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The Great Barrier , Environment and Management Second Edition

Pat Hutchings, Michael Kingsford, Ove Hoegh-Guldberg (Eds)

Describes the , and other of the reef, and the biological, chemical and physical processes that influence them.

The iconic and beautiful Park is home to one of the most diverse in the world. With contributions from international experts, this timely and fully updated second edition of The Great Barrier Reef describes the animals, plants and other organisms of the reef, as well as the biological, chemical and physical processes that influence them. It contains new chapters on shelf slopes and and addresses pressing issues such as change, acidification, bleaching and disease, and invasive species. The Great Barrier Reef is a must-read for the interested reef tourist, student, researcher and environmental manager. While it has an Australian focus, it can equally be used as a reference text for most Indo-Pacific coral reefs. February 2019 488pp, Paperback ISBN: 9781486308194 FEATURES $125.00 • Up-to-date and accessible account of the issues that face the world’s most complex publish.csiro.au/book/7763 marine • Examines the diversity, , and conservation of the reef, as well as its role in monitoring global warming Available in and New Zealand through • Written by leading authorities on the of the Great Barrier Reef CSIRO Publishing. and highly illustrated throughout Elsewhere, please contact CRC Press.

2 The GreaT Barrier reef 1 – INTRODUCTION TO THE GREAT BARRIER REEF 3 26 The GreaT Barrier reef 3 – THE GREAT BARRIER REEF IN TIME AND SPACE: AND PALEOBIOLOGY 27

experts to contribute to this book. The ACRS (which Australian and repeated episodes of 365–452 kya. This is coincident with Marine Isotope repeatedly during high level episodes associated changes in several parameters of ’s orbit (so started out as the Great Barrier Reef Committee) is global environmental change associated with the Stage (MIS) 11, perhaps the warmest interglacial of with major environmental fluctuations in sea level, called Milankovitch cycles). These cycles influ-

