Research Article Origins and Implications of a Primary ...

Research ArticleOrigins and Implications of a Primary Crown-of-Thorns StarfishOutbreak in the Southern Great Barrier Reef

Ian Miller Hugh Sweatman Alistair Cheal Mike Emslie Kerryn JohnsMichelle Jonker and Kate Osborne

Australian Institute of Marine Science PMB No 3 Townsville MC QLD 4810 Australia

Correspondence should be addressed to Ian Miller imilleraimsgovau

Received 16 October 2014 Revised 23 January 2015 Accepted 19 February 2015

Academic Editor Baruch Rinkevich

Copyright copy 2015 Ian Miller et alThis is an open access article distributed under theCreativeCommonsAttribution License whichpermits unrestricted use distribution and reproduction in any medium provided the original work is properly cited

The crown-of-thorns starfish (COTS) is a major predator of hard corals Repeated COTS outbreaks in the Cairns and Centralsections of the Great Barrier Reef (GBR) have been responsible for greater declines in coral cover than any other type of disturbanceincluding cyclones disease and coral bleaching Knowledge of the precise timing and location of primary outbreaks could revealthe initial drivers of outbreaks and so could indicate possiblemanagementmeasures In the central GBR COTS outbreaks appear tofollow major flooding events but despite many years of observations no primary outbreak has ever been unequivocally identifiedin the central and northern GBR Here we locate a primary outbreak of COTS on the southern GBR which is not correlated withflooding Instead it appears to have been the result of a combination of life history traits of COTS and prevailing oceanographicconditions The hydrodynamic setting implies that the outbreak could disperse larvae to other reefs in the region

1 Introduction

The crown-of-thorns starfish (COTS) (Acanthaster planci)is an obligate corallivore that causes dramatic losses ofcoral cover at high population densities and is a majorchallenge in coral reef management [1] Populations of COTSalternate periods of low density when individuals are sparselydistributed across large areas of reef with briefer periods ofmuchhigher densities [2] A single adult starfish can consumeapproximately 10m2 of coral per year and large ldquooutbreakrdquopopulations have a major impact on local cover [3] that canpersist for more than a decade [4] Surveys on the GreatBarrier Reef (GBR) span some 50 years in which time therehave been three series of outbreaks beginning in 1962 1979and 1993 Since the advent of systematic monitoring in 1986two outbreaks have appeared in the Cairns Section of theGBR in 1993 and 2009 before subsequently spreading throughthe Central Section of the GBR In 2012 coral cover onthe GBR had fallen to half the value when surveys beganand COTS outbreaks have been responsible for a greaterproportion of declines in coral cover on reefs in the Cairnsand Central sections of the GBR than any other type of

disturbance including cyclones diseases and coral bleaching[5 6] This means that COTS are of critical concern on theGBR particularly as a new series of outbreaks is currentlydeveloping on Cairns Section reefs and following the samepattern [7]

Outbreaks typically begin with an abrupt populationincrease from very low density This change in abundanceby orders of magnitude is termed a ldquoprimary outbreakrdquo if ithas not arisen through recruitment from other high-densitypopulations nearby [8] Explanations for primary outbreaksrevolve round two types of drivers natural (population fluc-tuations [9] aggregation behaviour of existing populations[10 11] andor food limitation [12]) and anthropogenic(enhanced larval survival due to nutrients in terrestrialrunoff [13 14] and increased postsettlement survival due tothe removal of COTS predators [8]) In practice assigningcause(s) for COTS outbreaks is difficult outbreaks maywell result from a combination of factors that vary withtime and location [15] or they may simply be an artefactof the dispersal of passive larva in the water column [16]However primary outbreaks originate secondary outbreaksare thought to result either from colonisation by immense

Hindawi Publishing CorporationJournal of Marine BiologyVolume 2015 Article ID 809624 10 pageshttpdxdoiorg1011552015809624

2 Journal of Marine Biology

numbers of larvae spawned by existing high-density out-breaking populations that are carried to reefs downstream byprevailing currents or from adultmigration between reefs [8]

The first widely documented outbreak population ofCOTS on the GBR was at Green Island in 1962 [17] A varietyof surveys weremade at different times and locations between1966 and 1974 to determine extent of COTS activity on theGBR [2] From these it became apparent that outbreaks beganin the Cairns section of the marine park and spread souththrough the central GBR with time [18 19] as would beexpected if planktonic larva dispersed passively in the watercolumn Systematic surveys over the next two decades con-firmed this pattern for a second wave of outbreaks beginningin the late 1970s and also identified a northward spread ofoutbreaks [20] This pattern had become well established bythe third series of outbreaks beginning in 1993 [21]

Despite the 50-year history of COTS surveys on theGBR the location of primary outbreaks has been difficultto pinpoint To date no primary outbreak on the GBR hasbeen located with any precision even though outbreaksin the Cairns Section first appear on reefs between 145∘and 17∘S [22] While outbreaks were first reported at GreenIsland (1675∘S) near Cairns in 1962 [17] further researchsuggested that this was a secondary outbreak and the originalprimary outbreak(s) had occurred north of Green Island inthe 1950s [23] In 1979 another large outbreak of starfishwas observed on Green Island [24] and survey results andreliable unpublished data indicated that this outbreak wasalso secondarily derived from others located further northaround 16∘S [20] Similarly a third series of outbreaks begin-ning in 1993 was first detected near Lizard Island (145∘S)[15 21] However the exact location of the primary outbreakin 1993 remained uncertain as COTS increased on a numberof nearby reefs about the same time [7] More recent reportsindicated increased COTS activity on reefs near Green Islandfrom around 2010 [25] however COTS numbers have beenslowly building on reefs further north in the vicinity of theLizard Island since 2005 [26] Again the exact location of anyprimary outbreak was uncertain Thus despite an extensivehistory of surveys primary outbreaks of COTS on the GBRremain a matter of conjecture

Information on the location and timing of primaryoutbreaks of COTS could give valuable clues as to the causesPrimary outbreaks are potentially a pressure point for man-aging COTS on the GBR as these initial aggregations lead tosubsequent wide-spread outbreaks that cause such extensivedestruction of coral communities over very large areas [27]Although outbreaks have occurred on the GBR for eightmillennia [28] the frequency of outbreaks in recent decades(in combination with other disturbances such as intensecyclones) is clearly unsustainable leading to speculation thatthe frequency of outbreaks has increased [22] possibly dueto enrichment of GBR waters with nutrients from agriculturein coastal watersheds [14] The evidence that primary COTSoutbreaks on the GBR are linked to enhanced nutrient inputto theGBR lagoon is circumstantial [25]TheGBR is amosaicof over 3000 individual coral reefs stretching nearly 2000 kmfrom Torres Strait in the north (9∘151015840S) to Lady Elliot Islandin the south (24∘071015840S) on the continental shelf of northeast

20∘S

22∘S

24∘S

150∘E 152

∘E 154∘E

N

S

W E

0 50

(km)

200100

Figure 1 The MackayCapricorn Section of the GBR showing thelocation of the Swain Reefs and Capricorn-Bunker Reefs in relationto the coast Isobaths represent depth in meters and show theedge of the continental shelf Arrows show the general position ofthe South Equatorial Current and the East Australian Current andCapricorn EddyThemouth of the Fitzroy River is located southeastof Rockhampton and discharges into Keppel Bay

Australia [29] and because of its large size outbreaksmaywelloccur independently and for different reasons in differentparts of the GBR Based on extensive surveys using mantatows to document the distribution of COTS outbreaks onthe GBR since 1986 [7] we describe a primary outbreakdiscovered on some reefs in the Capricorn-Bunker Group inthe southern GBR We examine the possible causes and weweigh the evidence for links to outbreaks in other parts of theGBR based on the history of outbreaks onCapricorn-BunkerReefs

2 Methods

21 Physical Setting Reefs in the Capricorn-Bunker Groupare the southernmost extent of theGBRand are located 80 kmoffshore on the mid to outer continental shelf [30] Thesereefs are distinctive being mostly medium sized outer-shelfplatform reefs often with lagoons and sand cays [29] To thenorth the continental shelf widens at about 23∘S from about80 km to bemore than 200 kmwide leaving a substantial gapof over 100 km named the Capricorn Channel [31] betweenthe northern extent of the Capricorn-Bunker Reefs and theSwain Reefs The Swain Reefs are contiguous with the restof the GBR and are a series of lagoonal and planar reefslocated offshore on the margin of the continental shelf [29](see Figure 1)

Reefs of the GBR are strongly influenced by the EastAustralian Current (EAC) which is derived from SouthEquatorial Current that crosses the Coral Sea to divide at

