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R E S E A R C H A R T I C L E

D. Lecchini R. Galzin

Spatial repartition and ontogenetic shifts in habitat

use by coral reef fishes (Moorea, French Polynesia)

Received: 18 March 2004 / Accepted: 30 November 2004/ Published online: 18 January 2005 Springer-Verlag 2005

Abstract This study explores the extent to which onto-genetic habitat shifts modify spatial patterns of fishestablished at settlement in the Moorea Island lagoon(French Polynesia). The lagoon of Moorea Island was

divided into 12 habitat zones (i.e. coral seascapes), whichwere distinct in terms of depth, wave exposure, andsubstratum composition. Eighty-two species of recentlysettled juveniles were recorded from March to June2001. Visual censuses documented changes in the dis-tribution of juveniles of each species over time amongthe 12 habitats. Two patterns of juvenile habitat usewere found among species. Firstly, some species settledand remained in the same habitat until the adoption ofthe adult habitats (i.e. recruitment; e.g. Chaetodon citr-inellus, Halichoeres hortulanus, Rhinecanthus aculeatus).Secondly, others settled to several habitats and thendisappeared from some habitats through differential

mortality and/or post-settlement movement (e.g. 6570 mm size class for Ctenochaetus striatus, 4045 mmsize class for Epinephelus merra, 5055 mm size class forScarus sordidus). A comparison of the spatial distribu-tion of juveniles to that of adults (61 species recorded atboth stages) illustrated four patterns of subsequent

recruitment in habitat use: (1) an increase in the numberof habitats used during the adult stage (e.g. H. hortul-anus, Mulloidichthys flavolineatus); (2) a decrease in thenumber of habitats adults used compared to recently

settled juveniles (e.g. Chrysiptera leucopoma, Stethojulisbandanensis); (3) the use of different habitat types(e.g. Acanthurus triostegus, Caranx melampygus); and(4) no change in habitat use (e.g. Naso litturatus, Steg-astes nigricans). Of the 20 most abundant speciesrecorded in Moorea lagoon, 12 species modified thespatial patterns established at settlement by an ontoge-netic habitat shift.

Introduction

The settlement of fish to benthic habitats at the end ofthe larval phase is a critical period in the life cycle ofdemersal fish species (for a recent review, see Leis andMcCormick 2002). Mortality is high during andimmediately after the settlement event and there isstrong selection pressure to choose a microhabitat thatpromotes survival (Connell and Jones 1991; McCor-mick and Hoey 2004). Coral reef fish often showmarked selectivity in the habitats they choose based onthe presence of specific benthic substrata or the pres-ence of conspecifics or other species (e.g. McCormickand Makey 1997; Ohman et al. 1998; Holbrook et al.

2002). Literature suggests that it is often these initialchoices at settlement that determine the patterns ofabundance among habitats for the species (for a recentreview, see Doherty 2002). These initial distributionpatterns established at settlement will be less important,however, when fishes exhibit movement after settlementor are subject to differential mortality related to habitattype, which will modify the spatial patterns of juveniles(e.g. Eggleston 1995; Finn and Kingsford 1996;McCormick and Makey 1997; Dahlgren and Eggleston2000).

Communicated by T. Ikeda, Hakodate

D. Lecchini (&) R. GalzinEcole Pratique des Hautes Etudes,UMR-CNRS 8046, Universite de Perpignan,

66860 Perpignan, FranceE-mail: lecchini@univ-perp.frTel.: +81-98-8952221Fax: +81-98-8958576

D. Lecchini R. GalzinCentre de Recherches Insulaires et Observatoirede lEnvironnement, BP 1013,Moorea, French Polynesia

Present address: D. LecchiniLaboratory of Ecology and Systematics,University of the Ryukyus, 1 Senbaru,Nishihara, 903-0213 Okinawa, Japan

