PRIMARY RESEARCH PAPER

Small-scale variability in the size structure of scleractiniancorals around Moorea, French Polynesia: patterns acrossdepths and locations

Mehdi Adjeroud Æ Morgan S. Pratchett ÆMarie C. Kospartov Æ Christophe Lejeusne ÆLucie Penin

Received: 6 March 2007 / Revised: 14 March 2007 / Accepted: 24 March 2007 / Published online: 13 June 2007� Springer Science+Business Media B.V. 2007

Abstract The size structure of coral populations

is influenced by biotic and physical factors, as well

as species-specific demographic rates (recruit-

ment, colony growth, mortality). Coral reefs

surrounding Moorea Island are characterized by

strong environmental gradients at small spatial

scales, and therefore, we expected that the size

structure of coral populations would vary greatly

at this scale. This study aimed at determining the

degree of spatial heterogeneity in the population

size structure of two coral taxa, Pocillopora

meandrina and massive Porites spp., among

depths (6, 12, and 18 m) and among locations

(Vaipahu, Tiahura and Haapiti) representing

different exposure to hydrodynamic forces. Our

results clearly underlined the strong heterogene-

ity in the size structure of both P. meandrina and

massive Porites spp., with marked variation

among depths and among locations. However,

the lack of any consistent and regular trends in

the size structure along depths or among loca-

tions, and the lack of correlation between size

structure and mean recruitment rates may suggest

that other factors (e.g., stochastic life history

processes, biotic interactions, and disturbances)

further modify the structure of coral populations.

We found that the size structure of P. meandrina

was fundamentally different to that of massive

Porites spp., reflecting the importance of life

history characteristics in population dynamics.

Keywords Coral reefs � Scleractinian corals �Spatial patterns � Population structure �Size-frequency distribution � Life histories

The abundance of scleractinian corals varies

greatly in time and space, and understanding

these patterns often requires an appreciation of

Handling editor: I. Nagelkerken

M. Adjeroud (&) � L. PeninEcole Pratique des Hautes Etudes, UMR 5244CNRS-EPHE-UPVD, Laboratoire ‘‘EcosystemesAquatiques Tropicaux et Mediterraneens’’, Universitede Perpignan, Avenue Paul Alduy, Perpignan Cedex66860, Francee-mail: adjeroud@univ-perp.fr

M. Adjeroud � L. PeninCentre de Recherches Insulaires et Observatoire del’Environnement, Papetoai, Moorea BP 1013, FrenchPolynesia

M. S. PratchettARC Centre of Excellence for Coral Reef Studies,James Cook University, Townsville Q4811, Australia

M. C. KospartovCenter for Marine Studies, The University ofQueensland, St Lucia Q4072, Australia

C. LejeusneCentre Oceanologique de Marseille, UMR 6540,DIMAR, rue de la batterie des lions, 13007 Marseille,France

123

Hydrobiologia (2007) 589:117–126

DOI 10.1007/s10750-007-0726-2

spatial and temporal variation in population

dynamics (Connell et al., 1997; Hughes et al.,

1999). However, scleractinian corals are very

slow-growing and long-lived organisms, such that

traditional census techniques used to measure

demographic rates (e.g., following the fate of

individuals over time) require long-term studies

(see Connell et al., 1997). Alternatively, informa-

tion on population dynamics may be extracted

from data on age- or size structure of local

populations (Ebert, 1981; Sebens, 1983; Lewis,

1989; Barry & Tegner, 1990; Vermeij & Bak, 2002).

For colonial organisms, such as scleractinian

corals, ecological and demographic processes such

as growth, mortality and fecundity are strongly

correlated with colony size (Hughes & Jackson,

1980, 1985; Soong, 1993), and variation in the size

structure of coral populations has been used to

infer spatial patterns in their population dynamics

(e.g., Done, 1988; Bak & Meesters, 1998; Fong &

Glynn, 1998, 2001; Meesters et al., 2001).