Journal of Marine Biology 3

the Queensland Plateau around 18∘S into two approximatelyequal branches [32] The latitude of bifurcation can varyannually between 15∘ and 20∘S [33 34] with a mean ofabout 155∘S [35] The northward branch circulates clockwisearound the northern Coral Sea and flows along the coastalline of the GBR (Cape York) The southern branch is thesource of the EAC which flows southward along the outerGBR and ultimately into the Tasman Sea (Figure 1) In generalthe southward flow strengthens and moves onto the coastat about 17∘S This southward flow could be responsible fortransport of COTS larvae between reefs contributing to thecascade of outbreaks originating from reefs in the CairnsSection of the GBR [36] With the sudden narrowing of thecontinental shelf as the EAC passes the Swain Reefs at thesouthern end of the GBR a clockwise eddy (the CapricornEddy) is entrained which drives a current northward alongthe continental slope adjacent to the Capricorn-Bunker Reefs[37] The strength of the eddy varies seasonally with thesoutherly flow of the EAC The northerly current resultingfrom the Capricorn Eddy is strongest during spring and earlysummer and circulates water around the Capricorn-BunkerReefs [30] Sea surface temperature in the Capricorn-BunkerGroup averages 20∘C in winter and 27-28∘C in summer [30]

The major source of terrestrial input to the region is theFitzroy River which flows into Keppel Bay southeast of Rock-hampton (Figure 1) The catchment extends over approxi-mately 142 500 km2 and is topographically and geologicallydiverse Land use is dominated by grazing and mining Thecatchment was once extensively vegetated with Brigalow(Acacia harpophylla) that was largely cleared for grazing inthe mid-20th century [38] The Fitzroy is the second largestriver that flows into the GBR lagoon [39] and has a meanannual discharge of approximately 59 times 106 megalitres [40]Discharge is highly variable both seasonally and annuallywith themagnitude and frequency of discharge dependent onthe influence of the ElNino SouthernOscillation (ENSO) andsummer cyclone activity [41] In a typical year the dischargeof the Fitzroy River is relatively small but large floods occuron average every 5ndash10 years [42]

22 Sampling Techniques The Australian Institute of MarineScience (AIMS) has used the manta tow technique to surveyreefs along the length of the GBR since 1986 as part ofan ongoing Long Term Monitoring Program (LTMP) Themanta tow technique involves towing a snorkel diver alongthe surface behind a small boat The method provides ageneral description of large areas of reef and can be usedto gauge broad changes in distribution and abundance oforganisms Manta tows are used to describe the broadscalepattern and extent of COTS activity on the GBR Mantatowing is used to survey entire perimeters of reefs suchthat a reef constitutes the unit of survey Two teams workin opposite directions around the reef to survey about halfthe perimeter each A team consists of a boat driver and anobserver who is towed along the surface slowly behind theboat holding on to a manta board at approximately 6 kmh(100mperminute)The boat driver tows the observer parallelto the reef crest and close enough for the observer to see asmuch of the shallow (generally lt10m) reef slope as possible

At two-minute intervals the boat stops allowing the observerto record observations including percent cover of living hardand soft coral counts of COTS (including an estimate of theirsize) and associated feeding scarsThe number ofmanta towsrequired to complete a survey of each reef perimeter will varydepending upon the size of the reef On average a reef inthe Swain region requires sim40 two-minute tows to completewhile a reef in the Capricorn-Bunker region requires sim60tows The method is documented in detail in a StandardOperational Procedure [43]

The AIMS LTMP has surveyed varying numbers of reefsin the Swain and Capricorn-Bunker regions each year since1986 (see Table 1) While some reefs have been surveyedannually or biennially (key reefs) other reefs have beensurveyed much less frequently Sampling is designed to rep-resent a broad overview of reefs in the two sectors (Figure 1)so the individual reefs that are surveyed within a sector havevaried from year to year More details of the surveys can befound in the Long TermMonitoring Report 2008 [4] Surveysare made over the Austral summer so the reporting periodruns from 1 July to 30 June for instance the 2012 survey yearincludes reefs surveyed between July 2011 and June 2012

The densities of COTS on a reef are expressed as the aver-age number of COTS per tow Two categories of outbreak aredistinguished Active Outbreak (AO) and Incipient Outbreak(IO) In principle an Active Outbreak occurs when starfishdensities reach levels where loss of coral tissue throughstarfish feeding is likely to be faster than the growth of thecoral Research into the feeding behaviour of COTS indicatesthat depletion of live coral cover on reefs will occur above athreshold of 1000COTS kmminus2 (10 haminus1) [3] while manta towdata taking into account coral cover indicated a threshold ofsim1500COTS kmminus2 [44] This corresponds to approximately022 COTS per two-minute tow After consideration of therelative costs of Type I and Type II errors the criterion foran Active Outbreak was revised upwards to approximately10 COTS per two-minute manta tow [45] This representsa starfish density that is highly likely to cause net declinein coral cover Less dense populations (022ndash10 COTS pertow) are referred to as Incipient Outbreak representing COTSdensities that may impact on coral cover

3 Results

For the first twodecades ofmanta tow surveys small numbersof COTS were seen intermittently mainly on reefs in thesouthern part of the Capricorn-Bunker region and no out-breaks were recorded (Table 1) More recently COTS num-bers in this sector have increased steadily since 2008 (Figures2 and 3) with the highest numbers recorded during surveysin 2014 The greatest numbers of COTS were seen at FairfaxIsland Reef COTS were first recorded below outbreak levelsat 011 plusmn 001 (mean plusmn SE) COTS per tow on Fairfax IslandReef in 2010When the reef was next surveyed in 2012 COTSnumbers had increased to 097 plusmn 097 COTS per tow and thereef was declared anActiveOutbreakThiswas the first ActiveOutbreak to be recorded using manta tow on any reef in theCapricorn-Bunker Group since AIMS surveys began in 1986The average size of COTS in all surveys exceeded 25 cm in


Table 1 Survey year and number of reefs surveyed including the number of reefs recorded with outbreaks in that year (figures given inbrackets and include both Incipient and Active Outbreaks combined) in the Swain and Capricorn-Bunker Group Mean COTS per two-minute tow and standard error (SE) are given for each survey year A different suite of reefs is sampled in each survey year depending onsample design weather and logistical considerations Survey years run from 1 July of the preceding calendar year to 30 June so for instancethe 2012 survey year represents reefs surveyed between July 2011 and June 2012

Year

Swain Reefs Capricorn-Bunker ReefsNumber of reefs surveyed(number of outbreaking

reefs)Mean COTS SE COTS

Number of reefs surveyed(number of outbreaking


1986 32 (5) 0134 0062 8 (0) 0012 00121987 30 (7) 033 0126 5 (0) 0 01988 28 (3) 0227 0124 6 (0) 0002 00021989 No survey1990 20 (4) 0272 0207 6 (0) 0 01991 21 (3) 0222 0158 5 (0) 0005 00051992 24 (4) 0340 0218 6 (0) 0 01993 9 (2) 0781 0671 4 (0) 0 01994 11 (2) 0120 0090 3 (0) 0 01995 15 (3) 0142 0079 3 (0) 0006 00061996 16 (3) 0107 0042 5 (0) 0010 00061997 8 (3) 0872 0445 5 (0) 0012 00121998 9 (4) 1230 0646 4 (0) 0 01999 7 (3) 0431 0225 4 (0) 0 02000 8 (3) 2094 0797 4 (0) 0 02001 8 (5) 18775 7371 4 (0) 0005 00052002 7 (4) 4879 2798 4 (0) 0 02003 7 (4) 4077 2227 4 (0) 0 02004 7 (3) 0649 0414 4 (0) 0005 00052005 13 (1) 0050 0033 4 (0) 0 02006 19 (1) 0030 0016 8 (0) 0 02007 6 (0) 0005 0004 4 (0) 0 02008 17 (0) 0 0 7 (0) 0 02009 7 (0) 0011 0011 4 (0) 0 02010 15 (0) 0003 0003 8 (0) 0018 00132011 7 (0) 0002 0002 3 (0) 0 02012 15 (0) 0003 0002 6 (1) 0162 01622013 7 (0) 0023 0018 4 (1) 0066 00662014 14 (0) 0005 0003 8 (2) 0201 0146

diameter indicating that the individuals were three or moreyears old [43] Subsequent surveys in 2014 showed ActiveOutbreak densities (116 plusmn 074 COTS per tow) persisted andthat the population was highly concentrated in the back reefarea An IncipientOutbreak (040plusmn016COTSper tow Figure2) was also recorded on Lady Musgrave Island Reef in 2014The outbreak was first observed on this reef in 2013 (026plusmn01COTS per tow) Prior to that small numbers of COTS hadbeen observed occasionally at Lady Musgrave Island Reefsince the beginning of surveys in 1986 [7] Once again starfishwere all greater than 25 cm in diameter As with the outbreakon Fairfax Island Reef COTS were concentrated on the shel-tered back reef areawhere coral coverwas the highest Prior to

the discovery of these two outbreaks reefs in the Capricorn-Bunker region of the GBR had only supported small COTSpopulations that were well below outbreak densities (Table 1)Other researchers recorded similarly low densities of COTSon reefs in the region for decades prior to the beginning ofAIMS surveys [28] and the sector as a whole appears to haveescaped the substantial effects of COTS outbreaks seen inother parts of the central and southern GBR [46]

