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Estuarine, Coastal and Shelf Science 92 (2011) 581e587

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Estuarine, Coastal and Shelf Science

journal homepage: www.elsevier .com/locate/ecss

Short-term residence and movement patterns of the annular seabream Diplodusannularis in a temperate marine reserve

David March a,*, Josep Alós a, Amàlia Grau b, Miquel Palmer a

a Instituto Mediterráneo de Estudios Avanzados, IMEDEA (UIB-CSIC), Miquel Marqués 21, 07190 Esporles, Islas Baleares, Spainb Laboratori d’Investigacions Marines i Aqüicultura, LIMIA, Eng. Gabriel Roca 69, 07157 Port d’Andratx, Islas Baleares, Spain

a r t i c l e i n f o

Article history:Received 24 March 2010Accepted 23 February 2011Available online 12 March 2011

Keywords:acoustic telemetrytaggingSparidaesite fidelityhoming behaviourPalma BayLongitude: 2�410000Ee2�480000ELatitude: 39�250000Ne39�300000

* Corresponding author.E-mail address: david.march@uib.es (D. March).

0272-7714/$ e see front matter � 2011 Elsevier Ltd.doi:10.1016/j.ecss.2011.02.015

a b s t r a c t

The short-term movements of a small temperate fish, the annular seabream Diplodus annularis (Linnaeus1758), were examined using standard tag-recapture and passive acoustic telemetry in Palma Bay (NWMediterranean), a marine protected area (MPA). The study aimed to provide valuable information forassessing the recreational fishery and its results suggest that MPAs can be used to protect the adult stockof D. annularis. All the fish tagged with standard tags were recaptured near the release locations, witha maximum distance of w300 m. The maximum time between release and recapture was 185 d. Twodifferent arrays of acoustic receivers were deployed, one in 2008 and another in 2009, within the MPA.Twenty adults were surgically tagged with acoustic transmitters. Fish monitored in 2008 (n ¼ 12) weretranslocated from the point of capture to analyse the movement behaviour after artificial displacement.Upon release at displaced locations, 67% of the fish moved towards the original capture location usinga time of return that ranged from 0.75 to 15.25 h. Fish monitored in 2009 (n ¼ 8) were released at thepoint of capture. They showed high site fidelity with a maximum period of 27 d between the first timeand the last time they were detected.

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1. Introduction

The spatio-temporal behaviour of fish is important for a numberof marine management and conservation issues (Pittman andMcAlpine, 2003; Botsford et al., 2009). For example, fish move-ments and home ranges are important factors to consider in thedevelopment of marine protected areas (MPAs). The benefits ofa marine reserve depend on the rate and scale of movements ofspecies in relation to the reserve size (Kramer and Chapman, 1999;Sale et al., 2005). To provide effective protection, MPAs must besufficiently large to enclose the appropriate habitats in order tocontain the regular movements of the target species (i.e., theirhome range), but also allow dispersal and cross-boundary move-ments of early life stages to fishing grounds (i.e., adult spillover:Kramer and Chapman,1999; Bartholomew et al., 2008). In addition,site fidelity is also related to the possibility of using fish as bio-indicators, as sedentary species are more suitable for representinglocal exposure to human impacts (Burger and Gochfeld, 2001).Examining how individuals use space can reveal the diversity ofbehaviours within a species and increase our understanding ofbasic ecological processes.

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Acoustic telemetry has been used as an alternative as well asa complementary method to conventional mark and recapturestudies for studying fish movements (Voegeli et al., 2001; Parsonsand Egli, 2005; Abecasis et al., 2009). For example, this tech-nology has allowed researchers to quantify fish home ranges withinMPAs (e.g., Lowe et al., 2003; Parsons and Egli, 2005), determinesite fidelity (e.g., Collins et al., 2007; Abecasis and Erzini, 2008) andobtain data on homing abilities (Kaunda-Arara and Rose, 2004;Jadot et al., 2006). This technique uses an acoustic transmitterattached to an individual and the acoustic signal is received bya hydrophone. In addition, the recent development of miniaturetags has allowed transmitters to be used on small fish, which hasincreased the size range of animals that can be tagged. From amongthe different systems available for acoustic telemetry, automatedsystems that use arrays of acoustic listening stations have becomea popular research tool (Heupel et al., 2006). These passive systemscan be designed to monitor various individuals’ movementssimultaneously in a broad range of spatial and temporal scales inorder to infer spatial (e.g., home range) and temporal (e.g., dielactivity) movement patterns. Previous studies using passiveacoustic telemetry in Palma Bay Marine Reserve with Serranusscriba (March et al., 2010) and Serranus cabrilla (Alós et al., in press)showed that this technology performswell in this environment andis appropriate for small fish.