Interpreting data on population size structure

of scleractinian corals and extracting meaningful

biological information can be difficult (Vermeij &

Bak, 2002). The size structure of coral popula-

tions is structured by a number of opposing

processes (e.g., recruitment rates versus whole

colony mortality, intrinsic growth rates versus

partial mortality) and different combinations of

demographic rates may produce essentially sim-

ilar size structures (Hughes & Jackson, 1980;

Hughes, 1984; Bak & Meesters, 1998). In general,

the size structure of coral populations is positively

skewed, where populations are comprised mainly

of smaller colonies and relatively few large

colonies (Hughes & Jackson, 1985; Babcock,

1991; Soong, 1993; Bak & Meesters, 1998). The

degree of skewness in coral populations varies

among species, reflecting differences in their

underlying life history characteristics (Hughes &

Jackson, 1985; Babcock, 1991) and most impor-

tantly, differences in their intrinsic growth rates

relative to probabilities of injury and mortality.

Size structure also varies among populations of

the same coral species, mostly due to differences

in disturbance regimes (Gilmour, 2004) and water

quality (e.g., nutrients and turbidity; Bak &

Meesters, 1999; Meesters et al., 2001; but see

Lewis, 1997). Although rarely considered, the size

structure of coral populations may also vary

among different reef habitats, independently of

water pollution (Vermeij & Bak, 2002).

The purpose of this study was to examine, for

the first time in the Central Pacific, small-scale

(within-reef) variation in the size structure of two

dominant groups of corals with highly contrasting

life history characteristics, Pocillopora meandrina

and massive Porites spp., among depths (6, 12,

and 18 m), and among locations (Vaipahu, Tiah-

ura and Haapiti) on outer reef slopes at Moorea

(17�30¢ S, 149�50¢ W), Society Islands, French

Polynesia. Pocillopora meandrina is characterized

by relatively high recruitment rates, fast-growth

rates and short generation time (Laboute, 1985;

Gleason 1996; Magalon et al., 2005; Adjeroud

et al., 2005, 2007). In contrast, massive Porites are

long-lived species with relatively low recruitment

rates and slow growth rates, but high resistance to

perturbations, long life duration and large colony

size (Gleason, 1996; Bessat et al., 1997; Adjeroud

et al., 2005, 2007; Penin et al., 2007). Several

species of massive Porites have been recorded

from reefs around Moorea, including P. austral-

iensis (Vaughan, 1918), P. lobata (Dana, 1846),

and P. lutea (Edwards and Haime 1851), which

are the dominant species on reef slopes. All

species of massive Porites, which cannot be

discriminated in situ, were pooled into a single

taxonomic group.

To explore spatial variation in size structure,

coral colonies of P. meandrina and massive

Porites spp. were measured in April–May 2001

along replicate belt transects at three different

depths (6, 12, and 18 m) at each of the three

locations (Vaipahu, Tiahura, and Haapiti), giving

nine study sites (Fig. 1). At each location, the

linear distance between depth sites was 50–70 m,

depending on the reef slope morphology. The

three locations vary greatly in their exposure to

hydrodynamic forces (Adjeroud, 1997; Laurent

et al., 2004; Fig. 1). Haapiti is the most exposed,

being directly exposed to south-western swells

which prevail throughout much of the year and

generate large swells and considerable wave

energy, whereas Vaipahu and Tiahura are shel-

tered from south-western swells, but are directly

exposed to the more moderate northern swells

which occur during summer months (Laurent

118 Hydrobiologia (2007) 589:117–126

123

et al., 2004). On the reef slope, exposure caused

by swell and wave energy decreases with increas-

ing depth (Adjeroud, 1997). Three replicate

10 m2 (10 · 1 m) belt transects were sampled at

each of the nine study sites. Transects were run

parallel to depth contours from haphazardly

selected starting points at each depth. All coral

colonies with >50% of their living tissue area

contained within each transect were measured to

determine colony size. For each colony, we

recorded maximum diameter (L), the perpendic-

ular diameter (l), and maximum colony height

(h). The surface area of each colony (i.e., colony

size) was calculated using maximum linear dimen-

sions (L, l, and h) of continuous areas of live

tissue. These measurements were used in simple

geometric formulae to calculate approximate

surface area (Meesters et al., 2001). Despite their

sub-branching form, P. meandrina colonies had a

very consistent gross morphology, resembling a

sphere. As such, total surface area was calculated

using the formulae 4p · ((L + l + h)/6)2. Mean-

while, colonies of Porites spp. were best repre-

sented by a cylinder with an ellipsoidal base. As

such, the surface area was estimated using the

formulae p/4 · L · l · h.

