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Spatial models (meta-population models) Readings • Hilborn R et al. (2004) When can marine reserves improve fisheries management? Ocean and Coastal Management 47:197-205 • Hilborn R et al. (2006) Integrating marine protected areas with catch regulation. CJFAS 63:642-649 Overview • Why worry about space? • General metapopulation model • One-dimensional model of marine protected areas Why worry about space? Dutch beam trawl fleet England Netherlands Rijnsdorp AD et al. (1998) Micro-scale distribution of beam trawl effort in the southern North Sea between 1993 and 1996 in relation to the trawling frequency of the sea bed and the impact on benthic organisms. ICES Journal of Marine Science 55:403-419 UK fisheries Beam trawlers Dredgers Netters Otter trawlers Potters All combined Jennings S & Lee J (2012) Defining fishing grounds with vessel monitoring system data. ICES Journal of Marine Science 69:51-63 Ideal free distribution Abundance (variable) CPUE (constant) Effort (variable) Swain DP & EJ Wade (2003) Spatial distribution of catch and effort in a fishery for snow crab (Chionoectes opilio): tests of predictions of the ideal free distribution. CJFAS 60:897-909 MPA MPA MPA Marine protected area (MPA) MPA Murawski SA et al. (2005) Effort distribution and catch patterns adjacent to temperate MPAs. ICES Journal of Marine Science 62:1150-1167 Reason for “fishing the line” Dollars per hour CV of dollars Hours trawled Murawski SA et al. (2005) Effort distribution and catch patterns adjacent to temperate MPAs. ICES Journal of Marine Science 62:1150-1167 Metapopulation models A metapopulation is a series of discrete populations that are isolated but have limited exchange. Generalized metapopulation model Emigration: movement out of area i into area j Numbers in area i in time t Ni ,t 1 f Ni ,t , ei ,t Eij E ji j Environmental conditions in area i in time t j Immigration: movement from area j into area i Important details • Population dynamics within an area • Models of dispersal • Environmental models, process error, catastrophic events Boundaries? • Absorbing boundary http://www.google.com/pacman/ – Disappear/die on hitting the boundary – E.g. another country, advection into open ocean, natural range bounds • Reflective boundary – Remain in the cell next to the boundary – E.g. mountain range, river barrier • Pac-man (circular coastline) – Reappear on the opposite side – E.g. circumpolar, islands One-dimensional logistic-growth model with harvesting Numbers in area i in time t Density dependence Harvest Logistic model Ni ,t Ni ,t 1 Ni ,t Ni ,t ri 1 Ki Exploitation rate ui ,t Ni ,t Immigration m [1 ui 1,t ]Ni 1,t [1 ui 1,t ]Ni 1,t Emigration 2m[1 ui ,t ]Ni ,t Cell on the left Only survivors of harvest will move Movement rate the same in all cells Cell on the right Harvest, then movement 16 spatial models animation.r Diffusion scenario • 21 areas, 50 time steps, migration rate 0.2, r = 0.2, K = 1000, reflective boundary • Diffusion scenario: starting population = K in center cell, 0 in all other cells, exploitation rate 0 16 spatial models animation.r Diffusion, no harvesting, N11 = K Abundance (black = first year, light gray = last year) Cell number 16 spatial models animation.r Marine protected area scenario • As before: 21 areas, 50 time steps, migration rate 0.2, r = 0.2, K = 1000, reflective boundary • MPA scenario: starting population = K in all cells, exploitation rate 0 in center 5 cells (numbers 9–13), exploitation rate 0.2 in all other cells 16 spatial models animation.r MPA: no harvest in center cells Abundance (black = first year, light gray = last year) Cell number 16 spatial models animation.r Do protected areas increase yields? • 51 areas, migration rate m = 0.2, r = 0.2, K = 1000, start population = K in all areas • Run with u = 0, 0.01, 0.02, …, 0.9 • After a large number of time steps (1000) the model is at equilibrium, and yield in the final time step is the equilibrium yield for each value of u • No MPA: all areas have harvest rate = u • MPA: 5 middle areas have zero harvest rate, other have harvest rate = u 16 MPA yield.r 16 MPA yield.r Yield in each year Time series of catches Initially the no MPA scenario has higher catches; after year 14 with high u = 0.25 the MPA has higher catches; with no MPA, the population can go extinct No MPA Almost at equilibrium yield u = 0.1 MPA u = 0.25 Year 16 MPA yield.r Do protected areas increase yield? Higher yield without MPA u = r = 0.2 uMSY = r/2 = 0.1 Equilibrium yield No MPA Yield maximized at higher harvest rate with MPA MPA At u > r, MPA halts collapse (insurance policy) Harvest rate (u) 16 MPA yield.r Break-even point uMSY with MPA uMSY without MPA Equilibrium yield Zoomed in No MPA MPA Harvest rate (u) 16 MPA yield.r Bigger MPA (close 25 cells) (previously, close 5 of 51 cells) Equilibrium yield No MPA MPA Closing more areas reduces the maximum yield more Maximum yield reduced from 2550 to 1377 in this simulation Harvest rate (u) 16 MPA yield.r For the same yield what is u? Equilibrium yield No MPA MPA MPA: u = 0.10 produces MSY No MPA: two values of u result in equivalent yield: u = 0.032 and u = 0.168 Harvest rate (u) 16 MPA yield.r For the same yield, what is biomass? Abundance No MPA u = 0.032 MPA u = 0.1 No MPA u = 0.168 Cell number 16 MPA yield.r For the same yield, what is CPUE? (assume effort is proportional to harvest rate u) No MPA u = 0.032, yield = 1370, CPUE = 1370/0.032 = 42800 MPA u = 0.1, yield = 1370, CPUE = 13700 No MPA u = 0.168, yield = 1370, CPUE = 8200 16 MPA yield.r Lessons • Closing areas reduces the maximum yield • The more areas closed, the lower the maximum yield • Closed areas provide insurance against high fishing pressure (bad management, lack of enforcement) • For every level of yield with an MPA there is an equivalent yield without an MPA which has: – – – – – higher biomass outside the MPA lower biomass inside the MPA lower harvest rate where fishing occurs higher CPUE and hence greater profits no completely protected areas Old target New target Percent of maximum Tradeoffs of fishing Exploitation rate Worm B et al. (2009) Rebuilding global fisheries. Science 325:578-585 Lessons • • • • Spatial scale and pattern matters Simple movement models Marine protected areas: insurance vs. yield Trade-offs between catch, profit, and biodiversity