D. March et al. / Estuarine, Coastal and Shelf Science 92 (2011) 581e587582

The annular seabream, Diplodus annularis (Linnaeus 1758), isa littoral benthic fish, common in the bottoms covered by seagrassbeds from0 to 50mdepth (Bauchot,1987). This small Sparid inhabitsthe Mediterranean and Black Sea coast, the Atlantic from the gulf ofBiscay to Gibraltar, and the Madeira and Canary Islands (Bauchot,1987). It is classified as a rudimentary hermaphrodite (Buxton andGarratt, 1990; Alós et al., 2010), with a spawning period betweenMay and June (Alonso-Fernández et al., in press). Total lengths (TL) at50% maturity are 9.0 cm for males and 10.0 cm for females (Matic-Skoko et al., 2007). In the Balearic Islands (NW Mediterranean),D. annularis is one of the species most frequently targeted by recre-ational boat and shore anglers (Morales-Nin et al., 2005), and hasaminimum legal size of 12 cm. Recent studies have usedD. annularisto assess the impacts of recreational fisheries (Cardona et al., 2007;Alós et al., 2008). Thus, information on the movements ofD. annularis is relevant for the management of recreational fisheries.

In this study, we used both conventional external tagging andpassive acoustic telemetry to examine the short-term movementpatterns of a small temperate fish, the annular seabream Diplodusannularis. The specific aims of the study were: (1) to quantify theshort-term movement patterns of D. annularis; (2) to determineshort-term site fidelity; and (3) to describe movement behaviourafter artificial displacement.

2. Methods

2.1. Study site

The study was carried out in Palma Bay and Palma Bay MarineReserve (PBMR), Mallorca Island (NWMediterranean, Fig. 1). Palma

Fig. 1. Map of the study area showing the locations of the acoustic receiver arrays for the yeZone (IZ) and the Buffer Zone (BZ). The lower inset represents the locations of capture and

Bay is located in the southern part of the island, and is one of theareas with the highest densities of recreational fishers (Morales-Nin et al., 2005). Bottom habitats from 0 to 35 m are dominatedby seagrass meadows of Posidonia oceanica and rocky bottoms.Palma Bay Marine Reserve is located in the eastern part of PalmaBay, and protects an open water area that extends from theshoreline to the 30 m isobath. This MPA is zoned into twomanagement areas with different levels of protection: (1) theIntegral Zone (w2 km2), where all fishing activity is prohibited, and(2) the Buffer Zone (w24 km2), where both artisanal and recrea-tional fisheries are permitted but with some management regula-tions (e.g., daily bag limits, minimum hook size and temporalclosures) (Fig. 1).

2.2. Conventional tagging experiment

Conventional tagging was conducted in different taggingsessions between February 2007 andMay 2007 in Palma Bay (Fig. 1,lower inset). Fish were captured with hook and line using largeJ-hooks in order to reduce hooking mortality (Alós et al., 2008).Only individuals displaying good physical condition (i.e., thosewithout hooking injuries or evidence of barotraumas after 10 minof visual inspection) were selected to be marked. After capture, fishwere tagged with external T-tags (Floy Tag) inserted bellow thedorsal fin and then released at the capture location. Each tag dis-played an ID and a telephone number. Total fish length (to thenearest mm), date and time, capture depth (m) and geographiccoordinates were recorded for each released individual. A specificcommunication programwas designed and implemented to inform

ars 2008 and 2009. Palma Bay Marine Reserve (PBMR) boundaries enclose the Integralrelease used in the conventional tagging experiment.

D. March et al. / Estuarine, Coastal and Shelf Science 92 (2011) 581e587 583

the recreational fishing community regarding the external taggingprogram (Cardona-Pons et al., 2010).

A preliminary study in tanks showed that mortality related tocapture and tagging was negligible. In addition, the 35% of taggedindividuals (n ¼ 20) shed their tags after 19 d, but this percentageremained constant up to 50 d, when the experiment finished (Alóset al., in press).