By contrast COTS were numerous in the Swain Reefsfor the first two decades after 1986 with Active Outbreaklevels of COTS recorded on one or more reefs in the regionin nearly every year (Table 1) In particular from 1999 to2003 COTS activity was high and greatly exceeded outbreak


Survey year2006 2008 2010 2012 2014

Mea

n CO

TS p

er m

anta

tow

00

02

04

06

08

10

12

Fairfax Island ReefLady Musgrave Island Reef

Figure 2 Recent change in abundance of COTS on Fairfax IslandReef (surveyed in 2006 2008 2010 2012 and 2014) and LadyMusgrave IslandReef (surveyed in 2006 2007 2009 2010 2012 2013and 2014) Survey years run from 1 July of the preceding calendaryear to 30 June so for instance the 2012 survey year represents datafrom reefs that were surveyed between July 2011 and June 2012

Survey (year)

1986

1988

1990

1992

1994

1996

1998

2000

2002

2004

2006

2008

2010

2012

2014

001

01

1

10

Swain Sector ReefsCapricorn-Bunker Sector Reefs

Mea

n CO

TS p

er m

anta

tow

Figure 3 Historic change in COTS abundance on Swain Reefscompared to Capricorn-Bunker Reefs Surveys conducted in bothregions annually except for 1989 Survey years run from 1 July to 30June so for instance the 2012 survey year represents reefs surveyedbetween July 2011 and June 2012119884-axis log scale the small line peakin 2013 represents small numbers of COTS recorded from SwainReef surveys in that year (see Table 1)

levels across the region (Figure 3) After 2004 COTS activitydeclined in the Swain Reefs and by 2007 no outbreaks wererecorded COTS activity has remained low on reefs in thisregion since with the odd individual COTS observed butalways well below outbreak densities (Table 1 Figure 3)

Tota

l yea

rly fl

ow (m

egal

itres

)

40

30

20

10

0

times106

1986

1988

1990

1992

1994

1996

1998

2000

2002

2004

2006

2008

2010

2012

Year

Figure 4 Total annual flow in megalitres for the Fitzroy Rivermeasured in ldquowater yearsrdquo at ldquoTheGaprdquomonitoring site Awater yearruns from 1October of the preceding calendar year to 30 Septemberfor instance the bar for 2012 water year in the figure represents totalflow measured at ldquoThe Gaprdquo between October 2011 and September2012

River flow data for the Fitzroy River for the period ofAIMS broadscale surveys (1986 to 2012) shows that majorflood events (where total annual river flow was more thandouble average total annual flow for the period 1986 to 2012)occurred in 1991 and 2011 with minor floods (where annualflow exceeded average annual flow for the period 1986 to2012) in 1988 1989 1991 2008 2010 and 2012 (Figure 4)COTS are thought to breed in early wet season (Novemberto January) and major flood events (where total early seasonriver flow was more than double average early season flowfor the period November to January) occurred at this timeof year in 1991 and 2011 with minor flood events (whereearly season flow exceeded average early season flow for theperiod November to January) in 1996 1999 2001 and 2008(Figure 5)

The outbreak at Fairfax Island Reef appears to be aprimary outbreak because there were no known high-densityCOTS populations in the Swain Reefs that could havebeen sources of larvae Unlike the repeated pattern in theCairns Section of the GBR this outbreak did not follow theoccurrence of large flood plumes in the region

4 Discussion

This is the first compelling example of a primary outbreakrecorded from theGBR and unlike outbreaks recorded in theCairns Section of theGBR the timing does not appear relatedto flooding in nearby catchments While large populationfluctuations are a characteristic of COTS [1 47ndash49] theorigins of outbreaks are often subtle and seldom due to asingle cause [50] COTS are among the most fecund of anymarine species with annual gamete output per female of tensto hundreds of millions of gametes [51ndash53] COTS also havethe highest fertilisation efficiency recorded for any marine

6 Journal of Marine BiologyTo

tal e

arly

seas

on fl

ow (m

egal

itres

)

40

30

20

10

0

times106

1986

1988

1990

1992

1994

1996

1998

2000

2002

2004

2006

2008

2010

2012

Year

Figure 5 Total ldquoearly season flowrdquo inmegalitres for the FitzroyRivermeasured at ldquoThe Gaprdquo Early season flow reflects measurement oftotal water flow from 1 November to 28 February correspondingto the putative COTS spawning season Thus the bar in the figurelabelled 2012 represents total flow measured at ldquoThe Gaprdquo betweenNovember 2011 and February 2012

invertebrate [54] predisposing COTS to extreme populationfluctuations [55] Large aggregations may spawn billions oflarvae so a small increase in larval survival could causepopulation explosions [56] Changes in oceanographic condi-tions such as prevailing water currents and upwelling havebeen correlated with increased survival and recruitment ofCOTS larvae [57ndash59] that have been linked to phytoplanktonblooms [60ndash62] Upwelling nutrients fuel the phytoplanktonblooms though eutrophication can also contribute [63ndash66]The intensity and duration of phytoplankton blooms canpotentially have profound impacts on survival of planktivo-rous larvae including COTS [67]

Since 1960 four series of outbreaks have begun in theregion north ofCairns 2ndash5 years after the occurrence of floodswhere aggregate discharges from northern rivers (Burdekinto Daintree Rivers) in the early wet season (NovemberndashFebruary) exceeded 10 times 106 megalitres [25] A plausibleexplanation is that phytoplankton blooms resulting fromincreased nutrient loading in reef waters from floods havebeen the proximate cause of primary outbreaks in theregion The COTS outbreaks on reefs in the Capricorn-Bunker region do not show such a relationship with floodingAssuming that COTS also spawn inDecember and January inthe southernGBR [54] there have been a number of occasionsover the past thirty years where floods particularly earlyseason floods from the Fitzroy River may have engulfedCapricorn-Bunker Reefs These include major early seasonfloods in 1991 and 2011 and minor flood events in 1996 1999and 2008Despite this history of floods COTSoutbreaks havenot been recorded from this sector until recently Capricorn-Bunker Reefs may be far enough off shore that they arerarely exposed to flood plumes [68] For example the majorriver flood of the Fitzroy River from Tropical Cyclone Joy

in 1991 generated a plume that reached the northern reefsof the Capricorn-Bunker region (75 km offshore) for a fewdays [69] This was an exceptional event only exceeded inrecent years by flooding following Tropical Cyclone Tasha in2011 While flood waters from the Fitzroy River may haveimpinged on Capricorn-Bunker Reefs in 2011 the COTSoutbreak at Fairfax Island Reef was first observed in 2012 andcomprised of starfish thatwere at least three years old (gt25 cmin diameter) meaning that they must have been spawned atleast two years prior to the floodThus these outbreaks did notbegin 2ndash5 years after early season floods unlike the pattern inthe Cairns Section of the GBRWhile there was aminor floodin 2008 the flow was moderate and only the largest plumesare known to have reached reefs in the southern Capricorn-Bunker region Thus enhanced recruitment from eutrophi-cation caused by early wet season floods is unlikely to havecaused the current outbreaks on theCapricorn-Bunker Reefs

Nutrient enrichment from upwelling can also lead toCOTS outbreaks on coral reefs [59 70] In the Capricorn-Bunker region the Capricorn Eddy draws cooler nutrient-enriched oceanic water up into the shallows and transportsit to the reefs particularly during the summer [37 71] Atwelve-year study of chlorophyll 119886 concentrations on theGBR recorded the highest values for outer-shelf reefs any-where on the GBR in the Capricorn-Bunker region (average0519 120583gLminus1 11987510 = 0165) [72] with significant variationamong years Chlorophyll 119886 concentrations above 025120583gLminus1can lead to amuch higher rate of development and survival ofCOTS larvae in laboratory studies [73] Similarly the optimaltemperature for survival of COTS larvae in the laboratoryis 25ndash32∘C [74] which falls within the range of summersea surface temperatures for the Capricorn-Bunker regionThis suggests that conditions for enhanced recruitment ofCOTS larvae occur on a regular basis though outbreaks havenot been observed previously This highlights the uncertainrelationship between phytoplankton densities and survivor-ship of mass-spawned invertebrate planktotrophs generallyUntil the relation between the two is better understoodthe suggestion that the release from food limitation is theprincipal cause of enhanced COTS recruitment should beinterpreted with caution [75]