the oldest society in the world and most late Tertiary and Pleistocene ice ages. The ‘reefal’ the past 450 000 years (ky), and one with climatic and dioxide (CO2) over the past ence the amount of from the sun received of the authors are members and associates of this GBR is a relatively young geological that conditions most similar to those we are now experi- several hundred thousand years. Moreover, these by the Earth. The cycles include obliquity (changes society. It was also intimately involved in the estab- was slow to respond to favourable environmental encing. Some workers believe that the ‘switching- reefs have maintained a similar coral and algal spe- in the angle of Earth’s axis of rotation with respect lishment of the world’s largest marine park, the conditions early on. In fact, the central on’ of the GBR was related to the mid-Pleistocene cies composition during their repeated formation to the sun), eccentricity (changes in the circularity GBR Marine Park in 1975. This huge park, with has enjoyed warm tropical transition from 41 ky to 100 ky-long climatic cycles, (see later section on Paleoecology) (Fig. 3.1B, C). of Earth’s orbit around the sun) and precession of some of the largest no-take areas in the global that could well have supported coral growth for the and to the development during MIS 11 of a marked the equinoxes (changes in the position of the ocean, was enlarged and rezoned in 2004 based on past 15 million years (My). However, the best evi- high stand that enabled sustenance of both a Earth in its orbit around the sun at the time of the our much increased scientific understanding of dence indicates that the initiation of the GBR did not corridor and a reef tract along a relatively wide and Reef growth and global sea level change equinox). The cycles are 41 000, 100 000 and 23 000 coral reefs and their challenges and . occur until around 600 thousand years ago (kya), deeper continental shelf (Fig. 3.1A). The growth and decay of ice sheets in the north- years, respectively. During the last 500 000 years, The book is divided into three sections. The first and the regional province of reef systems as we now Cores drilled through Ribbon Reef 5 have shown ern hemisphere were controlled by 104- to 105-year global sea level underwent at least 17 such cycles section focuses on the geomorphology, paleobiol- know them probably did not occur until around that the GBR has been able to re-establish itself scale climate changes forced by natural cyclic of rise and fall. Average rates of sea level change ogy and of the GBR. Here, various of the GBR are discussed, not only from (A) (A) (C) the point of view of coral-dominated ecosystems, but also with regard to the important associated inter-reefal areas (Chapters 2–9). These compo- nents of the GBR, along with catchments and off- deeper waters, are highly interconnected (Fig. 1.2). The second section of the book focuses on the major processes that are affecting the reef and includes the description of organisms and pro- cesses that contribute to photosynthetic activity Fig. 1.1. Quasi-true colour image from space showing the and , as well as the flow of Fig. 1.2. Crossing the Blue Highway. Designed and written by science communicator Russell Kelley and published by Great Barrier Reef along the continental shelf of north- energy and within coral reef ecosystems. eastern Australia from Cape York to Gladstone (~2000 km). the ACRS, the Blue Highway poster portrays the reefs of the Great Barrier Reef as part of a larger supporting system that A mosaic of reefs can be seen parallel to the mainland Other chapters deal with the major within includes the coastal catchments and connects numerous interacting ecosystems from ridge to reef. The poster illustrates how natural loads from runoff and ocean fuel a connected mosaic of ecosystems and the role inter- (B) (C) and extending hundreds of kilometres across the GBR. and around the reef, illustrating its inherent 18 The GreaT Barrier reef 2 – GEOMORPHOLOGY OF CORAL REEFS WITH SPECIAL REFERENCE TO THE GREAT BARRIER REEF 19 38 The GreaT Barrier reef 4 – OCEANOGRAPHY 39 (Source: Image generated from the Moderate Resolution dynamic . This section also reviews our cur- reef habitats play in supporting migrating species as they move from inshore nursery grounds to the outer reefs. The Lutjanus sebae Imaging Spectroradiometer (MODIS), data courtesy of rent understanding of how local challenges (i.e. model species of is (red emperor snapper) that near the shelf edge and recruit to as NASA/GSFC, image courtesy of Dr Scarla Weeks, Centre larvae. Juveniles move from recruitment to reefs and inter-reefal habitats, before they mature. Printed copies of (B) (D) declining water quality and over exploitation of for Marine Studies at The University of Queensland. Inset this poster are available from the GBRMPA (Artwork: G. Ryan). (ii) lagoonal reefs that form as the outer reef depth of Pleistocene foundations on the GBR was (A) (B) fisheries) as well as aspects of (e.g. shows a closer cross-shelf view of the reefs out from extends around the margins enclosing a not random, with shallow depths in the north and Box 4.1. Tropical boundary currents (D) Hinchinbrook, near ). ocean warming, acidification and intensifying through the unique and spectacular biodiversity of being severely degraded in the next few years if we that is slowly infilled by patch reef far south, a cross-shelf gradient in the southern- A warm, southward flowing tropical boundary storms) have changed the circumstances under coral reefs. By weaving the basic of these don’t understand the problems and apply evidence- growth and by transport of from central GBR, and generally depths greater than is also found on the west of Australia. This book is aimed at undergraduate and post- which coral reefs have otherwise prospered for groups together with fascinating details of their based solutions. the windward margins into the lagoon 20 m on the central GBR shelf. Reef types reflect This is the Leeuwin Current (LC) that is the graduate students, the informed public, as well as thousands if not tens of millions of years (Chapters , it is hoped that the interest of the reader will It has been 11 years since the first edition of this (iii) a final senile stage consisting of planar reefs, this variation with submerged reefs, patches and continuation of the ‘Indonesian Throughflow’ from researchers and managers who would like to famil- 10–14). be inspired to explore this incredible diversity. book was published (2008). With a new decade, the with the lagoon infilled and reef flat the Indonesian , and flows south down the crescentic reefs dominatingFig. 