2.3. Acoustic tagging

In addition, acoustic tagging was conducted in MarcheApril2008 and JulyeAugust 2009 in PBMR. We surgically tagged adultindividuals (n ¼ 20) with acoustic transmitters (Sonotronics, PT-3)following the procedure described in March et al. (2010). Wecombined transmitters that were turned on for 24 h d�1, withtransmitters that yield an active period of 12 h d�1 in order toincrease the battery life to approximately 35 d (Table 1). Thosetransmitters were programmed to ran intermittently their set ofpulses twice, and then added a period of silence for the sameamount of time. Transmitters did not exceed 2.27% of the bodyweight of the fish. A preliminary evaluation of the surgical proce-dure using ‘dummy’ transmitters with 6 individuals showed normalbehaviour 8e10 min after surgery, full cicatrisation without trans-mitter loss and 100% survival after 1 week (Grau, unpubl. data).

2.4. Acoustic monitoring system

We used fixed arrays of acoustic monitoring stations (Sono-tronics, SUR-1) to track the movement of Diplodus annularis withinPBMR. This system uses small multifrequency, omnidirectionalreceivers which measure and record the interval period betweensuccessive pings from individual transmitters. Receivers wereplaced at depths from 10 to 35 m, and the detection range wasw250 m (March unpubl. data). Two different receiver arrays wereused (Fig. 1). The first was deployed between March and April 2008using 22 receivers and covering a monitoring area of approximately4.7 km2. The distance between receivers (w500 m) allowed fish tobe monitored for most of the period that they were within themonitoring area. The main habitat found within this array wasseagrass meadows of Posidonia oceanica. The second array wasdeployed between July and August 2009 using 20 receivers andcovered a wider area of approximately 7.5 km2. The distancebetween receivers (w900 m) meant that their ranges did notoverlap. In this case, the habitat type was more heterogeneous,with both seagrass meadows and sandy bottoms.

2.5. Displacement experiment

We released tagged fish at the capture site, except those taggedwith acoustic transmitters in 2008, which were displaced to a loca-tion w1500 m from the capture point. The release sites hada benthic habitat similar to the capture sites (i.e., seagrassmeadows). We assessed whether a fish moved back to the capturelocation by recording the time that theywere first detected by eitherone of the two receivers that were closest to the capture location.

Table 1The number of Diplodus annularis tagged with standard T-tags and recaptured foreach length interval.

Length interval (cm TL) Tagged Recaptured

8e10 12 010e12 266 012e14 686 1314e16 175 2>16 10 0

2.6. Data analysis

We examined conventional tag-recapture data in order todetermine the number of days and distance between capture andrecapture locations. As fish were not released again, only single-tagrecovery data was available for this study. We used the MicrosoftAccess database and R program described in March et al. (2010) tomanage and analyse the detection data downloaded from thereceivers. The data were filtered to remove potentially spuriousdetections. We defined spurious detections as any single trans-mitter code detection that occurred alone within a 24 h period.Daily presence histories were plotted to inspect the fish presencetimelines visually. We calculated the total period between the firstand last detection (TP), the number of days detected (DD) andthe residence index for each fish (RI; Abecasis and Erzini, 2008). Forthe 2008 array, in which detection ranges overlapped, we used theNadaraya-Watson normal-kernel estimator to calculate the posi-tion estimates, or “centre of activity” locations (COA), as describedin March et al. (2010). Distances between consecutive 15 min COAlocationswere used tomeasure the linearity index (Li; Rechisky andWetherbee, 2003). The linearity index was calculated as follows:

Li ¼ ðFn � F1Þ=D (1)

where Fne F1 is the distance between the first and last COA, andD isthe total distance travelled. Fish with nomadic behaviour shouldhave Li values close to 1, while fish with strong site fidelity shouldhave Li values approaching 0.

3. Results

3.1. Tag-recapture

We tagged 1149 individuals of Diplodus annularis with externalT-tags. The total length (TL) ranged between 8.7 and 16.7 cm(Table 1). At the end of the project (August 2008) the recapture ratewas low (1.31%), as only 15 fish were recaptured. Four (26.7%) of thefish were recaptured by the research team at tagging sessions,whereas 11 fish were recaptured by recreational anglers. In bothcases, recaptures were conducted using hook and line. Most of therecaptures occurred in locations close to the release sites. Themaximum distance between the capture and recapture positionswas w300 m, and the time between recaptures ranged between 7and 185 d (Table 2).

3.2. Acoustic telemetry

The 2008 acoustic receiver array was able to detect all taggedfish (n ¼ 12), whereas two (25%) fish tagged with acoustic trans-mitters in 2009 (n ¼ 8) were not detected. Fish detections aresummarised in Table 3. The movement patterns of Diplodus annu-laris detected using acoustic telemetry were very different fordisplaced and non-displaced individuals. While displaced individ-uals were detected by several receivers for few hours and then getout of the acoustic array, most of the non-displaced individuals

Table 2The number of recaptured Diplodus annularis according to the distance from thetagging position and time between recaptures.