Marine Biology (2005) 147: 4758DOI 10.1007/s00227-004-1543-z

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As an individual grows, its morphology and behav-iour change. For some species this means that they canno longer obtain the resources they require for lifeprocesses and they are faced with one of two choices:they can either die or move to a location that supportstheir life processes. Many studies have documentedsome changes in the spatial distribution of juveniles overtime (e.g. Frederick 1997; Booth 2002), but there are fewstudies that demonstrate positive evidence of themovement from juvenile to adult habitats (see forexceptions Deegan 1990; Sponaugle and Cowen 1996;McCormick and Makey 1997). This lack of informationon ontogenetic habitat shifts between juvenile and adultlife stages may be in part due to the focus of studies onprocesses that influence either juvenile or adult stagesseparately, or to such studies simply not accounting forontogenetic stage (for a recent review, see Gillanders

et al. 2003). Documenting how species change in theirhabitat requirements with ontogeny is crucial to theunderstanding of the processes that will influence pop-ulation abundance patterns and community composi-tion.

The overall objective of the present study was toinvestigate the proportion of tropical reef-associated fishthat exhibit ontogenetic shifts in habitat use during thegrowth of juveniles (i.e. post-settlement movement) andbetween juvenile and adult life stages (i.e. recruitmentstrategy). Recruitment is defined as the integration ofjuveniles into the adult population (Shapiro 1987;Lefevre 1991; Vigliola and Harmelin-Vivien 2001). Thisintegration corresponds to a shift of habitat and/orbehaviour. For example, the juveniles of Rhinecanthusaculeatus settle near the beach, whereas adults livemainly on the barrier reef (data from the present study).

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In this example, recruitment corresponds to a shift ofhabitat. In contrast, both juveniles and adults of Chr-omis viridis live in the same habitat (micro-atoll ofPorites rus at Moorea Island), but the juveniles do notswim as far from their habitat as the adults (Juncker2001). Recruitment corresponds in this example to abehavioural shift related to swimming ability. In thesetwo examples, recruitment reflects a shift in the life cycleof fish that prompts juveniles to join the adult popula-tion.

Specifically we investigated the spatial structure ofjuvenile and adult fish communities on coral reefs atMoorea Island (1) to explore the use of space by a broadrange of fish juvenile taxa during their growth (post-settlement movements), and (2) to document the prev-alence of major habitat shifts between juvenile and adultlife stages (recruitment strategies).

Materials and methods

Study area

Moorea Island (1730S, 1495W) is a high volcanicisland surrounded by a coral reef that delimits a lagoon(Galzin and Pointier 1985). The present study was car-ried out on the Matautia sector (north coast of MooreaLagoon), which can be divided into several habitat zones(i.e. coral seascapes, Fig. 1) distinct in terms of depth,wave exposure, and substratum composition (Chance-relle 1996; Lecchini 2003).

The reef crest and the first two seascapes locatedbehind the reef crest (one characterised by coral rubbleand the other made up of dead coral masses) have notbeen recorded because they correspond to the breaking

zone of oceanic waves. Thus, the first sampled seascape(Sea1) is located at 27 m from reef crest and is made upof only coral rubble (Fig. 1). The second seascape (Sea2)is characterised by a high cover percentage (>30%) ofliving coral masses, Sea3 by a low cover percentage(2 m)and a weak current. Sea5 is defined by a high coverpercentage of dead coral masses (41%) and coral rubble(42%), and Sea6 is a huge living coral mass of Poritesrus (31 m long, 80 m wide, and 2 m high). Sea7 and Sea9are located around the channel (Sea8) with a high depth(2.5 m), a strong current, and a high cover percentage of

living coral (>70%). Lastly, Sea10 is constituted by

many small coral masses (cover percentage >30%),Sea11 by micro-atolls, and Sea12 by sand.

Recruitment size of coral reef fish

If the juveniles shift habitat at recruitment (e.g. Rhin-ecanthus aculeatus), the recruitment size is that of thelargest juvenile in the settlement habitat. If juveniles andadults live in the same habitat (e.g. Chromis viridis), therecruitment size is defined according to the behaviour ofthe fish. The recruitment size of coral reef fish speciesrecorded in the present study (Table 1) was determinedby David Lecchini from February 2001 to January 2002,the period in which he conducted his PhD research atMoorea (Lecchini 2003), and was validated by ProfessorRene Galzin, who has studied the coral reef fish atMoorea since 1975 (Galzin 1977). The size of larvae atcolonisation was also measured by the company Aqua-Fish Technology (director Vincent Dufour), whichcaptured larvae daily at Moorea with crest nets.

