Abstract
This study examined whether a measured increase in average body size of adult sea scallops inside three fishery closed areas on Georges Bank (GB), United States (US), was sufficient to increase larval supply to closed areas and open fishing areas in both US and Canadian areas of the Bank. The effects of adult scallop density-at-size and fecundity-at-size on egg production were compared among open and closed fishery areas, countries, and time periods before and after the closed areas were established. Estimated egg production was then used to define spawning conditions in a coupled biological–physical larval tracking model that simulated larval development, mortality, and dispersal. Results showed that order of magnitude increases in larval settlement after closure were facilitated by increases in size-dependant egg production inside and dispersal from Closed Areas I and II, but not Nantucket Lightship Closed Area. The distributions of both egg production and larval settlement became more uniform across the Bank, causing the relative contribution of Canadian larvae to US scallop aggregations to decrease after establishment of Closed Areas I and II. Decreases in small and medium-sized scallop density in Canada and decreases in large scallops over the US-Southern Flank after closure caused local declines in egg production but were not sufficient to negatively affect larval settlement at the regional scale. Our model suggests that the establishment of fishery closed areas on GB considerably strengthened larval supply and settlement within and among several adult scallop aggregations.
Similar content being viewed by others
References
Abesamis RA, Russ GR (2005) Density-dependent spillover from a marine reserve: long-term evidence. Ecol Appl 15:1798–1812
Abesamis RA, Russ GR, Alcala AC (2006) Gradients of abundance of fish across no-take marine reserve boundaries: evidence from Philippine coral reefs. Mar Freshw Ecosyst 16:349–371
Alcala AC, Russ GR, Maypa AP, Calumpong HP (2005) A long-term, spatially replicated, experimental test of the effect of marine reserves on local fish yields. Can J Fish Aquat Sci 62:98–108
Almany GR, Connolly SR, Heath DD, Hogan JD, Jones GP, McCook LJ, Mills M, Pressey RL, Williamson DH (2009) Connectivity, biodiversity conservation, and the design of marine reserve networks for coral reefs. Coral Reefs 28:339–351
Beardsley RC, Chapman DC, Brink KH, Ramp SR, Schlitz R (1985) The Nantucket Shoals Flux Experiments (NSFE79). 1. A basic description of the current and temperature variability. J Phys Ocean 15:713–748
Botsford LW, White JW, Coffroth M-A, Paris CB, Planes S, Shearer TL, Thorrold SR, Jones GP (2009) Measuring connectivity and estimating resilience of coral reef metapopulations in MPAs: matching empirical efforts to modelling needs. Coral Reefs 28:134–145
Butman B, Beardsley R (1987) The physical oceanography of Georges Bank: introduction and summary. In: Backus RH (ed) Georges Bank. The MIT Press, Cambridge, pp 88–98
Carey JD, Stokesbury KDE (2011) An assessment of juvenile and adult sea scallop, Placopecten magellanicus, distribution in the northwest Atlantic using high-resolution still imagery. J Shellfish Res 30:569–582
Carey MJ, Carr MH, Hixon MA, Hughes TP, Jones GP, Menge BA (1996) Recruitment and the local dynamics of open marine populations. Ann Rev Ecol Syst 27:477–500
Chen C, Liu H, Beardsley R (2003) An unstructured grid, finite-volume, three-dimensional, primitive equations ocean model: application to coastal ocean and estuaries. J Atmos Ocean Technol 20:159–186
Christie MR, Tissot BN, Albins MA, Beets JP, Jia Y, Ortiz DM, Thompson SE, Hixon MA (2010) Larval connectivity in an effective network of marine protected areas. PLoS One 5:1–8
Davies KTA, Gentleman WC, Johnson CL, DiBacco C (2014) Relative contribution of bi-seasonally spawned larvae to scallop population connectivity on Georges Bank: importance of the spring spawn. MEPS. doi:10.3354/meps10975
DiBacco C, Robert G, Grant J (1995) Reproductive cycle of the sea scallop, Placopecten magellanicus (Gemlin 1971) on northeastern Georges Bank. J Shellfish Res 14:56–69
Fenberg PB, Caselle JE, Claudet J, Clemence M, Gaines SD, García-Charton J, Gonçalves EJ, Grorud-Colvert K, Guidetti P, Jenkins SR (2012) The science of European marine reserves: status, efficacy, and future needs. Mar Pol 36:1012–1021
Gaines SD, Gaylord B, Largier JL (2003) Avoiding current oversights in marine reserve design. Ecol Appl 13:S32–S46
Gell FR, Roberts CM (2003) Benefits beyond boundaries: the fishery effects of marine reserves. Trends Ecol Evol 18:448–455