<100 m 100e500 m >500 m

<1 week 21e2 weeks 82 weeks to 1 month 11e3 months 2>3 months 2

Fig. 2. Abacus plots of daily presence detections of Diplodus annularis. Vertical linesindicate the dates on which tagged individuals were detected within the monitoringarea during the 2008 (a) and 2009 (b) experiments. (C) Transmitter deploymentdates; (B) predicted transmitter death dates.

D. March et al. / Estuarine, Coastal and Shelf Science 92 (2011) 581e587584

were detected by only one receiver and remained within itscoverage for a longer period (Table 3). Two fish (#11 and #D3) werenot detected by any receiver, andwe also considered not taking intoaccount fish #D1 as only four detections were recorded by only onereceiver. In addition, fish #D7 and #D9 were continuously detectedby only one receiver during consecutive days (up to 31 and 29 drespectively), which could suggest that they had either lost theirtag or died (e.g., Taylor et al., 2006; Abecasis et al., 2009). Incontrast, fish #D8 was also only detected by one receiver, but forthis fish the detection pattern was intermittent, which suggeststhat it often moved out of the range of the receivers. The detectionpatterns for fish #D5 and #D11 were also intermittent, but thesefish were detected by more than one receiver. However, thesereceivers were located close to each other, which suggests that fish#D5 and #D11 moved within a reduced area.

Short-term residence differed among tagged fish (Fig. 2). In the2008 experiment, most of the displaced fish left the monitoringarea one day or a few days post-release, and only some of themreturned to the edges of the monitoring array after some days(Fig. 2a). Fish were detected for total periods ranging between 1and 9 d, with a mean of 2.8 d (Table 3). However, in the 2009experiment, non-displaced fish remained within the monitoringarea for longer periods. Those fish were detected for total periodsranging between 1 and 27 d (with a mean of 15.5 d) (Table 3).Combining the RI scores from the two experiments resulted in highvalues (0.90 � 0.36, mean � SD), which indicates that most of thefish remained continuously within the monitoring area during themonitoring period. However, some fish (i.e., fish #173, #185 and#D5) exhibited an intermittent daily pattern throughout themonitoring period.

3.3. Displacement experiment

Eight individuals (67%) from the displaced fish showed evidenceof directional movement towards the capture location (Table 4 andFig. 3). The time taken to return to the capture site ranged between0.75 h and 15.25 h (4.66 � 4.64, mean � SD). High Li values(0.75 � 0.19, mean � SD) in most of the fish indicated that move-ment from the release site to the capture location tended to beunidirectional (Table 4). Capture locations were located at themargin of the acoustic array. After fish returned to the vicinities of

Table 3Summary of themonitoring data for the 21 tagged Diplodus annularis individuals. TL, totalof days detected; and RI, residence index. No RI value for fish #D1 is presented as only a

Fish code TL (mm) Weight (g) Release date (yy/mm/dd) First date of detect

36a 141 44 08/03/13 08/03/13185a 141 48 08/03/14 08/03/14168 142 48 08/03/14 08/03/14171a 143 46 08/03/13 08/03/1333 143 46 08/03/14 08/03/14137a 143 48 08/03/14 08/03/14169a 144 51 08/03/14 08/03/14173a 145 53 08/03/13 08/03/1367a 152 56 08/03/14 08/03/1434a 153 58 08/03/14 08/03/1431a 154 64 08/03/14 08/03/14136a 157 58 08/03/13 08/03/13D1 148 59 09/07/17 09/07/28D8 148 58 09/07/17 09/07/17D11 152 64 09/07/17 09/07/22D7 153 65 09/07/17 09/07/17D9 153 65 09/07/17 09/07/17D5 165 82 09/07/17 09/07/1811a 152 63 09/07/17 e

D3 148 58 09/07/17 e

a Fish tagged with transmitters that were programmed to run intermittently to yield

the capture site, most of them were no longer detected by anyreceiver. However, two individuals (#173 and #185) were inter-mittently detected some days after having returned to the capturesite by the receivers located in the same vicinity.

