 Cliff Bari )XQFKL .DUSDWD Salt City Margate Mi DushiMi Carl's Hill Carl's MarResN Wayaka II 18th Palm .HHSVDNH Angel City Vista Blue South Bay Tori's Reef Tori's Chachacha 3OD\D)UDQV 2LO6OLFN/HDS Bachelor Beach Something Special

Graph 4.2.6

 Gorgonian cove r 35 30 25 20 15 10 5 0 Cliff Bari )XQFKL .DUSDWD Salt City Margate Mi DushiMi Carl's Hill Carl's MarResN Wayaka II 18th Palm .HHSVDNH Angel City Vista Blue South Bay Tori's Reef Tori's Chachacha 3OD\D)UDQV 2LO6OLFN/HDS Bachelor Beach Something Special

Graph 4.2.7

7KHJUDSKVKRZVWKHFRYHURIGHDGFRUDOFRYHUE\VLWH'HDGFRUDOFRYHUUDQJHGIURPLQ :D\DND,,WRLQ%DUL0RVWVLWHVH[KLELWHGKLJKFRYHURIGHDGFRUDORYHUJURZQE\WXUIDOJDHDQGRQO\ 2 sites exhibited less than 20% cover of dead coral (Cliff and Bari) while half the sites exhibited dead coral FRYHURIRUDERYH7KLVKLJKFRUDOPRUWDOLW\LVVXVSHFWHGWRUHÀHFWLQFUHDVLQJGDPVHO¿VKSRSXODWLRQV VWRUPGDPDJHIURP+XUULFDQH2PDUDQGGHFOLQLQJZDWHUTXDOLW\

7KHJUDSKVKRZVJRUJRQLDQFRYHUE\VLWH*RUJRQLDQFRYHUYDULHVIURP 0DUJDWH%D\ WR  :D\DND ,, DQG 3OD\D )UDQV  *RUJRQLDQ FRYHU LV KLJKHVW LQ VRXWKHUQ VLWHV 0DUJDWH %D\ 9LVWD %OXH DQGORZHVWLQQRUWKHUQVLWHV :D\DND,,DQG3OD\D)UDQV LQGLFDWLQJDEURDGQRUWKVRXWKJUDGLHQWRI decline in gorgonian cover.

39 Detailed Results

Graph 4.2.8

Multi-Dimensional Scaling analysis shows Bari as an outlier due to its high macroalgae cover (Graph  ,WDOVRVKRZVKRZWKDWGLIIHUHQWO\PDQDJHGVLWHVDUHFKDUDFWHUL]HGE\GLIIHUHQWVXEVWUDWHFRYHUV )LVK 3URWHFWHG$UHDV DUH DVVLJQHG FORVH WR WKH XUEDQ FHQWUH .UDOHQGLMN ZKHUH WKH FRUDO UHHI LV PRVW impacted by anthropogenic stressors, and are thus characterized by higher macroalgae and rubble cover. No Diving Areas tend to have the highest coral cover, and also the highest cover of dead coral overgrown by turf algae.

Rate of calcification

0.9 0.8 0.7 0.6 0.5  0.3 0.2 0.1 0 Cliff Bari .DUSDWD Salt City Mi Dushi Carl’s Hill Carl’s Wayaka II Wayaka .HHSVDNH Vista Blue Vista 18th Palm Angel City South Bay Tori’s Reef Tori’s 3OD\D)UDQV 3OD\D)XQFKL Cha 2LO6OLFN/HDS Bachelor’s Beach Something Special Marine Reserve North Graph 4.2.9

7KHJUDSKVKRZVWKHFDOFL¿FDWLRQUDWHVFDOFXODWHGIRUGLIIHUHQWUHHIV7KHFDOFL¿FDWLRQUDWHUHSUHVHQWV WKHUDWLRRIFDOFLI\LQJYHUVXVQRQFDOFLI\LQJVXEVWUDWHVRQWKHUHHI$FDOFL¿FDWLRQUDWHKLJKHUWKHQZRXOG

