# Seascape attributes, at different spatial scales, determine settlement and post-settlement of juvenile fish

Amalia Cuadros, Joan Moranta, Luis Cardona, Pierre Daniel Thiriet, Jérémy Pastor, Nina Larissa Arroyo, Adrien Cheminée

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## To cite this version:

Amalia Cuadros, Joan Moranta, Luis Cardona, Pierre Daniel Thiriet, Jérémy Pastor, et al.. Seascape attributes, at different spatial scales, determine settlement and post-settlement of juvenile fish. Estuarine, Coastal and Shelf Science, 2017, 185, pp.120-129. 10.1016/j.ecss.2016.12.014. hal-01486006

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Accepted Manuscript

Seascape attributes, at different spatial scales, determine settlement and post-settlement of juvenile fish

Amalia Cuadros, Joan Moranta, Luis Cardona, Pierre Thiriet, Jérémy Pastor, Nina Larissa Arroyo, Adrien Cheminee

<figure><img src="image_3.png" /></figure>

PII: S0272-7714(16)30775-2

DOI: 10.1016/j.ecss.2016.12.014

Reference: YECSS 5344

To appear in: *Estuarine, Coastal and Shelf Science*

Received Date: 25 March 2016

Revised Date: 27 November 2016

Accepted Date: 23 December 2016

Please cite this article as: Cuadros, A., Moranta, J., Cardona, L., Thiriet, P., Pastor, J., Arroyo, N.L., Cheminee, A., Seascape attributes, at different spatial scales, determine settlement and post-settlement of juvenile fish, *Estuarine, Coastal and Shelf Science* (2017), doi: 10.1016/j.ecss.2016.12.014.

This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.
Seascape attributes, at different spatial scales, determine settlement and post-settlement of juvenile fish

Amalia CUADROS¹,⁶*, Joan MORANTA², Luis CARDONA³, Pierre THIRIET⁴,⁵, Jérémy PASTOR⁶, Nina Larissa ARROYO⁷, Adrien CHEMINEE⁶

1. Instituto Español de Oceanografía (IEO), Estació d'Investigació Jaume Ferrer, PO Box 502, 07701 Maó, Spain.

2. Instituto Español de Oceanografía (IEO), Centre Oceanogràfic de les Balears, Moll de Ponent s/n, 07015 Palma de Mallorca, Spain.

3. IRBio, Department of Animal Biology, Faculty of Biology, University of Barcelona, Avinguda Diagonal 643, 08028 Barcelona, Spain.

4. Université de Nice-Sophia Antipolis, EA 4228 ECOMERS, Parc Valrose, 06108 Nice Cedex 2, France.

5. UMR BOREA, Muséum National d'Histoire Naturelle, CRESCO, 38, rue de Port-Blanc, 35800 Dinard, France

6. Univ. Perpignan Via Domitia, Centre de Formation et de Recherche sur les Environnements Méditerranéens (CNRS), UMR 5110, F-66860, Perpignan, France.

7. Institut Mediterrani d'Estudis Avançats (CSIC/UIB), Miquel Marque's 21, 07190 Esporles, Mallorca, Spain.

Emails: amalia.cuadros1@gmail.com*, joan.moranta@ba.ieo.es, luis.cardona@ub.edu, pierre.d.thiriet@gmail.com, jeremy.pastor@univ-perp.fr, nlarroyo@imedea.uib-csic.es, adrien.cheminee@univ-perp.fr.

*Corresponding author: amalia.cuadros1@gmail.com; [add telephone number at proof stage]

**Abstract**

Concern has increased in recent decades regarding processes influencing fish juvenile
density distributions; indeed, juveniles determine the replenishment of populations and
their habitats are often found in shallow coastal areas, where human impacts are
concentrated. We aimed to measure the relative importance of seascape attributes at various spatial scales (from seascape to microhabitat) in fish settlement and post-settlement processes. Along the coast of Minorca Island (NW Mediterranean), *Diplodus sargus* settlement variability was higher among the southern coast compared to the variability in the northern one. Independently of coast location, sheltered nurseries presented lower settlement intensity and recruitment levels compared to exposed ones. Such patterns suggested differential larval supply according to exposure level. Furthermore, subsequent density-dependent post-settlement mortality reduced the cove-specific variability of initial settlement. In addition, inside each cove, juveniles displayed ontogenetic changes of microhabitat use: smaller juveniles were more abundant in the most heterogeneous microhabitat. Consequently, juvenile density distributions responded to seascape attributes at different spatial scales; this suggests that both lager scale attributes and microhabitat influence both settlement and post-settlement processes, and may be limiting for recruitment. Our study demonstrated the importance of a diversified seascape to promote fish population replenishment.

**Key words:** *Diplodus sargus*, juveniles, settlement, growth, survival, connectivity, exposure, habitat structure, Minorca Island, Mediterranean Sea.

**Highlights**

1. Levels of coast exposure controlled settlement intensity and consequent recruitment level.

2. Density-dependent post-settlement mortality levelled site-specific variability of initial settlement and consequent recruitment level.
3. Juveniles of different size-classes displayed different density distribution across microhabitats.

# 1. INTRODUCTION

Most benthic marine species have a pelagic stage which is potentially dispersive (eggs, larvae) (Christie et al., 2010; Di Franco et al., 2012). In such species, various processes act at nested spatial scales to shape their juvenile spatial distribution patterns (Anderson and Millar, 2004; Thiriet, 2014; Zhang et al., 2014). First, at a broad spatial scale, dispersal from breeding areas, and subsequent larval supply to juvenile habitats, i.e., pre-settlement processes, determine the number of planktonic larvae available near juvenile habitat in a given locality (Pineda et al., 2010) and shapes the initial density of pelagic individuals available for settlement (Asplin et al., 1999; Peck et al., 2012; Victor, 1986; Watson and Munro, 2004). Settlement is the transition from the planktonic to the benthic habitats, settlement success at a given locality and year for a given species is measured by the observed maximum number of recently-settled individuals (Levin, 1994; Macpherson et al., 1997; Macpherson and Zika, 1999). Finally, post-settlement processes in benthic habitats, such as juvenile growth and survival determine the number of juveniles available for recruitment. Recruitment into adult populations may require active movement and dispersion towards adult habitats. Frequently, recruitment is measured as the number of late juveniles remaining in the nursery areas after an arbitrary period of time following the main settlement event, i.e. the recruitment level (Levin, 1994; Macpherson et al., 1997; Macpherson and Zika, 1999).