The waves of outbreaks that have been observed to movethrough the central GBR are thought to be propagated bythe immense numbers of larvae from outbreak densities ofadults being carried to downstream reefs by the prevailingcurrents It is surprising that more outbreaks have not beenrecorded on reefs in the Capricorn-Bunker Group becausethe region is potentially well-connected to the Swain Reefswhere Active Outbreaks were consistently present for twodecades (1986ndash2006) By the time the first outbreak in theCapricorn-Bunker region was observed at Fairfax Island Reefin 2012 some six years had elapsed since the last outbreak hadbeen recorded from the Swain ReefsThis long intervalmakesit highly unlikely that the outbreak was the result of enhancedlarval recruitment from large populations in the Swains

Population release after removal of predators is anotherimplied cause of COTS outbreaks [8] Historically bothFairfax Island Reef and Lady Musgrave Island Reef have


been subject to commercial and recreational fishingHoweverFairfax Island Reef was rezoned to exclude fishing in 2006and Lady Musgrave Island Reef has had a large area of reefperimeter protected from fishing since 1983 Prior to thistime when the reefs were open for fishing no outbreakswere observed Fishery targeted reef fishes have been shownto make a rapid recovery on the GBR once protectionis introduced [76] and protection from fishing putativelyreduces the ability of COTS to form outbreaks on other partsof the GBR [77] However this link remains tenuous as morecommonly fished large predatory fish do not usually preyupon COTS [78] and to date no fish predator has beenidentified that can effectively regulate COTS populationsalthough predation on juveniles undoubtedly occurs [79]This evidence combined with the lack of outbreaking COTSpopulations prior to the fishing closures strongly suggestspredator removal is not an underlying cause for the outbreak

Individual starfish can persist on a reef for nearly adecade [56] Outbreaks in general and primary outbreaks inparticular do not need to be the result of a single recruitmentpulse numbers may build up over a number of years toreach outbreak densities [80]There are a number of examplesdescribed from the northern GBR that appear to follow thispattern [15 21 81] This fits with what we know about theoutbreak on Fairfax Island Reef

Here we have described a primary outbreak of COTS onthe GBR This event is exceptional because the majority ofoutbreaks on the GBR are secondarily derived from larvaethat spread among well-connected reefs after the initiationof one or more primary outbreaks [23] These patterns ofsecondary outbreaks are also known from southern Japan[82 83] but may occur amongst high island archipelagosin other parts of the Indo-Pacific [84] An analyses of allreported Indo-Pacific outbreaks (246) since 1990 showedthat while a majority of outbreaks (29 and 56 resp)were reported from high islands and continental shelves asignificant proportion (14) had also been reported fromlow islands or atolls [56] That outbreaks had occurred onso many Indo-Pacific low islands and atolls is telling This isbecause research into the genetic structure and connectivityamong outbreak populations at 23 sites across the PacificOcean indicated that larval dispersal is highly constrainedand that high densities of larvae do not spread across openocean expanses to initiate secondary outbreaks at distant reefs[49 85] Furthermore where an outbreak has occurred onan isolated atoll or low island it can be often difficult toprescribe any anthropogenic forcing leading to the outbreak[56] other than overfishing [86] Hence proposed mecha-nisms generally invoke other sources usually nutrients thatcause plankton blooms leading to enhanced larval survivalincluding upwelling [59 60] bioturbation and resuspensionof sediments by severe tropical storms and oceanographicfeatures that create high productivity fronts [58] Resultspresented here support the view that given the lack ofevidence for any clear anthropogenic drivers the outbreakon Fairfax Island Reef is likely a result of one or a numberof the myriad factors that affect larval survivorship anddispersal prior to settlement Superficially this appears similarto cases recorded from low islands and atolls throughout

the Indo-Pacific where outbreaks cannot be readily linkedto terrestrial runoff More generally the link between theoutbreak on Fairfax Island Reef and episodes of enhancednutrients from oceanographic processes is less clear thoughambient conditions would indicate they certainly exist Whatis clear is that there is little evidence of a direct relationshipwith human activity which supports the idea that in somecases outbreaks can arise spontaneously with no immediateapparent explanation [75 87] Given the emergent complexityresulting from interactions between larval ecology oceano-graphic and postsettlement processes this is not surprisingand the exact reason(s) for the Fairfax Island Reef outbreakremain enigmatic Whatever the cause the location of theprimary outbreak at the southern end of the Capricorn-Bunker region has implications for the other nearby reefsThe Capricorn Eddy contributes to a north-westward flowwithin the region [88]This flow is greater during the summermonths when the EAC is stronger and the southeast tradewinds are weak [37] This effectively puts other reefs inthe region ldquodownstreamrdquo to those reefs currently supportingoutbreaks This northward flow is most prevalent during thesummer months when COTS spawning is likely to occurDrifter studies show that the orientation of the reef matrix inthe Capricorn-Bunker region approximately perpendicularto the main tidal flow can result in trapping mechanismssuch as island wake eddies that could keep particles withinthe reef array This may represent a mechanism of bothincreasing dispersion of larvae along the reef matrix and atthe same time containing them within this region [89] Asa result other reefs in this region may be subject to COTSoutbreaks in the future

Conflict of Interests

The authors declare that there is no conflict of interestsregarding the publication of this paper

Acknowledgments

The authors would like to thank past members of the LTMPfor helping to collect the data The authors also thank theskippers and crews of the RV Sirius RV Harry Messel RVLady Basten RV Cape Ferguson MV New Horizons MVIron Joy MV Capricorn Star and MV Centurion for gettingthem to the survey reefs and for looking after their health andsafetywhilst working in the fieldThis researchwas sponsoredby theAustralian Institute ofMarine Science theCooperativeResearch Centre for the Great Barrier Reef World HeritageArea theMarine and Tropical Sciences Research Facility andthe National Environmental Research Program

References

[1] C Birkeland and J S Lucas Acanthaster planci Major Man-agement Problem of Coral Reefs CRC Press Boca Raton CalifUSA 1990

[2] P J Moran ldquoTheAcanthaster phenomenonrdquoOceanography andMarine Biology Annual Review vol 24 pp 379ndash480 1986


[3] J K Keesing and J S Lucas ldquoField measurement of feedingandmovement rates of the crown-of-thorns starfishAcanthasterplance (L)rdquo Journal of Experimental Marine Biology and Ecol-ogy vol 156 no 1 pp 89ndash91 94ndash104 1992

[4] H P A Sweatman A J Cheal G J Coleman et al ldquoLong-termmonitoring of the Great Barrier Reefrdquo Status Report 8 Aus-tralian Institute of Marine Science Townsville Australia 2008httpwwwaimsgovaudocuments30301e055df14-24d0-40ca-b46d-4ae1988da07c

[5] K Osborne A M Dolman S C Burgess and K A JohnsldquoDisturbance and the dynamics of coral cover on the GreatBarrier Reef (1995ndash2009)rdquo PLoS ONE vol 6 no 3 Article IDe17516 2011

[6] G DersquoAth K E Fabricius H Sweatman andM Puotinen ldquoThe27-year decline of coral cover on the Great Barrier Reef and itscausesrdquo Proceedings of the National Academy of Sciences of theUnited States of America vol 109 no 44 pp 17995ndash17999 2012

[7] I R Miller and H Sweatman ldquoThe status of crown-of-thornsstarfish populations on the Great Barrier Reef from AIMS sur-veysrdquo Report to theNational Environmental Research ProgramReef and Rainforest Research Centre Cairns Australia 2013httpwwwnerptropicaleduausitesdefaultfilespublicationsfilesThe20status20of20crown-of-thorns20starfish20Miller20et20al202013pdf

[8] R Endean ldquoAcanthaster planci infestations on reefs of the GreatBarrier Reefrdquo in Proceedings of the 3rd International Coral ReefSymposium D L Taylor Ed pp 185ndash191Miami Fla USA 1977

[9] P J Vine ldquoDensities of Acanthaster planci in the Pacific OceanrdquoNature vol 228 pp 341ndash342 1970

[10] J M Branham S A Reed J H Bailey and J Caperon ldquoCoral-eating sea stars Acanthaster planci in Hawaiirdquo Science vol 172no 3988 pp 1155ndash1157 1971

