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The Status of Sea Bass

The Status of Sea Bass

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The status of sea bass (Dicentrarchus labrax) stocks around England and Wales, derived using a separable catch

-at-age model, and implications for fisheries management
Mike G. Pawson, Sven Kupschus and Graham D. Pickett
The Centre for Environment, Fisheries and Aquaculure Science (Cefas), Pakefield Road, Lowestoft, Suffolk NR33 0HT, UK Correspondence to M. G. Pawson: tel: +44 1502 524436; fax: +44 1502 526351; e-mail: mike.pawson@cefas.co.uk

Pawson, M. G., Kupschus, S., and Pickett, G. D. 2007. The status of sea bass (Dicentrarchus labrax) stocks around England and Wales, derived using a separable catch-at-age model, and implications for fisheries management. – ICES Journal of Marine Science, 64: 346–356. The commercial fishery for sea bass (Dicentrarchus labrax) around the coasts of England and Wales developed rapidly in the late 1970s and 1980s and, by 1986, involved some 3000 fishers using more than 2000 boats to make landings worth £3–4 million. The estimated annual catch of sea bass by sport-anglers was similar to the level recorded from the commercial fishery. Sampling of landings showed strong evidence of growthoverfishing in many areas, and a package of technical measures was introduced in 1990 aimed at improving the exploitation pattern and yield per recruit. This paper describes how data collected through a fishery and biological sampling programme around England and Wales over the period 1985–2004 have been utilized in a separable catch numbers-atage model to assess the dynamics of sea bass stocks in English and Welsh coastal waters. The model output shows that recruitment improved during the 1990s and that moderate levels of fishing mortality allied to an exploitation pattern that largely avoids juvenile fish have enabled the bass population and its fishery to develop sustainably. Results are discussed in relation to model development and management of the bass fishery. Keywords: assessment, fishery development, management, sea bass
Received 1 August 2006; accepted 2 November 2006; advance access publication 18 December 2006.


Around England and Wales, sea bass are primarily exploited by Introduction commercial fishers using a variety of fishing methods in small Material and methods Results boats close to shore and by anglers operating from boats and the Discussion shore. Sea bass <32 cm in total length appear to remain in the References inshore nursery areas to which they are recruited as post-larvae (Pawson et al., 1987; Jennings and Pawson, 1992), whereas fish >36 cm tend to emigrate from these areas and disperse widely around the coast (Pickett et al., 2004). Once bass mature, they adopt the habit of migrating between summer feeding grounds and offshore prespawning and spawning areas to the south and west, where they have been increasingly targeted by French midwater pairtrawlers since the early 1980s and more recently by British vessels (ICES, 2002). Partly because of the high prices offered for the species, the commercial bass fishery developed rapidly in the late 1970s and 1980s, and annual landings into the UK reached more than 1000 t in 1983 and 1984 (Pickett, 1990). With an estimated first sale value of £3–4 million in 1986, the landings accounted for a high proportion of the earnings of around 400 full-time commercial fishers using 270 boats, and about 2500 other fishers using more than 1800 boats took bass as a valuable bycatch. Dunn et al. (1989) estimated that some 24 500 sea-anglers fished regularly for bass in the UK in 1986/1987 and, because of its good eating qualities and high market value, most sport-anglers retained some of their catch. The estimated annual catch of bass taken by anglers in 1987 (Dunn et al., 1989) was 415 t, compared with landings of 630 t estimated for the commercial fishery. Biological sampling of bass landings into England and Wales showed that the exploitation pattern had been shifting towards younger ages in the early to mid-1980s and that recruitment to the important fishery in the eastern English Channel (ICES Division VIId) was at a length of 32–36 cm, owing to local fishing patterns targeting bass 3– 5 years old in and around harbours and estuaries (Pawson et al., 2005). Yield-per-recruit analyses showed strong evidence of growth-overfishing in many areas and indicated that yields in the southern North Sea (ICES Division IVc) and around Devon and Cornwall (VIIe and f) would increase if the size at first recruitment to the fishery was delayed to around 45 cm total length (Pawson and Pickett, 1987). As a consequence, the management priority was to direct the bass fishery away from juvenile fish, i.e. to improve the exploitation pattern. In 1990, a management package was introduced in England and Wales, which made it illegal to land sea bass <36 cm long, banned mesh sizes (65–90 mm) at which juvenile bass were most vulnerable to capture by enmeshing gears, and prohibited fishing for bass from boats in 34 designated nursery areas. Pawson et al. (2005) showed that these technical measures increased protection of juvenile bass and helped safeguard the stock fished close inshore by small boats, and that their success was based on sound science and an understanding of the potential implications for the UK fishery.