Different factors operate influencing such pre-settlement or post-settlement processes.
Pre-settlement processes are highly influenced by seascape connectivity, which in
parallel to landscape connectivity (Brooks, 2003) may be defined as the degree to which
the seascape facilitates or impedes individual movement among resource patches
(Cheminée et al., 2014; Taylor et al., 1993). According to Calabrese and Fagan (2004),
connectivity has two components: the functional connectivity (dispersal abilities
between populations of individuals and species) and structural connectivity (the
physical connection of populations). Regarding functional connectivity, biological
processes have been highlighted as important for larval dispersal and subsequent larval
supply. For example, larval swimming capacities (Bellwood and Fisher, 2001; Stobutzki
and Bellwood, 1997) and larval habitat selection among settlement areas (Bell and
Westoby, 1986; Gerlach et al., 2006; Leis and Carson-Ewart, 1998; Montgomery et al.,
2006; Simpson et al., 2005; Stobutzki and Bellwood, 1997) regulate functional
connectivity. Structural connectivity may be mainly determined by currents, weather
conditions, seabed topography and coast morphology (Bradbury et al., 2008; Cowen
and Sponaugle, 2009; Gilg and Hilbish, 2003). Larval supply is often not proportional
to settlement, due for example to larval mortality, however it is a constraining variable
for such processes (Pineda et al., 2010).

Once larvae arrive near juvenile habitats, juvenile microhabitat availability (subsets of
biotic and abiotic habitat components (Chapman, 1995)) is very important to
accomplish settlement (Beck et al., 2001; Cheminée et al., 2014; Nagelkerken et al.,
2015; Parsons et al., 2014). Often, fish larvae have strict taxon-specific microhabitat
requirements for settlement (Bussotti and Guidetti, 2010; Cheminée et al., 2013; García-
Rubies and Macpherson, 1995; Harmelin-Vivien et al., 1995; Macpherson and Zika,
1999), that may then change as early juveniles grow at post-settlement (Dahlgren and
Eggleston, 2000; Macpherson, 1998). Post-settlement processes (such as juvenile
growth and survival in benthic habitats until juveniles are ready for recruitment) depend
on the habitat quality, i.e., the trade-off between food and shelter availability (Beck et al., 2001; Cheminée et al., 2013; Gibson, 1994; Hobbs et al., 2006; Thiriet et al., 2014).

The above supports the importance of seascape determining juvenile density distributions, however, the relative importance of seascape attributes at various spatial scales (from seascape to microhabitat) in fish settlement and post-settlement processes and how they influence recruitment level is poorly investigated, since usually efforts are focused in one scale (Macpherson, 1998; Vigliola et al., 1998). The present study aimed to further understand the relative importance of seascape attributes at different spatial scales in settlement and post-settlement processes and how they influence recruitment level. We focus on the case of the white seabream, *Diplodus sargus* (Linnaeus, 1758), an abundant species (Froese and Pauly, 2016), highly dispersive in its early phases, with a commercial and recreational interest (Lloret et al., 2008) and with a functional role in Mediterranean rocky communities (Giakoumi et al., 2012; Guidetti, 2006). Their populations have been suggested to be recruitment limited in some areas (Cardona et al., 2007) and different authors reported high spatial variability of its settlement rates and recruitment levels (Arceo et al., 2012; Cheminee et al., 2011; Macpherson et al., 1997; Vigliola et al., 1998). We quantified and compared *D. sargus* settlement and post-settlement densities under the influence of contrasted seascape attributes, at different spatial scales around Minorca Island (NW Mediterranean Sea). Firstly, we tested the effect of coastal location by comparing coves facing contrasted hydrographic regimes: north vs. south coast of the island. Secondly, and crossed with location, we tested the effect of cove configuration in terms of exposure level, and hence with contrasted hydrographic regimes. Furthermore, within each cove we measured the importance of post-settlement processes (growth and survival) to test whether such processes reduced
or influenced the variability of initial settlement. Finally, within each cove we tested the
effect of microhabitats features on juvenile size-class density distributions.

**2. MATERIAL AND METHODS**

**2.1. Studied area**

Minorca Island (North-western Mediterranean) (Fig. 1) spans about 50 km from west to east and about 20 km from north to south and its coastline measures 441 km (Sales, 2007). The island is an ideal study locality since its coasts present contrasted exposure and orientation conditions. Indeed, because of the geomorphology of the island, the coast can be divided into two differentiated regions with different wind and hydrographic influences: north and south, see Llompart et al. (1979). The north shore faces water masses originating from the Lion Gulf (López-Jurado et al., 2008). The south shore faces water masses originating from the Algerian Basin (Bethoux, 1980; Millot, 1999; Pinardi and Masetti, 2000). It is assumed that these water masses may carry different fish larval inputs. Furthermore, the shoreline displays series of coves with various levels of exposure (i.e. more or less enclosed). Based on a preliminary study (Cuadros, 2015), we selected coves with suitable nursery habitats for *D. sargus*: shallow (less than 2 m deep), with gently sloping substrates of sand, gravel, pebbles, and boulders (Bussotti and Guidetti, 2010; García-Rubies and Macpherson, 1995; Harmelin-Vivien et al., 1995; Macpherson, 1998).

