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Marine Biology (2005) 147: 4758 DOI 10.



D. Lecchini R. Galzin

Spatial repartition and ontogenetic shifts in habitat use by coral reef shes (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 ontogenetic habitat shifts modify spatial patterns of sh established at settlement in the Moorea Island lagoon (French Polynesia). The lagoon of Moorea Island was divided into 12 habitat zones (i.e. coral seascapes), which were distinct in terms of depth, wave exposure, and substratum composition. Eighty-two species of recently settled juveniles were recorded from March to June 2001. Visual censuses documented changes in the distribution of juveniles of each species over time among the 12 habitats. Two patterns of juvenile habitat use were found among species. Firstly, some species settled and remained in the same habitat until the adoption of the adult habitats (i.e. recruitment; e.g. Chaetodon citrinellus, Halichoeres hortulanus, Rhinecanthus aculeatus). Secondly, others settled to several habitats and then disappeared from some habitats through dierential mortality and/or post-settlement movement (e.g. 65 70 mm size class for Ctenochaetus striatus, 4045 mm size class for Epinephelus merra, 5055 mm size class for Scarus sordidus). A comparison of the spatial distribution of juveniles to that of adults (61 species recorded at both stages) illustrated four patterns of subsequent

recruitment in habitat use: (1) an increase in the number of habitats used during the adult stage (e.g. H. hortulanus, Mulloidichthys avolineatus); (2) a decrease in the number of habitats adults used compared to recently settled juveniles (e.g. Chrysiptera leucopoma, Stethojulis bandanensis); (3) the use of dierent habitat types (e.g. Acanthurus triostegus, Caranx melampygus); and (4) no change in habitat use (e.g. Naso litturatus, Stegastes nigricans). Of the 20 most abundant species recorded in Moorea lagoon, 12 species modied the spatial patterns established at settlement by an ontogenetic habitat shift.

The settlement of sh to benthic habitats at the end of the larval phase is a critical period in the life cycle of demersal sh species (for a recent review, see Leis and McCormick 2002). Mortality is high during and immediately after the settlement event and there is strong selection pressure to choose a microhabitat that promotes survival (Connell and Jones 1991; McCormick and Hoey 2004). Coral reef sh often show marked selectivity in the habitats they choose based on the presence of specic benthic substrata or the presence of conspecics or other species (e.g. McCormick and Makey 1997; Ohman et al. 1998; Holbrook et al. 2002). Literature suggests that it is often these initial choices at settlement that determine the patterns of abundance among habitats for the species (for a recent review, see Doherty 2002). These initial distribution patterns established at settlement will be less important, however, when shes exhibit movement after settlement or are subject to dierential mortality related to habitat type, which will modify the spatial patterns of juveniles (e.g. Eggleston 1995; Finn and Kingsford 1996; McCormick and Makey 1997; Dahlgren and Eggleston 2000).

Communicated by T. Ikeda, Hakodate D. Lecchini (&) R. Galzin Ecole Pratique des Hautes Etudes, UMR-CNRS 8046, Universite de Perpignan, 66860 Perpignan, France E-mail: Tel.: +81-98-8952221 Fax: +81-98-8958576 D. Lecchini R. Galzin Centre de Recherches Insulaires et Observatoire de lEnvironnement, BP 1013, Moorea, French Polynesia Present address: D. Lecchini Laboratory of Ecology and Systematics, University of the Ryukyus, 1 Senbaru, Nishihara, 903-0213 Okinawa, Japan


As an individual grows, its morphology and behaviour change. For some species this means that they can no longer obtain the resources they require for life processes and they are faced with one of two choices: they can either die or move to a location that supports their life processes. Many studies have documented some changes in the spatial distribution of juveniles over time (e.g. Frederick 1997; Booth 2002), but there are few studies that demonstrate positive evidence of the movement from juvenile to adult habitats (see for exceptions Deegan 1990; Sponaugle and Cowen 1996; McCormick and Makey 1997). This lack of information on ontogenetic habitat shifts between juvenile and adult life stages may be in part due to the focus of studies on processes that inuence either juvenile or adult stages separately, or to such studies simply not accounting for ontogenetic stage (for a recent review, see Gillanders

et al. 2003). Documenting how species change in their habitat requirements with ontogeny is crucial to the understanding of the processes that will inuence population abundance patterns and community composition. The overall objective of the present study was to investigate the proportion of tropical reef-associated sh that exhibit ontogenetic shifts in habitat use during the growth of juveniles (i.e. post-settlement movement) and between juvenile and adult life stages (i.e. recruitment strategy). Recruitment is dened as the integration of juveniles into the adult population (Shapiro 1987; Lefevre 1991; Vigliola and Harmelin-Vivien 2001). This integration corresponds to a shift of habitat and/or behaviour. For example, the juveniles of Rhinecanthus aculeatus settle near the beach, whereas adults live mainly on the barrier reef (data from the present study).