4. Discussion

Despite the effort made to inform the fishing community andthe rewards offered for recaptures (Cardona-Pons et al., 2010), only11 Diplodus annularis were recaptured by recreational anglers. Tagshedding could be one of the main factors explaining such a low

length; TP, total time period between the first and last date of detection; DD, numberfew detections occurred.

ion (yy/mm/dd) TP (d) DD (d) RI Total detections No of receivers

1 1 1.00 13 49 4 0.44 78 81 1 1.00 27 51 1 1.00 16 51 1 1.00 18 31 1 1.00 70 51 1 1.00 23 49 3 0.33 138 62 2 1.00 103 51 1 1.00 10 41 1 1.00 34 66 4 0.67 41 32 2 e 4 17 5 0.71 25 1

27 17 0.63 82 231 31 1.00 651 129 29 1.00 437 126 13 0.50 128 3e e e 0 0e e e 0 0

an active period of 12 h d�1.

Table 4Summary of tagging and return (homing) conditions for individuals of Diplodusannularis released w1500 m from the capture location. Distance travelled to returnand the linearity index are based on COA locations.

Fish code Time takento return (h)

Distance travelledto return (m)

Linearity index

36a 1.25 1328 0.818185a 15.25 1929 0.568168 2.5 1505 0.757171a 6.5 1555 0.90833 e e e

137a 4.25 2074 0.536169a e e e

173a 3.75 706 0.95267a e e e

34a 0.75 1001 0.97231a 3 2162 0.512136a e e e

a Fish tagged with transmitters that were programmed to run intermittently toyield an active period of 12 h d�1.

D. March et al. / Estuarine, Coastal and Shelf Science 92 (2011) 581e587 585

recapture rate. Other factors, previously mentioned in Abecasiset al. (2009), that could lead to low recapture rates are that: (1)anglers avoid declaring tagged fish below the minimum legallanding size of D. annularis (12 cm); and (2) tagging could increase

Fig. 3. Paths representing the movement patterns of the 12 displaced individuals of Diplo

vulnerability to predators (colonisation of external tags by algaewas frequent in our study). All these factors could be responsible forreducing the probabilities of recapturing tagged fish. However,inferences of movement rates that are not biased by tag loss effects(i.e. tag shedding, non-reporting rate and tag-induced mortality)can be determined by recapture-conditioned estimators based ononly-recapture data (McGarvey, 2009). The lower rate of tag-recapture (1.31%) was close to other similar studies. Based on anextended dataset (i.e. wider area, longer period and more taggedfish), the recapture rate of D. annularis in southern Mallorca was0.7% (Cardona-Pons et al., 2010). In addition, the same study alsoreported similar recapture rates for Coris julis (4.6%) and Serranusscriba (0.8%), which are two of the most targeted species by therecreational fishery. In a previous study conducted with otherspecies of the same genus, Abecasis et al. (2009) reported recapturerates of 3.13% for Diplodus vulgaris, and 4.08% for Diplodus sargus.

The information obtained from the external tags suggests thatDiplodus annularis does perform small displacements. The distancebetween mark and recapture may be biased when fishing effortand/or release locations are clustered. For example, releasing allfish near a location with high concentration of fishing effort couldcause underestimation of distances between release and recapture.In our study, tagging locations could be considered as randomly

dus annularis. Paths are based on COA locations and do not represent true locations.

D. March et al. / Estuarine, Coastal and Shelf Science 92 (2011) 581e587586

distributed within the suitable habitat for D. annularis (Fig. 1, lowerinset). In addition, although recreational fishing effort cannot beconsidered strictly homogeneous (March et al. unpubl. data), itscurrent spatial variation in Palma Bay is not sufficiently large toinduce a severe bias in the estimated distances (Palmer et al., inpress). Unlike acoustic data, which are limited due to the batterylife of transmitters, fish recaptures can provide information fora time span over 1 month. In our case, the four fish that wererecaptured after 1 month and within maximum distance ofw300 m provides valuable information to assess the high sitefidelity of this species.

