2 1.8 > 1.6 a!= 1-1 Ë 1.2 E1 E 0.8 E 0.6 2 0.4 0.2 0 rÞ"- ,Þù* .Ñ 9' \ù .ñr "ê" èo¡ *o**d M.C. Tynell et dl. / Fisheríes Research 108 (201 1) 1-8 :T$ ì,.:4 c:,-{ ,df . 0d 'lo \9' ,ao 'oò cl \o \0) à '.s Øo t &a ,Þo ¡le f"at o' ¿s( Þ" -^S' nd o' og. oâ^rf )' á' "i .
M.C.TyneII et al. / Fßheries Research 108 (2011) 1-8 s Table 2 Comparison ofMSY (maximum sustainable yield) based biological reference points (Busv: biomass at maximum sustainable yield, SSBusv: spawning stock biomass at maximum sustainable yield) produced by explicitly incorporåting predation (= ¡gvi..¿' numerator) and traditional (= ñshery; denominator) methods. Prey species, author Predators Revised Bs5y or SSBs5y/ Traditional Busv or SSBusv Atlantic herring, Overholtz et el. (2008) Atlantic herring, Tyrrell et al. (2008) Atlantic mackerel, Moustahfid et al, (2009a) Atlantic mackerel, Tyrrell et al, (2008) Longfin squid, Mouståhfid et al. (2fr)9b) 29 species inc. fish, mar mamm, seabirds .11 demersal fish species 13 demersal fish species 1 1 demersal fish species 15 demersal fish species ered its projected spawning biomass estimates. Similar results have also been observed for several species simulated for the southeast Australian fisheries ecosystem (Fulton et al., 2007). Yield projections for individual species can both increase and decrease when ecological interactions are taken into account, depending on the dynamics between the focal species and whether alternate prey are available. Under the assumption that gray seal, Halichoerus grypus, predation on Atlantic cod was additive to other predation mortality, Mohn and Bowen (1996) showed >50% reductions in yield for cod during a time of increased seal abundance. Similarly, in the Northeast US, yields of prey species such as herring will likely decline as their predator stocks recover from decades of overfi shing (Overholtz et al., 2008). For predators with limited foraging areas and little alternate prey, precautionary approaches to calculating fisheries yields are of critical importance. Everson and de la Mare (1996) incorporated the requirements of land based predators (seals, penguins, petrels and other birds) on Antarctic krill, Ëuphausia superba, around South Georgia Island. They suggested a 75% reduction in the precautionary catch limit to allow for acceptable impacts of the krill fishery for these predators with limited foraging ranges. 2.4. Biological reþrence points are difþrent wíth ecological considerations Biological reference points derived with multispecies models differ from their traditional single-species counterparts in that they generally result in more precautionary management advice (Hall, 1 999; ICES, 2008). ln addition to the studies described in some detail below, many other studies support the finding that BRPs change when predation is explicitly modeled (e.g., ICES, 1997; Collie and Delong, 1999; Hvingel and Kingsley, 2006). Table 2 summarizes a suite of studies that explicitly compared BRPs from traditional stock assessment model parameterization vs. a situation where consumption on the focal species was calculated and the resulting revised predation mortalities and biomasses were used for BRP estimation. An important example of changed BRPs from including predation comes from the Baltic Sea ecosystem, where Gislason (1999) reported complex relationships between reference limits for cod and herring and sprat, but with a straightforward overriding conclusion - BRPs of stocks that interact should not be considered in isolation. Similarly, under the adverse recruitment conditions of recent decades, Koster et al. (2009) found that incorporating cannibalism for Eastern Baltic cod lowered the estimated fishing mortality rate that was required to reach Bpa. Collie and Gislason (2001) concluded that fishing mortality reference points for prey such as sprat should be conditioned on changes in predator abundance andJurado-Molina and Livingston (2002) found that their three prey species were also sensitive to the harvest levels of their predators. For Barents Sea capelin Gjøsæter et al. (2002) advocated stochastic reference points should be developed to account for variable predation by cod and marine mammals. Accurate calculation of reference limits, especially for forage species, requires consid- 1.62 1.38 2.71 1.1 1 4.27 Revised MSY/ Traditional MSY 2.39 7.25 1.83 7.57 3.36 eration of the dynamic biomass levels of both predator and prey populations. Another example of changed BRPs is the northwest Atlantic herring fishery. Overholtz et al. (2008) used a delay difference model to calculate surplus production ofage 2+ herringwith predatory removals by demersal fishes, marine mammals, large pelagic fishes and seabirds. The B¡¡5y derived from the model with predation explicitly incorporated was higher than the fishery only Brursv by a factor of 1.6. Moustahfid et al. (2009b) incorporated predatory removals of longfin inshore squid, lolþo peøleii, using a surplus production model in a similar manner as Overholtz et al. (2008) and found that B¡aup (a proxy for B¡a5y, maximum usable production) increased by more than a factor of three when predation was explicitly accounted for. Similarly, MSY almost doubled and SSB¡y¡5y increased by almost three times when predation by 13 demersal fish species was explicitly incorporated into an age-structured assessment model for Atlantic mackerel (Scomber scombrus, Moustahñd et al., 2009a). In an MSVPA of 14 predator stocks and 2 age structured prey species (Atlantic herring and Atlantic mackerel) of the Northeast US Continental shelf ecosystem, Tyrrell et al. (2008) found that herring's MSY and Brvrsv increased in a multispecies vs. single species context, but not to as strongly as reported by Overholtz et al. (2008). For mackerel, the MSVPA biomass estimates also resulted in BRPs that were more conservative than the reference points produced by traditional single-species methods (Tyrrell et al., 2008). Different types of modeling approaches (e.g., age structured vs. non-age structured) and different suites of predators and input parameters have resulted in variable point estimates of BRPs (e.g., Overholtz et al., 2008 vs. Tyrrell et al., 2008). Despite variation in the absolute value of BRPs with and without predation incorporated, B¡a5y or SSBy5y increased by ,7O% for all situations where these types of comparisons were made (Table 2). For other reference points such as F.r"r¡ and Fe.1 (both based off of stock-recruitment relationships, with Fe.1 being the fishing mortality rate at 10% of the maximal yield per recruit rate, and F.rur6 being the fishing rate which will produce a longterm spawning biomass per recruit (S/R) equal to the inverse of the instantaneous rate of variation of R with the biomass, at the initial point (S = 0, R = 0)), a similar finding of more conservative reference points being calculated has been reported by other authors. For example, both Fo.l and F.r"r¡ were lower in a multispecies context for MSVPAs of the Barents Sea and the North Sea (ICES, 1 997). As the majority of the aforementioned studies show, inferences from a variety of models for various fisheries species indicate that BRPs for forage species are different and generally point to more conservative harvest rates when ecological considerations are accounted for. To broaden the applicability to EBFM, BRPS can also be calculated for a suite of species in addition to individual values for each species. Mueter and Megrey (2006) aggregated fisheries species into a surplus production model to calculate an ecosystem-level MSY (termed multi-species maximum surplus production) for the Gulf of Alaska and Bering Sea commercially exploited groundfish species. They found that in both ecosystems, this ecosystem-level MSY was smaller than the component sum of single species MSYs (Table 3) and furthermore, that incorporation
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