Chapter 9: Persistence of grouper (Serranidae) <strong>spawning</strong>aggregations at high levels of habitat disturbanceJan Robinson, Calvin Gerry and Jude BijouxIntroductionTargeted fishing constitutes the major threat to reef fish <strong>spawning</strong> aggregations (Sadovy andDomeier 2005; Sadovy de Mitcheson and Erisman 2012). As a consequence, the majority ofstudies conducted on these vulnerable life history stages have focused primarily on fishing impacts(Domeier and Colin 1997; Russell et al. 2012). In addition to fishing, the formation of <strong>spawning</strong>aggregations can also be compromised by degradation or loss of benthic reef habitat resulting fromthe use of destructive fishing gears, coastal development (e.g. reclamation), or natural disturbances,such as severe storms (Koenig et al. 2000; Sadovy and Domeier 2005; Robinson et al. 2007).However, few studies have explicitly documented the effects of habitat change, caused by naturalor anthropogenic disturbances, on <strong>spawning</strong> aggregation formation and status.The processes that determine why, where and when <strong>spawning</strong> aggregations form remain largelyunknown, although several hypotheses have been proposed (Colin 2012). Predator evasion(Shapiro et al. 1988), egg predation (Johannes 1978; Lobel 1978) and dispersal (Barlow 1981;Doherty et al. 1985), and larval retention (Johannes 1978; Jones et al. 2005; Almany et al. 2007;Karnauskas et al. 2011) and survival (Robertson 1990) have all been identified as mechanismsconferring selective advantage of aggregative <strong>spawning</strong> at specific locations and times (Claydon2004; Molloy et al. 2012). Specific timing and locations of <strong>spawning</strong> may also serve as simplecues to synchronise reproduction and confer no other selective advantage (Claydon 2004).Depending on the hypothesis, benthic habitat will act as a primary (e.g. in terms of predatorevasion), lesser or even negligible factor in aggregation site selection. However, it is importantto consider that different processes can operate on ecological and evolutionary time-scales (e.g.Colin 2012). The processes that confer selective advantage and lead to the establishment of stable<strong>spawning</strong> sites operate on evolutionary scales and are likely to differ from ecological processes thatmaintain persistence at established sites. Those ecological processes include social behaviour (e.g.learning: Warner 1988, 1990) that are thought to enable fish to migrate to and attend <strong>spawning</strong>events at established locations. Moreover, in certain species, benthic habitat complexity may playan important role in providing shelter (Beets and Friedlander 1992; Johannes et al. 1999) andsubstrate for territorial/courtship behaviour, influencing fine-scale spatial distribution, abundanceand density of aggregations and, potentially, their persistence.The existence of multi-species aggregation sites comprised of broad phylogenies highlights variationin the importance of benthic habitat among aggregative spawners. For example, at Gladden Spiton the Belize Barrier Reef, at least 17 species aggregate to spawn, including numerous speciesfrom the families Carangidae, Lutjanidae and Serranidae (Heyman and Kjerfve 2008). Thoughall aggregating species documented at Gladden Spit form aggregations associated with identicalreef geomorphology, namely a pronounced reef promontory with strong currents, they vary intheir association with benthic habitat, with some, such as the carangids, being semi-pelagic. Thus,while benthic habitat may provide aggregating serranids with shelter from predators (Beets andFriedlander 1992; Johannes et al. 1999), this is clearly not the case for many lutjanids and carangidsthat aggregate partially or fully well above the reef substrate.Manipulative experiments and natural disturbances operating on ecological time-scales may offeruseful insights on the mechanisms underlying <strong>spawning</strong> aggregation site selection and persistence.Controlled experiments are rare, but the widespread overfishing of <strong>spawning</strong> aggregations isunfortunately common (Sadovy de Mitcheson et al. 2008). Recovery from depletion or othernon-destructive effects of fishing is clearly possible (Beets and Friedlander 1998; Nemeth 2005),86
