Skip to main content Accessibility help

Spillover from marine reserves and the replenishment of fished stocks



No-take marine reserves are widely recognized as an effective conservation tool for protecting marine resources. Despite considerable empirical evidence that abundance and biomass of fished species increase within marine reserve boundaries, the potential for reserves to provide fisheries and conservation benefits to adjacent waters remains heavily debated. This paper uses statistical and population models to evaluate published empirical data on adult spillover from marine reserves and shows that spillover is a common phenomenon for species that respond positively to reserve protection, but at relatively small scales, detectable on average up to 800 m from reserve boundaries. At these small scales, local fisheries around reserves were likely unsustainable in 12 of 14 cases without the reserve, and spillover partially or fully offsets losses in catch due to reserve closure in the other two cases. For reserves to play a role in sustaining and replenishing larger-scale fished stocks, networks of reserves may be necessary, but as few exist this is difficult to evaluate. The results suggest reserves can simultaneously meet conservation objectives and benefit local fisheries adjacent to their boundaries.


Corresponding author

*Correspondence: Dr Benjamin S. Halpern Tel: +1 805 892 2531 Fax: +1 805 892 2510 e-mail:


Hide All
Abesamis, R.A. & Russ, G.R. (2005) Density-dependent spillover from a marine reserve: long-term evidence. Ecological Applications 15 (5): 17981812.
Abesamis, R.A., Alcala, A.C. & Russ, G.R. (2006 a) How much does the fishery at Apo Island benefit from spillover of adult fish from the adjacent marine reserve? Fishery Bulletin 104 (3): 360375.
Abesamis, R.A., Russ, G.R. & Alcala, A.C. (2006 b) Gradients of abundance of fish across no-take marine reserve boundaries: evidence from Philippine coral reefs. Aquatic Conservation: Marine and Freshwater Ecosystems 16 (4): 349371.
Alcala, A.C., Russ, G.R., Maypa, A.P. & Calumpong, H.P. (2005) A long-term, spatially replicated experimental test of the effect of marine reserves on local fish yields. Canadian Journal of Fisheries and Aquatic Sciences 62 (1): 98108.
Ashworth, J.S. & Ormond, R.F.G. (2005) Effects of fishing pressure and trophic group on abundance and spillover across boundaries of a no-take zone. Biological Conservation 121 (3): 333344.
Cantrell, R.S. & Cosner, C. (2003) Spatial Ecology via Reaction-diffusion Equations. New York, NY, USA: John Wiley and Sons.
Claudet, J., Osenberg, C.W., Benedetti-Cecchi, L., Domenici, P., Garcia-Charton, J.A., Perez-Ruzafa, A., Badalamenti, F., Bayle-Sempere, J., Brito, A., Bulleri, F., Culioli, J.M., Dimech, M., Falcon, J. M., Guala, I., Milazzo, M., Sanchez-Meca, J., Somerfield, P.J., Stobart, B., Vandeperre, F., Valle, C. & Planes, S. (2008) Marine reserves: size and age do matter. Ecology Letters 11 (5): 481489.
Costello, C., Gaines, S.D. & Lynham, J. (2008) Can catch shares prevent fisheries collapse? Science 321: 16781681.
Fisher, R.A. (1937) The wave of advance of advantageous genes. Annals of Eugenics 7: 353369.
Forcada, A., Bayle-Sempere, J.T., Valle, C. & Sanchez-Jerez, P. (2008) Habitat continuity effects on gradients of fish biomass across marine protected area boundaries. Marine Environmental Research 66 (5): 536547.
Forcada, A., Valle, C., Bonhomme, P., Criquet, G., Cadiou, G., Lenfant, P. & Sanchez-Lizaso, J.L. (2009) Effects of habitat on spillover from marine protected areas to artisanal fisheries. Marine Ecology Progress Series 379: 197211.
Francini-Filho, R.B. & Moura, R.L. (2008) Evidence for spillover of reef fishes from a no-take marine reserve: an evaluation using the before-after control-impact (BACI) approach. Fisheries Research 93: 346356.
Gell, F.R. & Roberts, C.M. (2003) Benefits beyond boundaries: the fishery effects of marine reserves. Trends in Ecology and Evolution 18 (9): 448455.
Goni, R., Adlerstein, S., Alvarez-Berastegui, D., Forcada, A., Renones, O., Criquet, G., Polti, S., Cadiou, G., Valle, C., Lenfant, P., Bonhomme, P., Perez-Ruzafa, A., Sanchez-Lizaso, J.L., Garcia-Charton, J.A., Bernard, G., Stelzenmuller, V. & Planes, S. (2008) Spillover from six western Mediterranean marine protected areas: evidence from artisanal fisheries. Marine Ecology-Progress Series 366: 159174.