In this example, recruitment corresponds to a shift of habitat. In contrast, both juveniles and adults of Chromis viridis live in the same habitat (micro-atoll of Porites rus at Moorea Island), but the juveniles do not swim as far from their habitat as the adults (Juncker 2001). Recruitment corresponds in this example to a behavioural shift related to swimming ability. In these two examples, recruitment reects a shift in the life cycle of sh that prompts juveniles to join the adult population. Specically we investigated the spatial structure of juvenile and adult sh communities on coral reefs at Moorea Island (1) to explore the use of space by a broad range of sh juvenile taxa during their growth (postsettlement movements), and (2) to document the prevalence of major habitat shifts between juvenile and adult life stages (recruitment strategies).

many small coral masses (cover percentage >30%), Sea11 by micro-atolls, and Sea12 by sand. Recruitment size of coral reef sh If the juveniles shift habitat at recruitment (e.g. Rhinecanthus aculeatus), the recruitment size is that of the largest juvenile in the settlement habitat. If juveniles and adults live in the same habitat (e.g. Chromis viridis), the recruitment size is dened according to the behaviour of the sh. The recruitment size of coral reef sh species recorded in the present study (Table 1) was determined by David Lecchini from February 2001 to January 2002, the period in which he conducted his PhD research at Moorea (Lecchini 2003), and was validated by Professor Rene Galzin, who has studied the coral reef sh at Moorea since 1975 (Galzin 1977). The size of larvae at colonisation was also measured by the company AquaFish Technology (director Vincent Dufour), which captured larvae daily at Moorea with crest nets. Sampling strategy Diurnal underwater visual counting of sh was carried out from March to June 2001 along ve transects (bands perpendicular to the coast and continuous from the coast to the reef crest), 1 km long and 1 m wide (Fig. 1). Fish were identied to species (all species were recorded except Gobiidae and Blenniidae) and size class (intervals of 5 mm). The abundance of each size class of each species was recorded for each shelter use of each seascape of each transect. Thus, the spatial repartition and ontogenetic shifts in habitat use by coral reef shes were investigated at macro- (transect), meso- (coral seascape), and micro-scale (structural characteristics of shelter). The dierent categories of shelter were sand, coral rubble, coral slab, anemone, sponge, algae (with distinction between macro-algae, encrusting calcareous algae, and turf), and coral (with distinction between the coral form, the coral genus, and its statedead or live). As sh juveniles can change habitat frequently during the benthic phase (McCormick and Makey 1997), each of the ve transects was counted once every semi-lunar cycle (cycle centred on full or new moon). A counting cycle with a rhythm of one transect per day every second day represented a period of 10 working days per lunar half-month. Counting of juveniles was carried out at the end of the colonisation season (March to June 2001; Dufour and Galzin 1993) with ve consecutive counting cycles (the only cycle not recorded was the full moon during the month of May). This sampling schedule allowed for the investigation of post-settlement movements of juveniles during their growth (comparison of habitat use according to the size of juveniles). To investigate the ontogenetic shifts in habitat use between juveniles and adults (recruitment strategies), adults were counted at the cycle of the new moon of May 2001.

Materials and methods

Study area Moorea Island (1730S, 1495W) is a high volcanic island surrounded by a coral reef that delimits a lagoon (Galzin and Pointier 1985). The present study was carried out on the Matautia sector (north coast of Moorea Lagoon), which can be divided into several habitat zones (i.e. coral seascapes, Fig. 1) distinct in terms of depth, wave exposure, and substratum composition (Chancerelle 1996; Lecchini 2003). The reef crest and the rst two seascapes located behind the reef crest (one characterised by coral rubble and the other made up of dead coral masses) have not been recorded because they correspond to the breaking zone of oceanic waves. Thus, the rst sampled seascape (Sea1) is located at 27 m from reef crest and is made up of only coral rubble (Fig. 1). The second seascape (Sea2) is characterised by a high cover percentage (>30%) of living coral masses, Sea3 by a low cover percentage (<30%) of living coral masses, and Sea4 by the same characteristics as Sea3 but with a high depth (>2 m) and a weak current. Sea5 is dened by a high cover percentage of dead coral masses (41%) and coral rubble (42%), and Sea6 is a huge living coral mass of Porites rus (31 m long, 80 m wide, and 2 m high). Sea7 and Sea9 are 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