The UK's strategy for bass conservation in the late 1980s did not include direct controls on the level of fishing for bass. Effort limitation and catch quotas were considered inappropriate in view of doubts about the efficacy of these types of control in such a fragmented multispecies fishery that employs mainly small, inshore boats, but also because there were insufficient assessment data on which to base quantitative recommendations. Nevertheless, there remained a requirement to follow the development of the bass resource, particularly in terms of spawning potential and recruitment in relation to exploitation rates and climate change (Pawson, 1992). Here we describe how a multi-métier, fully statistical, separable catch numbers-at-age model (Methot, 1990) was applied to 12 data sets for three fishery métier groups (i.e. trawls, nets, and lines; within which the species composition in the catch of a particular gear type is relatively consistent) in four stock areas (IVb and c, VIId, VIIe, VIIa, f, and g; Figure 1) for which sufficient biological sampling information was available over the period 1985–2004. These stocks are not biological entities, because there is likely to be mixing between them at some life stages (for example, through spawning migrations, Pawson et al., 1987; larval drift, Jennings and Pawson, 1992; and emigration of preadults, Pickett et al., 2004), but the bass populations within each area are exploited by relatively separate fisheries (Pawson et al., 2007). The model's outputs provide an evaluation of the dynamics of sea bass stocks and their fishery in English and Welsh coastal waters, and we discuss the implications for further model development as a source of quantitative advice on management of sea bass fisheries.

Figure 1. The sea areas (ICES divisions) in which stocks of sea bass have been assessed, and the location of the Solent pre-recruit survey area.

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Material and methods

Estimating catch and effort Introduction The sea bass is not a total allowable catch (TAC) species, and a Material and methods Results considerable proportion of the UK catch is taken by commercial Discussion small-boat fisheries and by the recreational sector, landing into References minor ports and beaches, and is often sold directly to restaurants or small merchants. As a consequence, landings are under-reported in the official statistics and not well (if at all) sampled through the usual market system for the main commercial quota species, which may record as little as 20% of actual landings of bass in the UK (Pickett, 1990; ICES, 2004). The best estimates of annual catch and effort have been obtained by integrating official statistics derived from landings declarations and local market sales at major ports with those from a voluntary logbook scheme administered by Cefas for the England and Wales <10 m fleet since 1984 (Pickett, 1990; Pickett et al., 1995). The scheme's design takes into account the characteristics of the fisheries that take sea bass, and provides daily catch records from a sample of 34–65 vessels each year. These are raised to the numbers of active vessels likely to be used to catch bass, derived from a bi-annual regional fleet census, to provide estimates of total landings, stratified by gear, boat type (full time, directed at sea bass in season, or part time and number of crew) and ICES division. Owing to the wide range of gears used, up to three per boat in one day, it is not possible to derive accurate gear-specific effort, and a distinction has also to be made whether the gear is targeted at bass. Therefore, the most reliable measure of effort for the UK <10 m fleet is the boat day, when bass is a target species (Pickett, 1990). Effort data (days on the ground) for UK >10 m vessels landing bass into England and Wales are obtained from the Defra Fisheries Activity Database. Good effort data are available for gillnets and longlines from 1985 onwards, for commercial rod and line and handlines from 1986, and for midwater (pelagic) trawls since UK vessels entered the fishery in 1995. The data for other trawling métiers are less robust. Catch rate [catch per unit effort (cpue)] series, expressed in kilogrammes of bass per boat day by gear type, are available separately for >10 m (mainly trawlers) and <10 m vessels in England and Wales (bass landings into Scotland or Northern Ireland are negligible) from 1985 to 2004 in four stock areas: ICES Divisions IVb and c, VIId, VIIe, and VIIa, f, and g. Limited discard data are available for angling and commercial handlines and, although there is no reliable information on discards for other métiers, bass discards are thought to be either insignificant or to survive on being returned to the water. Biological sampling The Cefas sea bass logbook scheme has been run in conjunction with a biological sampling programme to reveal the age structure of catches within each métier. Lengths of bass in commercial landings are measured from either the whole landing or samples of >50 fish from each market size category, against targets set by métier group (i.e. demersal trawls, gillnets, lines, and pelagic trawls). Half-yearly (quarters 1 and 4, non-growing period; quarters 2 and 3, growing period) length distributions by métier group and stock area are then raised to equivalent total landings. In each stock area, length-stratified "all