[11] R F G Ormond and A C Campbell ldquoObservations on Acan-thaster planci and other coral reef echinoderms in the SudaneseRed Seardquo in Regional Variation in Indian Ocean Coral Reefs SM Yonge and D R Stoddart Eds vol 28 of Symposia of theZoological Society of London pp 433ndash454 Academic Press 1971

[12] T F DanaW A Newman and EW Fager ldquoAcanthaster aggre-gations interpreted as primarily responses to natural phenom-enardquo Pacific Science vol 26 pp 355ndash372 1972

[13] C Birkeland ldquoTerrestrial runoff as a cause of outbreaks ofAcan-thaster planci (Echinodermata Asteroidea)rdquo Marine Biologyvol 69 no 2 pp 175ndash185 1982

[14] J Brodie K Fabricius G Dersquoath and K Okaji ldquoAre increasednutrient inputs responsible for more outbreaks of crown-of-thorns starfish An appraisal of the evidencerdquoMarine PollutionBulletin vol 51 no 1ndash4 pp 266ndash278 2005

[15] M S Pratchett ldquoDynamics of an outbreak population ofAcanthaster planci at Lizard Island northern Great Barrier Reef(1995ndash1999)rdquo Coral Reefs vol 24 no 3 pp 453ndash462 2005

[16] G L Eckert ldquoEffects of the planktonic period on marine popu-lation fluctuationsrdquo Ecology vol 84 no 2 pp 372ndash383 2003

[17] J H Barnes ldquoThe crown-of-thorns starfish as a destroyer ofcoralrdquo Australian Natural History vol 15 pp 257ndash261 1966

[18] F H Talbot and M S Talbot ldquoThe crown-of-thorns starfish(Acanthaster) and the Great Barrier Reefrdquo Endeavour vol 30pp 38ndash42 1971

[19] R G Pearson ldquoChanges in distribution of Acanthaster plancipopulations on theGreat Barrier ReefrdquoNature vol 237 no 5351pp 175ndash176 1972

[20] P J Moran G Dersquoath V J Baker et al ldquoPattern of outbreaksof crown-of-thorns starfish (Acanthaster planci L) along theGreat Barrier Reef since 1966rdquoAustralian Journal of Marine andFreshwater Research vol 43 no 3 pp 555ndash568 1992

[21] I Miller ldquoHistorical patterns and current trends in the broad-scale distribution of crown-of-thorns starfish in the northernand central sections of the Great Barrier Reefrdquo in Proceedingsof the 9th International Coral Reef Symposium vol 2 pp 1478ndash1484 Ministry of Environment and the Indonesian Institute ofScience Bali Indonesia 2000

[22] K E Fabricius K Okaji and G Dersquoath ldquoThree lines of evidenceto link outbreaks of the crown-of-thorns seastar Acanthasterplanci to the release of larval food limitationrdquo Coral Reefs vol29 no 3 pp 593ndash605 2010

[23] R A Kenchington ldquoGrowth and recruitment of Acanthasterplanci (L) on the Great Barrier Reefrdquo Biological Conservationvol 11 no 2 pp 103ndash118 1977

[24] R Endean ldquoCrown-of-thorns starfish on the Great BarrierReefrdquo Endeavour vol 6 no 1 pp 10ndash14 1982

[25] M Furnas R Brinkman K Fabricius H Tonin and B Schaf-felke ldquoLinkages between river runoff phytoplankton bloomsand primary outbreaks of crown-of-thorns starfish in theNorthern GBRrdquo in Assessment of the Relative Risk of WaterQuality to Ecosystems of the Great Barrier Reef SupportingStudies A Report to the Department of the Environment andHeritage Protection chapter 1 pp 1ndash299 Queensland Govern-ment Brisbane Australia 2013 TropWATER Report 1330Townsville Australia

[26] Australian Institute of Marine Science Long Term MonitoringProgram Report on Surveys of the CooktownmdashLizard IslandCairns and Innisfail Sectors of the Great Barrier Reef AustralianInstitute of Marine Science Townsville Australia 2009 httpwwwaimsgovaudocsresearchmonitoringreefltm200911-12html

[27] R E Reichelt W Greve R H Bradbury and P J MoranldquoAcanthaster planci outbreak initiation a starfish-coral sitemodelrdquo Ecological Modelling vol 49 no 3-4 pp 153ndash177 1990

[28] P D Walbran R A Henderson A J T Jull and J M HeadldquoEvidence from sediments of long-term Acanthaster plancipredation on corals of the Great Barrier Reefrdquo Science vol 245no 4920 pp 847ndash850 1989

[29] D Hopley The Geomorphology of the Great Barrier ReefQuaternary Development of Coral Reefs John Wiley amp SonsNew York NY USA 1982

[30] J S Jell and G E Webb ldquoGeology of Heron island and adjacentreefs Great Barrier Reef Australiardquo Episodes vol 35 no 1 pp110ndash119 2012

[31] J H Middleton P Coutis D A Griffin et al ldquoCirculation andwater mass characteristics of the Southern Great Barrier ReefrdquoAustralian Journal of Marine amp Freshwater Research vol 45 no1 pp 1ndash18 1994

[32] J C Andrews and S Clegg ldquoCoral Sea circulation and transportdeduced from modal information modelsrdquo Deep Sea ResearchPart A Oceanographic Research Papers vol 36 no 6 pp 957ndash974 1989

[33] J A Church ldquoEast Australian current adjacent to the GreatBarrier Reefrdquo Australian Journal of Marine and FreshwaterResearch vol 38 no 6 pp 671ndash683 1987

[34] K Wyrtki ldquoGeopotential topographies and associated circula-tion in the western South Pacific Oceanrdquo Australian Journal ofMarine and Freshwater Research vol 13 no 2 pp 89ndash105 1962


[35] K R Ridgway and J R Dunn ldquoMesoscale structure of themean East Australian Current System and its relationship withtopographyrdquo Progress in Oceanography vol 56 no 2 pp 189ndash222 2003

[36] I J Dight L Bode and M K James ldquoModelling the larvaldispersal of Acanthaster planci I Large scale hydrodynamicsCairns Section Great Barrier Reef Marine Parkrdquo Coral Reefsvol 9 no 3 pp 115ndash123 1990

[37] S J Weeks A Bakun C R Steinberg R Brinkman and OHoegh-Guldberg ldquoThe Capricorn Eddy a prominent driver ofthe ecology and future of the southernGreat Barrier ReefrdquoCoralReefs vol 29 no 4 pp 975ndash985 2010

[38] H C Bostock B P Brooke D A Ryan et al ldquoHolocene andmodern sediment storage in the subtropical macrotidal FitzroyRiver estuary Southeast Queensland Australiardquo SedimentaryGeology vol 201 no 3-4 pp 321ndash340 2007

[39] M Furnas Catchments and Corals Terrestrial Runoff to theGreat Barrier Reef Australian Institute of Marine Science andCRC Reef Research Centre Townville Australia 2003

[40] Queensland Department of Natural Resources and Mines(QDNRM) ldquoFitzroy basin draft resource operations planrdquoOverview Report 2013 httpswwwdnrmqldgovau dataassetspdf file0014104054fitzroy-draft-rop-overviewpdf

[41] J M Lough ldquoClimate variability and change on the Great Bar-rier Reefrdquo in Oceanographic Processes on Coral Reefs Physics-Biology Links in the Great Barrier Reef E Wolanski Ed pp269ndash300 CRC Press Boca Raton Fla USA 2001

[42] P Larcombe and R M Carter ldquoCyclone pumping sedimentpartitioning and the development of theGreat Barrier Reef shelfsystem a reviewrdquo Quaternary Science Reviews vol 23 no 1-2pp 107ndash135 2004

[43] I R Miller M Jonker and G Coleman Crown-of-ThornsStarfish and Coral Surveys Using the Manta Tow and ScubaSearch Techniques Long-Term Monitoring of the Great BarrierReef Standard Operation Procedure Number 9 Australian Insti-tute of Marine Science Townsville Australia 3rd edition 2009httpwwwaimsgovaudocuments3030120e3bf4f-4b3b-4808-ac02-c15c2912c3f2

[44] P J Moran and G Dersquoath ldquoEstimates of the abundance of thecrown-of-throns starfish Acanthaster planci in outbreaking andnon-outbreaking populations on reefs within the Great BarrierReefrdquoMarine Biology vol 113 no 3 pp 509ndash515 1992

[45] B R Lassig and U Engelhardt ldquoCOTS Commsrdquo Reef Researchvol 5 no 1 pp 18ndash23 1995

[46] R A Henderson ldquoAssessment of crown-of-thorns skeletalelements in surface sediment of the Great Barrier Reefrdquo CoralReefs vol 11 no 2 pp 103ndash108 1992