b Fig. 1 A Map of the north coast of Moorea Island (French Polynesia) showing the location of the study area (sector of Matautia). B Schematic view of the position of dierent seascapes and of ve transects sampled in the Matautia sector. C Schematic view of cross-reef prole showing the relative position of seascapes with the mean width and the standard error calculated on the ve transects. MSL Mean sea level

50 Table 1 In the Matautia sector, 95 species of coral reef sh were recorded at juvenile (J) and/or adult (A) stage by underwater visual counting from March to June 2001. The rst size (in millimeters) corresponds to the total length (TL) of sh at colonisation (estimated to the nearest millimeter from captures Muraenidae (2 species) Echnida nebulosa (63, ?, J) Gymnothorax javanicus (80, ?, J-A) Synodontidae (2 species) Saurida gracilis (?, 95, J-A) Synodus binotatus (56, 75, J) Aulostomidae (1 species) Aulostomus chinensis (134, 180, J-A) Fistulariidae (1 species) Fistularia commersonii (146, 185, J-A) Scorpaenidae (3 species) Pterois antennata (15, 60, J) Pterois radiata (15, 30, A) Scorpaenodes guamensis (11, 60, J) Serranidae (3 species) Cephalopholis argus (24, 80, J) Epinephelus hexagonatus (22, 65, J-A) Epinephelus merra (19, 65, J-A) Apogonidae (1 species) Cheilodipterus quinquelineatus (26, 50, J-A) Carangidae (2 species) Caranx melampygus (82, 140, J-A) Caranx sexfasciatus (80, 120, J) Lutjanidae (2 species) Lutjanus fulviamma (31, 60, A) Lutjanus fulvus (49, 70, J-A) Lethrinidae (2 species) Gnathodentex aurolineatus (45, 65, J-A) Monotaxis grandoculis (48, 80, J-A) Mullidae (5 species) Mulloidichthys avolineatus (69, 100, J-A) Parupeneus barberinus (69, 90, J) Parupeneus bifasciatus (68, 100, J-A) Parupeneus cyclostomus (40, 90, J-A) Parupeneus multifasciatus (57, 90, A) Chaetodontidae (14 species) Chaetodon auriga (19, 40, J-A) Chaetodon citrinellus (29, 55, J-A) Chaetodon ephippium (17, 40, A) Chaetodon lunula (21, 60, J-A) with crest nets) and the second size (in millimeters) corresponds to TL at recruitment [in intervals of 5 mm; (e.g. size 40=35 40 mm) estimated from in situ visual observations]. The symbol ? means that the size at colonisation or recruitment is unknown Labroides dimidiatus (7, 40, J-A) Novaculichthys taeniorous (7, 25, J) Pseudocheilinus hexataenia (7, 45, J-A) Stethojulis bandanensis (7, 50, J-A) Thalassoma amblycephalum (31, 50, J) Thalassoma hardwicke (7, 40, J-A) Xyrichtys pavo (7, 40, J) Scaridae (6 species) Leptoscarus vaigiensis (8, 50, J-A) Scarus altipinnis (8, 50, J) Scarus frenatus (8, 55, J) Scarus oviceps (8, 65, J-A) Scarus psittacus (8, 40, J) Scarus sordidus (8, 65, J-A) Pinguipedidae (1 species) Parapercis clathrata (?, 80, J-A) Acanthuridae (9 species) Acanthurus guttatus (35, ?, A) Acanthurus nigricauda (47, 80, J) Acanthurus triostegus (30, 65, J-A) Ctenochaetus striatus (47, 80, J-A) Naso brevirostris (39, 80, J) Naso lituratus (63, 90, J-A) Naso unicornis (54, 90, J-A) Zebrasoma scopas (26, 50, J-A) Zebrasoma veliferum (28, 45, J-A) Zanclidae (1 species) Zanclus cornutus (55, 80, A) Siganidae (1 species) Siganus spinus (61, 100, A) Balistidae (2 species) Rhinecanthus aculeatus (37, 60, J) Balistipus undulatus (30, 75, J-A) Ostraciidae (2 species) Ostracion cubicus (12, 25, J-A) Ostracion meleagris (15, 45, J-A) Tetraodontidae (3 species) Canthigaster janthinoptera (30, 55, J-A) Canthigaster solandri (21, 55, J-A) Canthigaster valentini (13, 45, J)