Gilbert CS, Gentleman WC, Johnson CL, DiBacco C, Pringle JM, Chen C (2010) Modelling dispersal of sea scallop (Placopecten magellanicus) larvae on Georges Bank: the influence of depth distribution, planktonic duration and spawning seasonality. Prog Oceanogr 87:37–48
Greene CH, Pershing AJ (2003) The flip-side of the North Atlantic Oscillation and modal shifts in slope-water circulation patterns. Limnol Oceanogr 48:319–322
Halpern BS, Warner RR (2002) Marine reserves have rapid and lasting effects. Ecol Lett 5:361–366
Hannah CG, Naimie CE, Loder JW, Werner FE (1998) Upper-ocean transport mechanisms from the Gulf of Maine to Georges Bank, with implications for Calanus supply. Cont Shelf Res 17:1887–1911
Harmelin-Vivien M, Le Diréach L, Bayle-Sempere J, Charbonnel E, García- Charton JA, Ody D, Pérez-Ruzafa A, Reñones O, Sánchez-Jerez P, Valle C (2008) Gradients of abundance and biomass across reserve boundaries in six Mediterranean marine protected areas: evidence of fish spillover. Biol Conserv 141:1829–1839
Hart DR, Chute AS (2004) Essential fish habitat source document: sea scallop, Placopecten magellanicus, life history and habitat characteristics, 2nd edn. NOAA Technical Memorandum, NMFS NE-189
Hart DR, Rago PJ (2006) Long-term dynamics of U.S. Atlantic sea scallop Placopecten magellanicus populations. N Am J Fish Manage 26:490–501
Houghton RW, Aikman F, Ou HW (1988) Shelf-slope frontal structure and cross-shelf exchange at the New England shelf-break. Cont Shelf Res 8:687–710
Johnson CL, Pringle J, Chen C (2006) Transport and retention of dormant copepods in the Gulf of Maine. Deep-Sea Res II 53:2520–2536
Jones GP, Almany GR, Russ GR, Sale PF, Steneck RS, van Oppen MJH, Willis BL (2009) Larval retention and connectivity among populations of corals and reef fishes: history, advances and challenges. Coral Reefs 28:307–325
Langton R, Robinson W, Schick D (1987) Fecundity and reproductive effort of sea scallops Placopecten magellanicus from the Gulf of Maine. MEPS 37:19–25
Lester SE, Halpern BS, Grorud-Colvert K, Lubchenco J, Ruttenberg BI, Gaines SD, Airamé S, Warner RR (2009) Biological effects within no-take marine reserves: a global synthesis. MEPS 384:33–46
Lindholm J, Auster P, Valentine P (2004) Role of a large marine protected area for conserving landscape attributes of sand habitats on Georges Bank (NW Atlantic). MEPS 269:61–68
Mapstone BD, Little LR, Punt AE, Davies CR, Smith ADM, Pantus F, McDonald AD, Williams AJ, Jones A (2008) Management strategy evaluation for line fishing in the great barrier reef: balancing conservation and multi- sector fishery objectives. Fish Res 94:315–329
McCook LJ, Ayling T, Cappo M, Choat JH, Evans RD, De Freitas DM, Heupel M, Hughes TP, Jones GP, Mapstone B, Marsh H, Mills M, Molloy FJ, Pitcher CR, Pressey RL, Russ GR, Sutton S, Sweatman H, Tobin R, Wachenfeld DR, Williamson DH (2010) Adaptive management of the great barrier reef: a globally significant demonstration of the benefits of networks of marine reserves. Proc Natl Acad Sci 107:18278–18285
McGarvey R, Serchuk FM, McLaren IA (1993) Spatial and parent-age analysis of stock recruitment in the Georges Bank sea scallop (Placopecten magellanicus) population. Can J Fish Aquat Sci 50:564–574
McLean DL, Harvey ES, Meeuwig JJ (2011) Declines in the abundance of coral trout (Plectropomus leopardus) in areas closed to fishing at the Houtman Abrolhos Islands, Western Australia. J Exp Mar Biol Ecol 406:71–78
Murawski SA, Brown R, Lai HL, Rago PJ, Hendrickson L (2000) Large-scale closed areas as a fishery-management tool in temperate marine systems: the Georges Bank experience. Bull Mar Sci 66:775–798
Naimie CE, Loder JW, Lynch DR (1994) Seasonal variation of the three dimensional residual circulation on Georges Bank. J Geophys Res 99:15967–15989
Naimie CE, Limeburner R, Hannah C, Beardsley R (2001) On the geographic and seasonal patterns of the near-surface circulation on Georges Bank—from real and simulated drifters. Deep-Sea Res II 48:501–518
New England Fisheries Management Council (NEFMC) (2010) Framework Adjustment 21 to the Atlantic Sea Scallop Fisheries Management Plan. Available from www.nefmc.org/scallops/index.html
Noble M, Butman B (1985) Wind-current coupling on the southern flank of Georges Bank: variation with season and frequency. J Phys Ocean 15:604–620
North EW, Gallego A, Petitgas P (2009) Manual of recommended practices for modelling physical-biological interactions during fish early life. ICES Cooperative Research Report (1017-6195) N. 295, P. pp. 1–111