gears" scale samples of at least 150 fish are used to provide corresponding age data, stratified to ensure that half-yearly age–length keys (ALKs) contain a minimum of five samples per 1 cm total length class across the available length range. Where sampling is sparse (usually because landings were made outside the main port market system), annual ALKs were used. Data on length and age obtained from recreational catches of bass taken by charter-boat skippers and boat anglers contribute some 20–30% of the "lines" métier group (it is assumed that selectivity patterns of commercial and recreational hook-andline boats are similar). Good length and age composition data are available for the main métier groups in Divisions VIId and VIIe from 1986 to 2004, with the exception of lines in 1989. In the North Sea, sampling was largely confined to IVc, where gillnets have been well sampled since 1987 and lines since 1988, although demersal trawl catches have been poorly sampled throughout (the best was just 154 lengths during 1994). Division VIIa has generally been poorly sampled and, because there appears to be a single west coast stock of sea bass (Pawson et al., 1987, 2007), the data have been combined for assessment purposes with those of VIIf and g, where sampling has been good since 1988 (except for lines in 1989 and trawls and nets in 1992). Over the period 1985–2004, annual UK sampling in each stock area averaged >500 age samples (all métiers combined), which appears to be sufficient for assessment purposes (with reference to the quality of model output). Although landings by UK vessels in the winter offshore pelagic fishery in Divisions VIId, e, f, and g (mainly VIIe) were well sampled between 1999 and 2004, French vessels account for the majority of landings from this fishery, for which age and length data are not available for most of the time-series. Moreover, these fisheries are considered to exploit a small but unknown proportion of the various inshore sub-stocks (Pawson et al., 2007), chiefly the older ages. Therefore, the offshore fishery has not been considered in these assessments. Sex ratio, maturity, and length–weight conversion factors are derived from biological sampling carried out between 1982 and 1990 (Pawson and Pickett, 1996). In the sea areas covered by these assessments, the biological parameters of sea bass are relatively consistent: male bass mature at a length of 31–35 cm, aged 4–7 y, and females at 40– 45 cm, aged 5–8 (proportion mature at age 3, 0.03; 4, 0.23; 5, 0.43; 6, 0.57; 7, 0.9; 8+, 1.0). The data for the period 1985–2004 were compiled for the four stock areas, and they include age–length distributions, mean fish weight by age and by year (converted from length), and annual age compositions by métier group. Recruitment Fishery-independent abundance indices for pre-recruit bass are available from a trawl survey that covers the main nursery areas for bass in the Solent (Division VIId; Figure 1), in which a standard grid of 35 stations is fished over a 4-d period at the same part of the tidal cycle (falling Springs) in May and September each year (Pickett et al., 2002). The same commercial inshore fishing vessel has been chartered since 1983 to fish with a highheadline trawl of standard pattern and size and with a mesh size of 80–85 mm in the wings and 60 mm in the codend, the latter fitted with a fine mesh (5 mm) liner. The same extent of ground is fished at each station each year, at a towing speed equivalent to 5–

7 km h–1 over the ground, and the duration of the individual tows varies from 10 to 20 min, depending on the strength of the tide. Numbers of each year class caught at ages 2, 3, and 4 in up to six successive surveys are used to derive abundance indices for sea bass (Pawson, 1992). Model basis The assessment model is based on the stage 1 stock synthesis framework of Methot (1990) and explicitly uses effort and catch numbers-at-age information, the sum of landings by métier being equal to the total landings. In this implementation, each métier is treated as a separable sub-model, assuming constant catchability-at-age across years, and a linear relationship between effort f in year t and the instantaneous rate of fishing mortality F for that métier. Catchability q is the proportion of the fully selected population taken by one unit of effort in a métier, and selectivity s is the age-dependent scalar to account for less than fully selected ages for each métier (relative to 1.0 for the age with the largest catchability). Unlike the application of Methot (1990), no functional relationship is assumed between age and selectivity, which is determined independently for each métier m and age a. Thus,

(1) Total mortality Z is the sum of the Fs-at-age of all métiers plus natural mortality (M; assumed to be 0.1, given the longevity of bass to 20 + y), and the predicted catch numbers-at-age (C) for a métier in a year is given by

(2) where N is the number of fish at age a in year t in the population. Catch numbers-at-age are assumed to contain sampling variability, so observed and predicted catches-at-age are not expected to match exactly, although age determination is assumed to have no errors. The number of fish in the plus group (age 12 and older) is calculated as the sum of the survivors from the previous year's plus group and the survivors of the 11-year-olds the previous year. Selectivity for the plus group is assumed to be the same as that for 11-yearolds.

