Movement Patterns of Four Coral Reef Fish Species in a Fragmented Habitat in New Caledonia: Implications for the Design of Marin

ICES Journal of Marine Science Advance Access published October 9, 2008

Page 1 of 6 Movement patterns of four coral reef ﬁsh species in a fragmented habitat in New Caledonia: implications for the design of marine protected area networks

Olivier Chateau and Laurent Wantiez

Chateau, O., and Wantiez, L. 2009. Movement patterns of four coral reef fish species in a fragmented habitat in New Caledonia: implications for the design of marine protected area networks. – ICES Journal of Marine Science, 66: 000–000. Acoustic telemetry was used to examine the patterns of fish movements between a marine reserve and two unprotected reefs sep- arated by large areas of lagoon soft bottoms (900–2000 m) in the South Lagoon Marine Park of New Caledonia. Four commercial fish species (Epinephelus maculatus, Plectropomus leopardus, Chlorurus microrhinos, and Scarus ghobban) were studied for 17 months (45 fish). Nine fish (20%) were detected on reefs other than the reef onto which they were released. Four patterns of inter-reef movement were identified during the survey, including home range relocations, movements larger than the scale of the study, many inter- reef movements within the study area, and punctual excursions outside the daily home range. The information gathered in this study demonstrates the ability of the fish to carry out medium-scale movements in a fragmented habitat. Consequently, the effectiveness of the reserve to protect the entire population is probably limited for these species. Because all the identified patterns implied at least one movement across the reserve boundaries, our results support the hypothesis that the Lare´gne`re Marine Reserve, part of the South Lagoon Marine Park, could benefit the adjacent fished area through spillover. Keywords: acoustic telemetry, fish behaviour, fragmentation, home range, inter-reef movements, marine reserve, South Pacific. Received 25 October 2007; accepted 21 May 2008. O. Chateau and L. Wantiez: LIVE University of New Caledonia, Aquarium des Lagons, BP R4, 98851 Noumea cedex, New Caledonia. Correspondence to L. Wantiez: tel: þ687 26 68 92; fax: þ687 27 32 72; e-mail: [email protected].

Introduction 1991; Chapman and Kramer, 2000; Zeller et al., 2003; Kulbicki, No-take marine reserves have been used as tools for managing 2007). However, the species studied, the sampling methods, and fishery-targeted species and buffering against broader ecosystem the spatial and temporal scales of the studies may bias the percep- effects of overfishing (Roberts and Polunin, 1993; Russ, 2002; tion of coral reef fish movements. With the development of acous- Halpern, 2003). To provide effective protection, marine reserves tic telemetry, accurate studies of the movement patterns of large must be large enough and enclose appropriate habitats to fish are now possible (Zeller, 1999; Heupel et al., 2006; Chateau contain regular movements of targeted species (Meyer and and Wantiez, 2007), demonstrating frequent fish movements Holland, 2005; Meyer et al., 2007a, b). However, empirical data over hundreds or even thousands of metres (Holland et al., 1996; on target species movements are generally lacking and, con- Zeller, 1997; Wetherbee et al., 2004; Meyer et al., 2007a, b). sequently, most marine reserves do not have appropriate designs However, most of these studies focused on transient species (Roberts and Polunin, 1993; Kramer and Chapman, 1999; Meyer (e.g. Caranx melampygus, Holland et al., 1996; Caranx ignobilis, et al., 2007a, b). Most existing marine reserves are relatively Wetherbee et al., 2004; C. ignobilis, Meyer et al., 2007a; Aprion small (74% are ,10 km2; Halpern, 2003) and probably provide virescens, Meyer et al., 2007b), on short-term patterns of fish move- scant protection for mobile target species (DeMartini, 1993; ment (e.g. up to 118 h of tracking over 9–22 d; Meyer and Holland, Meyer et al., 2007a). Another objective of fishery reserves is to 2005), and on relatively small areas within continuous habitats sustain adjacent fisheries through the net export of post-settlement (e.g. 500 m; Meyer and Holland, 2005). Quantitative information fish (spillover; Russ, 2002). Evidence of density-dependent home on fish movement patterns between reefs over greater spatial and range relocation from a small, no-take reserve has been reported temporal scales is scarce (see Meyer et al., 2007a, b). Several in the Philippines (Abesamis and Russ, 2005). Movements of studies suggest that, within fragmented coral reef habitats, the coral reef fish across reserve boundaries can affect the abundance potential for fish to move between reefs or to relocate to areas and distribution of fish within and outside the reserve, and there- beyond their home range is influenced by differences in habitat fore affect the ability of reserves to preserve fish populations and quality and the presence of physical barriers (Kramer and enhance surrounding fisheries (Kramer and Chapman, 1999; Chapman, 1999; Chapman and Kramer, 2000). Movements may Chapman and Kramer, 2000; Russ, 2002). be facilitated across continuous areas of favourable habitats and Post-settlement coral reef fish are generally considered seden- restricted by a high level of fragmentation, such as large areas of tary and highly site-attached (Sale, 1991; Roberts and Polunin, sandy substratum with low structural complexity (Barrett, 1995).