Owen EF, Rawson PD (2013) Small-scale spatial and temporal structure of the Atlantic sea scallop (Placopecten magellanicus) in the inshore Gulf of Maine revealed using AFLPs. Mar Biol 160:3015–3025
Pettigrew NR, Churchill JH, Janzen CD, Mangum LJ, Signell RP, Xue HJ (2005) The kinematic and hydrographic structure of the Gulf of Maine Coastal Current. Deep-Sea Res II:2369–2391
Russ GR (2002) Yet another review of marine reserves as reef fishery management tools. In: Sale PF (ed) Coral reef fishes: dynamics and diversity in a complex system. Academic Press, New York, pp 421–443
Russ GR, Alcala AC (2010) Decadal-scale rebuilding of predator biomass in Philippine marine reserves. Oecologia 163:1103–1106
Russ G, Cheal AJ, Dolman AM, Emslie MJ, Evans RD, Miller I, Sweatman H, Williamson DH (2008) Rapid increase in fish numbers follows creation of world’s largest marine reserve network. Curr Biol 18:R514–R515
Smith SJ, Denton C, Hubley B, Jonsen ID, Lundy MJ, Pezzack D, Sameoto JA, Tremblay MJ (2009) Scallop fishing area 29: stock status and update for 2009. Can Sci Advis Sec Res Doc 2009/38
Stokesbury KDE (2002) Estimation of sea scallop, Placopecten magellanicus, abundance in closed areas of Georges Bank. Trans Am Fish Soc 131:1081–1092
Stokesbury KDE (2012) Stock definition and recruitment: implications for the U.S. sea scallop (Placopecten magellanicus) fishery from 2003–2011. Rev Fish Sci 20:154–164
Tian RC, Chen CS, Stokesbury KDE, Rothschild BJ, Cowles G, Xu QC, Harris BP, Marino MC (2009) Dispersal and settlement of sea scallop larvae spawned in the fishery closed areas on Georges Bank. ICES J Mar Sci 66:2155–2164
Tremblay MJ, Sinclair M (1990) Sea scallop Placopecten magellanicus larvae on Georges Bank: vertical distribution in relation to water column stratification and food. MEPS 61:1–15
Tremblay MJ, Sinclair M (1992) Planktonic sea scallop larvae (Placopecten magellanicus) in the Georges Bank region: broadscale distribution in relation to physical oceanography. Can J Fish Aquat Sci 49:1597–1615
Tremblay MJ, Loder JW, Werner FE, Naimie CE, Page FH, Sinclair MM (1994) Drift of sea scallop larvae Placopecten magellanicus on Georges Banks: a model study of the roles of mean advection, larval behavior and larval origin. Deep Sea Res. II 41:7–49
Visser AW (1997) Using random walk models to simulate the vertical distribution of particles in a turbulent water column. MEPS 158:275–281
Wen CKC, Almany GR, Williamson DH, Pratchett MS, Mannering TD, Evans RD, Leis JM, Srinivasan M, Jone GP (2013) Recruitment hotspots boost effectiveness of no-take marine reserves. Biol Conserv 166:124–131
Williamson DH, Russ GR, Ayling AM (2004) No-take marine reserves increase abundance and biomass of reef fish on inshore fringing reefs of the great barrier reef. Environ Conserv 31:149–159
Xue H, Incze L, Xu D, Wolff N, Pettigrew N (2008) Connectivity of lobster populations in the coastal Gulf of Maine Part I: circulation and larval transport potential. Ecol Model 210:193–211
Zimmerman JTF (1979) On the Euler–Lagrange transformation and the Stokes’ drift in the presence of oscillatory and residual currents. Deep-Sea Res I 26:505–520
Acknowledgments
Thank you to the scallop stock assessment division at the Bedford Institute of Oceanography, Stephen Smith (Fisheries and Oceans Canada), Brad Hubley (Fisheries and Oceans Canada), Deborah Hart (National Marine Fisheries Service), and Penny Kuhn for providing scallop tow data and discussion on the manuscript, as well as two anonymous referees for providing comments. Thank you to Brittney Vujcich and Jesse Burgess for assisting in model development. We are grateful to Jamie Pringle and Changsheng Chen for producing and providing Lagrangian physical fields from the FVCOM hydrodynamic model, to Chad Gilbert for developing an earlier iteration of the scallop biological model and data fields, and to anonymous reviewers for providing comments. Funding provided by NSERC Discovery Grant to WCG and by Fisheries and Oceans Canada’s Ecosystem Research Initiative to CDB and CLJ.
Author information
Authors and Affiliations
Corresponding author
Electronic supplementary material
Below is the link to the electronic supplementary material.
Rights and permissions
About this article
Cite this article
Davies, K.T.A., Gentleman, W.C., DiBacco, C. et al. Fisheries Closed Areas Strengthen Scallop Larval Settlement and Connectivity Among Closed Areas and Across International Open Fishing Grounds: A Model Study. Environmental Management 56, 587–602 (2015). https://doi.org/10.1007/s00267-015-0526-9
Received:
Accepted:
Published:
Issue Date:
DOI: https://doi.org/10.1007/s00267-015-0526-9