Other parameters estimated by the model are recruitment for each year and the numbersat-age in the first year, so numbers-at-age in the following year can be calculated from

(3) Landings (L, assuming no discards) by métier are given by

(4) where W is the weight-at-age a in year t. As with the catch numbers-at-age data, landings are assumed to contain normally distributed stochastic variability. Landings information is used in addition to the catch numbers-at-age not only to better scale the stock numbers, but also to allow for the inclusion of information for years where catch numbers-at-age information is not available for a given métier. In those cases, predicted catches-at-age are summed and expressed as landings, taking into account the differences between the expected and observed landings. Residuals of landings are minimized in the same way as residual catches-at-age. The model reverts to a stock-reduction model in order to bridge the gap in stock numbers-at-age, thus allowing a much longer time-series to be used and giving better information regarding trends in stock dynamics than that would be possible with either model separately. The model estimates selectivities, catchability, recruitment, and numbers-at-age in the first year. The numbers of parameters estimated by the model are 56 for each of Divisions IVb and c, VIId, and VIIe, and 60 for VIIa, f, and g, because it includes one additional age. Parameter estimation is implemented in AD Model Builder (Fournier, 1996), utilizing the auto-differentiating functions to maximize the Gaussian likelihood based on the differences between observed and predicted catch numbers and landings, wherein the catch numbers-at-age carry the greater weight in the analysis because they better inform on stock numbers-at-age. In addition to maximizing the likelihood based on catch-at-age and landings, it was found necessary to constrain the model to a realistic range of fishing mortalities. As with most statistical catch-at-age models, F and recruitment are confounded, which frequently leads to high biomass estimates and low F, or vice versa. To attain model convergence, an additional constraint was implemented as a penalty on F, using the sum of squares on the deviation of the average F (ages 4–6 over the whole time-series) from the average F estimated by catch curves for the same ages and period, multiplied by a scaling multiplier.

The suitability of the chosen scaling multiplier was evaluated from the proportion of the penalized likelihood attributable to the F-penalty, and the ratio of the landings and catchat-age squared deviances, given the data. Estimates of spawning-stock biomass (SSB) were calculated based on the estimated numbers-at-age a in year t assuming no temporal trends in the proportion mature-at-age p, but weight-at-age W was variable across years:


Top Introduction Material and methods Results Discussion References

Fleet size The numbers of UK vessels involved in fishing for sea bass in each stock area are estimated every 2 y from a fleet census carried out since 1985 (Table 1). In Divisions IVb and c, vessels fishing for bass peaked in 1994 (at 451), fell in 1995 and 1996, then rose again to 625 by 2004. In VIId, the numbers of vessels fishing for bass also peaked in 1994 (at 786), fell in 1995 and 1996, then averaged 416 between 2000 and 2004. In VIIe and h, the bass "fleet" declined to a low of 202 vessels in 1991, then rose steadily to 584 by 2004; and in VIIa, f, and g, the fleet increased, with some fluctuations, from 491 vessels in 1985 to an average of 875 between 2000 and 2004. Between 20 and 40 French midwater trawlers have targeted prespawning and spawning bass in the winter offshore fishery in VIIe each year since the early 1980s and, from 1995, up to five pairs of UK vessels have also participated in that fishery. Overall, the number of UK boats fishing for sea bass peaked in 1994 (2282) and again in 2002 (2328). Table 2 shows the bass fishing effort associated with these UK vessels, estimated from the effort census and the days fished for bass by boats sampled through the logbook scheme.

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Table 1. Mean numbers (and annual range) of UK boats fishing for bass and numbers of bass sampled for length and age each year, by sea area, 1985–2004.

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Table 2. Nominal fishing effort (days fished, by gear) by UK (England and Wales) vessels fishing for sea bass, 1985–2004.

Catch, effort, and cpue The "best estimates" of commercial bass landings in England and Wales, derived by combining the official statistics and estimates derived from the Cefas voluntary logbook scheme for each métier/year/area (and choosing the higher, if two estimates were available for the same combination), were between two and six times larger than those recorded in official statistics between 1985 and 2004. Landings of bass into England and Wales remained around 600 t between 1985 and 1992, rose rapidly to 2200 t in 1994, then fluctuated between 1050 and 1900 t (mean around 1500 t) until exceeding 2200 t in 2004 (Table 3). During that period, landings were mainly from netting and line métiers, whereas UK pairtrawlers targeting bass in winter and spring contributed some 5–13% of total landings between 1996 and 2004. French landings from the same fishery are also shown in Table 3. Estimates of recreational catch and effort for 1986/1987 (Dunn et al., 1989) and 1992/1993 (Dunn and Potten, 1994) show similar values (around 415 t), which suggest that commercial landings account for most of the variability in the fishery.

