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Marine Reserve Networks Conserve Biodiversity by Stabilizing Communities and Maintaining Food Web Structure

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Marine Reserve Networks Conserve Biodiversity by Stabilizing Communities and Maintaining Food Web Structure Marine reserve networks conserve biodiversity by stabilizing communities and maintaining food web structure STEPHEN R. WING AND LUCY JACK Department of Marine Science, University of Otago, 310 Castle Street, Dunedin, New Zealand 9054 Citation: Wing, S. R., and L. Jack. 2013. Marine reserve networks conserve biodiversity by stabilizing communities and maintaining food web structure. Ecosphere 4(10):XXX. http://dx.doi.org/10.1890/ES13-00257.1 Abstract. Theory predicts that networks of fully protected marine reserves conserve biodiversity by stabilising communities and maintaining food web structure in the face of inadequately constrained fishery exploitation. To test these ideas we examine trends in species incidence, community and trophic structure of temperate reef fishes over an eight year period within the Fiordland no-take marine reserve network, at management zones subject to commercial fishing and at those closed to commercial exploitation but open to recreational fishers. We use information from extensive stratified subtidal surveys of the reef fish community and abundance of macroalgae, as well as oceanographic data collected in 2002, 2006 and 2010. Our analyses indicate a regional decline in species richness of exploited reef fish in areas open to fishing between 2002 and 2010. Following implementation of spatial management (2006–2010), richness of ‘exploited’ species increased within marine reserves, but remained unchanged in areas open to fishing. Further, analysis of differences in community structure in this time period (2006–2010) indicate that both ‘exploited’ and ‘non-target’ groups were more stable within marine reserves than were those within fished areas. Consequentially average trophic level of the community remained stable within marine reserves but declined sharply in areas open to fishing, indicating both declines in large omnivorous species and increases in forage fish within exploited regions. These analyses offer an important test of the direct and indirect effects of marine reserve networks on the dynamics of reef fish communities at the landscape scale. We demonstrate the potential for multiple no-take reserves spread over a heterogeneous marine landscape to maintain biodiversity by stabilizing community structure and preserving intact food webs on a regional scale. Key words: conservation; diversity; fishing; marine reserve network; reef fish community; species richness; trophic level. Received 19 August 2013; accepted 16 September 2013; published 00 Month 2013. Corresponding Editor: S. Cox. Copyright: Ó 2013 Wing and Jack. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. http://creativecommons.org/licenses/by/3.0/ E-mail: [email protected] INTRODUCTION genic perturbation. Early theoretical studies predicted that restricted diet within a food web Successful ecosystem management puts mea- lowers community stability (MacArthur 1955). sures in place that guard against the erosion of Indeed virtual food webs with large numbers of ecological processes and promotes the mainte- weak, ephemeral linkages are more stable, and nance of biodiversity. Ecologists have long been thus more persistent, than those with few strong, fascinated by the interaction between the com- requisite linkages (Polis 1994, McCann and plexity of food web architecture and community Hastings 1997). Omnivory, where a consumer stability, where more stable communities have feeds at multiple trophic levels (Pim and Lawton greater resistance to environmental or anthropo- 1978), results in a fractional trophic position of v www.esajournals.org 1 Month 2013 v Volume 4(10) v Article 0 //xinet/production/e/ecsp/live_jobs/ecsp-04-10/ecsp-04-10-13/layouts/ecsp-04-10-13.3d Thursday, 10 October 2013 9:49 pm Allen Press, Inc. Page 1 ECSP ES13-00257R Jack WING AND JACK the consumer (Levine 1980), calculated as an spatial closures to conserve or stimulate recovery average of the constituent trophic levels of the of intact communities. The Fiordland Marine prey base. The degree of omnivory of a species Area (Te Moana o Atawhenua) (FMA) is a region may change during ontogeny (Abrams 2011), of globally significant natural and cultural vary spatially within a population (Parsons and heritage and of great economic importance to LeBrasseur 1970) or vary according to competi- New Zealand. In recognition of this, two no-take tive interactions within linked food webs (Holt marine reserves were established in 1993, in and Polis 1997). Food webs with a high degree of Doubtful and Milford Sounds. Furthermore, in