Goni, R., Quetglas, A. & Renones, O. (2006) Spillover of spiny lobsters Palinurus elephas from a marine reserve to an adjoining fishery. Marine Ecology Progress Series 308: 207219.
Guenette, S., Pitcher, T.J. & Walters, C.J. (2000) The potential of marine reserves for the management of northern cod in Newfoundland. In: Essential Fish Habitat and Marine Reserves, Bulletin of Marine Science Volume 66, ed. EDITORS, pp. 831–852. Miami, FL, USA: University of Miami, Rosenthiel School of Marine and Atmospheric Science.
Guidetti, P. (2007) Potential of marine reserves to cause community-wide changes beyond their boundaries. Conservation Biology 21 (2): 540545.
Halpern, B.S. (2003) The impact of marine reserves: do reserves work and does reserve size matter? Ecological Applications 13 (1): S117S137.
Halpern, B.S., Gaines, S.D. & Warner, R.R. (2004) Confounding effects of the export of production and the displacement of fishing effort from marine reserves. Ecological Applications 14 (4): 12481256.
Halpern, B.S., Walbridge, S., Selkoe, K.A., Kappel, C.V., Micheli, F., D'Agrosa, C., Bruno, J., Casey, K.S., Ebert, C., Fox, H.E., Fujita, R., Heinemann, D., Lenihan, H.S., Madin, E.M.P., Myers, R., Perry, M., Selig, E., Spalding, M., Steneck, R. & Watson, R. (2008) A global map of human impact on marine ecosystems. Science 319: 948952.
Harmelin-Vivien, M., Le Direach, L., Bayle-Sempere, J., Charbonnel, E., Garcia-Charton, J.A., Ody, D., Perez-Ruzafa, A., Renones, O., Sanchez-Jerez, P. & Valle, C. (2008) Gradients of abundance and biomass across reserve boundaries in six Mediterranean marine protected areas: evidence of fish spillover? Biological Conservation 141 (7): 18291839.
Hastings, A. & Botsford, L.W. (1999) Equivalence in yield from marine reserves and traditional fisheries management. Science 284 (5419): 15371538.
Hilborn, R., Stokes, K., Maguire, J.J., Smith, T., Botsford, L.W., Mangel, M., Orensanz, J., Parma, A., Rice, J., Bell, J., Cochrane, K.L., Garcia, S., Hall, S.J., Kirkwood, G.P., Sainsbury, K., Stefansson, G. & Walters, C. (2004) When can marine reserves improve fisheries management? Ocean and Coastal Management 47 (3–4): 197205.
Kaplan, D.M. & Botsford, L.W. (2005) Effects of variability in spacing of coastal marine reserves on fisheries yield and sustainability. Canadian Journal of Fish and Aquatic Science 62: 905912 [www document]. URL
Kaunda-Arara, B. & Rose, G.A. (2004) Effects of marine reef national parks on fishery CPUE in coastal Kenya. Biological Conservation 118 (1): 113.
Kellner, J.B., Nisbet, R.M. & Gaines, S.D. (2008) Spillover from marine reserves related to mechanisms of population regulation. Theoretical Ecology 1: 117127.
Kellner, J.B., Tetreault, I., Gaines, S.D. & Nisbet, R.M. (2007) Fishing the line near marine reserves in single and multispecies fisheries. Ecological Applications 17 (4): 10391054.
Kelly, S. & MacDiarmid, A.B. (2003) Movement patterns of mature spiny lobsters, Jasus edwardsii, from a marine reserve. New Zealand Journal of Marine and Freshwater Research 37 (1): 149158.
Le Quesne, W.J.F. & Codling, E.A. (2009) Managing mobile species with MPAs: the effects of mobility, larval dispersal, and fishing mortality on closure size. ICES Journal of Marine Science 66 (1): 122131.
Lester, S.E., Halpern, B.S., Grorud-Colvert, K., Lubchenco, J., Ruttenberg, B.I., Gaines, S.D., Airame, S. & Warner, R.R. (2009) Biological effects within no-take marine reserves: a global synthesis. Marine Ecology Progress Series 384: 3346.
Lubchenco, J., Palumbi, S.R., Gaines, S.D. & Andelman, S. (2003) Plugging a hole in the ocean: the emerging science of marine reserves. Ecological Applications 13: S3S7.
McClanahan, T.R. & Mangi, S. (2000) Spillover of exploitable fishes from a marine park and its effect on the adjacent fishery. Ecological Applications 10: 17921805.
McClanahan, T.R. & Mangi, S. (2001) The effect of a closed area and beach seine exclusion on coral reef fish catches. Fisheries Management and Ecology 8 (2): 107121.
Micheli, F., Halpern, B.S., Botsford, L.W. & Warner, R.R. (2004) Trajectories and correlates of community change in no-take marine reserves. Ecological Applications 14 (6): 17091723.