View this Table 3. Best estimates of sea bass catch (t) for UK inshore fisheries and UK and French offshore pair-trawls fished in ICES Divisions IVb table: [in this and c, VIId, VIIe and h, and VIIa, f, and g, 1985–2004. window] [in a new


Total international landings of sea bass from the southern North Sea and eastern English Channel (Divisions IVb and c, and VIId) were relatively stable at around 500 t over the period 1985–1990, peaked at 1900 t in 1994, and fluctuated between 1210 and 1824 t until 2004 (Table 4). During that period, landings of bass by English vessels constituted some 50–60% of the total until 1999, when the proportion fell to 30–40%, with France accounting for most of the balance.

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Table 4. Nominal landings (t) and additional UK catch (according to the Cefas logbook scheme) of bass by country and stock area, 1985– 2004.

In the Western Channel and Western Approaches (Divisions VIIe and h), total international landings of bass fluctuated between 260 and 470 t over the period 1985– 1993 (except for 980 t in 1987), rose to around 1100 t between 1996 and 2002, then to 1600 t in 2003 and 2004 (Table 4). English vessels generally accounted for between 30 and 40% of the annual landings. On the west coast of England and Wales (Divisions VIIa, f, and g), total international landings of bass fluctuated around 210 t over the period 1985–1992, peaked at 854 t in 1994, averaged 500 t until 2003, and rose to 800 t in 2004 (Table 4). English vessels generally accounted for some 90% of the annual landings. Note that there has been no commercial bass fishery in the Republic of Ireland since 1990 (ICES, 2002). Table 5 gives the cpue (kg d–1) of bass by gear in the English and Welsh fisheries for the years 1985–2004. Most series show a declining trend from 1985 to 1992, followed by a strong increase to generally high but fluctuating catch rates between 1994 and 2004. These biomass indices suggest that the production of bass around England and Wales remained higher in the mid-to-late 1990s and early 2000s than in the late 1980s.

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Table 5. Catch of sea bass per unit of effort from the Cefas logbook

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scheme for inshore England and Wales fishing vessels.

Recruitment The abundance indices derived from the Solent survey in VIId are compared with the modelled recruitment estimates for the four stocks shown in Figure 2. The model estimates of recruitment are thought to be the most reliable in the year range indicated between vertical lines shown in Figure 2, which differ in position because of the variation in data availability and the age at first recruitment among the four stocks. Strong year classes are indicated for all four stocks in 1989, and for three of the stocks in 1995, 1997, and 1999, whereas the 1985 and 1986 year classes are weak in all areas and the 1984 and 1996 year classes weak in three areas. Overall, recruitment has been higher from 1989 onwards in all areas. The pattern of relative year-class strength over the period 1984– 2002 is similar among all four stocks, whilst that for VIId closely matches the Solent index (located in VIId). Although the survey indicates the strongest year class of the timeseries in 1983, this is not evident in the recruitment estimate for any of the four stocks. Bass were exploited heavily at ages 3 and 4 in the early 1980s (Pawson et al., 2005) and, as the model implies low selectivities for these ages and catchability is assumed to be constant, model estimates of recruitment early in the time-series are not reliable.

Figure 2. Recruitment estimates for sea bass at age 3 in Divisions IVb and c, VIId, VIIe and h, and VIIa, f, and g, and abundance indices (age 2–4) derived from the Solent survey in VIId, 1983–2001 year classes (black bars). Estimates of recruitment are most reliable in the year range indicated between vertical dashed lines.

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Landings Model estimates of landings for the English and Welsh inshore fisheries are generally consistent with those observed, as one might expect given that it is part of the likelihood function used in the model fitting (Figure 3). The model, however, consistently underestimates landings for all stocks over the period 1987–1993, most obviously in VIIe (where the offshore fishery mainly takes place). Overall, it is apparent that bass landings by UK vessels increased considerably from 1993 on, and remained higher, on average, than in the mid-to-late 1980s.

Figure 3. Model estimates (black bars) and observed (open bars) landings (t) of sea bass in English inshore fisheries in IVb and c, VIId, VIIe, and VIIa, f, and g, 1985–2004.