**2.2. Sampling design**

The study was conducted in twelve coves selected according to crossed factors: location (two levels: north and south) and cove configuration in terms of exposure (two levels: exposed and sheltered) (Fig. 1). To define the two exposure levels we adapted Miller's
index of exposure (1985) to Minorca coves: sheltered and exposed coves were
respectively without and with line of sight to the open ocean (Fig. 1).

<figure><img src="image_4.png" /><figcaption>Fig. 1. The twelve sampled coves, according to crossed factors “location” (two levels: north and south) and “exposure” (two levels: exposed (gray squares) and sheltered (black squares)). 1= “Estancats”, 2= “Rotja”, 3= “Cabra Salada”, 4= “S’Esmolador”, 5= “Sa Mesquida”, 6= “Taulera”, 7= “Sa Caleta”, 8= “Es Talaier”, 9= “Mitjana”, 10= “Cales Coves”, 11= “Biniparratx”, 12= “Binibeca”.</figcaption></figure>

We additionally differentiated three microhabitat types inside each cove (Macpherson,
1998): ‘beach area’ (BA) usually characterized by sandy sediments and a depth of 0-50
cm; ‘mixed areas’ (MA) with the presence of rocks, boulders or pebbles in the sand,
ranging 50-100 cm in depth (Supplemental material #1); and ‘rocky areas’ (RA) with
rocky substratum ranging 50-200 cm in depth.

**2.3. Data collection**

A total of 144 surveys were carried out from 17 th May to 18 th July, in 2013, during the
known *D. sargus* settlement period (Harmelin-Vivien et al., 1995) (Supplemental
material #2). We sampled all coves at three different times, every two (or three) weeks.
Additionally, exposed coves (which displayed higher juvenile densities, see results) were sampled weekly, when weather permitted, for the analysis of the microhabitat effect. Sampling days were randomly chosen among days presenting good weather conditions, since weather is known to affect density estimations (Harmelin-Vivien et al., 1985). Surveys were conducted between 7 am and 11 am, avoiding the time of higher bathing activity (Juaneda and Roig, 2002; Munar and Xavier, 2003).

In each cove, sampling was carried out snorkelling parallel to the shoreline, covering the entire suitable habitat. Underwater Visual Census (UVC) was performed along transects within the three defined microhabitats (mean of 30 m²) by a single diver. The diver recorded abundances and sizes of juvenile *D. sargus*. Total length (TL) was estimated with the help of fish silhouettes of different sizes pictured on the slate (5 mm TL size-classes from 10-15 mm to 80-85 mm TL) (Cheminee et al., 2011, 2013; Pastor et al., 2013; Vigliola et al., 1998). The approximate TL of *Diplodus sargus* at settlement is 10 mm. The precision of this size estimating method was estimated about ± 3.5 mm for *Diplodus* species in a previous study (Macpherson, 1998). For the analyses, juvenile fish densities were standardised to one linear metre of shoreline (ind.m⁻¹) (Arceo et al., 2012; Cheminee et al., 2011; Harmelin-Vivien et al., 1985; Pastor et al., 2013).

**2.4. Statistical analyses**

In order to study juvenile population dynamics according to the three seascape attributes (i.e. location, exposure, and microhabitat type, thereafter named as treatments), five response variables were analyzed: 1) multivariate density patterns across surveys; 2) univariate density at each of the three simultaneous surveys; 3) univariate daily mean total length; 4) univariate daily mean total density, and 5) univariate mean juvenile density by size-class.
In order to test the crossed influence of location and exposure, fish sufficiently large to
decrease their fidelity to nursery areas and then more likely to disperse were excluded
(i.e. for *D. sargus*, >40 mm TL; Vigliola (1998)) and only density data corresponding to
the three surveys conducted simultaneously at all coves were considered (hereafter
referred as ‘first’, ‘second’ and ‘third’ surveys). First we applied a multivariate
PERMutational ANalysis Of VAriance (PERMANOVA) (Anderson, 2001) to test for
differences of mean density patterns through surveys between groups of sites. If
differences were detected, univariate PERMANOVAs were applied to each survey
individually to test for survey-specific mean density differences between groups of
sites. Thus, more precisely, first, we used the multivariate data-frame displaying
densities per surveys (as columns) and per sites (as rows): juvenile multivariate density
patterns across the three surveys were compared between studied factors by means of a
multivariate PERMANOVA Type III, based on the Binomial deviance (scaled) measure
of distance (Anderson and Gorley, 2008). Crossed terms included in the model were:
location as a fixed factor (two levels: north vs. south) and exposure as a fixed factor
(two levels: exposed vs. sheltered). Furthermore, when interactions of factors were
significant we applied pair-wise comparison tests. Secondly, for each individual survey,
a univariate PERMANOVA Type III, using Euclidean distances, was applied in order to
compare density between treatments levels (location, exposure). First and second
surveys corresponded respectively to census from days 2/06 to 8/06 and days 20/06 to
23/06. The first survey was considered as a proxy of settlement peak (i.e. 10-15 mm TL
fish peak density) since when sampling started, juveniles of 10-15 mm were already
present (Cheminee et al., 2011). The third survey corresponded to days 16/07-18/07 and
was considered as a measure of the recruitment level (Macpherson and Zika, 1999). For
these permutational analyses and when modelling our various response variables as a
function of the treatments, P-values were obtained by 999 permutations of residuals under a reduced model. Monte Carlo P-values were considered when there were insufficient possible permutations (<200). Terms were pooled as suggested by Anderson et al. (2008).