To examine the size structure of coral popula-

tions, we constructed size-frequency distributions

based on estimates of the surface area of living

tissue (SA) for each colony. Colony surface-area

data were log10 transformed to normalize size-

frequency distributions and increase resolution

among smaller size classes, following Bak &

Meesters (1998), and Vermeij & Bak (2002). We

then compared the overall shape of size-fre-

quency distributions between populations using

2-sample Kolmogorov–Smirnov (KS) tests. We

also compared variation in colony sizes (coeffi-

cient of variation, CV), skewness (g1 describes the

relative abundance of colonies that are smaller or

larger than the geometric mean) and kurtosis (g2

describes the concentration of data around the

central mode of a distribution) among popula-

tions, following Bendel et al. (1989). Variation in

CV, g1, g2 and mean size of P. meandrina and

Porites spp. were analyzed using 2-way analysis of

variance (ANOVA), testing for variation among

depths and among locations.

A total of 1,922 colonies of P. meandrina and

712 colonies of massive Porites spp. were counted

at the nine study sites, corresponding with mean

densities of 71.1 (±4.5 SE) and 26.3 (±4.4 SE)

colonies per 10 m2 transect, respectively. The

abundance of both P. meandrina and Porites spp.

varied greatly among depths and among locations

(Table 1). For P. meandrina, lowest densities

were recorded in 12 m depth at Tiahura and

Vaipahu, whereas at Haapiti, the highest density

was recorded in 12 m depth. Densities of Porites

spp. decreased with depth at Vaipahu and Tiah-

ura, but this pattern was not recorded at Haapiti,

where lowest values were found at 6 m depth, and

highest values at 12 m depth (Table 1).

For P. meandrina, the surface area of colonies

ranged from 0.78 cm2 up to 3,996.46 cm2, and the

mean size of colonies was 445.82 cm2 ± 11.98 SE.

The majority (86%) of P. meandrina colonies had

a surface area less than 1,000 cm2, and nearly half

of all colonies were less than 200 cm2. The mean

size of P. meandrina varied significantly among

depths, among locations, and also among depths

within locations (Table 2). Among depths, the

mean size of colonies was much smaller in 6 m

(304.50 cm2 ± 14.86 SE), compared to 12 m

(563.63 cm2 ± 21.90 SE), and 18 m (498.92 cm2

Fig. 1 Map of Moorea showing the nine study sites(represented by black dots). Sampling was conducted atthree different depths (6, 12, and 18 m) at each of the threelocations (Vaipahu, Tiahura and Haapiti). Arrows show thepercentage of days throughout 1993, that swells approachedMoorea from each direction (data courtesy of MeteoF-rance, Centre de Polynesie francaise; Laurent et al., 2004)

Hydrobiologia (2007) 589:117–126 119

123

± 25.02 SE). Moreover, the mean size of

P. meandrina at Vaipahu (396.42 cm2 ± 20.06

SE) was smaller than at Tiahura (552.64 cm2 ±

26.66 SE) or Haapiti (418.15 cm2 ± 17.11 SE).

For massive Porites spp., the mean size of

colonies (6,200.95 cm2 ± 1,958.77 SE) was much

higher than for P. meandrina. In 18 m depth at

Vaipahu and Haapiti, populations of Porites spp.

comprised relatively few, but very large (up to

1,067,316 cm2) colonies. By comparison, local

populations of Porites spp. comprised high den-

sities of relatively small (minimum size: 1.17 cm2,

but mostly <2,000 cm2) colonies in 6 m and 12 m.

There was a significant variation in the mean size

of Porites spp. among depths, but the patterns of

depth variation varied among locations (Table 2).

The mean colony size of Porites spp. increased

consistently with increasing depth at Vaipahu and

Haapiti, but this trend was not observed at Tiahura

(Table 1). Among depths, the mean size of Porites

spp. was greatly reduced at 12 m (1,477.85 cm2 ±

270.05 SE) and 6 m (2,581.76 cm2 ± 397.73 SE),

compared to 18 m (35,383.92 cm2 ± 15,124.36

SE). The mean size recorded at Tiahura

(2,968.30 cm2 ± 563.10 SE) was much smaller

than at Haapiti (5,865.40 cm2 ± 2,798.70 SE) and

Vaipahu (8,040.10 cm2 ± 3,771.98 SE).