Chaetodon ornatissimus (?, 50, J-A) Chaetodon pelewensis (36, 60, J) Chaetodon reticulatus (?, 55, J-A) Chaetodon trifascialis (12, 55, J) Chaetodon trifasciatus (15, 50, J-A) Chaetodon ulietensis (21, 35, A) Chaetodon unimaculatus (19, 60, J-A) Chaetodon vagabundus (23, 50, J-A) Forcipiger longirostris (?, 50, J-A) Heniochus chrysostomus (47, 60, A) Pomacanthidae (1 species) Centropyge avissimus (21, 50, J-A) Pomacentridae (13 species) Abudefduf sexfasciatus (16, 40, J-A) Abudefduf sordidus (16, 30, A) Chromis atripectoralis (10, 30, J-A) Chromis margaritifer (11, 35, J-A) Chromis viridis (8, 25, J-A) Chromis iomelas (11, ?, A) Chrysiptera leucopoma (19, 35, J-A) Dascyllus aruanus (7, 20, J-A) Dascyllus reticulatus (?, 40, J-A) Dascyllus trimaculatus (16, 30, A) Pomacentrus pavo (18, 35, J-A) Stegastes albifasciatus (21, 45, J-A) Stegastes nigricans (18, 40, J-A) Cirrhitidae (2 species) Neocirrhites armatus (?, 55, J) Paracirrhites arcatus (20, 50, J-A) Labridae (16 species) Cheilinus chlororous (7, 55, J-A) Coris aygula (7, 35, J-A) Coris gaimard (7, 45, J-A) Gomphosus varius (7, 40, J-A) Halichoeres hortulanus (7, 50, J-A) Halichoeres margaritaceus (7, 40, J-A) Halichoeres marginatus (7, 25, J-A) Halichoeres trimaculatus (7, 40, J-A) Labroides bicolor (7, 25, J)

Statistical analysis Spatial repartition of seascapes of each transect (sampling stations) was analysed by using hierarchical clustering, which searched the data for distinct clusters of stations with similar sh communities. The clustering algorithm was based on Wards method of minimum variance clustering. Distribution of sh species was analysed by using correspondence analysis (CA; length of gradient from the initial detrended CA screening step was over 3; Benzecri 1973), which compared the relative abundance of dierent sh species among seascapes of each transect (56 sampling stations). To identify the sh communities on CA, Wilks test (Wilks 1932) was used. This statistical test compared variation between communities to variation within each community. If variation between communities was signicantly greater than variation within communities, then the designated communities provided signicant information on the

spatial structure of coral reef sh; if not, then communities were considered invalid and the respective communities merged.

Spatial pattern in juvenile communities A total of 16,958 juveniles belonging to 82 species (ve cycles 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 the lagoon with 40% of species settled in one or two seascapes and only 7% of species settled in eight or more seascapes. The repartition of stations in a clustering algorithm (Fig. 3) was characterised by four clusters at a Ward distance of 1.4 (channel seascape excluded from data, no juvenile recorded). The rst cluster was con-

51 Fig. 2 Variation in density (A) and species richness (B) of coral reef sh at juvenile and adult stages among coral seascapes. Error bars represent one standard error

stituted by the stations of seascape 1, the second by the stations of seascapes 2 to 6. A third cluster was constituted by the stations of seascapes 7 to 11, and a fourth by the stations of seascape 12. The distribution of species from a CA was characterised by a Guttman eect that can be divided into four communities (Fig. 4). The Wilks test validated the distribution of coral reef sh species in four communities, corresponding with the four clusters of stations, more for some ubiquitous species (lambda<0.0001, F=5.26, P=0.042). Thus the reef crest community (cluster of stations dened by seascape 1) was characterised by species either present only on seascape 1 (coordinates inferior at 1.1 on axis 1 and superior at 1.2 on axis 2Coris gaimard, Thalassoma amblycephalum) or present on several barrier reef seascapes but with higher density on seascape 1 (e.g. Balistipus undulatus, Centropyge avissimus). The barrier reef community (cluster of stations dened by seascapes 2 to 6) and the fringing reef community (cluster of stations dened by seascapes 7 to 11) were characterised by the most abundant species (e.g. Gomphosus varius, Thalassoma hardwicke for barrier reef community, and Scarus sordidus, Stegastes albifasciatus for fringing reef