Sampling strategy

Diurnal underwater visual counting of fish was carriedout from March to June 2001 along five transects (bandsperpendicular to the coast and continuous from thecoast to the reef crest), 1 km long and 1 m wide (Fig. 1).Fish were identified to species (all species were recordedexcept Gobiidae and Blenniidae) and size class (intervalsof 5 mm). The abundance of each size class of eachspecies was recorded for each shelter use of each sea-scape of each transect. Thus, the spatial repartition andontogenetic shifts in habitat use by coral reef fishes were

investigated at macro- (transect), meso- (coral seascape),and micro-scale (structural characteristics of shelter).The different categories of shelter were sand, coralrubble, coral slab, anemone, sponge, algae (with dis-tinction between macro-algae, encrusting calcareous al-gae, and turf), and coral (with distinction between thecoral form, the coral genus, and its statedead or live).

As fish juveniles can change habitat frequently duringthe benthic phase (McCormick and Makey 1997), eachof the five transects was counted once every semi-lunarcycle (cycle centred on full or new moon). A countingcycle with a rhythm of one transect per day every secondday represented a period of 10 working days per lunar

half-month. Counting of juveniles was carried out at theend of the colonisation season (March to June 2001;Dufour and Galzin 1993) with five consecutive countingcycles (the only cycle not recorded was the full moonduring the month of May). This sampling scheduleallowed for the investigation of post-settlement move-ments of juveniles during their growth (comparison ofhabitat use according to the size of juveniles). Toinvestigate the ontogenetic shifts in habitat use betweenjuveniles and adults (recruitment strategies), adults werecounted at the cycle of the new moon of May 2001.

Fig. 1 A Map of the north coast of Moorea Island (FrenchPolynesia) showing the location of the study area (sector ofMatautia). B Schematic view of the position of different seascapesand of five transects sampled in the Matautia sector. C Schematicview of cross-reef profile showing the relative position of seascapeswith the mean width and the standard error calculated on the fivetransects. MSL Mean sea level

b

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Statistical analysis

Spatial repartition of seascapes of each transect (sam-pling stations) was analysed by using hierarchical clus-tering, which searched the data for distinct clusters ofstations with similar fish communities. The clusteringalgorithm was based on Wards method of minimumvariance clustering. Distribution of fish species was

analysed by using correspondence analysis (CA; lengthof gradient from the initial detrended CA screening stepwas over 3; Benzecri 1973), which compared the relativeabundance of different fish species among seascapes ofeach transect (56 sampling stations). To identify the fishcommunities on CA, Wilks test (Wilks 1932) was used.This statistical test compared variation between com-munities to variation within each community. If varia-tion between communities was significantly greater thanvariation within communities, then the designatedcommunities provided significant information on the

spatial structure of coral reef fish; if not, then commu-nities were considered invalid and the respective com-munities merged.

Results

Spatial pattern in juvenile communities

A total of 16,958 juveniles belonging to 82 species (fivecycles of counting) were recorded in Matautia sector,corresponding to a density of 0.67 juveniles m2 (Fig. 2).The 82 species sampled had a selective distribution in thelagoon with 40% of species settled in one or two sea-scapes and only 7% of species settled in eight or moreseascapes. The repartition of stations in a clusteringalgorithm (Fig. 3) was characterised by four clusters at aWard distance of 1.4 (channel seascape excluded fromdata, no juvenile recorded). The first cluster was con-

Table 1 In the Matautia sector, 95 species of coral reef fishwere recorded at juvenile (J) and/or adult (A) stage by under-water visual counting from March to June 2001. The first size(in millimeters) corresponds to the total length (TL) of fish atcolonisation (estimated to the nearest millimeter from captures

with crest nets) and the second size (in millimeters) correspondsto TL at recruitment [in intervals of 5 mm; (e.g. size 40=3540 mm) estimated from in situ visual observations]. The symbol? means that the size at colonisation or recruitment is un-known