Overall size-frequency distributions for both

P. meandrina and massive Porites spp. were

strongly and positively skewed, with a prepon-

derance of colonies in the smallest size-classes

(Fig. 2a, 3a). For P. meandrina, log-transformation

Table 1 Size-frequency distribution parameters for (a) Pocillopora meandrina and (b) massive Porites spp., from threedepths (6, 12, and 18 m) at each of three locations (Vaipahu, Tiahura and Haapiti)

Vaipahu Tiahura Haapiti

6 m 12 m 18 m 6 m 12 m 18 m 6 m 12 m 18 m

(a) Pocillopora meandrinan 248 178 234 229 97 176 253 345 162�v 2.1 (0.2) 6.7 (0.4) 5.1 (0.3) 4.7(0.3) 4.8 (0.6) 6.9 (0.5) 2.4 (0.2) 5.3 (0.3) 4.6 (0.5)

Pnorm 0.12 0.00 0.14 0.00 0.10 0.00 0.05 0.00 0.14CV 0.45 0.27 0.35 0.35 0.27 0.27 0.37 0.23 0.37g1 0.09 –1.19 –0.46 –0.77 –0.56 –0.62 –0.37 –0.87 –0.53g2 –0.77 0.64 –0.42 –0.59 –0.30 –0.66 –0.68 0.23 –0.82

(b) Massive Porites spp.n 119 74 15 122 46 1 41 220 74�v 19.4 (5.0) 25.3 (94.6) 534.6 (375) 39.3 (7.6) 4.9 (1.4) 2.4 (na) 4.5 (1.3) 13.3 (3.9) 321.9 (168)

Pnorm 0.75 0.45 0.67 0.89 0.94 – 0.78 0.66 0.17CV 0.41 0.32 0.37 0.30 0.25 – 0.49 0.29 0.57g1 0.03 –0.87 1.29 –0.52 –0.03 – 0.05 –0.22 0.97g2 –0.25 0.69 1.33 0.24 0.89 – –0.86 0.34 0.79

n: total number of colonies measured, �v: mean surface area (·102 cm2) of colonies (±SE), Pnorm: probability that data arenormally distributed (KS test), CV: coefficient of variation, g1: skewness, g2: kurtosis

Table 2 ANOVA results for comparisons of size-frequency distributions for Pocillopora meandrina and massive Por-ites spp., among depths (6, 12, and 18 m) and among locations (Vaipahu, Tiahura and Haapiti)

Pocillopora meandrina Massive Porites spp.

Location Depth Location · Depth Location Depth Location · Depth

�v F2/1913 = 7.69** F2/1913 = 34.67*** F4/1913 = 5.62** F2/703 = 1.37 F2/703 = 19.64** F4/703 = 38.05***CV F2/18 = 11.45** F2/18 = 54.75*** F4/18 = 8.00** F2/16 = 10.12** F2/16 = 9.15** F3/16 = 5.43**g1 F2/18 = 1.56 F2/18 = 21.54** F4/18 = 12.00*** F2/16 = 3.34 F2/16 = 3.28 F3/16 = 2.81g2 F2/18 = 0.46 F2/18 = 7.66** F4/18 = 0.20 F2/16 = 7.97 F2/16 = 1.58 F3/16 = 5.75**

�v: mean surface area (·102 cm2) of colonies, CV: coefficient of variation, g1: skewness, g2: kurtosis. ‘*’ indicates significanteffects: * = P < 0.05, ** = P < 0.01, *** = P < 0.001

120 Hydrobiologia (2007) 589:117–126

123

of colony surface area greatly increased centering

of the size-frequency distribution around the

mean (Fig. 2b), though the log-transformed dis-

tributions were deviated significantly from nor-

mality (KS test, P < 0.01). After transformation,

the distribution was actually negatively skewed

(g1 = –0.59) and platikurtic or flattened

(g2 = –0.55). The overall size-frequency distribu-

tion of Porites spp. was found to be normal (KS

test, P = 0.81), but slightly positively skewed

(g1 = 0.07) and highly leptokurtic (g2 = 0.43).

The overall size-frequency distribution for P.

meandrina was significantly different to that of

Porites spp. (KS test, P < 0.01).