there an interaction between the marine reserve and adjacent unprotected reefs? (ii) if so, what was the nature of the interaction? (iii) was it possible to relate the detected movements across marine reserve boundaries to ecological processes or behaviour traits of fish? Material and methods The study area includes a marine reserve (Lare´gne`re islet), part of a reserve network within the South Lagoon Marine Park, and two unprotected reefs (Lare´gne`re reef and Crouy reef; Figure 1). Lare´gne`re Marine Reserve (8.5 km2) is a coralline island located in the middle of the lagoon, 10.5 km from the coast and 7.4 km from the barrier reef (Figure 1). Fishing and collecting have been prohibited within the reserve, with active enforcement of park regulations, since 1990. Lare´gne`re reef and Crouy reef are two unprotected reefs located at 900 and 2000 m, respectively, from the reserve (Figure 1). From July 2005 to January 2007, 23 VR2 omnidirectional hydrophones (Vemco) were deployed 4–13 m deep around the three reefs (Figure 1). A minimum of 200 m radius functional range was measured in situ during test operations (OC, pers. comm.). All the hydrophones were located 100 m away from the reef slope, on soft bottoms to minimize acoustic shadows that might occur in structurally complex areas. The detection range of each receiver included the outer part of the reef flat, the entire reef slope, and a strip of 300 m of soft bottoms (OC, pers. comm.). The network of hydrophones covered 276 ha:143 ha in the reserve, 85 ha at Crouy reef, and 48 ha at Lare´gne`re reef. The automated VR2 tracking system provides continuous data on fish presence through an array of hydrophones. The fish were tagged with a Vemco ultrasonic V9-2L-R04K coded transmitter (frequency: 69 kHz; power output: 142 dB; delay between two pulses: 40–120 s; length: 29 mm; diameter: 9 mm; weight in water: 2.9 g). The approximate life of a transmitter is 374 d. In all, 45 fish (23–65 cm FL) were monitored: 7 E. maculatus (Serranidae), 12 P. leopardus (Serranidae), 13 C. microrhinos (Scaridae), and 13 S. ghobban (Scaridae). Serranidae were caught with barbless hook and lines and Scaridae with a landing net. Figure 1. Map of the study area, with the location and the detection After their capture, fish were kept in 2000-l tanks (mean duration: area of the receivers. 5.6 + 4.8 d). Then, the fish were anaesthetized with clove oil at 0.2 ml l21 (Wagner and Cooke, 2005). Acoustic transmitters were surgically implanted in the abdominal cavity (Zeller, 1999; The South Lagoon Marine Park of New Caledonia was created Parsons et al., 2003). After application of a healing agent in 1990. It includes a network of one temporary and nine perma- (Lotagen), the incision was sutured with two stitches of synthetic nent no-take marine reserves (Figure 1). Coral reefs within the absorbable surgical suture (Wagner and Cooke, 2005). For all fish, park have been surveyed regularly since its creation. The effects the transmitter weight in the water never exceeded 1% of body of the marine reserves were analysed using before–after dataseries weight. The fish were retained for a post-operation monitoring (Wantiez et al., 1997) and protected–unprotected comparisons period (mean duration: 4.3 + 2.7 d) before their release in the (Sarrame´gna, 2000; Chateau and Wantiez, 2005). These studies study area. No mortality occurred during this process. Divers indicate an increase in the density, biomass, and species richness released the fish near reef shelters to limit their stress and possible after protection, and the consistency of these changes over time attacks by large predators. Of the fish, 35 (78%) were released at (Wantiez et al., 1997; Chateau and Wantiez, 2005). The inter- their capture site (28 fish in the reserve and 7 fish in an unpro- actions between the reserves and the adjacent unprotected reefs tected reef). Ten fish (22%) were caught in an unprotected reef have to be studied to optimize the design of the marine reserve and released in the reserve. network. Data collected during the first 24 h were considered to repre- Within this context, movements of four major commercial reef sent an acclimatizing period and were not included in the sub- species, two Serranidae [Epinephelus maculatus (Bloch, 1790) and sequent analyses (Zeller, 1997). Patterns of inter-reef movements Plectropomus leopardus (Lace´pe`de, 1802)], and two Scaridae were identified according to the number of detected reef [Chlorurus microrhinos (Bleeker, 1854) and Scarus ghobban changes, the number (or duration) of detection in the entire (Forsska˚l, 1775)], were analysed between a marine reserve and study area, the area of origin, and/or the destination area. The two unprotected reefs. We focused on three questions: (i) was absence of detection was not used to identify the patterns, Movement patterns of four coral reef fish species in a fragmented habitat Page 3 of 6