Since sheltered coves supported very few fish (see results), only exposed coves were considered for subsequent analyses. Daily juvenile mean total length (MTL) (mm) and daily total density (ind·m⁻¹) of each cove were used in Least squared regression analyses to calculate regression slopes ('b') and obtain respectively juvenile growth rates (mm day⁻¹) and density decrease rates (ind·m⁻¹·day⁻¹) for each cove. These rates were used to describe the post-settlement population dynamics (Vigliola, 1998). Regressions were calculated using the daily MTL of all the juveniles present at each cove, from the first to the last sampling day, and density decrease rates were calculated considering the density of individuals from the first to the last day of sampling and excluding fish capable to decrease their level of fidelity to nursery areas (i.e. for *D. sargus*, >40 mm TL, Vigliola (1998)). Analysis of covariance (ANCOVA) was used to test the effect of cove (included as a categorical factor in the model) on MTLs and densities, considered as dependent variables, while controlling for the effect of time, considered as a continuous co-variable. Tukey HSD post-hoc comparisons between coves were performed for MTL and density when appropriate.

Finally, ontogenetic changes in habitat use were assessed by comparing the density of juveniles of different size-classes occurring at the three microhabitats within each cove. All the juveniles observed at the nurseries were considered and they were split in five size-classes (in mm TL): [10-25], [25-40], [40-55], [55-70]. The [70-85] mm TL size class was not considered for further analyses due to the low number of fish (n<10).
Each size class was analysed separately in a univariate PERMANOVA Type III, using Euclidean distances. In the model, factor microhabitat had three levels (BA, MA and RA) and was fixed; and factor cove had six levels (Estancats, Rotja, Sa Mesquida, Es Talaier, Mitjana, Binibeca) and was fixed. Terms were pooled as suggested by Anderson et al. (2008). P-values were obtained by 999 permutations of residuals under a reduced model. Monte Carlo P-values were considered when there were insufficient possible permutations (<200).

Since ecological data are by nature highly variable, in this work terms were considered significant for P-values < 0.1. All data treatment and analyses were performed using the R 2.15.0 statistical software (R_Development_Core_Team, 2013) and PERMANOVA+ add on package for PRIMER software (Anderson and Gorley, 2008; Clarke and Gorley, 2006).

3. RESULTS

**3.1. Effect of location and exposure on juvenile density temporal patterns**

As a general pattern, a decreasing density trend was observed in the all exposed coves,
from the beginning to the end of the sampling period. Conversely density remained
close to zero ind.m⁻¹ through the whole sampling period in the sheltered coves (Fig. 2).
Density trends differed between exposed and sheltered coves, whatever the location,
although the magnitude of the difference was greater in the south coves, which resulted
in a significant interaction term (PERMANOVA, Table 1). At each survey, densities
differed according to exposure and not according to location (PERMANOVA, Table 1)
and juveniles were systematically more abundant in exposed coves (Fig. 3).
<figure><img src="image_5.png" /><figcaption>Fig. 2. *Diplodus sargus* juvenile density (ind.m⁻¹) in each cove (coves 1 to 12) through time, according to location (north vs south) and exposure (exposed vs sheltered). Coves: 1= "Estancats", 2= "Rotja", 3= "Cabra Salada", 4= "S'Esmolador", 5= "Sa Mesquida", 6= "Taulera", 7= "Sa Caleta", 8= "Es Talaier", 9= "Mitjana", 10= "Cales Coves", 11= "Biniparratx", 12= "Binibeca".</figcaption></figure>

<table><caption>Table 1. PERMANOVA table of results: effect of location and exposure on juvenile *Diplodus sargus* multivariate density patterns across surveys (ind.m⁻¹) and on univariate juvenile *Diplodus sargus* density at each of the three simultaneous surveys at all the coves (ind.m⁻¹); the first survey is a proxy of the settlement peak and third survey a proxy of the recruitment level. Significance: *P*≤0.1; **P** ≤ 0.05; ***P** ≤ 0.01.</caption><thead><tr><th colspan="5">Multivariate PERMANOVA of juvenile density patterns across surveys</th></tr><tr><th>Source of variation</th><th>Df</th><th>MS</th><th>Pseudo-F</th><th>P(perm)</th></tr></thead><tbody><tr><td>Location (lo)</td><td>1</td><td>0.58</td><td>2.03</td><td>0.131</td></tr><tr><td>Exposure (ex)</td><td>1</td><td>6.26</td><td>21.94</td><td><strong>0.003**</strong></td></tr><tr><td>lo x ex</td><td>1</td><td>0.91</td><td>3.18</td><td><strong>0.052</strong></td></tr><tr><td>Residuals</td><td>8</td><td>0.29</td><td></td><td></td></tr><tr><td>Total</td><td>11</td><td></td><td></td><td></td></tr></tbody></table><table><thead><tr><th colspan="5">Univariate PERMANOVA of juvenile density at the three individual surveys</th></tr><tr><th colspan="5">First survey (proxy of settlement peak)</th></tr><tr><th>Source of variation</th><th>Df</th><th>MS</th><th>Pseudo-F</th><th>P(perm)</th></tr></thead><tbody><tr><td>Location (lo)</td><td>1</td><td>2.76</td><td>3.19</td><td>0.105</td></tr><tr><td>Exposure (ex)</td><td>1</td><td>10.25</td><td>11.86</td><td><strong>0.009**</strong></td></tr><tr><td>lo x ex</td><td>1</td><td>1.26</td><td>1.45</td><td>0.292</td></tr><tr><td>Residuals</td><td>8</td><td>0.86</td><td></td><td></td></tr><tr><td>Total</td><td>11</td><td></td><td></td><td></td></tr></tbody></table><table><thead><tr><th colspan="5">Second survey</th></tr><tr><th>Source of variation</th><th>Df</th><th>MS</th><th>Pseudo-F</th><th>P(perm)</th></tr></thead><tbody><tr><td>Location (lo)</td><td>1</td><td>4.76</td><td>2.47</td><td>0.151</td></tr><tr><td>Exposure (ex)</td><td>1</td><td>13.42</td><td>6.96</td><td><strong>0.019*</strong></td></tr><tr><td>lo x ex</td><td>1</td><td>3.77</td><td>1.95</td><td>0.184</td></tr><tr><td>Residuals</td><td>8</td><td>1.93</td><td></td><td></td></tr><tr><td>Total</td><td>11</td><td></td><td></td><td></td></tr></tbody></table><table><thead><tr><th colspan="5">Third survey (proxy of recruitment level)</th></tr><tr><th>Source of variation</th><th>Df</th><th>MS</th><th>Pseudo-F</th><th>P(perm)</th></tr></thead><tbody><tr><td>Location (lo)</td><td>1</td><td>0.18</td><td>0.84</td><td>0.384</td></tr><tr><td>Exposure (ex)</td><td>1</td><td>1.19</td><td>5.72</td><td><strong>0.039*</strong></td></tr><tr><td>lo x ex</td><td>1</td><td>6.86.10-2</td><td>0.33</td><td>0.576</td></tr></tbody></table>
<table><tr><td>Residuals</td><td>8</td><td>0.21</td></tr><tr><td>Total</td><td>11</td><td></td></tr></table>