[47] S Uthicke B Schaffelke and M Byrne ldquoA boom-bust phylumEcological and evolutionary consequences of density variationsin echinodermsrdquo Ecological Monographs vol 79 no 1 pp 3ndash242009

[48] K Narvaez and F A Zapata ldquoFirst record and impact ofthe crown-of-thorns starfish Acanthaster planci (SpinulosidaAcanthasteridae) on corals of Malpelo Island ColombianPacificrdquo Revista de Biologia Tropical vol 58 supplement 1 pp139ndash143 2010

[49] M A Timmers C E Bird D J Skillings P E Smouse and RJ Toonen ldquoTherersquos no place like home crown-of-thorns out-breaks in the central pacific are regionally derived and indepen-dent eventsrdquo PLoS ONE vol 7 no 2 Article ID e31159 2012

[50] Australian Academy of Science ldquoAcanthaster planci (crown-of-thorns starfish) and the Great Barrier Reefrdquo Reports of

the Australian Academy of Science 11 Australian Academy ofScience Canberra Australia 1970

[51] J S Lucas ldquoReproductive and larval biology of Acanthasterplanci (L) in Great Barrier Reef watersrdquoMicronesica vol 9 pp197ndash203 1973

[52] C Conand ldquoDistribution reproductive cycle and morpho-metric relationship of Acanthaster planci (EchinodermataAsteroidea) in New Caledonia western tropical Pacificrdquo inProceedings of the 5th International Echinoderm Conference onEchinodermata B F Keegan and B D S OrsquoConnor Eds pp499ndash506 Balkema Rotterdam The Netherlands 1985

[53] E Ramırez-Llodra ldquoFecundity and life-history strategies inmarine invertebratesrdquo Advances in Marine Biology vol 43 pp88ndash170 2002

[54] R C Babcock and C NMundy ldquoReproductive biology spawn-ing and field fertilization rates ofAcanthaster plancirdquoAustralianJournal of Marine and Freshwater Research vol 43 no 3 pp525ndash534 1992

[55] C Birkeland ldquoThe Faustian traits of the crown-of-thornsstarfishrdquo American Scientist vol 77 no 2 pp 154ndash163 1989

[56] M S Pratchett C F Caballes J A Rivera-Posada and H P ASweatman ldquoLimits to understanding and managing outbreaksof crown-of-thorns starfish (Acanthaster spp)rdquo in Oceanogra-phy and Marine Biology An Annual Review R N Hughes DJ Hughes and and I P Smith Eds vol 52 pp 133ndash200 CRCPress Boca Raton Fla USA 2014

[57] M Yamaguchi ldquoAcanthaster planci infestations of reefs andcoral assemblages in Japan a retrospective analysis of controleffortsrdquo Coral Reefs vol 5 no 1 pp 23ndash30 1986

[58] P Houk S Bograd and R van Woesik ldquoThe transition zonechlorophyll front can triggerAcanthaster planci outbreaks in thePacific Ocean historical confirmationrdquo Journal of Oceanogra-phy vol 63 no 1 pp 149ndash154 2007

[59] P Houk and J Raubani ldquoAcanthaster planci outbreaks inVanuatu coincide with ocean productivity furthering trendsthroughout the pacific oceanrdquo Journal of Oceanography vol 66no 3 pp 435ndash438 2010

[60] V A Mendonca M M Al Jabri I Al Ajmi M Al MuharramiM Al Areimiand and H A Al Aghbari ldquoPersistent andexpanding population outbreaks of the corallivorous starfishAcanthaster planci in the North western Indian Ocean are theyreally a consequence of unsustainable starfish predator removalthrough overfishing in coral reefs or a response to a changingenvironmentrdquo Zoological Studies vol 49 no 1 pp 108ndash1232010

[61] J C Brock andC RMcClain ldquoInterannual variability in phyto-plankton blooms observed in the northwestern ArabianSea during the southwest monsoonrdquo Journal of GeophysicalResearch vol 97 no 1 pp 733ndash750 1992

[62] S Shang L Li J Li Y Li G Lin and J Sun ldquoPhytoplanktonbloom during the northeast monsoon in the Luzon Straitbordering the Kuroshiordquo Remote Sensing of Environment vol124 pp 38ndash48 2012

[63] P R F Bell ldquoEutrophication and coral reefsmdashsome examples inthe Great Barrier Reef lagoonrdquo Water Research vol 26 no 5pp 553ndash568 1992

[64] P Dufour and B Berland ldquoNutrient control of phytoplanktonicbiomass in atoll lagoons and Pacific Ocean waters studies withfactorial enrichment bioassaysrdquo Journal of ExperimentalMarineBiology and Ecology vol 234 no 2 pp 147ndash166 1999


[65] P Douillet S Ouillon and E Cordier ldquoA numerical model forfine suspended sediment transport in the southwest lagoon ofNew Caledoniardquo Coral Reefs vol 20 no 4 pp 361ndash372 2001

[66] J-P Torreton E Rochelle-Newall O Pringault S Jacquet VFaure and E Briand ldquoVariability of primary and bacterialproduction in a coral reef lagoon (New Caledonia)rdquo MarinePollution Bulletin vol 61 no 7ndash12 pp 335ndash348 2010

[67] T Platt S Sathyendranath G N White et al ldquoDiagnostic pro-perties of phytoplankton time series from remote sensingrdquoEstuaries and Coasts vol 33 no 2 pp 428ndash439 2010

[68] M J Devlin L W McKinna J G Alvarez-Romero et alldquoMapping the pollutants in surface riverine flood plume watersin the Great Barrier Reef Australiardquo Marine Pollution Bulletinvol 65 no 4ndash9 pp 224ndash235 2012

[69] MDevlin JWaterhouse J Taylor and J Brodie ldquoFlood plumesin the Great Barrier Reef spatial and temporal patterns incomposition and distributionrdquo GBRMPA Research Publication68 Great Barrier Reef Marine Park Authority TownsvilleAustralia 2001

[70] V M Mendonca M M Al Jabri I Al Ajmi M Al MuharramiM Al Areimi and H A Al Aghbari ldquoPersistent and expand-ing population outbreaks of the corallivorous starfish Acan-thaster planci in the Northwestern Indian Ocean are theyreally a consequence of unsustainable starfish predator removalthrough overfishing in coral reefs or a response to a changingenvironmentrdquo Zoological Studies vol 49 no 1 pp 108ndash1232010

[71] J A Kleypas andDM Burrage ldquoSatellite observations of circu-lation in the southern Great Barrier Reef Australiardquo Interna-tional Journal of Remote Sensing vol 15 no 10 pp 2051ndash20631994

[72] J Brodie G Dersquoath M Devlin M Furnas and M WrightldquoSpatial and temporal patterns of near-surface chlorophyll a inthe Great Barrier Reef lagoonrdquoMarine and Freshwater Researchvol 58 no 4 pp 342ndash353 2007

[73] T Ayukai K Okaji and J Lucas ldquoFood limitation in the growthand development of crown-of-thorns starfish larvae in theGreatBarrier Reefrdquo in Proceedings of the 8th International Coral ReefSymposium vol 1 pp 621ndash626 Smithsonian Tropical ResearchInstitute Panama City Panama 1997

[74] M Lamare D Pecorino N Hardy M Liddy M Byrne andS Uthicke ldquoThe thermal tolerance of crown-of-thorns (Acan-thaster planci) embryos and bipinnaria larvae implications forspatial and temporal variation in adult populationsrdquoCoral Reefsvol 33 no 1 pp 207ndash219 2014

[75] D J W Lane ldquoAcanthaster planci impact on coral communitiesat permanent transect sites on Bruneian reefs with a regionaloverview and a critique on outbreak causesrdquo Journal of theMarine Biological Association of the United Kingdom vol 92 no4 pp 803ndash809 2012

[76] G R Russ A J Cheal A M Dolman et al ldquoRapid increase infish numbers follows creation of worldrsquos largest marine reservenetworkrdquo Current Biology vol 18 no 12 pp 514ndash515 2008

[77] H Sweatman ldquoNo-take reserves protect coral reefs from preda-tory starfishrdquo Current Biology vol 18 no 14 pp R598ndashR5992008

[78] H P Sweatman ldquoA field study of fish predation on juvenilecrown-of-thorns starfishrdquo Coral Reefs vol 14 no 1 pp 47ndash531995

[79] J K Keesing and A R Halford ldquoImportance of postsettlementprocesses for the population dynamics of Acanthaster planci

(L)rdquo Australian Journal of Marine amp Freshwater Research vol43 no 3 pp 635ndash651 1992

[80] C Johnson ldquoReproduction recruitment and hydrodynamics inthe crown-of-thorns phenomenon on the Great Barrier Reefintroduction and synthesisrdquo Australian Journal of Marine andFreshwater Research vol 43 pp 517ndash523 1992