community). The beach community (seascapes 12) was dened by four species: Acanthurus triostegus, Caranx melanpygus, Chaetodon lunula, and Rhinecanthus aculeatus. Lastly, some species were not included in these four communities. Cheilinus chlorourus, Ctenochaetus striatus, Halichoeres trimaculatus, Leptoscarus vaigiensis, Naso lituratus, and Scarus frenatus were ubiquitous species that settled in eight or more seascapes. Spatial pattern in adult communities The adult sh population was represented by 5,551 individuals and 74 species (one cycle of counting) corresponding to a density of 1.16 sh m2 (Fig. 2). The 74 species sampled had an area of distribution less selective than that of the juveniles, with only 6% of species present in one seascape and 64% present in four to nine seascapes. The repartition of stations in a clustering algorithm (Fig. 5), once seascapes 8 and 12 were excluded from the data matrix (too few adults recorded in these seascapes), highlighted three clusters at a Ward distance of 4. The distribution of species (Fig. 6) was

52 Fig. 3 Hierarchical clustering analysis showing the spatial repartition of 56 stations when the juveniles were recorded. Stations are coded according to the abbreviation of seascapes (Sea1 to Sea12) followed by their position on the transects (from 1 to 5)

reected by three adult communities: reef crest, barrier reef, and fringing reef (Wilks test, lambda<0.0001, F=18.35, P=0.047). Ontogenetic shifts in habitat use by coral reef sh according to their development stage Only the 20 most abundant species were studied to give credence to the analysis (to avoid species with a too low density). The comparison of spatial distribution of juveniles versus adults highlighted four strategies of recruitment (Table 2): (1) some species increased the number of seascapes used during the adult stage (e.g. Halichoeres hortulanus, Mulloidichthys avolineatus). (2) Other species decreased the number of seascapes they used compared to recently settled juveniles (e.g. Chrysiptera leucopoma, Stethojulis bandanensis). (3) The third recruitment strategy was the use of dierent seascape types between juvenile and adult life stages. Abudefduf sexfasciatus juveniles settled in micro-atolls of seascapes 10 and 11 and as adults they lived on the barrier reef. Caranx melampygus changed from the beach community

at the juvenile stage (Fig. 4) to the barrier reef community at the adult stage (Fig. 6). (4) Lastly, some species lived on the same seascape(s) with the same shelter (e.g. Stegastes albifasciatus, Stegastes nigricans) or with a dierent shelter (e.g. Naso litturatus, Thalassoma hardwicke). Thalassoma hardwicke lived around the coral masses of Porites rus and Pocillopora verrucosa, but juveniles lived as resident sh in a living branching coral colony of P. verrucosa, whereas adults lived as transient sh around coral masses (see raw data of Lecchini 2003). Stegastes nigricans lived at both juvenile and adult stages in branched masses of Porites with algal turf, located between seascapes 1 and 7. Only the behaviour of sh allowed us to dierentiate the juvenile from the adult stage for this species (Table 1). Ontogenetic shifts in habitat use by coral reef sh juveniles during their growth From March to June, the distribution of species size classes along the dierent seascapes was recorded on ve counting cycles. Two post-settlement patterns were


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

found among species (Fig. 7). Firstly, some species settled and remained in the same habitat until the adoption of the adult habitats (e.g. Chaetodon citrinellus, Halichoeres hortulanus, Rhinecanthus aculeatus, Thalassoma hardwicke). Secondly, others settled to several seascapes but disappeared, from a determined size, from some seascapes through dierential mortality and/or postsettlement movement (e.g. 6570 mm for Ctenochaetus striatus, 4045 mm for Epinephelus merra, 5055 mm for Scarus sordidus). The juveniles of C. striatus from the size class 47 (colonisation size) to 70 mm settled on all seascapes (except the channel). The juveniles from the size class 70 to 80 mm (recruitment size) were only present from seascapes 1 to 9, probably due to dierential mortality and/or post-settlement movement. For E. merra (from 4045 mm size class), there was post-settlement movement (from fringing to barrier reef), possibly associated with a dierential mortality. From March to April, the juveniles of E. merra were mainly recorded on the fringing reef with a size range from 21 (colonisation size) to 40 mm, and from May to June, they were mainly recorded on the barrier reef with a size over 40 mm.