Within taxa, size-frequency distributions varied

among depths and among locations. For example,

strong differences in size-frequency distributions

of P. meandrina occurred at Vaipahu, between

populations in 6 m vs. 12 m (Fig. 4). Size-fre-

quency distributions of P. meandrina were mostly

significantly different from normality (Table 1),

and the proportion of colonies in the smallest

size-class (< 200 cm2) varied greatly among depths

and among locations, ranging from 26% to 75%. In

contrast, size-frequency distributions of Porites

spp. populations were all normal (Table 1). For

Porites spp., the typical size-frequency distribution

(e.g., in 6 m at Vaipahu) comprised large numbers

of very small (< 800 cm2) as well as few large

colonies (up to 1,067,316 cm2). However, in 12 m

at Tiahura and in 6 m at Haapiti, the largest

colonies of Porites spp. were only 6,446 cm2 and

3,534 cm2, respectively, and virtually all colonies

were less than 800 cm2 (Fig. 5).

Significant spatial variation in size-frequency of

P. meandrina and massive Porites spp. was also

apparent from comparisons of coefficient of

variation (CV), skewness (g1) and kurtosis (g2;

Table 1). At Moorea, CV was very similar

between P. meandrina (0.34) and Porites spp.

(0.37). ANOVA revealed that CV varied signif-

icantly among locations, among depths, and also

among depths within locations, for both taxa

(Table 2). Among locations, CV was highest at

Vaipahu for P. meandrina, but highest at Haapiti

Fig. 2 Size structure for Pocillopora meandrina at Moo-rea. Size-frequency distributions are shown for (a) un-transformed estimates of colony surface area and (b)log10 transformed estimates of colony surface area. Datawere pooled across depths and locations. n = 1,922colonies

Fig. 3 Size structure for massive Porites spp. at Moorea.Size-frequency distributions are shown for (a) untrans-formed estimates of colony surface area and (b) log10

transformed estimates of colony surface area. Data werepooled across depths and locations. n = 712 colonies

Hydrobiologia (2007) 589:117–126 121

123

for Porites spp. Among depths, CV tended to be

lowest in 12 m depth, compared to 6 m or 18 m

depth, for both taxa (Table 1). For P. meandrina,

size-frequency distributions of local populations

were strongly negatively skewed (g1 < 0), except

in 6 m at Vaipahu, and skewness varied among

depths but not locations (Table 2). In contrast,

the skewness (g1) of size distributions for Porites

spp. were highly varied, ranging from –0.86 to

1.33, and exceeded differences between taxa

(Table 1). Nevertheless, skewness in Porites spp.

did not significantly vary among depths nor

locations (Table 2). Size-frequency distributions

were mostly platikurtic (g2 < 0) for P. meandrina,

but leptokurtic (g2 > 0) for Porites spp. (Table 1).

For P. meandrina, kurtosis was much higher in

12 m, compared to 6 m or 18 m, at every location.

Overall, kurtosis varied significantly among

depths, but there was no variation among loca-

tions (Table 2). Kurtosis of Porites spp. increased

with depth at all locations, though there was no

overall variation among depths or among loca-

tions (Table 2).

This study has shown that the size structure of

P. meandrina was fundamentally different to that

of massive Porites spp. Most notably, the mean

and maximum sizes of massive Porites spp. were

much larger than for P. meandrina, reflecting

Fig. 4 Within-reef variation in size structure of Pocillo-pora meandrina. n: number of colonies counted at eachstudy site. Sites with significantly different distributions

(Kolmogorov–Smirnov tests, P > 0.05) are indicated oneach graph. V: Vaipahu, T: Tiahura, H: Haapiti, 6: 6 m, 12:12 m, 18: 18 m

122 Hydrobiologia (2007) 589:117–126

123

their increased persistence and greater longevity

(Potts et al., 1985; Bessat et al., 1997). As in other

coral reefs around the world (Endean & Camer-

on, 1990; Wittenberg & Hunte, 1992), massive

Porites in French Polynesia are highly persistant

species, characterized by lower recruitment rates

but higher resistance to perturbations (Gleason,

1996; Adjeroud et al., 2005, 2007; Penin et al.,

2007). On the other hand, the more even

size-frequency distributions of P. meandrina

(platikurtic distribution) reflects their faster

growth and higher population turnover (Grigg

& Maragos, 1974). Pocillopora meandrina was

also characterized by a lower proportion of small

colonies that may result from higher post-settle-

ment mortality. As in other Indo-Pacific coral

reefs (Grigg & Maragos, 1974; Tomascik et al.,

1996; Connell et al., 1997; Bianchi et al., 2006;