Acoustic telemetry data complemented conventional tagging byproviding new information on the short-term movements of Dip-lodus annularis. However, as in other studies, the low number oftagged individuals by acoustic tags represents a limitation(Abecasis and Erzini, 2008; Abecasis et al., 2009; March et al., 2010).In addition, a commonproblemwhenworkingwith small fish is thedifficulty in capturing individuals large enough to be fitted withacoustic transmitters. In our case, this factor was especially relevantin the displacement experiment: after receiver deployment therewas a lack of availability of fish with an adequate size in the studyarea. This fact compromised the experimental design and we wereunable to capture fish in locations with a better coverage ofreceivers (i.e., not so close to the edge). It compromised the capa-bility of the monitoring system to detect whether fish presenteda homing behaviour, as most of the fish were no longer detectedonce they had pass through to the capture site. Even if fish wouldhave established their home range close to the capture location(e.g., Kaunda-Arara and Rose, 2004), such pattern could not beeasily detected by the receivers, as the capture locationwas close tothe edge of the acoustic array. Nevertheless, despite of these limi-tations, we consider some direct and indirect evidences that couldbe consistent with a homing behaviour: 1) directional navigationmovement towards the capture locationwas not random (8 from 12displaced fish); 2) one individual (#173) was detected intermit-tently by the receivers near to the capture location; 3) results ofconventional tagging and non-displaced fish in 2009 (i.e., theycould be considered as a control group) suggest a high site fidelitybehaviour that has been related previously with homing abilities(e.g., Kaunda-Arara and Rose, 2004), and 4) homing abilities havebeen documented in other species of the Sparidae family, such asSarpa salpa (Jadot et al., 2006) and Sparus aurata (Abecasis andErzini, 2008). Spatial learning through different homing mecha-nisms (i.e., cognitive mapping, social transmission and landmarkorientation) may allow fish to navigate back to resting, foraging orspawning sites (Kaunda-Arara and Rose, 2004).

The information derived from the acoustic array deployed in2009 contrasts with the data obtained from the 2008 array. Theperformance of the acoustic array in this study is relevant. Theseparation between receivers produced areas of non-continuousdetection. This fact compromised the capability of the system todiscriminate between fish with a reduced home range (i.e., lessthan the distance between receivers) and fish that could have beendead or lost their tags. In a previous work (March et al., 2010), wedetermined that the detection of additional patterns (e.g. dielperiodicities in the number of detections) could suggest that fishcould be alive when detected by only one receiver. However, thatwas not the case for fish #D7 and #D9 and we did not considerthem for analysis. Moreover, due to fish availability, the releaselocations were in shallow waters (<10 m) where the detectionrange may be greatly reduced due to signal absorption and back-ground noise. Fish that were not displaced were rarely detected bydifferent receivers. Although the spacing between receivers wasgreater than the array deployed in 2008, these results suggesta high site fidelity. This pattern is consistent with the results

reported by conventional tagging. In addition, similar results werereported for individuals of Diplodus vulgaris, where fish weredetected by only one receiver during long periods of time (Abecasiset al., 2009).

The results of this study provide valuable information for themanagement of recreational fisheries and MPAs. Diplodus annularishas been used in recent studies to evaluate the direct and indirecteffects of the recreational fishery in the Balearic Islands (Cardonaet al., 2007; Alós et al., 2008, 2009; Cerdà et al., 2010). Theimportance of site fidelity relies on the exposure of this species tothe local environment over time. Some evidence can be extrapo-lated from a previous study in which D. annularis showed differ-ences in diet between artificial reef and control habitats inPosidonia oceanica meadows, in which blocks were deployed in anarea of similar size to our acoustic arrays (Sánchez-Jerez et al.,2002). Thus, if D. annularis shows high site fidelity, it could bea good bioindicator for assessing local exposure to environmentalfactors (e.g., fishing overexploitation). Our results provide differentinformation sources that could be used to complement each otherin MPA management. Within MPAs, site fidelity may enhance thelikelihood of sustaining locally reproducing adults that couldprovide sources of dispersing larval recruits to adjacent fishinggrounds (Kaunda-Arara and Rose, 2004). The evidence of high sitefidelity in the present work indicates that MPAs could play a vitalrole in conserving these fish and managing the recreational fishery.

Acknowledgments

We thank the many people that collaborated in this project,particularly AM. Grau, B. Morales-Nin and all the voluntary anglers.The habitat map was obtained from the LIFE Posidonia program,Government of the Balearic Islands. Aerial photography wasprovided by the �Instituto Geográfico Nacional de España. Thisstudy was financed by the projects ROQUER (CTM2005-00283) andCONFLICT (CGL2008-958) funded by the Spanish Ministry ofResearch and Science, and by the research project ACOUSTICTRACKING (UGIZC) funded by the Government of the BalearicIslands. D.M. was awarded a fellowship by the Spanish Ministry ofResearch and Science (BES-2006-13252). This studywas carried outwith permission from the local fisheries administration, Govern-ment of the Balearic Islands, for fish tagging and receiver deploy-ment in Palma Bay Marine Reserve.

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