Spatial structure of juvenile sh communities In the Matautia sector, the juveniles of coral reef sh were dispatched among four communities spatially

distributed according to the distance from the reef crest. Hypothetically, this pattern could be explained by current, depth, or turbidity gradients. Due to the shallowness in Matautia lagoon (maximum of 3 m depth; Galzin and Pointier 1985), the depth hypothesis is rejected. Similarly, the studied area has no turbidity gradient because water quality on the north coast of Moorea is very good due to the absence of major human pollution, the shallowness of the lagoon, and numerous passes that maintain a quick renewal of water in the lagoon (from 0.4 to 1.3 m s1; Renon 1989). As a consequence, current gradient (current speed is high near the reef crest and decrease towards the beach) should be the best hypothesis to explain the spatial distribution of the juveniles. Nevertheless, changes among seascapes in some biological factors, such as percentage of living cover, percentage cover of each substratum type, and substratum diversity and complexity, could increase or decrease the inuence of current on juvenile distribution. Thus, the dierentiation of the reef crest community (vs barrier reef community) would be due to the topographic characteristics of seascape 1, which is made of coral rubble. The absence of coral reef patches on this seascape, dierent from other barrier reef seascapes (barrier reef community), means that high current action from oceanic waves is not obstructed, allowing mainly the smaller species (which hide themselves among coral rubble) and species with high swimming abilities (which ght against the current without protecting themselves behind a coral reef patch) to occupy the habitat. The main target species of the reef crest community meeting these criteria are Halichoeres margaritaceus, Coris aygula, C. gaimard, and Thalassoma amblycephalum, In

54 Fig. 5 Hierarchical clustering analysis showing the spatial repartition of 56 stations when the adults were recorded

addition, the Pomacentrid Chrysiptera leucopoma, which has limited swimming abilities (personal observation), was also found. Subsisting near the bottom, this species needs less speed to swim against the current due to friction eects (Wood and Bain 1995). This physical phenomenon allows some species to occupy seascapes at high current speeds when their swimming abilities would not normally allow them to do so (Gerstner 1998). The dierentiation of the beach community (vs the fringing reef community) is also related to the low structural complexity of seascape 12 but the current action is not accentuated here. This seascape is a shallow sandy area without coral shelter (dierent from other fringing reef seascapes) and with low current. These characteristics favour the presence of transient species with high swimming abilities to ee predators. The main target species of this community are Acanthurus triostegus, Caranx melampygus, and Rhinecanthus aculeatus. To conclude, the communities structure, spatially ordered 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 decreases towards the beach), the topographic characteristics of some coral seascapes (Sea1 and Sea12), and the position of the species in the water column. Other factors could, nevertheless, inuence where a sh settles, and where the juveniles live, such as morphological/functional components of species, components involving the food they can exploit, and ecological factors (e.g. facilitation, intra/interspecic competition, and predation). Ontogenetic shifts in habitat use Two patterns of shifts in habitat use were found during the growth of juveniles at Moorea Island. Firstly, some species settled and remained in the same habitat until the adoption of the adult habitats. Secondly, others settled to several habitats but disappeared from some habitats through dierential mortality and/or post-settlement movement. Thus, post-settlement behaviour of sh has the potential to alter the distribution of juveniles