Table 1. Summary of the patterns of inter-reef movements of the telemetry-monitored fish. All fish studied E. maculatus P. leopardus C. microrhinos S. ghobban Parameter (45) (7) (12) (13) (13) Number of fish with inter-reef movements 9 (20%) 3 (43%) 1 (8%) 2 (15%) 3 (23%) ...... Number of fish without inter-reef movements 36 (80%) 4 (57%) 11 (92%) 11 (85%) 10 (77%) ...... Home range relocations 2 1 – – 1 ...... Movements larger than the scale of the study 3 – 1 1 1 ...... Many inter-reef movements 2 2 – – – ...... Excursions 4 1 – 1 2

because an undetected fish could be in or outside the reserve. The the study area just after release and passed Lare´gne`re reef (Figure 3a), distances were calculated using ArcGIS and high-resolution cali- and one S. ghobban, which remained in the marine reserve for 9 d brated satellite photos (Quickbird). The calculated distances repre- before leaving the area (Figure 3a). The latter fish was spearfished sent the minimal linear distance between the detection areas of the 12 d later near an unprotected reef (Prony reef), located 6 km hydrophones. from the reserve (Figure 3a). A departure was also observed for one P. leopardus caught and released in the reserve (Figure 3b). This fish was detected for 218 d in the reserve before leaving the Results study area and passing Lare´gne`re reef (one detection). It covered Nine fish (20% of the specimens studied), at least one of 3500 m in 66 h (Figure 3b). each species, carried out inter-reef movements. Four patterns of inter-reef movements were identified (Table 1). All the patterns involved at least one movement across the reserve boundaries. The first pattern corresponded to home range relocation. It was characterized by a low detection at origin (1052 detections) and a high detection at destination (8734 detections) for one E. maculatus (Figure 2). This fish was caught at Crouy reef and released in the marine reserve. It was detected in the reserve for 2 d before moving to Lare´gne`re reef, where it was regularly detected for the next 64 d (Figure 2). This change of reef corresponded to a minimum linear movement of 1000 m. This distance was covered in 1 h 55 min. The pattern was also characterized by a high detection both at origin (5774 detections) and destination areas (6255 detections) for one S. ghobban caught and released in the reserve (Figure 2). This fish was detected in the reserve for 86 d, then relocated its home range to Lare´gne`re reef, where it was detected for the next 150 d (Figure 2). This change of reef corresponded to a minimum linear movement of 510 m. The distance was covered in 56 min. The second pattern corresponded to movements larger than the scale of the study. It was observed for one C. microrhinos,whichleft