Murawski, S.A., Wigley, S.E., Fogarty, M.J., Rago, P.J. & Mountain, D.G. (2005) Effort distribution and catch patterns adjacent to temperate MPAs. ICES Journal of Marine Science 62 (6): 11501167.
Neubert, M.G. (2003) Marine reserves and optimal harvesting. Ecology Letters 6 (9): 843849.
Pauly, D., Christensen, V., Dalsgaard, J., Froese, R. & Torres, F. (1998) Fishing down marine food webs. Science 279 (5352): 860863.
Perez-Ruzafa, A., Martin, E., Marcos, C., Zamarro, J. M., Stobart, B., Harmelin-Vivien, M., Polti, S., Planes, S., Garcia-Charton, J. A. & Gonzalez-Wanguemert, M. (2008) Modelling spatial and temporal scales for spill-over and biomass exportation from MPAs and their potential for fisheries enhancement. Journal of Nature Conservation 16 (4): 234255.
Pitcher, T.J., Buchary, E.A. & Hutton, T. (2002) Forecasting the benefits of no-take human-made reefs using spatial ecosystem simulation. ICES Journal of Marine Science 59: S17S26 [www document]. URL
R (2006) R: a Language and Environment for Statistical Computing. Vienna, Austria: R Development Core Team.
Rakitin, A. & Kramer, D.L. (1996) Effect of a marine reserve on the distribution of coral reef fishes in Barbados. Marine Ecology Progress Series 131 (1–3): 97113.
Roberts, C.M., Bohnsack, J.A., Gell, F., Hawkins, J.P. & Goodridge, R. (2001) Effects of marine reserves on adjacent fisheries. Science 294 (5548): 19201923.
Roberts, C.M., Hawkins, J.P. & Gell, F.R. (2005) The role of marine reserves in achieving sustainable fisheries. Philosophical Transactions of the Royal Society B. Biological Sciences 360 (1453): 123132.
Rodwell, L.D., Barbier, E.B., Roberts, C.M. & McClanahan, T.R. (2003) The importance of habitat quality for marine reserve: fishery linkages. Canadian Journal of Fisheries and Aquatic Sciences 60: 171181.
Rowley, R.J. (1994) Marine reserves in fisheries management. Aquatic Conservation, Marine and Freshwater Ecosystems 4 (3): 233254.
Russ, G.R., Alcala, A.C., Maypa, A.P., Calumpong, H.P. & White, A.T. (2004) Marine reserve benefits local fisheries. Ecological Applications 14 (2): 597606.
Samoilys, M.A. (1997) Movement in a large predatory fish: coral trout, Plectropomus leopardus (Pisces: Serranidae), on Heron Reef, Australia. Coral Reefs 16 (3): 151158.
Shigesada, N. & Kawasaki, K. (1997) Biological Invasions: Theory and Practice. Oxford, UK: Oxford University Press.
Smith, M.D. & Wilen, J.E. (2003) Economic impacts of marine reserves: the importance of spatial behavior. Journal of Environmental Economics and Management 46: 183206.
Topping, D.T., Lowe, C.G. & Caselle, J.E. (2005) Home range and habitat utilization of adult California sheephead, Semicossyphus pulcher (Labridae), in a temperate no-take marine reserve. Marine Biology 147 (2): 301311.
Tupper, M.H. (2002) Marine reserves and fisheries management. Science 295 (5558): 12331233.
Williams, I.D., Walsh, W.J., Claisse, J.T., Tissot, B.N. & Stamoulis, K.A. (2009) Impacts of a Hawaiian marine protected area network on the abundance and fishery sustainability of the yellow tang, Zebrasoma flavescens. Biological Conservation 142 (5): 10661073.
Worm, B., Barbier, E.B., Beaumont, N., Duffy, J.E., Folke, C., Halpern, B.S., Jackson, J.B.C., Lotze, H. K., Micheli, F., Palumbi, S.R., Sala, E., Selkoe, K.A., Stachowicz, J.J. & Watson, R. (2006) Impacts of biodiversity loss on ocean ecosystem services. Science 314 (5800): 787790.
Zeller, D. & Reinert, J. (2004) Modelling spatial closures and fishing effort restrictions in the Faroe Islands marine ecosystem. Ecological Modelling 172 (2–4): 403420.
Zeller, D., Stoute, S.L. & Russ, G.R. (2003) Movements of reef fishes across marine reserve boundaries: effects of manipulating a density gradient. Marine Ecology Progress Series 254: 269280.


Type Description Title
Supplementary materials

Halpern et al. supplementary material
Supplementary tables

 Word (359 KB)
359 KB


Altmetric attention score

Full text views

Total number of HTML views: 0
Total number of PDF views: 0 *
Loading metrics...

Abstract views

Total abstract views: 0 *
Loading metrics...

* Views captured on Cambridge Core between <date>. This data will be updated every 24 hours.

Usage data cannot currently be displayed