[81] R Stump ldquoAn investigation to describe the population dynam-ics of Acanthaster planci (L) around Lizard Island Cairns Sec-tion Great Barrier Reef Marine Parkrdquo CRC Reef Research Cen-tre Technical Report 10 CRC Reef Research Centre TownsvilleAustralia 1996

[82] N Yasuda S Nagai M Hamaguchi K Okaji K GErard andK Nadaoka ldquoGene flow of Acanthaster planci (L) in relationto ocean currents revealed bymicrosatellite analysisrdquoMolecularEcology vol 18 no 8 pp 1574ndash1590 2009

[83] D C Potts ldquoCrown-of-thorns starfish man induced pest ornatural phenomenonrdquo inThe Ecology of Pests Some AustralianCase Histories R E Kitching and R E Jones Eds pp 55ndash86CSIRO Melbourne Australia 1981

[84] N Yasuda C Taquet S Nagai T Yoshida and M AdjeroudldquoGenetic connectivity of the coral-eating sea star Acanthasterplanci during the severe outbreak of 2006ndash2009 in the SocietyIslands French PolynesiardquoMarine Ecology 2014

[85] C Vogler J A H Benzie K Tenggardjaja P H Barber andGWorheide ldquoPhylogeography of the crown-of-thorns starfishgenetic structure within the Pacific speciesrdquo Coral Reefs vol 32no 2 pp 515ndash525 2013

[86] N K Dulvy R P Freckleton and N V C Polunin ldquoCoralreef cascades and the indirect effects of predator removal byexploitationrdquo Ecology Letters vol 7 no 5 pp 410ndash416 2004

[87] M Kayal J Vercelloni T L de Loma et al ldquoPredator crown-of-thorns starfish (Acanthaster planci) outbreak mass mortality ofcorals and cascading effects on reef fish and benthic communi-tiesrdquo PLoS ONE vol 7 no 10 Article ID e47363 2012

[88] D A Griffin J H Middleton and L Bode ldquoThe tidal andlonger-period circulation of Capricornia Southern Great Bar-rier ReefrdquoAustralian Journal ofMarine and Freshwater Researchvol 38 no 4 pp 461ndash474 1987

[89] A Mantovanelli M L Heron S F Heron and C R SteinbergldquoRelative dispersion of surface drifters in a barrier reef regionrdquoJournal of Geophysical Research C Oceans vol 117 no 11 ArticleID C11016 2012

Submit your manuscripts athttpwwwhindawicom

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Anatomy Research International

PeptidesInternational Journal of


Hindawi Publishing Corporation httpwwwhindawicom

International Journal of

Volume 2014

Zoology


Molecular Biology International

GenomicsInternational Journal of


The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014


BioinformaticsAdvances in

Marine BiologyJournal of



Signal TransductionJournal of


BioMed Research International

Evolutionary BiologyInternational Journal of



Biochemistry Research International

ArchaeaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014


Genetics Research International


Advances in

Virolog y

Hindawi Publishing Corporationhttpwwwhindawicom

Nucleic AcidsJournal of

Volume 2014

Stem CellsInternational



Enzyme Research



Microbiology


numbers of larvae spawned by existing high-density out-breaking populations that are carried to reefs downstream byprevailing currents or from adultmigration between reefs [8]

The first widely documented outbreak population ofCOTS on the GBR was at Green Island in 1962 [17] A varietyof surveys weremade at different times and locations between1966 and 1974 to determine extent of COTS activity on theGBR [2] From these it became apparent that outbreaks beganin the Cairns section of the marine park and spread souththrough the central GBR with time [18 19] as would beexpected if planktonic larva dispersed passively in the watercolumn Systematic surveys over the next two decades con-firmed this pattern for a second wave of outbreaks beginningin the late 1970s and also identified a northward spread ofoutbreaks [20] This pattern had become well established bythe third series of outbreaks beginning in 1993 [21]

Despite the 50-year history of COTS surveys on theGBR the location of primary outbreaks has been difficultto pinpoint To date no primary outbreak on the GBR hasbeen located with any precision even though outbreaksin the Cairns Section first appear on reefs between 145∘and 17∘S [22] While outbreaks were first reported at GreenIsland (1675∘S) near Cairns in 1962 [17] further researchsuggested that this was a secondary outbreak and the originalprimary outbreak(s) had occurred north of Green Island inthe 1950s [23] In 1979 another large outbreak of starfishwas observed on Green Island [24] and survey results andreliable unpublished data indicated that this outbreak wasalso secondarily derived from others located further northaround 16∘S [20] Similarly a third series of outbreaks begin-ning in 1993 was first detected near Lizard Island (145∘S)[15 21] However the exact location of the primary outbreakin 1993 remained uncertain as COTS increased on a numberof nearby reefs about the same time [7] More recent reportsindicated increased COTS activity on reefs near Green Islandfrom around 2010 [25] however COTS numbers have beenslowly building on reefs further north in the vicinity of theLizard Island since 2005 [26] Again the exact location of anyprimary outbreak was uncertain Thus despite an extensivehistory of surveys primary outbreaks of COTS on the GBRremain a matter of conjecture

Information on the location and timing of primaryoutbreaks of COTS could give valuable clues as to the causesPrimary outbreaks are potentially a pressure point for man-aging COTS on the GBR as these initial aggregations lead tosubsequent wide-spread outbreaks that cause such extensivedestruction of coral communities over very large areas [27]Although outbreaks have occurred on the GBR for eightmillennia [28] the frequency of outbreaks in recent decades(in combination with other disturbances such as intensecyclones) is clearly unsustainable leading to speculation thatthe frequency of outbreaks has increased [22] possibly dueto enrichment of GBR waters with nutrients from agriculturein coastal watersheds [14] The evidence that primary COTSoutbreaks on the GBR are linked to enhanced nutrient inputto theGBR lagoon is circumstantial [25]TheGBR is amosaicof over 3000 individual coral reefs stretching nearly 2000 kmfrom Torres Strait in the north (9∘151015840S) to Lady Elliot Islandin the south (24∘071015840S) on the continental shelf of northeast

20∘S

22∘S

24∘S

150∘E 152

∘E 154∘E

N

S

W E

0 50

(km)

200100

Figure 1 The MackayCapricorn Section of the GBR showing thelocation of the Swain Reefs and Capricorn-Bunker Reefs in relationto the coast Isobaths represent depth in meters and show theedge of the continental shelf Arrows show the general position ofthe South Equatorial Current and the East Australian Current andCapricorn EddyThemouth of the Fitzroy River is located southeastof Rockhampton and discharges into Keppel Bay

Australia [29] and because of its large size outbreaksmaywelloccur independently and for different reasons in differentparts of the GBR Based on extensive surveys using mantatows to document the distribution of COTS outbreaks onthe GBR since 1986 [7] we describe a primary outbreakdiscovered on some reefs in the Capricorn-Bunker Group inthe southern GBR We examine the possible causes and weweigh the evidence for links to outbreaks in other parts of theGBR based on the history of outbreaks onCapricorn-BunkerReefs

2 Methods

21 Physical Setting Reefs in the Capricorn-Bunker Groupare the southernmost extent of theGBRand are located 80 kmoffshore on the mid to outer continental shelf [30] Thesereefs are distinctive being mostly medium sized outer-shelfplatform reefs often with lagoons and sand cays [29] To thenorth the continental shelf widens at about 23∘S from about80 km to bemore than 200 kmwide leaving a substantial gapof over 100 km named the Capricorn Channel [31] betweenthe northern extent of the Capricorn-Bunker Reefs and theSwain Reefs The Swain Reefs are contiguous with the restof the GBR and are a series of lagoonal and planar reefslocated offshore on the margin of the continental shelf [29](see Figure 1)

Reefs of the GBR are strongly influenced by the EastAustralian Current (EAC) which is derived from SouthEquatorial Current that crosses the Coral Sea to divide at