Muraenidae (2 species) Chaetodon ornatissimus (?, 50, J-A) Labroides dimidiatus (7, 40, J-A)Echnida nebulosa (63, ?, J) Chaetodon pelewensis (36, 60, J) Novaculichthys taeniorous (7, 25, J)Gymnothorax javanicus (80, ?, J-A) Chaetodon reticulatus (?, 55, J-A) Pseudocheilinus hexataenia (7, 45, J-A)Synodontidae (2 species) Chaetodon trifascialis (12, 55, J) Stethojulis bandanensis (7, 50, J-A)Saurida gracilis (?, 95, J-A) Chaetodon trifasciatus (15, 50, J-A) Thalassoma amblycephalum (31, 50, J)

Synodus binotatus (56, 75, J) Chaetodon ulietensis (21, 35, A) Thalassoma hardwicke (7, 40, J-A)Aulostomidae (1 species) Chaetodon unimaculatus (19, 60, J-A) Xyrichtys pavo (7, 40, J)Aulostomus chinensis (134, 180, J-A) Chaetodon vagabundus (23, 50, J-A) Scaridae (6 species)Fistulariidae (1 species) Forcipiger longirostris (?, 50, J-A) Leptoscarus vaigiensis (8, 50, J-A)Fistularia commersonii (146, 185, J-A) Heniochus chrysostomus (47, 60, A) Scarus altipinnis (8, 50, J)Scorpaenidae (3 species) Pomacanthidae (1 species) Scarus frenatus (8, 55, J)Pterois antennata (15, 60, J) Centropyge flavissimus (21, 50, J-A) Scarus oviceps (8, 65, J-A)Pterois radiata (15, 30, A) Pomacentridae (13 species) Scarus psittacus (8, 40, J)Scorpaenodes guamensis (11, 60, J) Abudefduf sexfasciatus (16, 40, J-A) Scarus sordidus (8, 65, J-A)Serranidae (3 species) Abudefduf sordidus (16, 30, A) Pinguipedidae (1 species)Cephalopholis argus (24, 80, J) Chromis atripectoralis (10, 30, J-A) Parapercis clathrata (?, 80, J-A)Epinephelus hexagonatus (22, 65, J-A) Chromis margaritifer (11, 35, J-A) Acanthuridae (9 species)Epinephelus merra (19, 65, J-A) Chromis viridis (8, 25, J-A) Acanthurus guttatus (35, ?, A)Apogonidae (1 species) Chromis iomelas (11, ?, A) Acanthurus nigricauda (47, 80, J)Cheilodipterus quinquelineatus (26, 50, J-A) Chrysiptera leucopoma (19, 35, J-A) Acanthurus triostegus (30, 65, J-A)Carangidae (2 species) Dascyllus aruanus (7, 20, J-A) Ctenochaetus striatus (47, 80, J-A)

Caranx melampygus (82, 140, J-A) Dascyllus reticulatus (?, 40, J-A) Naso brevirostris (39, 80, J)Caranx sexfasciatus (80, 120, J) Dascyllus trimaculatus (16, 30, A) Naso lituratus (63, 90, J-A)Lutjanidae (2 species) Pomacentrus pavo (18, 35, J-A) Naso unicornis (54, 90, J-A)Lutjanus fulviflamma (31, 60, A) Stegastes albifasciatus (21, 45, J-A) Zebrasoma scopas (26, 50, J-A)Lutjanus fulvus (49, 70, J-A) Stegastes nigricans (18, 40, J-A) Zebrasoma veliferum (28, 45, J-A)Lethrinidae (2 species) Cirrhitidae (2 species) Zanclidae (1 species)Gnathodentex aurolineatus (45, 65, J-A) Neocirrhites armatus (?, 55, J) Zanclus cornutus (55, 80, A)Monotaxis grandoculis (48, 80, J-A) Paracirrhites arcatus (20, 50, J-A) Siganidae (1 species)Mullidae (5 species) Labridae (16 species) Siganus spinus (61, 100, A)Mulloidichthys flavolineatus (69, 100, J-A) Cheilinus chlororous (7, 55, J-A) Balistidae (2 species)Parupeneus barberinus (69, 90, J) Coris aygula (7, 35, J-A) Rhinecanthus aculeatus (37, 60, J)Parupeneus bifasciatus (68, 100, J-A) Coris gaimard (7, 45, J-A) Balistipus undulatus (30, 75, J-A)Parupeneus cyclostomus (40, 90, J-A) Gomphosus varius (7, 40, J-A) Ostraciidae (2 species)Parupeneus multifasciatus (57, 90, A) Halichoeres hortulanus (7, 50, J-A) Ostracion cubicus (12, 25, J-A)Chaetodontidae (14 species) Halichoeres margaritaceus (7, 40, J-A) Ostracion meleagris (15, 45, J-A)Chaetodon auriga (19, 40, J-A) Halichoeres marginatus (7, 25, J-A) Tetraodontidae (3 species)