<figure><img src="image_6.png" /><figcaption>Fig. 3. *Diplodus sargus* mean density (ind.m⁻¹) per exposure level (exposed vs. sheltered coves) the three sampled dates for all coves: first survey was employed as a proxy of settlement peak, and third survey was considered a proxy of recruitment level. Mean and standard error (SE) values are given.</figcaption></figure>

## 3.2. Post-settlement dynamics in exposed coves

For exposed coves, MTLs changed through time similarly in all coves (ANCOVA, Table 2) and corresponding growth rates ranged from 0.375±0.043 mm.day⁻¹ (in Mitjana) to 0.486±0.054 mm.day⁻¹ (in Binibeca). Furthermore, south coves, Talaier and Mitjana, presented significantly different density trends than the others, with higher decrease rates (ANCOVA, Table 2; Tukey HSD post-hoc, Supplemental material 3#; and Fig. 4).
<table><caption>Table 2. Post-settlement dynamics in exposed coves. Results of the predicted relationship between A) Mean Total Lenghts (MTLs, mm) and B) densities of juvenile *Diplodus sargus* with time in each cove and results of the analysis of covariance (ANCOVA) comparing the A) MTLs (mm) and B) densities (ind.m⁻¹) according to time and coves (interaction cove-day was removed when not significant). a: intercept: b: slope; r²: correlation coefficient; Df: degrees of freedom; MS: mean squared; F: F statistic with its Df (Degree of freedom); and its significance (P≤0.1; *P ≤ 0.05; **P ≤ 0.01; ***P ≤ 0.001).</caption><thead><tr><th colspan="7">A) MTL</th></tr><tr><th>Exposure</th><th>Location</th><th>Linear Regression</th><th>a</th><th>b</th><th>r<sup>2</sup></th><th>F</th><th>P</th></tr></thead><tbody><tr><td rowspan="6">Exposed</td><td rowspan="3">North</td><td>Estancats</td><td>8.557±1.576</td><td>0.438±0.038</td><td>0.956</td><td>131.9 (1-5 DF)</td><td><strong>&lt;0.001***</strong></td></tr><tr><td>Rotja</td><td>9.558±1.387</td><td>0.400±0.037</td><td>0.944</td><td>119.4 (1-6 DF)</td><td><strong>&lt;0.001***</strong></td></tr><tr><td>Sa Mesquida</td><td>11.938±2.408</td><td>0.384±0.063</td><td>0.839</td><td>37.45 (1-6 DF)</td><td><strong>&lt;0.001***</strong></td></tr><tr><td rowspan="3">South</td><td>Es Talaier</td><td>8.248±1.258</td><td>0.412±0.031</td><td>0.967</td><td>178.1 (1-5 DF)</td><td><strong>&lt;0.001***</strong></td></tr><tr><td>Mitjana</td><td>10.709±1.763</td><td>0.375±0.043</td><td>0.924</td><td>74.26 (1-5 DF)</td><td><strong>&lt;0.001***</strong></td></tr><tr><td>Binibeca</td><td>5.802±2.163</td><td>0.486±0.054</td><td>0.931</td><td>81.94 (1-5 DF)</td><td><strong>&lt;0.001***</strong></td></tr><tr><th colspan="2">ANCOVA</th><th>Df</th><th>MS</th><th>F</th><th>P</th><th></th></tr><tr><td colspan="2">Date</td><td>1</td><td>2645.8</td><td>483.31</td><td>&lt;0.001***</td><td></td></tr><tr><td colspan="2">Cove</td><td>5</td><td>7.3</td><td>1.34</td><td>0.297</td><td></td></tr><tr><td colspan="2">Residuals</td><td>37</td><td>5.5</td><td></td><td></td><td></td></tr><tr><th colspan="7">B) Densities</th></tr><tr><th>Exposure</th><th>Location</th><th>Linear Regression</th><th>a</th><th>b</th><th>r<sup>2</sup></th><th>F</th><th>P</th></tr><tr><td rowspan="6">Exposed</td><td rowspan="3">North</td><td>Estancats</td><td>1.156±0.151</td><td>-0.012±0.004</td><td>0.603</td><td>10.12 (1-5 DF)</td><td><strong>0.025*</strong></td></tr><tr><td>Rotja</td><td>1.460±0.293</td><td>0.003±0.008</td><td>-0.131</td><td>0.19 (1-6 DF)</td><td>0.678</td></tr><tr><td>Sa Mesquida</td><td>1.432±0.316</td><td>-0.019±0.008</td><td>0.367</td><td>5.05 (1-6 DF)</td><td><strong>0.066*</strong></td></tr><tr><td rowspan="3">South</td><td>Es Talaier</td><td>8.894±1.201</td><td>-0.103±0.029</td><td>0.652</td><td>12.26 (1-5 DF)</td><td><strong>0.017*</strong></td></tr><tr><td>Mitjana</td><td>4.799±0.518</td><td>-0.060±0.013</td><td>0.777</td><td>21.87 (1-5 DF)</td><td><strong>0.005**</strong></td></tr><tr><td>Binibeca</td><td>0.952±0.378</td><td>-0.001±0.009</td><td>-0.198</td><td>0.01 (1-5 DF)</td><td>0.928</td></tr><tr><th colspan="2">ANCOVA</th><th>Df</th><th>MS</th><th>F</th><th>P</th><th></th></tr><tr><td colspan="2">Date</td><td>1</td><td>11.85</td><td>23.50</td><td><strong>&lt;0.001***</strong></td><td></td></tr><tr><td colspan="2">Cove</td><td>5</td><td>21.33</td><td>42.32</td><td><strong>&lt;0.001***</strong></td><td></td></tr><tr><td colspan="2">Date x cove</td><td>5</td><td>4.36</td><td>8.64</td><td><strong>&lt;0.001***</strong></td><td></td></tr><tr><td colspan="2">Residuals</td><td>32</td><td>0.50</td><td></td><td></td><td></td></tr></tbody></table>