55 Fig. 6 Correspondence analysis showing the spatial distribution of adult species

established at settlement. The post-settlement behaviour can, nevertheless, vary according to coral reef sh families. In our study, the 4 Pomacentridae species belonging to the 20 most abundant species (Abudefduf sexfasciatus, Chrysiptera leucopoma, Stegastes albifasciatus, and S. nigricans) did not distort their settlement signal by post-settlement movement. Among Labridae, 3 of 5 species belonging to the 20 most abundant species distorted their settlement signal (Gomphosus varius, Halichoeres margaritaceus, H. trimaculatus). The low post-settlement dispersal rate seems to be characteristic of Pomacentridae (Sponaugle and Cowen 1996; Booth 2002) and may help them reduce predation risk (Beukers and Jones 1998). Sponaugle and Cowen (1996) showed that older juvenile distribution of the damselsh Stegastes partitus closely reected settlement patterns, while older juveniles of the surgeonsh Acanthurus bahianus were most abundant at sites of lower settlement, which had a more suitable habitat. Whatever the sh family studied (Pomacentridae, Acanthuridae, Labridae, etc.), dierences in the distribution of size classes among different habitats have been frequently used to infer postsettlement movement (see Gillanders et al. 2003). However, other explanations may also account for spatially explicit patterns of organism size distribution among habitats. For example, dierences in growth rates (i.e. growth could have an inuence through post-settlement survival in the face of a selective mortality regime) or dierential mortality among habitats may result in patterns similar to those attributable to post-settlement movement. Our data on Ctenochaetus striatus, Epinephelus merra, and Scarus sordidus were not able to highlight if post-settlement movement or dierential

mortality is more important (Fig. 7). But, in both cases (dierential mortality and/or post-settlement movement), the settlement signal is lost before recruitment and post-settlement behaviour alters the distribution of juveniles and subsequently adults. The settlement signal can also be lost by the recruitment strategy of sh. Deegan (1990) sampled four locations in juvenile habitat (marsh) and four locations in adult habitat (open bay). He found that density of Gulf menhaden Brevoortia patronus was the highest in tidal creeks (average sh length 3040 mm) and that abundances in creeks declined when sh grew to 4550 mm, when they likely moved to an open bay area and then oshore. The comparative analysis of spatial distribution of juveniles and adults in the present study highlighted four recruitment strategies: (1) an increase in the number of habitats used during the adult stage; (2) a decrease in the number of habitats adults used compared to recently settled juveniles; (3) no change in habitat use; and (4) use of nursery areas by juveniles followed by extensive movement to an 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 temperate and tropical species (e.g. Parrish 1989; Nagelkerken et al. 2002). Among the 20 most abundant species (Table 2), the 5 species that used dierent habitat types for juvenile and adult stages (Abudefduf sexfasciatus, Acanthurus triostegus, Caranx melampygus, Gnathodentex aurolineatus, and Rhinecanthus aculeatus) settled on seascapes 10, 11, and 12, dened by Lefevre (1991) as the nursery area on the north coast of Moorea Island.

56 Table 2 Distribution in density (number of sh per square meter) along the dierent seascapes of the 20 most abundant sh species sampled in the Matautia sector, to explore ontogenetic shifts in habitat use by juveniles at recruitment (comparison of habitat use of juveniles and adults). Four recruitment strategies have been Species highlighted: an increase in the number of seascapes used during the adult stage (Inc); a decrease in the number of seascapes adults used compared to recently settled juveniles (Dec); the use of different seascape types (Dif); no change in seascape use by species (NC) Channel Sea9 Sea10 Sea11 Sea12 Strategy of recruitment 0.053 0.051 0.02 0.002 0.105 0.003 0.006 0.029 0.005 0.124 0.015 0.534 0.041 0.002 0.003 0.002 0.095 0.011 0.044 0.008 0.166 0.007 0.030 0.008 0.001 0.025 0.006 0.003 0.004 0.369 0.048 0.031 0.003 0.002 0.004 0.591 0.059 0.003 0.007 0.008 0.012 0.007 0.086 0.011 Dif 0.011 0.002 0.003 0.018 Dif 0.002 0.001 0.001 0.005 0.003 0.003 NC Dec 0.122 0.202 0.189 0.130 Dec 0.031 0.013 0.023 0.024 NC 0.002 0.001 0.003 0.053 0.145 Dif 0.009 0.007 0.003 NC 0.008 Inc 0.003 0.002 NC 0.022 0.008 0.03 0.013 0.004 0.004 0.013 0.002 0.01 0.008 0.003 0.060 0.002 NC 0.011 0.038 0.032 Inc 0.013 0.006 0.01 NC 0.008 0.051 Dif Dec Dif

Sampling No. sh Sea1 Sea2 Sea3 Sea4 Sea5 Sea6 Sea7 stage counted 86 34 31 111 51 12 199 34 243 18 5,017 80 149 12 210 18 322 26 154 53 279 8 558 45 290 45 194 30 32 11 1,013 52 1,072 25 203 44 249 56 1,177 83