Chaetodon citrinellus (29, 55, J-A) Halichoeres trimaculatus (7, 40, J-A) Canthigaster janthinoptera (30, 55, J-A)Chaetodon ephippium (17, 40, A) Labroides bicolor (7, 25, J) Canthigaster solandri (21, 55, J-A)Chaetodon lunula (21, 60, J-A) Canthigaster valentini (13, 45, J)

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stituted by the stations of seascape 1, the second by thestations of seascapes 2 to 6. A third cluster was consti-tuted by the stations of seascapes 7 to 11, and a fourthby the stations of seascape 12. The distribution of spe-cies from a CA was characterised by a Guttman effectthat can be divided into four communities (Fig. 4). TheWilks test validated the distribution of coral reef fishspecies in four communities, corresponding with thefour clusters of stations, more for some ubiquitousspecies (lambda

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reflected by three adult communities: reef crest, barrierreef, and fringing reef (Wilks test, lambda

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found among species (Fig. 7). Firstly, some species set-tled and remained in the same habitat until the adoptionof the adult habitats (e.g. Chaetodon citrinellus, Hali-choeres hortulanus, Rhinecanthus aculeatus, Thalassomahardwicke). Secondly, others settled to several seascapesbut disappeared, from a determined size, from some

seascapes through differential mortality and/or post-settlement movement (e.g. 6570 mm for Ctenochaetusstriatus, 4045 mm for Epinephelus merra, 5055 mm forScarus sordidus). The juveniles ofC. striatus from the sizeclass 47 (colonisation size) to 70 mm settled on all sea-scapes (except the channel). The juveniles from the sizeclass 70 to 80 mm (recruitment size) were only presentfrom seascapes 1 to 9, probably due to differential mor-tality and/or post-settlement movement. For E. merra(from 4045 mm size class), there was post-settlementmovement (from fringing to barrier reef), possibly asso-ciated with a differential mortality. From March toApril, the juveniles ofE. merra were mainly recorded onthe fringing reef with a size range from 21 (colonisationsize) to 40 mm, and from May to June, they were mainlyrecorded on the barrier reef with a size over 40 mm.

Discussion

Spatial structure of juvenile fish communities

In the Matautia sector, the juveniles of coral reef fishwere dispatched among four communities spatially

distributed according to the distance from the reef crest.Hypothetically, this pattern could be explained by cur-rent, depth, or turbidity gradients. Due to the shallow-ness in Matautia lagoon (maximum of 3 m depth;Galzin and Pointier 1985), the depth hypothesis is re-jected. Similarly, the studied area has no turbidity gra-dient because water quality on the north coast ofMoorea is very good due to the absence of major humanpollution, the shallowness of the lagoon, and numerouspasses that maintain a quick renewal of water in the

lagoon (from 0.4 to 1.3 m s

1

; Renon 1989). As a con-sequence, current gradient (current speed is high nearthe reef crest and decrease towards the beach) should bethe best hypothesis to explain the spatial distribution ofthe juveniles.