<figure><img src="image_7.png" /><figcaption>Fig. 4. Observed data and predicted relationship of densities (ind.m⁻¹) of *Diplodus sargus* with time in each cove. Observed data=dots; predicted relationship =lines.</figcaption></figure>

### 3.3. Juvenile ontogenetic changes in microhabitat use
For most of juvenile size-classes, in each cove, densities differed between the three microhabitats considered (PERMANOVA, Table 3), except for juveniles in the size-class [55, 70]. Furthermore, the interaction term between microhabitat and cove was not significant for each size class separately, thus indicating that tendencies of density distribution among microhabitats within the nursery were consistent across all coves (Fig. 5).

<table><caption>Table 3. PERMANOVAs table of results for *Diplodus sargus* juvenile densities in exposed coves in different microhabitats inside the nursery, for each size classes (mm TL): [10, 25), [25, 40), [40, 55), [55, 70). Significance: · P ≤ 0.1; * P ≤ 0.05; ** P ≤ 0.01; *** P ≤ 0.001.</caption><thead><tr><th colspan="5">[10, 25) mm TL</th></tr><tr><th>Source</th><th>df</th><th>MS</th><th>Pseudo-F</th><th>P(perm)</th></tr></thead><tbody><tr><td>Microhabitat (mi)</td><td>2</td><td>63.42</td><td>5.64</td><td><strong>0.008**</strong></td></tr><tr><td>Cove (co)</td><td>5</td><td>36.01</td><td>3.20</td><td><strong>0.016*</strong></td></tr><tr><td>mi x co</td><td>9</td><td>12.53</td><td>1.11</td><td>0.335</td></tr><tr><td>Residuals</td><td>372</td><td>11.24</td><td></td><td></td></tr><tr><td>Total</td><td>388</td><td></td><td></td><td></td></tr></tbody><tbody><tr><th colspan="5">[25, 40) mm TL</th></tr><tr><th>Source</th><th>df</th><th>MS</th><th>Pseudo-F</th><th>P(perm)</th></tr><tr><td>Microhabitat (mi)</td><td>2</td><td>12.39</td><td>4.86</td><td><strong>0.008**</strong></td></tr><tr><td>Cove (co)</td><td>5</td><td>18.32</td><td>7.19</td><td><strong>0.001***</strong></td></tr><tr><td>mi x co</td><td>9</td><td>2.94</td><td>1.16</td><td>0.327</td></tr><tr><td>Residuals</td><td>372</td><td>2.55</td><td></td><td></td></tr><tr><td>Total</td><td>388</td><td></td><td></td><td></td></tr></tbody><tbody><tr><th colspan="5">[40, 55) mm TL</th></tr><tr><th>Source</th><th>df</th><th>MS</th><th>Pseudo-F</th><th>P(perm)</th></tr><tr><td>Microhabitat (mi)</td><td>2</td><td>1.40</td><td>3.92</td><td><strong>0.024*</strong></td></tr><tr><td>Cove (co)</td><td>5</td><td>0.81</td><td>2.25</td><td><strong>0.056*</strong></td></tr><tr><td>mi x co</td><td>9</td><td>0.47</td><td>1.30</td><td>0.218</td></tr><tr><td>Residuals</td><td>372</td><td>0.36</td><td></td><td></td></tr><tr><td>Total</td><td>388</td><td></td><td></td><td></td></tr></tbody><tbody><tr><th colspan="5">[55, 70) mm TL</th></tr><tr><th>Source</th><th>df</th><th>MS</th><th>Pseudo-F</th><th>P(perm)</th></tr><tr><td>Microhabitat (mi)</td><td>2</td><td>2.62·10<sup>-3</sup></td><td>0.68</td><td>0.519</td></tr><tr><td>Cove (co)</td><td>5</td><td>1.09·10<sup>-2</sup></td><td>2.80</td><td><strong>0.028*</strong></td></tr><tr><td>mi x co</td><td>9</td><td>2.00·10<sup>-3</sup></td><td>0.51</td><td>0.828</td></tr><tr><td>Residuals</td><td>372</td><td>3.88·10<sup>-3</sup></td><td></td><td></td></tr><tr><td>Total</td><td>388</td><td></td><td></td><td></td></tr></tbody></table>
<figure><img src="image_8.png" /><figcaption>Fig. 5. *Diplodus sargus* juvenile densities among microhabitats (BA: Beach Area, MA: Mixed Area, RA: Rocky Area) for each size-classes: [10, 25], [25, 40], [40, 55) and [55, 70) mm TL; Corresponding number of individuals (n) and pair-wise tests results between microhabitat levels are given (different lower case characters indicate significant density differences between microhabitats).</figcaption></figure>

Pair-wise comparisons of juvenile density between microhabitats types (Fig. 5) revealed that juveniles of the [10, 25] and [25, 40] size classes were more abundant in mixed areas (MA) than in the beach areas (BA) or rocky areas (RA) (pair-wise habitat tests, p<0.05). Fish of the [40-55] size-class were more abundant in both MA and BA than in RA (p<0.01). Finally, fish of the [55, 70] size-class were homogeneously distributed in the three microhabitats (pair-wise test p>0.1).