3.1. the(A) central GBR and iarise themselves with the complexity of coral reefs The third and final section of this book deals Throughout this book we come back to the on the GBR have increased and trends Plot for the past 1.5 million years showing the change in frequency and amplitude of the climatic, and by coast of . It turns east at Cape extending across the entire reef. Sediment and most planar reefs foundinference, -levelof fluctuations or south ofafter the mid-Pleistocene transition (MPT) at 0.9–0.6 Mya (MPT, a shift in the periodicity such as the GBR. The project arose out of an with the diversity of organisms that live in and such as increasing and ocean pH Leeuwin, where it becomes the South Australian major challenges that reefs face in our changing beds or may dominate of radiative forcing by atmospheric that caused higher amplitude climate periodicities). Growth of the Mackay. Understanding this sequence, its controls, Current, and flows eastwards below most of South advanced undergraduate course that has been held around coral reefs (Fig. 1.3). In this section, the world. For this reason, our book is unique in that it continue to warrant responses at a global and local GBR occurred after the MPT (Source: after Larcombe and Carter 2004). (B) Photographs from the Ribbon Reef 5 core the reef flat and coral cays are common on and the speed with which reefs may proceed from Australia to , where it is known as the on the GBR for almost 20 years and which gener- reader is introduced to broad categories of - reviews the past, current and future trajectories level; you will find examples of this in all the chap- showing the major coral components of ‘Assemblage A’. These include robust branching of species from the such reefs, including low wooded . juvenile to senile states is critical for conservation Zeehan Current. The LC creeps closest to the coast (C) ated extensive discussions on the need for a book isms (e.g. ) as well as the basic taxonomy and possible management responses of the GBR. ters. We have added two extra chapters one on Acropora humilis group (AH), the Acropora robusta group (AR), Acropora palifera (AP), Stylophora pistillata (S) and Fig. 3.2. (A) Sea level, temperature and greenhouse gas fluctuations over the past 650 ky from the EPICA ice core from and management of the GBR, as habitats for a vast (Source: Brook 2005). (B) View of the Pleistocene and Holocene raised reef terraces at Huon Peninsula, PNG near Ningaloo, ~1200 km north of , where such as this at Coral Reef Society of the major groups of organisms and their biology Two dominant factors determine the stage a Pocillopora (P). Much of the coral framework is encrusted with coralline . (C) Photographs from the Ribbon Reef 5 Globally, coral reefs are at risk and knowledge- mesophotic reefs (Chapter 7) and another on fish- (C) the current may be only 10 km off the coast. The mid-shelf reef has reached. The first is the depth to number of organismscore become showing available the major and coral are componentslost of ‘Assemblage B1’ and ‘B2’. These include massive Porites (e.g. Porites cf (Photo: R. Kelley). Sea level curve for the past 150 ky derived from Huon Peninsula, supplemented with observations (ACRS). Through the Society and the course, we and ecology (Chapters 15–32). We believe that this based management is absolutely critical. The GBR eries of the GBR (Chapter 10). Chapter 7 reveals lutea) (PO), encrusting Porites (EPO) and massive faviids such as Favites (FA) and Plesiastrea versipora (PE). Again, from Bonaparte Gulf, Australia (Source: from Lambeck et al. 2002). (D) Pleistocene reef terrace from the 125 kya reef at thermal signal of the LC can be readily detected at the Pleistocene antecedent reef (that influences the as reefs progress through this geomorphological were able invite the appropriate international provides the reader with a fascinating journey is at the forefront of this, with a growing risk of amazing diversity in habitats and associated extensive coralline algal rims encrust much of the coral framework (Images: B and C from Webster and Davies 2003). Exmouth, Ningaloo, Western Australia (Photo: R. Kelley). Fig. 4.1. 3-D rendering of the of the GBR . sequence. depth of lagoon to be infilled), with reefs growing and the adjoining . The South Equatorial Current The EAC and LC both carry warm and clear from shallow foundations most quickly reaching transports waters from the oceanic surface well mixed waters that are nutrient poor (oligotrophic, modern sea level and progressing to a mature or outer shelf reefs layer that overlies waters that are 4000 m deep to the Chapter 8). The direction of transport and the oligotrophic waters have a great influence on the senile stage. The second is the size of the reef, Most reefs on the shoulder of the continental shelf GBR on a continental shelf 30 m to 100 m deep (Source: After Wolanski 2001). of the east and west . Corals because this determines the ratio of highly produc- (with the exception of the detached reefs in the are found as far south as Rottnest in the west