View larger version (22K): [in this window] [in a new window] [Download PowerPoint slide] Fishing mortality Because the model assumes a linear relationship between fishing mortality and effort exerted by the métiers that exploit a particular stock, estimated trends in F differ between the stocks along with fluctuations in the effort of the different métiers (bearing in mind that some métiers have a higher catching efficiency than others per boat day). Model estimates for IVb and c indicate levels of F between 0.1 and 0.2 prior to 1991, a rapid increase to 0.5 in 1993, then lower levels of around 0.2–0.4 since 1995 (Figure 4). A similar pattern is evident for VIId, F averaging around 0.25 in the mid-to-late 1980s, increasing sharply in 1990 and 1991 to around 0.4, then declining again in 1995 and 1996 to fluctuate around 0.2. Model estimates of F for VIIe have been more constant, at around 0.2 for most of the period 1985–2004, rising to peak at 0.4 in 1997, then returning to previous lower levels in 1998. There has been a slowly increasing trend in F estimates for VIIa, f, and g over the time-series, from around 0.15 to around 0.25, with some minor fluctuations including a peak at nearly 0.5 in 2000. These patterns in F are consistent with the changes in the numbers of vessels fishing for bass in the four areas.

Figure 4. Model estimates of fishing mortality (y–1) on bass in IVb and c, VIId, VIIe, and VIIa, f, and g, 1985–2004.

View larger version (16K): [in this window] [in a new window] [Download PowerPoint slide] Spawning-stock biomass (SSB) The model indicates that the SSB of bass around the coasts of England and Wales generally increased between 1985 and 2004, although the pattern and magnitude of the increase differed between areas (Figure 5). Putting aside the SSB estimates in the early part of the time-series (for which estimated numbers-at-age in the population are less robust owing to sampling and model uncertainties), the IVb and c spawning stock appears to have increased slightly from about 600 to 800 t. In Division VIId, SSB was relatively stable at around 1500 t until 1998, after which it increased linearly to >3000 t in 2004 (although this continued expansion is associated with some uncertainty in effort levels towards the end of the time-series). SSB in VIIe quadrupled, from around 500 t in 1985 to >2000 t in 2003/2004, though note the absence of the offshore fishery's catches in this assessment. SSB in VIIa, f, and g appears to have followed a pattern similar to that in VIIe and h, rising from 600 to 800 t prior to 1993, then to around 1200 t in the years 1997–2002, after which it appears to have continued to increase.

Figure 5. Model estimates of SSB (t) of sea bass in IVb and c, VIId, VIIe, and VIIa, f, and g, 1985– 2004.

View larger version (13K): [in this window] [in a new window] [Download PowerPoint slide] Selectivity The estimated gear selectivity patterns for the different stock areas are similar (Figure 6), remain relatively high at the older ages in the western areas (Divisions VIIa, f, and g, and VIIe), reduce rapidly above age 8 in Division VIId, and diverge most strongly in Divisions IVb and c, particularly for nets and lines. In general, selectivity increases strongly at ages 4 and 5, is at its maximum at ages 5–8, depending on area and gear, and is most consistent in the netting métier between areas (where gears, mesh sizes, and types of ground fished are probably most uniform) and most varied for the line métier. Other than in Division VIId, the netting and line métiers appear to have complementary selectivity patterns; these gears often being used by the same vessels to exploit bass in different habitats.

Figure 6. Model estimates of selectivity-at-age for sea bass in trawls, nets, and lines in IVb and c, VIId, VIIe, and VIIa, f, and g, 1985–2004. Values for catchability q are given to indicate the relative potency of each gear.

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Top Introduction Material and methods Results Discussion References

Model performance This study provides the first formal quantification of the dynamics of bass stocks around the coasts of England and Wales, using an assessment model that was developed taking into account the data availability, the known characteristics of the fisheries, and the biology of sea bass. A separable model using gear-dependent effort data to scale fishing mortality (Methot, 1990) is ideally suited to examining the trends in bass stock dynamics, because there are large fluctuations in effort between métiers, and reliable effort data are available through the Cefas logbook scheme. Although model convergence was attained for the IVb and c and VIIa, f, and g stocks without a constraint on F, it was necessary to implement such a constraint via a penalty function for both VIId and VIIe stocks. Although this represents only a small portion of the penalized likelihood (2.41 x 10–7 for VIId, 0.011 for VIIe), it implies that the model has some difficulty in dealing with the data. The raw data show clear year-class signals, so it is likely that the difficulties arise from process error or parameter confounding. Methot's (1990) model requires less precision in the catch numbers-at-age information than most age-based models, because it assumes error in the catch-at-age matrix, and the inclusion of landings data and the large number of year classes in bass fisheries tend to smooth out variability. However, the model fits selectivities independently at each age and assumes no relationship between ages (other than between age 11 and the plus group). Thus, strong year classes, changes in the character of each métier, or changes in fishery regulations will all contribute to noise in the selectivity curves. The model's assumption that catchability-at-age is constant for each métier through the assessment period (1985–2004) is questionable for métiers such as netters, which specifically target strong year classes (e.g. 1983, 1989) in the fishery (Reis and Pawson, 1992). The implementation of a management package in 1990, aimed at raising the size of recruitment by the equivalent of at least 1 y growth (Pawson et al., 2005), might be expected to have reduced the fisheries' selectivity for younger sea bass. However, analysis of separate cpue trends pre- and post-1990 indicated a small effect in comparison