4. DISCUSSION

At the island scale (several tens of km), location of the coves in the north or the south coast, which *a priori* face different hydrographic conditions, did not have a major impact on the settlement peak nor on the recruitment level or on the growth rate of juvenile *D. sargus*. Nevertheless, coves in the south coast exhibited a higher variability
in the density trends than those in the north, because of the very high settlement peak in
two south exposed coves (Es Talaier and Mitjana). Site-specific variability of settlement
peak of such species has been attributed in the literature mainly to habitat structure
(which influences habitat quality, see e.g. Anderson and Millar, 2004; García-Charton et
al., 2004; Thiriet et al., 2014) although there are no studies which test this (Cheminee et
al., 2011; Vigliola et al., 1998). The two coves in the south shore with the highest
settlement peaks displayed a highly heterogeneous habitat, with the patches of the three
defined microhabitats lying in close quarters. Since habitat complexity and
heterogeneity (sensu August 1983) determine shelter and food availability and hence
subsequent mortality and growth, it is plausible that observed differences in settlement
peak were due to such heterogeneity (Anderson and Millar, 2004; García-Charton et al.,
2004). Furthermore, in view of the fact that *D. sargus* spawns in areas far from coast
and larvae spend on average 28 days among the plankton (Vigliola, 1998), these
differences may be also related to the relative location and availability of nurseries
*versus* spawning sites, coupled with differences in local hydrodynamics (Pineda et al.,
2010; Roy, 2012). Indeed, in the south, where high settlement peaks were recorded both
in 2012 (Cuadros, 2015) and 2013, the coast displays a smaller amount of available
nursery habitat than in the north part (Cuadros, 2015): this may lead juveniles to
concentrate in the few available habitats, leading to the observed high densities.
However there is no precise information about the spawning areas for this species in the
Mediterranean and specifically Minorca, and detailed hydrodynamics in the locality are
still poorly known.

At lower seascape scale (a few km), settlement peak and density trends were controlled
by exposure, as both the settlement and the recruitment peak were significantly higher at
the exposed coves, independently on the location in the coastline. The density of
juvenile *D. sargus* in the sheltered coves remained close to zero ind.m⁻¹ throughout the
whole sampling period. Although we did not accomplish a direct examination of larval
abundances, this result suggests a positive influence of cove exposure on larval supply
and subsequent settlement intensity. In this sense larval supply may have been
functioning as a limiting variable (Pineda et al., 2010). Some studies have coupled
hydrological regimes with bathymetry, topography and the distribution of fish eggs and
larvae: available data indicate a match between the distribution of these spawning
products and hydrodynamics (Brown et al., 2005; Jenkins et al. 1998; Martins et al.,
2007; Pepin et al., 1995). It usually results into a low egg/larval supply to inner parts of
the shoreline, e.g. the inner parts of the coral reefs systems (D'Alessandro et al., 2007),
the upper reaches of the estuaries (Bell et al., 1988; Martins et al. 2007) and the
innermost part of bays. For this reason, settlement is favoured in areas close to open
waters, if suitable habitats exist (Brown et al., 2005; Jenkins et al., 1998; Pastor et al.,
2013). This conceptual framework is consistent with the results reported here.
Nevertheless, although a given degree of exposure may be favourable, it is notable that
previous research on *D. sargus* also showed the preference of settlers for coast not only
with gentle slope and heterogeneous boulders and pebbles bottoms, but as well
protected from swell and strong hydrodynamics (i.e. coves) (Harmelin-Vivien et al.,
1995). Vigliola (1998) even suggested that survival of *D. sargus* juveniles was favoured
in the most enclosed coves, through enhanced water temperature and consequent faster
growth.

Once larvae have reached the nursery habitat and settlement is accomplished, post-
settlement processes determine their survival and growth (Vigliola, 1998). In our study,
recruitment level was compared using a crossed analysis (location x exposure), but very
low numbers of juveniles occurred in sheltered coves, so the analyses of post-settlement
dynamics were conducted only at the exposed coves. There, the growth of juveniles was similar between north and south coves and within the range reported by previous research for this species (Planes et al., 1999; Vigliola, 1998). This indicates that juvenile growth rate was independent on juvenile density, since large differences existed among coves in settlement peaks but juveniles grew at a similar rate everywhere. This is consistent with a previous work in 2012 in the area (Cuadros, 2015) but in contrast with the density-dependent growth patterns reported by Planes et al. (1998) for this species. This lack of density dependence for growth may suggest that food resources for juveniles were not a limiting factor, discarding a possible intraspecific competition for food resources (Hazlerigg et al., 2012; Shephard and Jackson, 2009).

Contrastingly, the rate of density decrease exhibited a strong density-dependent pattern, as juvenile density decreased faster in those coves where the settlement peak was the highest. As a result the recruitment level was similar at the six exposed coves. Density-dependent mortality has been described in previous studies about this species (Macpherson et al., 1997; Planes et al., 1998). Competition for food resources is not a likely cause, because of the density-independent growth rate reported above, and hence mortality by predation combined with competition for shelter stands as the main causes for the density dependent depletion of juveniles (Forrester and Steele, 2000; Hixon and Jones, 2005; Anderson, 2001). Decoupling larval supply, settlement and recruitment, has been frequently reported but there is no consensus regarding the relative importance of pre-settlement, settlement and post-settlement processes in determining recruitment (Di Franco et al., 2013; Félix-Hackradt et al., 2013; Jenkins et al., 1998; Macpherson and Zika, 1999; Pineda et al., 2010). In this sense, topography may play a pivotal role decoupling and modulating the relative importance of these processes. In fact, in this study, although post-settlement processes, i.e. density-dependent mortality certainly
resulted into major changes in juvenile density, seascape attributes in terms of cove
exposure restricted settlement (and likely pre-settlement) and stand as the main process
limiting the recruitment of *D. sargus*.