40 Detailed Results

indicate a growing reef, with more calcifying organisms (live coral) than non-calcifyers (other substrate W\SHV ZKLOHDFDOFL¿FDWLRQUDWHRIOHVVWKDQZRXOGLQGLFDWHDUHHIWKDWLVQRWJURZLQJDWDQRSWLPDOOHYHO GXH WR WKH GRPLQDQFH RI QRQFDOFLI\HUV RYHU FDOFLI\HUV 3OD\D )UDQV DQG 6RXWK %D\ KDYH WKH KLJKHVW FDOFL¿FDWLRQUDWHV DQGUHVSHFWLYHO\ ZKLOH:D\DND,,DQG&KDFKDFKDDQG:D\DND,,KDYHWKH ORZHVWFDOFL¿FDWLRQUDWHV DQGUHVSHFWLYHO\ 

4.3 Coral population structure

Number 6 All corals Area 300 5

 200 3

2 Area(m2) 100 Numbercolonies of 1

0 -

6-10' 0-2.5' 2.6-5' 11-20'   >320 81-160'161-320 Graph 4.3.1

The distribution of size classes is shown by number of colonies, and by area of colonies for all size classes *UDSK 2QDYHUDJHWKHUHZHUHFRORQLHVLQDQDUHDRIP2, corresponding to 15.6 m2 of coral colony surface.

7KHGRPLQDQWVL]HFODVVE\DUHDZDVFP7KHORZFRQWULEXWLRQRIFRORQLHVODUJHUWKDQPLQGLFDWHV that colonies are not reaching extreme old age due to some stress, or the low number of large colonies could indicate a mass mortality event in the past and that the reef is in a recovery phase.

The low number of coral recruits (colonies 0-2.5 cm in size) is indicative of low coral reproduction and UHFUXLWPHQWIRUWKLVFRKRUWDQGFRXOGEHOLQNHGWRDUHFHQWVWUHVVDQGPRUWDOLW\HYHQW HJ+XUULFDQH2PDU in 2008).

41 Detailed Results

6,000 Numcols Area (m2) 35 5,000 30 25  20 3,000 15 2,000 10

1,000 5 0 0 Cliff Bari )XQFKL .DUSDWD Salt City Margate Mi DushiMi Carl's Hill Carl's MarResN Wayaka II 18th Palm .HHSVDNH Angel City Vista Blue South Bay Tori's Reef Tori's Chachacha 3OD\D)UDQV 2LO6OLFN/HDS Bachelor Beach Something Special Graph 4.3.2

 Numcols Area (m2) 35

30

25

20

15

10

5

0 Dip cli Dip Ste int Dip str Dip Sid sid lab Dip Mil sppMil nat Col Por ast Por por Myc sp Acr pal Den cyl Den Eus fas Acr cer Mad mir Mon fra Mon fav Aga lam Mon ann Aga aga Mon cav Mad dec Mea mea Graph 4.3.3

7KHWDEOHVKRZVWKHWRWDOQXPEHURIFRORQLHVDQGWKHWRWDOFRUDOFRYHUDUHDSHUVLWH7KHKLJKHVW FRUDOFRYHUDQGQXPEHURIFRORQLHVDUHLQQRUWKHUQVLWHV 3OD\D)UDQV0DULQH5HVHUYH1RUWK.DUSDWD  .OHLQ%RQDLUHVLWHV 6RXWK%D\DQG.HHSVDNH DQGDFRXSOHRIVRXWKHUQVLWHV $QJHO&LW\DQG0DUJDWH Bay). Sites are ordered from north to south and it is clear to see that sites closest to the urban centre .UDOHQGLMN LQGLFDWHGZLWKDWKLFNKRUL]RQWDOEDU DQGWKXVLPSDFWIURPFRDVWDOGHYHORSPHQWKDYHWKH lowest coral covers (Bari, Something Special, Chachacha, 18th 3DOP %DFKHORU¶V %HDFK  .HHSVDNH exhibits the highest number of coral colonies despite not having the largest area, indicating a dominance of smaller class sizes.