Figure 3. Pattern of large-scale movements (S. ghobban, white stars; Figure 2. Patterns of home range relocation (E. maculatus, black C. microrhinos, black stars; P. leopardus, grey stars). Stars indicate sites stars; S. ghobban, white stars). Stars indicate sites of ﬁsh detection; of ﬁsh detection; numbers within stars indicate the chronology of numbers within stars indicate the chronology of the detections. the detections. Page 4 of 6 O. Chateau and L. Wantiez

Figure 5. Pattern of excursion outside an established home range (E. maculatus). Stars indicate sites of ﬁsh detection; numbers within stars indicate the chronology of the detections.

physical barriers to their movements between coral reefs (Kramer and Chapman, 1999; Chapman and Kramer, 2000). A home range relocation process (Robertson, 1988) was clearly demonstrated for two fish (E. maculatus and S. ghobban), which set a stable home range in the Lare´gne`re reef. The distinction between Figure 4. Pattern of multiple inter-reef movements (E. maculatus). relocation and migration depended on time, because fish may have Stars indicate sites of fish detection; numbers within stars indicate returned to their original home range. Many species migrate to the chronology of the detections. reproduce or after environmental change (McFarland, 2001). In this study, relocation was during the spawning season of these two species in New Caledonia (Kulbicki, 2007). However, The third pattern corresponded to many inter-reef movements these fish were detected at destination (Lare´gne`re reef) for 2 (4–5 changes in 9–23 d) and was characterized by low detections, (E. maculatus) and 3 months (S. ghobban), with no return to both at origin and at destination (,110 detections; Figure 4). This origin (reserve) after the spawning season. These observations pattern corresponded either to fish with a large home range, which support the hypothesis that this pattern corresponded to the estab- included at least two reefs, or to fish unable to set a stable home lishment of a new home range on an unprotected reef. Home range range within the study area. It was observed for two E. maculatus relocation is an adaptive behaviour when the benefits of moving caught and released at Crouy reef. The minimum linear distances are greater than the benefits of staying (Kramer and Chapman, covered by these fish were 9640 and 16 900 m. The fish may carry 1999). The costs and benefits associated with different locations out several inter-reef movements the same day. For example, one are related to the physical and chemical environment, food fish made three of its four changes the same day, covering supply, density of predators and parasites, availability of shelter 7400 m in ,10 h. and breeding sites, and abundance of competitors and cooperators The fourth pattern corresponded to 1–5 excursions of the fish (Kramer and Chapman, 1999). Changes in competitive and from their home range to another reef with a return to origin cooperative environment or in levels of predation can reduce the (Figure 5). This pattern was characterized by a high detection at benefits of staying and produce opportunities for moving to origin (93–99% of detections) and a low detection at destination, other locations. Kramer and Chapman (1999) indicated that indicating that the excursions were limited in time. It was observed many of these effects would be density-dependent. This suggests for one E. maculatus, two S. ghobban, and one C. microrhinos. Most that the high density of fish in the reserve (Chateau and of the excursions (88%) were carried out from the reserve to the Wantiez, 2005) may favour home range relocation towards the Lare´gne`re reef, where the fish were detected for a few minutes to unprotected area. This process was probably enhanced within 3 h. The E. maculatus was the only fish to carry out an excursion our system by a significant loss of habitat caused by Cyclone (4 d) from Lare´gne`re reef to the reserve (Figure 5). Erica in the reserve (Wantiez