with the overall contrast in the data over the time period 1985–2004. The occurrence of three consecutive weak year classes from 1984 to 1986 led to an upward shift of the exploitation pattern in many bass fisheries before 1990, because they directed effort at the relatively strong 1982 and 1983 year classes. As a consequence, the selectivity values for fish aged 3 and 4 in the period 1985–1989 were quite low, and the model had little difficulty in coping with any violation of the constant selectivity assumption for at least two of the stocks. Trawl gear used inshore is most likely to take sea bass as bycatch, so that its selectivity is generally attributable to gear effects in relation to fish size rather than to fisher behaviour. As a consequence, the catching characteristics of that gear group are likely to be the most consistent between areas, and trawl selectivities were most similar, although they did decrease after age 8 in the eastern English Channel and the North Sea. The two peaks in selectivity at ages 5/6 and 8/9 for the net fishery in VIIe (Figure 6) are likely a consequence of the use of gears with different mesh sizes to target small bass inshore and larger fish offshore within this métier group (Pickett and Pawson, 1994). Whereas the patterns in fishing mortality and recruitment revealed by this study are consistent with the evidence provided by the Cefas logbook scheme and the fisheryindependent recruit survey, the model links effort directly to fishing mortality, and forward-propagating virtual population analyses, such as the stock synthesis approach, are generally very good at estimating strong and weak year classes (Walters and Martell, 2004). There are, however, potential problems with the scaling of F, and hence biomass estimates. Two other fisheries take bass and are not included in the assessments. One is the (mainly French) winter offshore pelagic trawl fishery mainly in VIIe, although it accounts for a relatively small proportion of sea bass that are otherwise exploited in the UK inshore fishery (Pawson et al., 2007), and there is insufficient age composition data available for it to be included in the models. The other fishery is by shore-based recreational anglers, which may be a substantial contributor to bass mortality inshore but was not monitored in any of the stock areas. Although the recreational fishery might be expected to take largely younger, smaller fish, so influencing estimates of recruitment to the commercial fishery, these are consistent between areas and highly correlated with fishery-independent indices of bass abundances in the Solent nurseries. We presume, therefore, that the effect of recreational fishing has been relatively consistent over time. The pelagic fishery, on the other hand, largely takes older fish (>6 y) that are probably under-represented in the catch numbers-at-age matrix. Although omitting these constituents of the catches clearly influences scaling of the biomass estimates, we suggest that the model reflects reasonably the dynamics of the inshore stock both in stock numbers and in F, but that the estimates of the absolute size of the spawning stock could be subject to error. Migration between stocks must also be taken into account (Dunn and Pawson, 2002). Tagging information suggests that the migrations at ages >4 are substantial, particularly between Divisions VIId and VIIe and between VIIe and VIIf, g, and a as late juveniles recruiting to the fishery (ages 4–6; Pickett et al., 2004) and as migrating pre and postspawning adults (Pawson et al., 1987, 2007). These studies also suggest that the migration pattern appears to have changed over time, with proportionally more adult fish migrating between IVb and c and VIId and VIIe prior to the mid-1980s than in recent