7KH ¿JXUH  VKRZV WKH WRWDO QXPEHU RI FRORQLHV DQG WRWDO FRYHU DUHD SHU FRUDO VSHFLHV$UHD LV dominated by the slow-growing massive Montastrea annularis and Montastrea faveolata. The number of colonies is dominated by Agaricites agaricites. No Acropora cervicornis or Acropora palmata colonies ZHUHUHFRUGHGLQGLFDWLQJWKHKLJKLPSDFWRI+XUULFDQH2PDURQWKHVHIUDJLOHEUDQFKLQJVSHFLHV

42 Detailed Results

600

500 '0-2.5



300

200

Density (per 100 m2) (per 100 Density 100

0 Cliff Bari )XQFKL .DUSDWD Salt City Margate Mi DushiMi Carl's Hill Carl's MarResN Wayaka II 18th Palm .HHSVDNH Angel City Vista Blue South Bay Tori's Reef Tori's Chachacha 3OD\D)UDQV 2LO6OLFN/HDS Bachelor Beach

Something Special Site Graph 4.3.4

70

60 0-2.5

50



30

20 Density (per 100 m2) (per 100 Density 10

0 Dip cli Dip Ste int Dip str Dip Sid sid lab Dip Mil sppMil Col nat Col Por ast Por por Acr pal Den cyl Den Lob nat Mad mir Eus fas Acr cer Mon fra Aga lam Mon fav Aga aga Mon ann Mad dec Mon cav Myc spp Mea mea Site Graph 4.3.5

7KHWDEOHVKRZVWRWDOGHQVLW\RIFRUDOUHFUXLWV FRORQLHVVL]HGFP SHUVLWH.HHSVDNHVWDQGV RXWDVKDYLQJWKHKLJKHVWGHQVLW\RIFRUDOUHFUXLWV UHFUXLWVSHUP ZKLOH2LO6OLFN/HDSDQG Mi Dushi have the lowest recruitment rates (50 recruits per 100 m2). In general, there is a gradient of recruitment from north to south, with lower recruitment in northern sites and higher recruitment in southern sites. This is because there is a higher abundance of brooding species (corals that reproduce with internal fertilization, releasing only sperm which is negatively buoyant and can harbor unfertilized eggs for weeks) LQWKHVRXWK.HHSVDNHLVORFDWHGRQWKHHDVWHUQVLGHRI.OHLQ%RQDLUHLQWKHFKDQQHOEHWZHHQ%RQDLUH DQG.OHLQ%RQDLUHDQGWKLVORFDWLRQFRXOGSURPRWHKLJKUHFUXLWPHQWGXHWRHGGLHVWKDWFRXOGRFFXULQWKH channel leading to higher retention time and recruitment success.

7KHWDEOHVKRZVWRWDOGHQVLW\RIUHFUXLWPHQWSHUFRUDOVSHFLHV5HFUXLWPHQWLVODUJHO\GRPLQDWHGE\ Agaricites agaricites, followed by Porites asteroids.

43 Detailed Results

Graph 4.3.6

7KLVEXEEOHSORW  WRWKHOHIWVKRZVWKHGHQVLW\RIVXVFHSWLEOHFRUDOVSHFLHV IRUWKHSXUSRVHRIWKLV survey as follows, Acropora palmata, Acropora cervicornis, Eusmilia fastigata, Meandrina meandrites, Montastrea annularis, Montastrae faveolata, Montastrea franksii  LQ GLIIHUHQW VLWHV 7KH PRUH µSULVWLQH¶ VLWHVWKDWKDYHKLJKHUFRYHURIVXVFHSWLEOHVSHFLHVDUH3OD\D)UDQV0DULQH5HVHUYH1RUWK6RXWK%D\ and Margate Bay, all sites away from impact from urban and coastal development. The high impact site Chachacha is clearly an outlier due to its low coral cover.

4.4 Algae Populations

All algae 60 50 

% 30 20 10 0 Turf NCC Jania Padina Valonia Dictyota Caulerpa Halimeda Amphiroa Corallines Wrangelia Lobophora Chlorophyta Anadyomene Cyanobacteria Dictyosphaeria Graph 4.4.1

7KHJUDSKDERYH  VKRZVWKDWDOJDOSRSXODWLRQVLQ%RQDLUH¶VUHHIVDUHGRPLQDWHGE\WXUIDOJDHZKLFK FRYHUVRIWKHVXEVWUDWH7KLVLVODUJHO\GXHWRFRUDOPRUWDOLW\DQGGDPVHO¿VKFUHDWLQJWXUIDOJDH µ\DUGV¶RQFRUDOKHDGV7KHPRVWFRPPRQPDFURDOJHLVDictyota, a brown algae covering 8.0% of the substrate. Coralline algae cover is low at 3.6%.