et al., 2006). Three fish left the reserve, moving outside the study area Discussion (P. leopardus, S. ghobban,andC. microrhinos). Because the departure This study gives evidence of the ability of the species studied to of P. leopardus occurred outside the spawning season of this species carry out medium-scale movements (510–6000 m) over bare (Kulbicki, 2007), this movement could be a home range relocation soft bottoms between coral reef habitats. The adult fish popu- outside the study area. The two other fish (S. ghobban and C. micro- lations of the three reefs studied are connected, despite their rhinos) left the reserve shortly after their release and were not separation by large areas of lagoon soft bottom. It contrasts with detected in the study area for the rest of the period monitored. the commonly documented hypotheses that reef-associated The high density of fish (Chateau and Wantiez, 2005) and the sig- species demonstrate strong site-fidelity at medium spatial scale nificant loss of habitat caused by Cyclone Erica (Wantiez et al., (Sale, 1991; Zeller, 1997; Chapman and Kramer, 2000; Willis 2006) may create conditions unfavourable for new adults inside et al., 2001) and that large areas of sandy substratum constitute the reserve (higher rate of competition or predation) and favour Movement patterns of four coral reef fish species in a fragmented habitat Page 5 of 6 their departure to the unprotected area (Kramer et al., 1997; Acknowledgements Kramer and Chapman, 1999). Financial support for this study was provided by the ZoNeĆo pro- Multiple inter-reef movements were observed for two fish gramme. We thank Richard Farman (Aquarium Director) for his (E. maculatus). They may have large home ranges, which could assistance, and the south province of New Caledonia, the include areas of lagoon soft bottoms and at least two reefs in the ADECAL, and the SMAI for their administrative and technical study area. This hypothesis is consistent with Lieske and Myers support. (2001), who explain that E. maculatus tends to live around isolated coral heads located in sandy areas, seagrass beds, or at the base of References reefs. However, the size of the area frequented, the number of Abesamis, R. A., and Russ, G. R. 2005. Density-dependent spillover detected reef changes (up to three changes in 1 d), and the scale from a marine reserve: long-term evidence. Ecological of the movements (up to 7 km in 10 h) were rather unexpected. Applications, 15: 1798–1812. These observations contrast with the documented assumption Barrett, N. S. 1995. Short- and long-term movement patterns of six that this species is sedentary at this medium spatial scale temperate reef fishes (families Labridae and Monacanthidae). (Kulbicki, 2007). The movements could also correspond to a suc- Marine and Freshwater Research, 46: 853–860. cession of home range relocations; however, this seems unlikely Chapman, M. R., and Kramer, D. L. 2000. Movements of fishes within because the inter-reef movements were in the short term (up to and among fringing reefs in Barbados. Environmental Biology of three changes in 1 d). This pattern could also correspond to fish Fishes, 57: 11–24. unable to set a stable home range within the study area because Chateau, O., and Wantiez, L. 2005. Comparaison de la structure des communaute´s de poissons coralliens entre une re´serve marine et of unfavourable habitats, insufficient resources, or negative inter- deux zones proches non prote´geés dans le Parc du Lagon Sud de actions with conspecifics or other species. Nouvelle-Cale´donie. Cybium, 29: 159–174. Short excursions outside an established home range were Chateau, O., and Wantiez, L. 2007. Site fidelity and activity patterns of observed for four fish (E. maculatus, C. microrhinos, and two a humphead wrasse, Cheilinus undulatus (Labridae), as determined S. ghobban). Fish can make exploratory movements outside their by acoustic telemetry. Environmental Biology of Fishes, 80: areas of normal activity (Kramer and