omnivory support fewer strong interactions and 2005, the Fiordland Marine Management Act have a reduced likelihood of dramatic indirect closed the inner regions of eleven fjords to effects such as trophic cascades (Polis 1994). commercial fishing (46,002 ha; 59% of the FMA) Consequentially, complex communities with high and established a network of eight new marine trophic level omnivores can maintain more stable reserves nested within commercial exclusion temporal and spatial dynamics (Polis and Strong zones, bringing the total no-take areas to 10, 1996). These studies form the basis for asking a and covering 10,421 ha or 13.11% of the FMA question of great importance for successful (Fig. 1). To meet fisheries and biodiversity management of exploited systems: What hap- conservation objectives in this highly subdivided pens to a community subject to selective removal habitat (Lubchenco et al. 2003) efforts were taken of top trophic level omnivores? to distribute fully protected marine reserves Omnivory is widespread in marine systems widely across the region among fjords and where large bodied, generalist predators have within representative habitat types (Wing and been heavily exploited by fishing, in many cases Jack 2010). All except one of the no-take marine resulting in a simplification of food webs (Pauly reserves were nested in a buffer zone, where et al. 1998) and in changes to the trophic commercial fishing was closed and recreational dynamics of communities (Frank et al. 2011). fishing limits were reduced. Temperate reef fish communities contain a wide Testing the ecological consequences of a diversity of omnivorous species. Here selective protected area network is implicit to a successful removal of high trophic level omnivores has adaptive management strategy whose goal is to provided an explicit test of system stability in optimize regional biodiversity patterns. Com- their absence. The effects of localized fishing monly employed impact-control studies measure mortality on community dynamics may be spatial management outcomes by comparing relatively elusory in open systems where subsi- regions that are under different management dies and rescue effects from nearby refuge regimes (no-take reserve, commercial exclusion populations can dampen local exploitation-driv- zones, open fished areas) but that are otherwise en changes in spatial dynamics (Wing and Wing comparable, before and after implementation. 2001, Kritzer and Sale 2006). However in more However the New Zealand fjords are character- insular systems, the effects of exploitation may be ized by strong physical environmental gradients locally enhanced (Roberts 1995). In theory, containing the extremes of salinity, wave action networks of no-take marine reserves may stabi- and irradiance that together influence patterns of lize communities by providing a direct refuge to pelagic productivity and density of habitat- the exploited component of a community, typi- providing macroalgae along the axis of each cally omnivorous and top predatory fishes fjord (Goebel et al. 2005, Wing et al. 2007). The (Fogarty 1999). The present study investigated resulting gradient in composition of organic this idea by examining patterns in reef fish matter source pools can be directly linked to community and food web structure within a diet, sub-population structure, growth and fe- new marine reserve network established in cundity of reef fish (Wing et al. 2012, Beer and Fiordland, southwest New Zealand. Wing 2013). Thus comparable control sites in The network of marine protected areas in differing management zones are rare or nonex- Fiordland offers a model system for testing the istent. effects of selective removal of high trophic level To overcome this limitation, in the present omnivores in nature and for testing the ability of study we make use of a time series from three v www.esajournals.org 2 Month 2013 v Volume 4(10) v Article 0 //xinet/production/e/ecsp/live_jobs/ecsp-04-10/ecsp-04-10-13/layouts/ecsp-04-10-13.3d Thursday, 10 October 2013 9:49 pm Allen Press, Inc. Page 2 ECSP ES13-00257R Jack WING AND JACK Fig. 1. Map of the Fiordland Marine Area in southwestern New Zealand with marine reserves (dark gray), commercial exclusion zones (light gray) and open areas (white) indicated. Positions of long term monitoring sites surveyed in 2002, 2006 and 2010 are indicated by black circles. Additional sites surveyed only in 2006 and 2010 are indicated by open circles. Fiordland-wide subtidal surveys conducted over changes in diversity, community composition an eight-year period to compare changes in the and food web structure are coincident with species composition and trophic structure of the spatial management of fishing. fish community within regions under different management regimes. We use information
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