Nakamura & Nakamori, 2006), P. meandrina thus

represent an opportunistic species (Laboute

1985), with high recruitment rates, high potential

for larval dispersal and colonization, but low

resistance to perturbations such as cyclones and

bleaching events (Magalon et al., 2005; Adjeroud

et al., 2005, 2007). These results demonstrate the

overriding importance of intrinsic processes, such

Fig. 5 Within-reef variation in size structure of massivePorites spp. n: number of colonies counted at each studysite. Sites with significantly different distributions (Kol-

mogorov–Smirnov tests, P > 0.05) are indicated on eachgraph. V: Vaipahu, T: Tiahura, H: Haapiti, 6: 6 m, 12:12 m, 18: 18 m

Hydrobiologia (2007) 589:117–126 123

123

as reproductive strategies and colony growth, in

structuring coral populations (Hughes & Jackson,

1985; Bak & Meesters, 1999; Meesters et al.,

2001).

This study has also shown that the size struc-

ture of coral populations can vary enormously

over relatively small spatial scales, within reefs.

Variation in the size structure of P. meandrina

and Porites spp. at Moorea was much greater than

for any of the 11 species studied by Meesters

et al. (2001), in which spatial variation in size

structure of populations was attributed to heavy

urbanization, and associated environmental deg-

radation at some locations. Locations selected in

the present study were far removed from sites of

heavy urbanization and there were no obvious

direct anthropogenic impacts (dredging, construc-

tion activities, sewage discharge) at any of the

sampling locations (Adjeroud et al., 2002). Thus,

our results underlined that large differences in the

population size structures may be independent of

any major differences in environmental degrada-

tion (cf. Meesters et al., 2001), and suggest that

researchers must be very cautious when inter-

preting differences in size-frequency distributions

among coral populations in different locations or

over time.

While there was significant spatial variation in

the size structure of both P. meandrina and

massive Porites spp., we failed to detect any clear

and regular patterns of variation among depths

for the three locations examined, as well as no

consistent differences between exposed (Haapiti)

and less exposed locations (Tiahura and Vai-

pahu). This lack of any consistent trends may

suggest that other factors (e.g., stochastic life

history processes, biotic interactions, and distur-

bances) further modify the structure of coral

populations. Spatial variation in the size structure

of corals may also be caused by variation in rates

of recruitment, colony growth, injury and/or

mortality, all of which can vary over relatively

small spatial scales (Baird & Hughes, 1997).

Strong heterogeneity in rates of recruitment was

found among the nine study sites around Moorea,

with significant variation across depths and loca-

tions (Adjeroud et al., 2007). However, we failed

to detect any correlation between recruitment

rates (annual or seasonal mean rates) and

size-frequency distribution parameters for the

two taxa examined (Pearson correlation coeffi-

cient, P > 0.01).

Variation in the size structure of local coral

populations has important implications for the

subsequent fate (persistence and resilience) of

taxa in different reef environments (Done, 1988).

Since fecundity is positively correlated to colony

size (e.g., Hughes & Jackson, 1985; Babcock,

1991), populations comprised entirely of small

colonies would be less likely to reproduce and

persist through time, compared to populations

comprising large numbers of colonies with high

reproductive output. In this study, we found that

some localized populations comprise many large

colonies, and probably make a disproportionate

contribution to reproductive output, compared to

other populations that are comprised exclusively

of small, and probably non-reproductive, colo-

nies. At the scale of 10s to 100s of kilometers,

dispersal of larvae may provide continual input of

new recruits, as found for P. meandrina in French

Polynesia (Magalon et al., 2005) and thus, facil-

itate the occurrence of corals in even the most

marginal reef environments (Meesters et al.,

2001). It is important, therefore, to consider

population dynamics, and not just patterns of

abundance, when studying ecological processes

(e.g., disturbances) and their effects on popula-

tions. Multi-scale studies of population size struc-

tures may provide valuable insights into the scale

at which ecological processes affect populations

(e.g., Done, 1988; Bak & Meesters, 1999).

Acknowledgements This manuscript was greatlyimproved following discussions with DC Afzal, AHBaird, RPM Bak and TP Hughes, and critical commentsmade by anonymous reviewers. This research wassupported by grants from the Plan Pluri FormationEPHE, from ‘‘La Fondation d’Entreprise Total’’, fromthe Ministere de l’Outre-Mer, and from the ResearchProgrammes BioAqua (Delegation a la Recherche de laPolynesie francaise, DRRT, ANR) and CoralDiv (ANR).

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