44 Detailed Results

Trididenmnum 5   3.5 3 2.5 2 % cover % 1.5 1 0.5 0 Cliff Bari Tori's .DUSDWD Salt City Mi DushiMi Carl's Hill Carl's Wayaka II 18th Palm .HHSVDNH Angel City Vista Blue South Bay 3OD\D)UDQV ChaChaCha 3OD\D)XQFKL Margate Bay 2LO6OLFN/HDS Marine reserve Bachelor Beach Something Special Graph 4.4.2

Graph 4.4.3

It is worth noting the cover of Lobophora (3.0%) a brown algae, and Trididemnum (2.8%) a tunicate that competes with corals, which although small at this sampling depth (10m) have been noted to be LQFUHDVLQJHVSHFLDOO\LQGHHSHUVHFWLRQVRIWKHUHHI7KHWDEOHVKRZVWKHFRYHURI7ULGLGHPQXPD WXQLFDWHWKDWFRPSHWHVZLWKRYHUJURZVKDUGFRUDO7KHEXEEOHSORWSURYLGHVDYLVXDOUHSUHVHQWDWLRQ of relative Trididemnum cover. It is clear that Trididemnum is affecting only northern sites at this sampling GHSWKZLWKKLJKHVWFRYHUVLQ0DULQH5HVHUYH1RUWK  DQG:D\DND,,  DQGQR7ULGLGHPQXP SUHVHQWVRXWKRI.DUSDWD7KHUHLVDQHFGRWDOHYLGHQFHWKDW7ULGLGHPQXPFRYHUVLQFUHDVHZLWKGHSWKDQG are smothering corals.

45 Detailed Results

Lobophora 7 6 5  3 % cover % 2 1 0 Cliff Bari Tori's .DUSDWD Salt City Mi DushiMi Carl's Hill Carl's Wayaka II 18th Palm .HHSVDNH Angel City Vista Blue South Bay 3OD\D)UDQV ChaChaCha 3OD\D)XQFKL Margate Bay 2LO6OLFN/HDS Marine reserve Bachelor Beach Something Special Graph 4.4.4

Graph 4.4.5

7KHJUDSKVKRZV/RERSKRUDFRYHUDEURZQPDFURDOJDHZKLFKFRPSHWHVZLWKDQGRYHUJURZVFRUDOV 7KHEXEEOHSORWVKRZVUHODWLYH/RERSKRUDFRYHUVEHWZHHQVLWHV7KHVLWHZLWKWKHKLJKHVW/RERSKRUD FRYHULV.HHSVDNH  RQ.OHLQ%RQDLUHIROORZHGE\2LO6OLFN/HDS  DQG&DUO¶V+LOO  7KHUH ZDVQR/RERSKRUDIRXQGDWVLWHVFORVHWR.UDOHQGLMNVXFKDV%DUL6RPHWKLQJ6SHFLDODQG&KDFKDFKDDQG there was also very low cover at 18th Palm (1.0%).

46 Detailed Results

4.5 Fish community structure

7KHJUDSK EHORZ VKRZVKHUELYRURXVDQGSUHGDWRU\¿VKELRPDVVIRUDOOVLWHVVXUYH\HG7KHEXEEOH SORWRQWKHERWWRPOHIW  VKRZVUHODWLYHELRPDVVRIKHUELYRUHVSHUVLWHZKLOHWKHEXEEOHSORWRQWKH ERWWRPULJKW  VKRZVUHODWLYHELRPDVVRISUHGDWRUVSHUVLWH%LRPDVVRIKHUELYRUHVLVKLJKHUWKDQ predators in most sites except for a few exceptions, notably Margate Bay and Vista Blue where large VFKRROVRIJUXQWVFDXVHGKLJKQXPEHUVIRUSUHGDWRUELRPDVV2YHUDOODYHUDJHELRPDVVIRUKHUELYRUHV was 7319 g/m2 compared to 5290 g/m2 for predators. Sites with highest herbivore biomass were Bari 5HHI DQ)3$ 6RXWK%D\0L'XVKLDQG6DOW&LW\0DUJDWH%D\DQG9LVWD%OXHKDGWKHKLJKHVWSUHGDWRU biomass. The horizontal dotted lines represent average values.