Nevertheless, changes among seascapes in some bio-logical factors, such as percentage of living cover, per-centage cover of each substratum type, and substratumdiversity and complexity, could increase or decrease theinfluence of current on juvenile distribution. Thus, thedifferentiation of the reef crest community (vs barrierreef community) would be due to the topographiccharacteristics of seascape 1, which is made of coralrubble. The absence of coral reef patches on this sea-scape, different from other barrier reef seascapes (barrierreef community), means that high current action fromoceanic waves is not obstructed, allowing mainly thesmaller species (which hide themselves among coralrubble) and species with high swimming abilities (whichfight against the current without protecting themselvesbehind a coral reef patch) to occupy the habitat. Themain target species of the reef crest community meetingthese criteria are Halichoeres margaritaceus, Corisaygula, C. gaimard, and Thalassoma amblycephalum, In

Fig. 4 Correspondence analysis (CA) showing spatial distributionof juvenile species. Plots of the CA show ordination along the firsttwo axes (the inertia of each axis is given). Species are coded by thefirst two letters of name of genus and species

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addition, the Pomacentrid Chrysiptera leucopoma, whichhas limited swimming abilities (personal observation),was also found. Subsisting near the bottom, this speciesneeds less speed to swim against the current due tofriction effects (Wood and Bain 1995). This physicalphenomenon allows some species to occupy seascapes athigh current speeds when their swimming abilities wouldnot normally allow them to do so (Gerstner 1998).

The differentiation of the beach community (vs thefringing reef community) is also related to the low

structural complexity of seascape 12 but the currentaction is not accentuated here. This seascape is a shallowsandy area without coral shelter (different from otherfringing reef seascapes) and with low current. Thesecharacteristics favour the presence of transient specieswith high swimming abilities to flee predators. The maintarget species of this community are Acanthurus trios-tegus, Caranx melampygus, and Rhinecanthus aculeatus.

To conclude, the communities structure, spatiallyordered according to their distance from the reef crest,could be explained primarily by a current gradient

(current speed is high near the reef crest and decreasestowards the beach), the topographic characteristics ofsome coral seascapes (Sea1 and Sea12), and the positionof the species in the water column. Other factors could,nevertheless, influence where a fish settles, and where thejuveniles live, such as morphological/functional com-ponents of species, components involving the food theycan exploit, and ecological factors (e.g. facilitation,intra/interspecific competition, and predation).

Ontogenetic shifts in habitat use

Two patterns of shifts in habitat use were found duringthe growth of juveniles at Moorea Island. Firstly, somespecies settled and remained in the same habitat until theadoption of the adult habitats. Secondly, others settledto several habitats but disappeared from some habitatsthrough differential mortality and/or post-settlementmovement. Thus, post-settlement behaviour of fish hasthe potential to alter the distribution of juveniles

Fig. 5 Hierarchical clusteringanalysis showing the spatialrepartition of 56 stations whenthe adults were recorded

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established at settlement. The post-settlement behaviourcan, nevertheless, vary according to coral reef fish fam-ilies. In our study, the 4 Pomacentridae species belong-ing to the 20 most abundant species (Abudefdufsexfasciatus, Chrysiptera leucopoma, Stegastes albifasci-atus, and S. nigricans) did not distort their settlementsignal by post-settlement movement. Among Labridae, 3of 5 species belonging to the 20 most abundant speciesdistorted their settlement signal (Gomphosus varius,

Halichoeres margaritaceus, H. trimaculatus). The lowpost-settlement dispersal rate seems to be characteristicof Pomacentridae (Sponaugle and Cowen 1996; Booth2002) and may help them reduce predation risk (Beukersand Jones 1998). Sponaugle and Cowen (1996) showedthat older juvenile distribution of the damselfish Steg-astes partitus closely reflected settlement patterns, whileolder juveniles of the surgeonfish Acanthurus bahianuswere most abundant at sites of lower settlement, whichhad a more suitable habitat. Whatever the fish familystudied (Pomacentridae, Acanthuridae, Labridae, etc.),differences in the distribution of size classes among dif-ferent habitats have been frequently used to infer post-

settlement movement (see Gillanders et al. 2003). How-ever, other explanations may also account for spatiallyexplicit patterns of organism size distribution amonghabitats. For example, differences in growth rates (i.e.growth could have an influence through post-settlementsurvival in the face of a selective mortality regime) ordifferential mortality among habitats may result in pat-terns similar to those attributable to post-settlementmovement. Our data on Ctenochaetus striatus, Epi-nephelus merra, and Scarus sordidus were not able tohighlight if post-settlement movement or differential

mortality is more important (Fig. 7). But, in both cases(differential mortality and/or post-settlement move-ment), the settlement signal is lost before recruitmentand post-settlement behaviour alters the distribution ofjuveniles and subsequently adults.