Abudefduf sexfasciatus Acanthurus triostegus Caranx melampygus Chaetodon citrinellus Chrysiptera leucopoma Ctenochaetus striatus Epinephelus merra Gnathodentex aurolineatus Gomphosus varius Halichoeres hortulanus Halichoeres margaritaceus Halichoeres trimaculatus Mulloidichthys avolineatus Naso litturatus

Juvenile Adults Juvenile Adult Juvenile Adult Juvenile Adult Juvenile Adult Juvenile Adult Juvenile Adult Juvenile Adult Juvenile Adult Juvenile Adult Juvenile Adult Juvenile Adult Juvenile Adult Juvenile Adult Rhinecanthus Juvenile aculeatus Adult Scarus sordidus Juvenile Adult Stegastes Juvenile albifasciatus Adult Stegastes Juvenile nigricans Adult Stethojulis Juvenile bandanensis Adult Thalassoma Juvenile hardwicke Adult

0.008 0.022 0.011 0.005 0.007 0.003 0.006 0.009 0.01 0.006 0.003 0.001 0.403 0.027 0.01 0.001 0.004 0.033 0.005 0.003 0.003 0.356 0.032 0.005 0.001 0.156 0.502 0.009 0.042 0.014 0.012 0.008

0.097 0.006 0.022 0.004 0.026

0.003 0.115 0.003 0.011 0.008

0.003 0.017 0.013 0.004 0.008 0.003 0.014 0.004 0.014 0.006 0.003 0.007

0.033 0.007 0.01 0.002 0.011 0.008 0.006 0.006 0.016 0.005 0.047 0.015 0.035 0.016 0.013 0.007 0.009 0.012 0.004 0.023 0.004 0.004 0.001 0.009 0.048 0.002 0.023 0.007 0.013 0.029 0.066 0.044 0.063 0.023 0.018 0.017 0.007 0.018 0.020 0.012 0.047 48.000 0.029 0.037 0.009 0.014 0.011 0.014 0.037 0.011 0.017 0.024 0.032 0.068 0.015 0.032 0.001 0.05 0.033 0.013 0.017 0.029 0.003 0.007 0.008 0.002 0.011 0.005 0.005 0.009 0.055 0.183 0.108 0.213 0.085 0.024 0.02 0.032 0.023 0.022 0.013 0.026 0.027 0.026

0.006 0.013 0.016 0.019 0.007 0.010 0.004 0.005

0.050 0.107 0.013

0.078 0.067 0.117 0.018 NC 0.026 0.036 0.027 NC 0.007 0.003 0.009 0.003 0.018 0.005 0.002 0.02 0.019 0.009 Dec NC

To conclude, the present study is the rst to explore the use of space by a broad range of sh taxa from settlement to recruitment (i.e. at the juvenile stage), and to document the prevalence of major habitat shifts between juvenile and adult life stages. This multi-species perspective oers a broader survey of the varied ways in which sh can redistribute themselves post-settlement than has been done in previous work (see Gillanders et al. 2003). Thus among the 20 most abundant species recorded in Moorea lagoon, 12 species distorted spatial patterns of sh established at settlement by some ontogenetic habitat shifts (two patterns of post-settlement movement, four recruitment strategies). Although this study was carried out at a single site transect across the Moorea lagoon, other studies support the generality of ontogenetic habitat use (see Gillanders et al. 2003). Thus

the prevalence of ontogenetic habitat shifts highlighted in the present study has a substantial implication in terms of interpreting data on sh settlement to the reef. Measuring and describing settlement rates and patterns (i.e. selective choice of habitat, high mortality, strong competition between settling sh and resident sh) has long been a focus of reef sh ecology (see Doherty 2002). Our results suggest, however, that the settlement patterns can be misleading over time when sh survive dierentially or redistribute themselves among habitats.
Acknowledgements The authors wish to thank C. Rua, A. Lo-Yat, and S. Planes for their comments on the rst draft, J. Algret and Y. Chancerelle for their assistance in the eld, and M. McCormick for having checked the English language. The rst author also thanks the French Ministry of Foreign Aairs for the Lavoisier Fellowship.


Fig. 7 Distribution in density (number of sh per square meter) of juvenile species by size class among coral seascapes from March to June 2001. Errors bars represent one standard error. Sizes (total length in millimeters) at colonisation and recruitment are given in parentheses

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