30000

Herbivores 25000 Predators )

2 20000

15000

Biomass (g/100m 10000 7319 5290 5000

0 07. Cliff All Sites .DUSDWD 2LO6OLFN 6DOW&LW\ 19. Mi Dushi 19. Mi 08. Bari Reef 18. Carl's Hill 02. Wakaya II .HHSVDNH 17. Vista Blue 13. Angel City 20. South Bay 15. Tori's Reef 3OD\D)UDQV 16. Margate Bay 3OD\D)XQFKL 1RUWK5HVHUYH 10. Cha Cha Beach 11. Eighteenth Palm 12. Bachelor's Beach 09. Something Special Graph 4.5.1

Graph 4.5.2 Graph 4.5.3

47 Detailed Results

2WKHU&DUQLYRUHV 22000 Jacks 20000 Groupers Grunts 18000 Snappers 16000 ) 2 

12000

10000

8000 Biomass (g/100m 6000



2000

0 07. Cliff .DUSDWD 2LO6OLFN 6DOW&LW\ 19. Mi Dushi 19. Mi 08. Bari Reef 18. Carl's Hill 02. Wakaya II .HHSVDNH 17. Vista Blue 13. Angel City 20. South Bay 15. Tori's Reef 3OD\D)UDQV 16. Margate Bay 3OD\D)XQFKL 1RUWK5HVHUYH 10. Cha Cha Beach 11. Eighteenth Palm 12. Bachelor's Beach 09. Something Special

Graph 4.5.4

Denuding (Acanthurids, Microspathadon) 12000 Non-denuding (Territorial Damsels) 11000 Excavating (Scarids)

10000

9000

8000 ) 2 7000

6000

5000

Biomass (g/100m 

3000

2000

1000

0 07. Cliff .DUSDWD 2LO6OLFN 6DOW&LW\ 19. Mi Dushi 19. Mi 08. Bari Reef 18. Carl's Hill 02. Wakaya II .HHSVDNH 17. Vista Blue 13. Angel City 20. South Bay 15. Tori's Reef 3OD\D)UDQV 16. Margate Bay 3OD\D)XQFKL 1RUWK5HVHUYH 10. Cha Cha Beach 11. Eighteenth Palm 12. Bachelor's Beach 09. Something Special Graph 4.5.5

7KHJUDSKRQWKHDERYHOHIWVKRZVWKHELRPDVVRIGLIIHUHQWSUHGDWRU\¿VKW\SHVE\VLWH7KHJUDSKRQ WKHDERYHULJKWVKRZVWKHELRPDVVRIGLIIHUHQWKHUELYRURXV¿VKIXQFWLRQDOJURXSVE\VLWH7KHELRPDVVRI predators in most sites is dominated by snappers and grunts, and large populations of these predators give Margate Bay (over 21,000 g/100m2) and Vista Blue (over 15,000 g/100m2) in the south the largest SUHGDWRUELRPDVVRIDOOVLWHV6LWHVZLWKWKHORZHVWSUHGDWRUELRPDVVDUH2LO6OLFN/HDS0DULQH5HVHUYH North and Mi Dushi. Sites with the highest grouper biomass are South Bay, Margate Bay, Bachelor’s %HDFK DQG 3OD\D )UDQV7KH ODUJH PDMRULW\ RI KHUELYRURXV ELRPDVV LV PDGH XS RI H[FDYDWLQJ VFDULGV SDUURW¿VK %DUL5HHI6RXWK%D\0L'XVKLDQG6DOW&LW\KDYHWKHKLJKHVWVFDULGELRPDVVDOORYHU JUDPVSHUP+RZHYHU 2LO6OLFN /HDS  JP  DQG6RXWK%D\  JP  KDYH WKH KLJKHVWGDPVHO¿VKELRPDVV