tive perimeter to the volume of lagoon to be infilled. north) are linear, parallel to the shelf edge and have Large-scale net currents and extend to Sydney in the east. The larvae of Larger reefs, once they reach sea level, progress originated as fringing reefs at earlier low sea levels. tropical species are advected south. A host of newly Fig. 4.2. (A) Large scale circulation in the Coral Sea derived from ship-borne CTD data (plotted as contours of volume The strong, westward flowing, South Equatorial 3 much more slowly. It was shown earlier that the Shelf edge morphology and tidal range are the settled tropical fishes and often arrive transport (in Sv; 1 Sv, 1 million m /s) for the top 1000 m (positive for clockwise flow, negative for counter-clockwise Current (SEC), that generates the East Australian at high latitude towards the end of the summer, but flow). (Source: Adapted from Andrews JC, Clegg S (1989) Coral Sea circulation and transport from modal information Current (EAC), prevails in the Coral Sea and is few survive the cold of winter. As Wernberg and models. Deep-Sea Research 36, 957–974). (B) The net circulation in the south-east trade wind season in the GBR and surrounding seas (Source: Adapted from Wolanski E, Lambrechts J, Thomas C, Deleersnijder E (2013) The net water (A) (B) (C) directed towards the GBR (Fig. 4.2A). The strength colleagues showed in 2016, the northern extent of macroalgae (e.g. Macrocystis and Ecklonia) is also circulation through Torres . Continental Shelf Research 64, 66–74). (C) The SLIM model-predicted average circulation of this flow is modulated by the Southern Oscilla- (D) affected by warm currents, as these algae cannot in the southern GBR. The symbols are various net currents that are explained in the text. Satellite image of the GBR near 14°S showing the effective blockage by reefs forming a barrier that largely blocks the currents and restricts tion Index (SOI). The SOI is the difference in atmos- persist in warm and nutrient poor waters. pheric between Tahiti and Darwin. The connectivity between the central and the northern of the GBR (Source: Map data from Google, SIO, NOAA, US Navy, NGO, GEBCO). SOI also influences the strength of the other major tropical boundary current around Australia, (Fig. 4.2C). The EAC rotates around the Swain Reefs (Fig. 4.2D). Thus the NQCC does not spread over namely the Leeuwin Current on the west coast and creates an eddy (E in Fig. 4.2C), that occasion- the northern GBR, where the net circulation is a (Box 4.1). On meeting the GBR continental shelf, (D) (E) (F) ally sheds to help create the transient Capricorn wind-driven current (WC) that is facilitated by the the SEC splits into two oceanic currents, namely Eddy (CE in Fig. 4.2B). In turn, the Capricorn Eddy oceanic waves breaking over the outer reef crest; the southward flowing East Australian Current Fig. 2.4. modulates the strength of the CSLC. In the south- this generates an inflow of Coral Sea water into the An evolutionary classification for mid-shelf reefs growing from Pleistocene reefal foundations (Source: Hopley (EAC) south of the bifurcation point and the north- 1982). ern GBR, the wind also drives an inflowing north- northern GBR, which is the WW current shown in ward flowing Coastal Current ward longshore coastal boundary layer (CBL) Fig. 4.2B. A juvenile stage capturing growth between ini- (iii) irregular reef patches, formed when coral (NQCC) north of the bifurcation point (Fig. 4.2A). current (Fig. 4.2B, C) that turns into a Cross-Current The water circulation becomes extremely com- tiation and reaching sea level consisting of: growth first reaches sea level. The bifurcation point fluctuates between Towns- (CC) that feeds into the CSLC (Fig. 4.2B). The influ- plex on reaching where the waters are ville and . The EAC does enter the ence of the EAC extends over a 1000 km south of very shallow and the net currents are diminished A mature stage where reef flats are present con- Fig. 2.5. Reef types of the Great Barrier Reef. (A) Reef patches, the youngest stage of reef development: Barnett Patches (i) the drowned Pleistocene reef with possibly off . (B) Crescentic reef with hard-line growth on the windward side: Eagle Reef, off Bowen. reef mosaic of the GBR and generates the Coral Sea the GBR and veers eastwards, forming a tempera- by friction that is enhanced by interacting with sisting of: only Halimeda growth (C) Enclosed lagoonal reefs, infilling from internal patch reef growth and transport of sediment from the productive Lagoonal Current (CSLC) from the Central GBR to ture discontinuity called the Tasman Front. strong tidal currents. In the south-east wind margins: Heron and Wistari Reefs, Bunker Capricorn Group. (D) A small senile planar reef with flat top (still with a living (ii) submerged shoal reefs, with coral growth (i) crescentic reefs formed when reef patches the Swains (Fig. 4.2A). This flow is weak and epi- There is little oceanographic exchange of water season, northern GBR waters flows out to the Gulf coral veneer) and no lagoon: Wheeler Reef off Townsville (note the small unvegetated sand cay on the leeward side). I mage of sea surface temperature showing the warm Leeuwin Current flowing southwards and into the usually restricted to the higher parts of the coalesce into more extensive reef flat on the (E) The continuous line of outer shelf ribbon reefs: Yonge Reef near Lizard Island. (F) Typical high island fringing reef: sodic compared with the EAC flow on the outside between the northern and the central GBR, because of Carpentaria through the Through Torres Strait Great Australian Bight (Image: CSIRO). older reef foundation windward margins, with a hard-line reef front (Photos: D. Hopley). of the reef that can reach speeds of 0.5 m/s of the blockage of the GBR shelf by reefs near 14°S Current (TTS in Fig. 4.2B) and to the Gulf of Papua