Chapman, 1999). This 503–508. process was clearly demonstrated for one E. maculatus, which DeMartini, E. E. 1993. Modeling the potential of fishery reserves for carried out a 4-d excursion in the reserve before returning to managing Pacific coral reef fisheries. Fishery Bulletin US, 91: 1–7. Lare´gne`re reef. Because this movement was in February, it was Halpern, B. 2003. The impact of marine reserves: do reserves work and does reserve size matter? Ecological Applications, 13: S117–S137. not correlated with reproduction in New Caledonia (Kulbicki, Heupel, M. R., Semmens, J. M., and Hobday, A. J. 2006. Automated 2007). This pattern could also correspond to a combination of acoustic tracking of aquatic animals: scales, design and develop- home range characteristics (size and shape) and the location of ment of listening station arrays. Marine and Freshwater Research, the hydrophones in the area. In this case, the fish used large 57: 1–13. home ranges including at least all the hydrophones where they Holland, K. N., Lowe, C. G., and Wetherbee, B. M. 1996. Movements were detected. For example, the length of the home range of the and dispersal patterns of blue trevally (Caranx melampygus)ina E. maculatus was at least 4 km. Although this is a great distance, fisheries conservation zone. Fisheries Research, 25: 279–292. we cannot exclude the possibility of a large home range for this Kramer, D. L., and Chapman, M. R. 1999. 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PhD thesis, Ecole Pratique des Hautes by the temporal succession of two patterns for one S. ghobban, Etudes, Perpignan, France. 195 pp. which carried out an excursion from the reserve before relocating Lieske, E., and Myers, R. F. 2001. Coral Reef Fishes. Indo-Pacific and its home range to Lare´gne`re reef. Caribbean. Harper Collins, London. 400 pp. Because all the identified patterns implied at least one move- McFarland, D. 2001. Le comportement animal, psychobiologie, e´tho- ment across the reserve boundaries, our results have important logie et evolution. De Boeck Universite´, Paris. 613 pp. implications for the design of marine reserve networks. If the Meyer, C. G., and Holland, K. N. 2005. Movement patterns, home range size and habitat utilization of the bluespine unicornfish, objective of the network is to protect resident populations, our Naso unicornis (Acanthuridae) in a Hawaiian marine reserve. results suggest that larger reserves, including several reefs, spawn- Environmental Biology of Fishes, 73: 201–210. ing areas, and lagoon soft bottoms, would be necessary. Another Meyer, C. G., Holland, K. N., and Papastamatiou, Y. P. 2007a. Seasonal objective of marine reserves is the maintenance of adjacent and diel movements of giant trevally Caranx ignobilis at remote fisheries through the net export of fish biomass from the reserve Hawaiian atolls: implications for the design of marine protected (Russ, 2002). Our results demonstrate that 20% of the fish areas. Marine Ecology Progress Series, 333: 13–25. studied were more or less exposed to fishing according to Meyer, C. G., Papastamatiou, Y. P., and Holland, K. N. 2007b. the pattern involved. Even if net export was not clearly demon- Seasonal, diel, and tidal movements of green jobfish (Aprion virescens, Lutjanidae) at remote Hawaiian atolls: implications for strated, our results support the hypothesis that the Lare´gne`re marine protected area design. Marine Biology, 151: 2133–2143. Marine Reserve could benefit the adjacent fished area. Parsons, D. M., Babcock, R. C., Hankin, R. K. S., Willis, T. J., Aitken, J. P., Irrevocable evidence was the fishing of one S. ghobban (originally O’Dor, R. K., and Jackson, G. D. 2003. Snapper Pagrus auratus resident in the reserve) near an unprotected reef located 6 km (Sparidae) home range dynamics: acoustic tagging studies in a from the study area. marine reserve. Marine Ecology Progress Series, 262: 253–265. Page 6 of 6 O. Chateau and L. Wantiez

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