Responses to marine reserves: Decreased dispersion of the sparid Pagrus auratus (snapper)
Introduction
Understanding the movement dynamics of exploited species is important for the design and justification of marine reserves. Accordingly, many animal movement studies have focused on marine reserves (Attwood and Bennett, 1994, Meyer et al., 2000, Eristhee and Oxenford, 2001, Kelly and MacDiarmid, 2003, Lowe et al., 2003, Parsons et al., 2003, Starr et al., 2004, Edgar et al., 2004, Egli and Babcock, 2004), while other studies have attempted to justify reserves as a conservation tool by investigating the potential for movement from marine reserves into adjacent areas (spill-over) (Russ and Alcala, 1996, Kramer and Chapman, 1999, McClanahan and Mangi, 2000, Roberts et al., 2001, Zeller et al., 2003, Russ et al., 2004, Abesamis and Russ, 2005). Few studies, however, have concurrently tagged fish inside and adjacent to marine reserves, and as a result we have a poor understanding of how factors unique to the marine reserve environment may influence the movement dynamics of individual fish. Some factors that are known to differ inside reserves which may be potentially important include highly elevated con-specific densities (Willis et al., 2003), habitat differences resulting from trophic cascades (Shears and Babcock, 2002), reduced human disturbance (Eggleston and Parsons, 2008) and the removal (or reduction) of fisheries-induced selection (Biro and Post, 2008). If the behavior of individuals or the behavioral composition of populations is influenced by the above factors then this may be an important consideration for conservation managers designing reserves.
The few studies that have concurrently tagged animals inside and adjacent to marine reserves produced inconsistent results. Attwood and Cowley (2005) found a higher frequency of long distance movements of dart tagged galjoen (Dichistius capensis) from a fished site, in contrast to galjoen tagged inside a marine reserve. Conversely, Cole et al. (2000) obtained fewer re-sightings of blue cod (Parapercis colias) tagged inside a marine reserve, consistent with larger scale dispersal of blue cod from the marine reserve. Through the tagging of lobsters (Jasus edwardsii) inside and outside of a reserve Linnane et al. (2005) observed greater movements of lobsters tagged inside a marine reserve. Finally, Zeller and Russ (1998) observed greater movements of freeze branded coral trout (Plectropomus leopardus) inside a marine reserve, but found no difference in the mean distance moved per day (inside vs. outside), when the same comparison was made with acoustic telemetry.
If managers are to make informed decisions regarding the design of marine reserves then the expected response to protection, which is dependent on animal behavior, needs to be known. The studies listed above and the likely environmental changes expected within reserves both suggest that behavioral responses to marine reserves are not well understood and that either increased or decreased mobility may be expected. If an increase in mobility occurs, this has the potential to reduce the recovery of exploited species but increase supplementation to adjacent fisheries. In this instance managers may therefore wish to scale reserves to ensure that some central portion of the reserve provides complete protection in spite of the higher mobility of the species concerned. Alternatively, a decrease in mobility may suggest that exploited species recovery can be expected in even the smallest reserves. In this situation managers may see benefit in implementing more but smaller reserves. Behavioral responses to reserve implementation may also be opposite for different species or even interact with the habitat in which a reserve is emplaced. Managers may therefore need to prioritize the species for which the reserve is designed or adjust reserve design according to the habitat where the reserve will be located.
In the current study, we examined the movement dynamics of snapper Pagrus auratus inside, and adjacent to, the Cape Rodney to Okakari Point (CROP) Marine Reserve, near Leigh in north-eastern New Zealand. Populations of legal sized snapper (>27 cm Fork Length (FL) for recreational and >25 cm FL for commercial fishers) are estimated to be 14 times greater within the reserve than in adjacent fished areas (Willis et al., 2003). The snapper fishery is open to both commercial and recreational fishers throughout the year, although recreational fishing effort has a seasonal pattern, peaking over the austral summer (Hartill, Fishery Scientist, NIWA, unpublished data). Recreational fishers concentrate most of their effort in coastal areas and as such the majority of the snapper catch from the areas around the reserve boundaries is recreational. While accurate estimates of the catch from around the reserve are not available it is likely in the order of multiple tons annually (Hartill pers. comm.). The effect of this exploitation also impacts on lower trophic levels. Algal abundance on exploited reefs is thought to be suppressed by high abundances of herbivorous urchins, Evechinus chloroticus that are free of dominant reef carnivores such as snapper and spiny rocky lobster (Jasus edwardsii) (Shears and Babcock, 2002).
Multiple tagging studies have been conducted on snapper within the CROP reserve. These studies have confirmed that snapper within the reserve are capable of restricting their movements to small areas of rocky reef (at a scale of hundreds of meters), explaining the higher abundance compared to the adjacent fished waters (Willis et al., 2001, Parsons et al., 2003, Egli and Babcock, 2004). Conversely, movement studies outside the marine reserve have been restricted to broad spatial scale mark recapture experiments (but see Hartill et al., 2003), focused on snapper from deeper, soft sediment dominated environments (Paul, 1967, Crossland, 1976, Gilbert and McKenzie, 1999). These later studies provided only modest movement information, with poor associated spatial resolution, but did show that snapper can move over areas of many tens of kilometers. Therefore, we made direct comparisons of snapper movement inside vs. outside a marine reserve, by deploying an acoustic tracking array over both a portion of the CROP reserve, and the adjacent coastline, and tagged and released snapper in each. We then assessed the potential for differential movement dynamics between the two areas. Results suggest that reserves do have the potential to alter the behavior or the behavioral make up of individuals or populations, with likely consequences for the design of reserves and the maintenance of biodiversity within exploited populations.
Section snippets
Study area and receiver set up
This study was conducted inside and adjacent to the CROP Marine Reserve (Fig. 1) from November 2007 to April 2008. We specifically chose the eastern end of the reserve and the immediately adjacent fished coastline to compare snapper movements, as both of these sections of coast are formed from the same rock type, and have similar topographic relief, bathymetry and exposure to the prevailing wind and swell (predominantly on a SW by NE axis). An array of 30 omni-directional hydrophones (Vemco VR2
Results
Thirty-nine snapper were tagged inside and adjacent to the CROP Marine Reserve in November 2007. The movements of these fish were monitored by an array of acoustic receivers for 5 months, although only the first 2 months of tracking were completed with a full array of receivers. Individual movement behavior of tagged snapper varied, but fell into three distinct categories. The first category was fish detected frequently throughout the tracking period (present for >65% of available half hour time
Discussion
This study is one of the first to explicitly compare the movement behavior of an animal inside and adjacent to a marine reserve concurrently. Our results show that some tagged snapper from non-reserve areas utilized space in a different manner from that of snapper tagged in the marine reserve. Specifically, of the resident snapper tagged in the non-reserve area, half had home ranges with more than one main area of use, and as a result their home ranges spanned a linear distance of ∼2127 m on
Acknowledgements
Thanks to Richard Griffiths, Drew Lorrey, Matt Smith and James Williams for assistance with field work as well as Murray Birch, Arthur Cozens, Brady Doak and Daniel Egli of the Leigh Marine Laboratory for providing research facilities, equipment loans, boat usage and mooring construction. I am extremely grateful for advice on this manuscript received from Bruce Hartill and Nick Tolimieri, as well as technical assistance from Dale Webber of VEMCO and Greg Urbahn. Thanks to Dan Breen and Thelma
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