The settlement signal can also be lost by therecruitment strategy of fish. Deegan (1990) sampledfour locations in juvenile habitat (marsh) and fourlocations in adult habitat (open bay). He found that

density of Gulf menhaden Brevoortia patronus was thehighest in tidal creeks (average fish length 3040 mm)and that abundances in creeks declined when fish grewto 4550 mm, when they likely moved to an open bayarea and then offshore. The comparative analysis ofspatial distribution of juveniles and adults in thepresent study highlighted four recruitment strategies:(1) an increase in the number of habitats used duringthe adult stage; (2) a decrease in the number of hab-itats adults used compared to recently settled juveniles;(3) no change in habitat use; and (4) use of nurseryareas by juveniles followed by extensive movement toan adult habitat. This last ontogenetic habitat shift, to

settle to the shallows and then move as they got older,is a general trend in the literature for many temperateand tropical species (e.g. Parrish 1989; Nagelkerkenet al. 2002). Among the 20 most abundant species(Table 2), the 5 species that used different habitattypes for juvenile and adult stages (Abudefduf sexfas-ciatus, Acanthurus triostegus, Caranx melampygus,Gnathodentex aurolineatus, and Rhinecanthus aculeatus)settled on seascapes 10, 11, and 12, defined by Lefevre(1991) as the nursery area on the north coast ofMoorea Island.

Fig. 6 Correspondenceanalysis showing the spatialdistribution of adult species

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To conclude, the present study is the first to explorethe use of space by a broad range of fish taxa fromsettlement to recruitment (i.e. at the juvenile stage), andto document the prevalence of major habitat shifts be-tween juvenile and adult life stages. This multi-species

perspective offers a broader survey of the varied ways inwhich fish can redistribute themselves post-settlementthan has been done in previous work (see Gillanderset al. 2003). Thus among the 20 most abundant speciesrecorded in Moorea lagoon, 12 species distorted spatialpatterns of fish established at settlement by some onto-genetic habitat shifts (two patterns of post-settlementmovement, four recruitment strategies). Although thisstudy was carried out at a single site transect across theMoorea lagoon, other studies support the generality ofontogenetic habitat use (see Gillanders et al. 2003). Thus

the prevalence of ontogenetic habitat shifts highlightedin the present study has a substantial implication interms of interpreting data on fish settlement to the reef.Measuring and describing settlement rates and patterns(i.e. selective choice of habitat, high mortality, strong

competition between settling fish and resident fish) haslong been a focus of reef fish ecology (see Doherty 2002).Our results suggest, however, that the settlement pat-terns can be misleading over time when fish survivedifferentially or redistribute themselves among habitats.

Acknowledgements The authors wish to thank C. Rua, A. Lo-Yat,and S. Planes for their comments on the first draft, J. Algret and Y.Chancerelle for their assistance in the field, and M. McCormick forhaving checked the English language. The first author also thanksthe French Ministry of Foreign Affairs for the Lavoisier Fellow-ship.

Table 2 Distribution in density (number of fish per square meter)along the different seascapes of the 20 most abundant fish speciessampled in the Matautia sector, to explore ontogenetic shifts inhabitat use by juveniles at recruitment (comparison of habitat useof juveniles and adults). Four recruitment strategies have been

highlighted: an increase in the number of seascapes used duringthe adult stage (Inc); a decrease in the number of seascapes adultsused compared to recently settled juveniles (Dec); the use of dif-ferent seascape types (Dif); no change in seascape use by species(NC)

Species Samplingstage

No. fishcounted

Sea1 Sea2 Sea3 Sea4 Sea5 Sea6 Sea7 Channel Sea9 Sea10 Sea11 Sea12 Strategyofrecruitment