ut the potential for recovery at the same site following ‘complete’ eradication remains unknown(Sadovy and Domeier 2005) and may operate on time-scales not suited for study. By contrast, themechanisms underlying the persistence of established <strong>spawning</strong> aggregations are more amenableto study. Here, we report on the impacts of a natural disturbance on the persistence of grouper<strong>spawning</strong> aggregation site at Farquhar atoll in Seychelles.In December 2006, a cyclone passed directly over Farquhar atoll in the southwest corner of theSeychelles archipelago at which <strong>spawning</strong> aggregations are known to form in the months ofDecember to February. Research conducted at a key <strong>spawning</strong> site at the atoll between 2003 and2006 verified <strong>spawning</strong> aggregation by Epinephelus polyphekadion and Epinephelus fuscoguttatus(Robinson et al. 2008). The objectives of this study were to (1) assess the impacts of the 2006 cycloneon <strong>spawning</strong> aggregation habitat at the site and (2) determine whether <strong>spawning</strong> aggregations ofEpinephelus polyphekadion and Epinephelus fuscoguttatus continue to form at the site following thedisturbance.MethodsFamiliarisation dives conducted at the study site in January 20<strong>10</strong> on lunar days (LD) 18 and 19revealed that aggregation habitat had changed dramatically since the most recent site survey ofNovember 2006 (Robinson et al. 2008). Using a November 2006 geo-referenced Google Earthimage imported into ArcGIS, changes in <strong>spawning</strong> habitat and area usage between the 2003-2006 period and 20<strong>10</strong> were examined. GPS positions that marked the perimeter of <strong>spawning</strong>aggregation areas during previous assessments (i.e. between 2003 and 2006; Robinson et al.2008b) were reconciled with the outline of reefs in the 2006 imagery. Extrapolations were madewhere cloud cover obscured reefs. Over 2 days (LDs 28 and 29) in January 20<strong>10</strong>, a period close tothe known <strong>spawning</strong> time for both species (Robinson et al. 2008b), the core <strong>spawning</strong> areas werereassessed by divers. As with earlier surveys (Robinson et al. 2008b), we considered the core of theaggregations to be areas where obvious signs of <strong>spawning</strong> behaviour were observed, as opposed toareas (‘boundary areas’; Robinson et al. 2008b) that had high densities of aggregating fish but nosigns of <strong>spawning</strong> behaviour. Signs of <strong>spawning</strong> behaviour were primarily the presence of gravidfemales being guarded and courted by territorial males. The extent of boundary reef areas was notdetermined in 20<strong>10</strong>.Fixed transects were used to survey aggregations between 2003 and 2006 (Robinson et al. 2008b).Following the disturbance, one of three fixed transects in the core was completely lost, while asecond was partially lost. As a result, random point counts (7-m radius) were introduced in 20<strong>10</strong> inorder to confirm that aggregations of E. fuscoguttatus and E. polyphekadion still formed following thedisturbance. Ten counts were conducted per census across reefs where signs of <strong>spawning</strong> behaviourwere observed. In order to use this method for E. fuscoguttatus, censuses were only performed afew days (LD 27-29) before <strong>spawning</strong>, when gravid females were present and territorial maleswere generally tolerant of divers. Censuses of E. polyphekadion, which allow divers to approachclosely, began earlier on LD 20. In 20<strong>10</strong>, bad weather prevented diving between LD 23 and 26,inclusive. Density estimates of E. fuscoguttatus and E. polyphekadion aggregations made in 20<strong>10</strong>were compared qualitatively with density estimates from aggregations observed in January 2004,since formal statistical comparisons were invalidated by the change in sampling methodology (i.e.from fixed to random sampling units).To determine if reproductive behaviour was affected by the habitat disturbance, the relativefrequency of occurrence (RFOO) of <strong>spawning</strong>-related signs and behaviours, i.e. aggression,courtship, gravid females and gamete release, was assessed during the census. RFOO of behaviour(e.g. courtship) is the ratio of the number of fish showing that behaviour to the total number offish observed in the point count (Pet et al. 2005).87
- Page 4:
The designation of geographical ent
- Page 9:
Chapter 1: IntroductionJan Robinson
- Page 12 and 13:
limited, subsistence levels of expl
- Page 14:
NTRs for spawning aggregations usin
- Page 17 and 18:
al. 2003). Verification may include
- Page 19 and 20:
a fraction of spawning sites are pr
- Page 21 and 22:
Chapter 3: Targeted fishing of the
- Page 23 and 24:
verifying spawning aggregations, we
- Page 25 and 26:
ecorded from inshore close to the c
- Page 27 and 28:
(a)(b)Fig. 3. Spatial patterns ofca
- Page 30 and 31:
pooled sizes of the three spawning
- Page 32 and 33:
found S. sutor contributed up to 44
- Page 34 and 35:
2011b). However, observations of fi
- Page 36 and 37:
MethodsTo identify seasonal and lun
- Page 38 and 39:
n=199Females GSI (mean ± SE)2.521.