CONTACT US CSIRO Publishing, Locked Bag 10, Clayton South VIC 3169, Australia ABN: 41 687 119 230 Banner Image © Flickr.com/Tchami/CC BY-SA 2.0 Local: +(61 3) 9545 8400 or 1300 788 000 This brochure is printed on paper with fibres sourced Email: [email protected] from certified, responsibly managed forests. CONTENTS SECTION 3: OVERVIEW OF REEF ABOUT THE EDITORS BIODIVERSITY AND ORGANISMS Foreword 15. Biodiversity Pat Hutchings is a Senior Fellow at the Abbreviations 16. Plankton Australian Museum Research Institute, Author biographies having retired in 2016 as a Senior Principal 17. Macroalgae Research at the Australian 1. Introduction to the Great Barrier Reef 18. Mangroves and Museum. She has spent her research 19. SECTION 1: NATURE OF THE REEF career working on the and 20. Pelagic and comb jellies 2. Geomorphology of coral reefs with ecology of , especially on special reference to the GBR 21. Hexacorals 1: sea anemones and coral reefs. Her contributions were anemone-like animals (Actiniaria, 3. The GBR in time and space: eologyg Zoantharia, Corallimorpharia, recognised by the Australian Coral Reef and paleobiology Ceriantharia and Antipatharia) Society with honorary membership. 4. Oceanography 22. Hexacorals 2: reef-building or hard Mike Kingsford is a Distinguished 5. Coral reef habitats and their influence corals (Scleractinia) on reef assemblages Professor in the and 23. Octocorals group of the College of 6. Seabed environments, habitats and 24. Worms biological assemblages Science and Engineering at James 25. : and 7. The GBR outer-shelf Cook University, and has over 30 years' pycnogonids research experience on the Great Barrier 8. Primary production, nutrient recycling 26. Molluscs and energy flow through coral reef Reef. He is also a Chief Investigator ecosystems 27. with the ARC Centre of Excellence for 9. Calcification, erosion and the 28. Innovative Coral Reef Studies. establishment of the framework of 29. (ascidians and their allies) Ove Hoegh-Guldberg is Professor of coral reefs 30. The fish assemblages of the GBR: their diversity and origin Marine Science and Director of the SECTION 2: FACTORS AFFECTING Global Change Institute at the University 31. Marine mammals and THE REEF of Queensland. An internationally 32. 10. Fisheries of the GBR recognised expert in his field, he has 11. Disturbances and pressures to coral reefs Epilogue and acknowledgements received numerous prizes including a 12. Coral reefs in a changing world Index Eureka Prize, Thomson Reuters Citation 13. Terrestrial runoff to the GBR and Award, an Australian Research Council the implications for its long term Laureate Fellowship and the Climate Change ecological status Prize from HSH Prince Albert II of Monaco. 14. Planning and managing the GBR Marine Park

EXCERPT FROM THE FOREWORD "In its second edition, this book on the Reef builds on the tradition of the first edition and has expanded the already amazing array of authors and expertise to tackle a number of new and current issues to do with the Reef. Stretching from the geology, biology and oceanography of the Reef, to modern issues such as water quality, climate change and socio- economics of the Reef, this book enables the reader to get a total insight to how the world’s largest coral reef functions." Honourable Robert Hill AC – Former Minister for the Environment and for Defence, Ambassador for Australia to the United Nations

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