48 Detailed Results

Graph 4.5.6

Graph 4.5.7 (from Vermeij et al in press) Graph 4.5.8 (from Vermeij et al in press)

7KHEXEEOHSORWVKRZVUHODWLYHDEXQGDQFHRIGDPVHO¿VKE\VLWH2LO6OLFN/HDS.HHSVDNH0DULQH 5HVHUYH1RUWKDQG6RXWK%D\VKRZWKHKLJKHVWUHODWLYHDEXQGDQFHRIQRQGHQXGLQJGDPVHO¿VKZKLOH :D\DND,,DQG3OD\D)XQFKLH[KLELWWKHORZHVWUHODWLYHDEXQGDQFHV

7KHJUDSKVKRZVWKHUHODWLRQVKLSEHWZHHQDYHUDJHELRPDVVRIGDPVHO¿VKDQGDYHUDJHELRPDVVRI SUHGDWRUV7KHUHLVDFOHDUQHJDWLYHFRUUHODWLRQEHWZHHQWKHWZRVXJJHVWLQJWKDWGDPVHO¿VK 6WHJDVWHV SODQLIURQV  ELRPDVV LV FRQWUROOHG E\ GLUHFW FRQVXPSWLRQ E\ SUHGDWRUV 7KH JUDSK  VKRZV WKH relationship between S. Planifrons and their preferred habitat (Montastrea coral species). There is no clear correlation between the two, again suggesting that it is the predator biomass which is controlling the GDPVHO¿VKSRSXODWLRQ

49 Detailed Results

Graph 4.5.9

Graph 4.5.10

Graph 4.5.11

7KH EXEEOH SORWV DERYH   DQG   VKRZ WKH UHODWLYH ELRPDVV RI GLIIHUHQW KHUELYRURXV functional groups by site. To the far left are excavating scarids, in the middle are browsers (Chaetodons, Pomacanthids) and to the far right are denuding families (Acanthurids, Microspathadon). Most sites had relatively abundant excavating scarid populations, but sites with relatively smaller scarid biomass SRSXODWLRQVDUHVHHQWRWKHULJKWRIWKHSORW 3OD\D)XQFKL0DUJDWH%D\6RPHWKLQJ6SHFLDODQG(LJKWHHQWK 3DOP 7KHVLWHVZLWKWKHKLJKHVWUHODWLYHELRPDVVRIEURZVHUVDUH0DUJDWH%D\DQG2LO6OLFN/HDS0DQ\ sites had high relative biomass of denuding species, but sites with lower biomass can be seen to the top RIWKHSORWLQFOXGLQJ%DUL2LO6OLFN/HDS.HHSVDNH&DUO¶V+LOO9LVWD%OXHDQG6RPHWKLQJ6SHFLDO

50 References

References

2EXUD'DQG*ULPVGLWFK*  5HVLOLHQFHDVVHVVPHQWRIFRUDOUHHIV,8&1SS

Steneck, R, Mumby, P and Arnold, S (2007) A report on the status of the coral reefs of Bonaire in 2007 with UHVXOWVIURPPRQLWRULQJ8QLYHUVLW\RI0DLQHSS

STINAPA (2006) Bonaire National Marine Park Management Plan. STINAPA, 107pp.

STINAPA (2008) Construction handbook Bonaire. 29pp.

9HUPHLM0GH%H\+6DQGLQ6*ULPVGLWFK*DQGGH/HRQ5 LQSUHSDUDWLRQ 3UHGDWRUDEXQGDQFH determines the relative importance of top down and bottom up processes in controlling the abundance of an unwanted habitat engineer on Caribbean reefs.

Wilkinson, C (2002) Status of the coral reefs of the world 2002. AIMS.

51 References

INTERNATIONAL UNION FOR CONSERVATION OF NATURE

:25/'+($'48$57(56 Global Marine and Polar Programme Rue Mauverney 28 1196 Gland Switzerland

7HO )D[ [email protected] www.iucn.org/marine

52