- Page 40 and 41:
The estimate of size at maturity in
- Page 42 and 43:
This study was designed to verify S
- Page 44 and 45: were selected. Fish selected for ta
- Page 46 and 47: The number of traps increased on th
- Page 48 and 49: Of the 9 tagged fish detected by re
- Page 50 and 51: Fig. 7. Diel patterns ofdetection f
- Page 52 and 53: Spawning aggregation site fidelity
- Page 54 and 55: Chapter 6: Shoemaker spinefoot rabb
- Page 56 and 57: anterior of the anus and below the
- Page 58 and 59: A high percentage (80.8%) of depart
- Page 60 and 61: arrivals and departures at these tw
- Page 62 and 63: are typically applied for reef fish
- Page 64 and 65: (a)(b)(c)Chapter 3, Figure 3. Spati
- Page 66 and 67: (1)(2)(3)(4)(5)(6)Chapter 7, Table
- Page 68 and 69: Chapter 12, Fig. 1 Fraction of fema
- Page 70 and 71: Plates 8. Selected photographs from
- Page 72 and 73: MethodsStudy sitesThe study area wa
- Page 74 and 75: which shelved gently ( ca. 25 o ) t
- Page 76 and 77: Fig. 4. Lunar periodicity in number
- Page 78 and 79: Behaviour and appearanceDescription
- Page 80 and 81: eported aggregations forming betwee
- Page 82 and 83: The sizes of E. fuscoguttatus aggre
- Page 84 and 85: Materials and methodsStudy area and
- Page 86 and 87: TL. All fish tagged were considered
- Page 88 and 89: Lunar timing of arrivals and depart
- Page 90 and 91: Fig. 8. The presence and absence of
- Page 92 and 93: aggregation fishing. This critical
- Page 96 and 97: ResultsBetween 2003 and 2006, the c
- Page 98 and 99: Fig. 2. Mean (± standard error, SE
- Page 100 and 101: A few species (e.g. Epinephelus gut
- Page 102 and 103: (a)(b)Fig. 1. Map of (a) study site
- Page 104 and 105: Fig. 2. Number of E. lanceolatus ob
- Page 106 and 107: was having any impact on the popula
- Page 108 and 109: A spawning aggregation is said to o
- Page 110 and 111: Chapter 11: Evaluation of an indica
- Page 112 and 113: Table 1 Aggregation fisheries asses
- Page 114 and 115: the lists of Jennings et al. (1999)
- Page 116 and 117: with the more vulnerable labrids an
- Page 118 and 119: The remaining serranid populations
- Page 120 and 121: Spawning aggregation behaviour is c
- Page 122 and 123: tiger grouper, Mycteroperca tigris:
- Page 124 and 125: of protecting the normal residence
- Page 126 and 127: • Since grouper males are afforde
- Page 128 and 129: Fig. 2 Yield-per-recruit normalized
- Page 130 and 131: The approaches identified above are
- Page 132 and 133: during full moon periods. Siganus s
- Page 134 and 135: model, many parameter estimates are
- Page 136 and 137: ReferencesAbunge C (2011) Managing
- Page 138 and 139: Cox DR (1972) Regression models and
- Page 140 and 141: Grüss A, Kaplan DM, Hart DR (2011b
- Page 142 and 143: Kaunda-Arara B, Rose GA (2004a) Eff
- Page 144:
Newcomer RT, Taylor DH, Guttman SI
- Page 147 and 148:
Sancho G, Petersen CW, Lobel PS (20
- Page 149 and 150:
Appendix 1. QuestionnaireMASMA SPAW
- Page 151 and 152:
8. Spawning aggregation knowledgeUs
- Page 153 and 154:
Example items KSh Furthest site Clo
- Page 155 and 156:
Appendix II. Experimental testing o
- Page 157 and 158:
Clove oil concentrationAt a concent
- Page 159 and 160:
Appendix III. Application of acoust
- Page 161 and 162:
153