Abudefdufsexfasciatus

Juvenile 86 0.053 0.051 Dif Adults 34 0.02 0 .002 0.003 0.007 0.008 0.012

Acanthurustriostegus

Juvenile 31 0.011 Dif Adult 111 0.105 0.003 0.007 0.086 0.011 0.002

Caranxmelampygus

Juvenile 51 0.003 0.018 Dif Adult 12 0.006 0.001 0.008

Chaetodoncitrinellus

Juvenile 199 0.029 0.025 0.031 0.022 0.011 0.005 0.007 0.002 0.001 0.001 NCAdult 34 0.005 0.006 0.003 0.003 0.006 0.009 0.005 0.003 0.003

Chrysipteraleucopoma

Juvenile 243 0.124 0.003 0.002 0.01 0.006 0.003 0.001 DecAdult 18 0.015 0.004 0.004

Ctenochaetusstriatus

Juvenile 5,017 0.534 0.369 0.591 0.403 0.356 0.156 0.502 0.122 0.202 0.189 0.130 DecAdult 80 0.041 0.048 0.059 0.027 0.032 0.009 0.042 0.031

Epinephelusmerra

Juvenile 149 0.002 0.01 0.005 0.014 0.012 0.013 0.023 0.024 NCAdult 12 0.003 0.001 0.001 0.008 0.002 0.001 0.003

Gnathodentexaurolineatus

Juvenile 210 0.053 0.145 Dif Adult 18 0.002 0.003 0.004 0.003 0.017 0.009 0.007

Gomphosus

varius

Juvenile 322 0.095 0.097 0.115 0.033 0.013 0.004 0.008 0.003 NC

Adult 26 0.011 0.006 0.003 0.005 0.003 0.014 0.004 0.008Halichoeres

hortulanusJuvenile 154 0.044 0.022 0.011 0.003 0.014 IncAdult 53 0.008 0.004 0.008 0.003 0.006 0.003 0.007 0.003 0.002

Halichoeresmargaritaceus

Juvenile 279 0.166 0.026 NCAdult 8 0.007

Halichoerestrimaculatus

Juvenile 558 0.030 0.006 0.013 0.016 0.019 0.033 0.022 0.013 0.060 0.002 NCAdult 45 0.008 0.007 0.010 0.004 0.005 0.007 0.008 0.002 0.011

Mulloidichthysflavolineatus

Juvenile 290 0.01 0.03 0.01 0.038 0.032 IncAdult 45 0.002 0.011 0.008 0.006 0.006 0.016 0.005 0.013 0.008 0.013

Naso litturatus Juvenile 194 0.047 0.015 0.035 0.016 0.013 0.007 0.009 0.004 0.003 0.006 0.01 NCAdult 30 0.012 0.004 0.023 0.004 0.004 0.001 0.009 0.004 0.008

Rhinecanthusaculeatus

Juvenile 32 0.051 Dif Adult 11 0.048 0.002

Scarus sordidus Juvenile 1,013 0.023 0.007 0.013 0.029 0.066 0.044 0.063 0.050 0.107 0.013 DecAdult 52 0.023 0.018 0.017 0.007 0.018 0.020

Stegastes

albifasciatus

Juvenile 1,072 0.012 0.047 48.000 0.078 0.067 0.117 0.018 NC

Adult 25 0.029 0.037 0.026 0.036 0.027Stegastes

nigricansJuvenile 203 0.009 0.014 0.011 0.014 0.037 0.011 0.017 NCAdult 44 0.024 0.032 0.068 0.015 0.032 0.001

Stethojulisbandanensis

Juvenile 249 0.05 0.033 0.013 0.017 0.029 0.003 0.007 0.007 0.003 0.009 DecAdult 56 0.008 0.002 0.011 0.005 0.005 0.009 0.003

Thalassomahardwicke

Juvenile 1,177 0.055 0.183 0.108 0.213 0.085 0.024 0.02 0.018 0.005 0.002 NCAdult 83 0.032 0.023 0.022 0.013 0.026 0.027 0.026 0.02 0.019 0.009

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