3 Results




Year





1986 32 (5) 0134 0062 8 (0) 0012 00121987 30 (7) 033 0126 5 (0) 0 01988 28 (3) 0227 0124 6 (0) 0002 00021989 No survey1990 20 (4) 0272 0207 6 (0) 0 01991 21 (3) 0222 0158 5 (0) 0005 00051992 24 (4) 0340 0218 6 (0) 0 01993 9 (2) 0781 0671 4 (0) 0 01994 11 (2) 0120 0090 3 (0) 0 01995 15 (3) 0142 0079 3 (0) 0006 00061996 16 (3) 0107 0042 5 (0) 0010 00061997 8 (3) 0872 0445 5 (0) 0012 00121998 9 (4) 1230 0646 4 (0) 0 01999 7 (3) 0431 0225 4 (0) 0 02000 8 (3) 2094 0797 4 (0) 0 02001 8 (5) 18775 7371 4 (0) 0005 00052002 7 (4) 4879 2798 4 (0) 0 02003 7 (4) 4077 2227 4 (0) 0 02004 7 (3) 0649 0414 4 (0) 0005 00052005 13 (1) 0050 0033 4 (0) 0 02006 19 (1) 0030 0016 8 (0) 0 02007 6 (0) 0005 0004 4 (0) 0 02008 17 (0) 0 0 7 (0) 0 02009 7 (0) 0011 0011 4 (0) 0 02010 15 (0) 0003 0003 8 (0) 0018 00132011 7 (0) 0002 0002 3 (0) 0 02012 15 (0) 0003 0002 6 (1) 0162 01622013 7 (0) 0023 0018 4 (1) 0066 00662014 14 (0) 0005 0003 8 (2) 0201 0146





Survey year2006 2008 2010 2012 2014

Mea

n CO

TS p

er m

anta

tow

00

02

04

06

08

10

12



Survey (year)

1986

1988

1990

1992

1994

1996

1998

2000

2002

2004

2006

2008

2010

2012

2014

001

01

1

10


Mea

n CO

TS p

er m

anta

tow



Tota

l yea

rly fl

ow (m

egal

itres

)

40

30

20

10

0

times106

1986

1988

1990

1992

1994

1996

1998

2000

2002

2004

2006

2008

2010

2012

Year




4 Discussion



tal e

arly

seas

on fl

ow (m

egal

itres

)

40

30

20

10

0

times106

1986

1988

1990

1992

1994

1996

1998

2000

2002

2004

2006

2008

2010

2012

Year















Acknowledgments


References






































































































Volume 2014

Zoology






















Advances in

Virolog y



Volume 2014




Enzyme Research



Microbiology








3 Results




Year





1986 32 (5) 0134 0062 8 (0) 0012 00121987 30 (7) 033 0126 5 (0) 0 01988 28 (3) 0227 0124 6 (0) 0002 00021989 No survey1990 20 (4) 0272 0207 6 (0) 0 01991 21 (3) 0222 0158 5 (0) 0005 00051992 24 (4) 0340 0218 6 (0) 0 01993 9 (2) 0781 0671 4 (0) 0 01994 11 (2) 0120 0090 3 (0) 0 01995 15 (3) 0142 0079 3 (0) 0006 00061996 16 (3) 0107 0042 5 (0) 0010 00061997 8 (3) 0872 0445 5 (0) 0012 00121998 9 (4) 1230 0646 4 (0) 0 01999 7 (3) 0431 0225 4 (0) 0 02000 8 (3) 2094 0797 4 (0) 0 02001 8 (5) 18775 7371 4 (0) 0005 00052002 7 (4) 4879 2798 4 (0) 0 02003 7 (4) 4077 2227 4 (0) 0 02004 7 (3) 0649 0414 4 (0) 0005 00052005 13 (1) 0050 0033 4 (0) 0 02006 19 (1) 0030 0016 8 (0) 0 02007 6 (0) 0005 0004 4 (0) 0 02008 17 (0) 0 0 7 (0) 0 02009 7 (0) 0011 0011 4 (0) 0 02010 15 (0) 0003 0003 8 (0) 0018 00132011 7 (0) 0002 0002 3 (0) 0 02012 15 (0) 0003 0002 6 (1) 0162 01622013 7 (0) 0023 0018 4 (1) 0066 00662014 14 (0) 0005 0003 8 (2) 0201 0146





Survey year2006 2008 2010 2012 2014

Mea

n CO

TS p

er m

anta

tow

00

02

04

06

08

10

12



Survey (year)

1986

1988

1990

1992

1994

1996

1998

2000

2002

2004

2006

2008

2010

2012

2014

001

01

1

10


Mea

n CO

TS p

er m

anta

tow



Tota

l yea

rly fl

ow (m

egal

itres

)

40

30

20

10

0

times106

1986

1988

1990

1992

1994

1996

1998

2000

2002

2004

2006

2008

2010

2012

Year




4 Discussion



tal e

arly

seas

on fl

ow (m

egal

itres

)

40

30

20

10

0

times106

1986

1988

1990

1992

1994

1996

1998

2000

2002

2004

2006

2008

2010

2012

Year















Acknowledgments


References






































































































Volume 2014

Zoology






















Advances in

Virolog y



Volume 2014




Enzyme Research



Microbiology



Year





1986 32 (5) 0134 0062 8 (0) 0012 00121987 30 (7) 033 0126 5 (0) 0 01988 28 (3) 0227 0124 6 (0) 0002 00021989 No survey1990 20 (4) 0272 0207 6 (0) 0 01991 21 (3) 0222 0158 5 (0) 0005 00051992 24 (4) 0340 0218 6 (0) 0 01993 9 (2) 0781 0671 4 (0) 0 01994 11 (2) 0120 0090 3 (0) 0 01995 15 (3) 0142 0079 3 (0) 0006 00061996 16 (3) 0107 0042 5 (0) 0010 00061997 8 (3) 0872 0445 5 (0) 0012 00121998 9 (4) 1230 0646 4 (0) 0 01999 7 (3) 0431 0225 4 (0) 0 02000 8 (3) 2094 0797 4 (0) 0 02001 8 (5) 18775 7371 4 (0) 0005 00052002 7 (4) 4879 2798 4 (0) 0 02003 7 (4) 4077 2227 4 (0) 0 02004 7 (3) 0649 0414 4 (0) 0005 00052005 13 (1) 0050 0033 4 (0) 0 02006 19 (1) 0030 0016 8 (0) 0 02007 6 (0) 0005 0004 4 (0) 0 02008 17 (0) 0 0 7 (0) 0 02009 7 (0) 0011 0011 4 (0) 0 02010 15 (0) 0003 0003 8 (0) 0018 00132011 7 (0) 0002 0002 3 (0) 0 02012 15 (0) 0003 0002 6 (1) 0162 01622013 7 (0) 0023 0018 4 (1) 0066 00662014 14 (0) 0005 0003 8 (2) 0201 0146





Survey year2006 2008 2010 2012 2014

Mea

n CO

TS p

er m

anta

tow

00

02

04

06

08

10

12



Survey (year)

1986

1988

1990

1992

1994

1996

1998

2000

2002

2004

2006

2008

2010

2012

2014

001

01

1

10


Mea

n CO

TS p

er m

anta

tow



Tota

l yea

rly fl

ow (m

egal

itres

)

40

30

20

10

0

times106

1986

1988

1990

1992

1994

1996

1998

2000

2002

2004

2006

2008

2010

2012

Year




4 Discussion



tal e

arly

seas

on fl

ow (m

egal

itres

)

40

30

20

10

0

times106

1986

1988

1990

1992

1994

1996

1998

2000

2002

2004

2006

2008

2010

2012

Year















Acknowledgments


References






































































































Volume 2014

Zoology






















Advances in

Virolog y



Volume 2014




Enzyme Research



Microbiology


Survey year2006 2008 2010 2012 2014

Mea

n CO

TS p

er m

anta

tow

00

02

04

06

08

10

12



Survey (year)

1986

1988

1990

1992

1994

1996

1998

2000

2002

2004

2006

2008

2010

2012

2014

001

01

1

10


Mea

n CO

TS p

er m

anta

tow



Tota

l yea

rly fl

ow (m

egal

itres

)

40

30

20

10

0

times106

1986

1988

1990

1992

1994

1996

1998

2000

2002

2004

2006

2008

2010

2012

Year




4 Discussion



tal e

arly

seas

on fl

ow (m

egal

itres

)

40

30

20

10

0

times106

1986

1988

1990

1992

1994

1996

1998

2000

2002

2004

2006

2008

2010

2012

Year















Acknowledgments


References






































































































Volume 2014

Zoology






















Advances in

Virolog y



Volume 2014




Enzyme Research



Microbiology


tal e

arly

seas

on fl

ow (m

egal

itres

)

40

30

20

10

0

times106

1986

1988

1990

1992

1994

1996

1998

2000

2002

2004

2006

2008

2010

2012

Year















Acknowledgments


References






































































































Volume 2014

Zoology






















Advances in

Virolog y



Volume 2014




Enzyme Research



Microbiology








Acknowledgments


References






































































































Volume 2014

Zoology






















Advances in

Virolog y



Volume 2014




Enzyme Research



Microbiology




































































































Volume 2014

Zoology






















Advances in

Virolog y



Volume 2014




Enzyme Research



Microbiology

Research Article Origins and Implications of a Primary ...

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