Coast configuration and thus suitable habitat availability is also likely to cause the
decoupling observed between the high settlement peaks reported in some coves in this
study and the global low density of adult *D. sargus* in suitable habitats in Minorca as
compared with other adjoining regions (Cardona et al., 2013, 2007; Coll et al., 2012).
These low adult densities were attributed to the oligotrophy of the coastal waters of the
island and a limited larval supply (Cardona et al., 2007). However, as regards to the
high settlement values reported here in some coves, it is more plausible that settlement
and subsequent recruitment are limited due to the scarcity of suitable coves as respect to
the coastline along most of the south and northwest coast of Minorca. Indeed, Cuadros
(2015) highlight that these nursery habitats are scarce since these coves with suitable
habitats for juveniles represent less than 9% of the shoreline. Finally, at the nursery
scale (tens of m), juveniles of different size classes were distributed differently across
microhabitats. Smaller juveniles (i.e. settlers and juveniles <40 mm TL) mainly
occurred in the mixed areas. As they grew up, they also used the beach area in the same
proportion and finally sub-adult fishes larger than >55 mm used equally all the
microhabitats present inside the nurseries, without any preference. Since 'mixed areas'
usually occur in the more sheltered parts of the coves, Macpherson (1998) suggested for
*D. sargus* a larval distribution mechanism related to slower water flow, similar to that
reported by Breitburg et al. (1996). In this sense, early juveniles, with limited swimming
capabilities, were dragged to those backwaters. However, for other species, Dalgreen
and Eggleston (2000) suggested that ontogenetic changes of microhabitats aim to switch
to the habitat displaying the better compromise between food availability and predation
risk. Our observations might consequently also be related to shelter availability, especially important in early juvenile stages (Sogard, 1997): more shelter may be provided by the more structured habitat (i.e. the mixed areas) at the beginning of *D. sargus* development in the nurseries. Furthermore, early individuals may feed mainly on harpacticoids (Christensen, 1978); some of our own data of juvenile stomach content and microhabitat food availability indicate a clear preference of smaller juveniles for harpacticoids which were usually found in the seaweed in the pebbles and rocks, rather than interstitial harpacticoids which inhabited the sand (Cuadros, 2015). Later, older juvenile fish, that are less vulnerable (>40 mm) may explore new habitat with less shelter (Anderson, 1988; Sogard, 1997). They may begin to move to other microhabitats inside the cove (such as beach or rocky areas), displaying a home-range expansion, habitat diversification, diet diversification, and starting their emigration from the nursery (Christensen, 1978; Macpherson, 1998; Vigliola and Harmelin-Vivien, 2001). This supports the importance of the presence of various microhabitats for accomplish the development of *D. sargus* juveniles.

From a coastal management perspective, our results underline the importance of a multi-scale approach to identify important areas for the development of *Diplodus sargus* juveniles: from coast configuration, taking into account cove exposure, to habitat and microhabitat availability. Furthermore, in order to protect such nursery areas it is recommended to avoid human activities involving the homogenisation of the coastal seabed. Indeed, in the Mediterranean Sea, nurseries of such species tend to be physically destroyed even in protected areas, due to coastal development projects and physical construction on them (Cheminée et al., 2014; Meinesz et al., 2006). Furthermore, beach management planning through all Mediterranean coasts usually do not take into account the nursery habitats of this species. Indeed, classical beach management is restricted to
the protection of dune systems, and possible associated wet-lands, the limitation of anchoring in seagrass meadows, and the removing by heavy machinery (tracks, tractors, etc.) of the cast material of seagrass deposited along the shoreline (Borum et al., 2004; Silva et al., 2007; Van der Meulen and Salman, 1996). Beach nourishment projects are a usual practice in Europe and worldwide (Davison et al., 1992; Hanson et al., 2002) and may potentially affect broadly the replenishment of *Diplodus sargus* species, since it homogenises the sea bed in coves, and consequently causes the disappearance of the juvenile micro-habitat, i.e., the heterogeneous mixed bottoms with pebbles, boulders and rocks.

## 5. CONCLUSIONS

Seascape attributes at different spatial scales influence settlement and post-settlement and determine the density distributions and recruitment levels of juvenile *D. sargus*. Among coves displaying suitable microhabitats for *D. sargus* juveniles, exposed ones are the main nursery areas for *D. sargus*, since sheltered ones reported a very low recruitment peak preceded by very low settlement rates. Density-dependent mortality modulates and determines the recruitment peak in exposed coves with a high supply of settlers, thus resulting in partial decoupling between settlement and recruitment. Finally, at a finer scale inside suitable coves, microhabitats characteristics drive ontogenetic changes in microhabitats preferences. In addition, in Minorca, total recruitment might be limited by the scarcity of suitable habitats, mainly along the south and northwest coast of the island. From a coastal management perspective, we may conclude that a multiscale-approach is necessary to identify key areas for *Diplodus sargus* juveniles; furthermore, it is recommended to avoid human activities involving the homogenisation of the coastal seabed such as coastal development projects or beach nourishments.
Acknowledgements

This study was performed at the Estació d'Investigació Jaume Ferrer (Jaume Ferrer Research Station) from the Govern de les Illes Balears (Balearic Island Government). It was part of ACU's PhD thesis, co-founded by the Govern de les Illes Balears and the Centre Oceanografic de les Balears (Oceanographic Center of Balearic Islands). Authors are grateful to the members from CEFREM lab which provided helpful criticism of the manuscript in various stages of completion. Finally, we would also like to thank the anonymous reviewers of this paper for constructive comments.

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