years. A future development would be to incorporate several stocks into a single model capable of parameterizing the extent of migrations between them and changes in spatial and temporal exploitation. Methot's (1990) stock-synthesis framework is expandable and flexible with regard to its implementation and incorporation of additional data sources, and it also allows for a realistic evaluation of the error structures, based on both the dynamics of the stocks and the sampling design used to observe them. Therefore, uncertainty regarding age and length sampling can be included in further model development, using multinomial likelihood penalty functions in the more advanced models (stage 2/3 of Methot, 1990). Better estimates of absolute values for stock size/production and mortality might be obtained if the confounding between the two parameters was reduced through reparameterization of the model. One possibility is to use the Solent recruitment index in the model to better tie down recruitment strength. For the present paper, however, it was deemed necessary to use the fishery-independent Solent recruitment index to evaluate the model's suitability for assessing the fishery and to determine whether the Solent index could be used for stocks other than in VIId. Stock dynamics The model estimates for F and SSB trends are consistent with our understanding of sea bass stocks and fisheries in the four stock areas, in particular, the boost to production and the increase in F in all areas associated with the recruitment of the very strong and fastgrowing 1989 year class at ages 4 and 5. In addition, the recruitment patterns generally match the independently derived Solent recruitment index. As recruitment to the fishery corresponds to the age at first maturity, year-class signals based on catch numbers-at-age per unit of effort provide a good indication of recruiting year-class strength, both to the fishery and to the spawning stock. The respective SSB values estimated for the assessed "stocks" are based on the premise that the bass populations in Divisions IVb and c, VIId, VIIe and h, and VIIa, f, and g represent separate entities. Division VIId appears to hold the largest stock, estimated at around 3400 t in 2004, followed by VIIe and h, and VIIa, f, and g at 2000 and 1500 t, respectively, with IVb and c estimated at around 1000 t. For the reasons discussed above, these SSB values are better taken as indices of abundance rather than absolute values, and they are not entirely additive, because we know that populations of sea bass around the UK mix considerably. In addition, by not including the estimates of the bass catch by the UK recreational fishery (around 415 t in both 1987 and 1993) or of the offshore fishery in the analysis, biomass is probably underestimated. Nevertheless, the long-term trends and levels of F over the time-series are consistent between stocks, and it appears that the biomass of adult bass around the coasts of England and Wales approximately doubled between 1985 and 2004. The most substantial increase in SSB was in the late 1990s and early 2000s, and it is associated with an increased level of recruitment from 1989 on and a coincident range expansion (as far as southern Norway) (J. E. Colman, pers comm).

Directed commercial fisheries for sea bass have been established since the early 1980s in the English Channel and on the west coast of the UK, where they are known to respond quickly to changes in the availability of recruits (Pawson et al., 2005). Therefore, modelled trends in stock biomass are considered to be representative for the populations in these areas. However, other than in the Thames Estuary, there is no tradition of commercial fishing for sea bass in the North Sea (Pickett and Pawson, 1994). Our effort census suggests that the bass fishery on both the English and continental coasts expanded from IVc into IVb in the late 1990s, rather later than the boost to production effected by recruitment of the 1989 year class. It is therefore possible that the uptake of the North Sea fishery in response to stock growth may have been much slower than elsewhere, and that increased local recruitment and production of bass is not so well reflected in landings data collected from the North Sea fishery. Management considerations Although we consider that the separable model is a suitable candidate for evaluation of the dynamics in bass fisheries around the UK, the current model assumes closed populations, which these are not, given the offshore fishery and known migration patterns. Consequently, the present assessments are probably unsuitable for short-term forecasts, setting TACs, or the evaluation of uncertainty. Nevertheless, the output can still be used to give sound precautionary management advice in relation to stock status and sustainability. The results of these assessments and those reported by Pawson et al. (2005) suggest that bass stocks in UK coastal waters are being exploited sustainably, at a moderate level of F and with an exploitation pattern that gives a near maximum yield per recruit, and that there has been an increase in exploitable biomass since the early 1990s. It is interesting that an increase in fishing mortality in 1992 and 1993 preceded the recruitment of the strong 1989 year class in Divisions IVb and c, VIId, and VIIa, f, and g, when biomass estimates were not higher than those in the 1980s. Subsequent levels of F have not increased in line with either recruitment or SSB suggesting, possibly, that the package of management measures introduced in 1990 has been effective in directing fishing activity away from the vulnerable juveniles (Pawson et al., 2005). The estimated recruitment time-series in Divisions IVb and c, VIId, VIIe and h, and VIIa, f, and g show common features regarding year-class strength, indicating that stocks of sea bass around England and Wales are linked biologically (Pickett et al., 2004) and/or that recruitment is controlled by large-scale environmental patterns. Warm or cold summers and winters can have a considerable influence on the survival of 0-group bass through their first year (Kelley, 1986), and strong year classes are associated with higher temperatures (Pawson, 1992). Although there is a link between recruitment and stock, it is driven by increased recruitment levels raising SSB since the early 1990s and is not a conventional stock–recruit relationship. Consequently, it has not been possible to set absolute or even relative F reference points based on stock–recruit considerations. There is, however, no sign of recruitment-overfishing. If the status of bass around England and Wales and the large variability in recruitment strength are representative of the population in Northwest Europe as a whole, it is possible that fisheries could continue to operate sustainably, provided reproductive

success continues to be favoured by climatic conditions (warming) and that recruitment of the resulting strong year classes to the spawning stock is protected by adequate technical measures. Forecasts of the potential changes in yield, however, indicate gains in some areas with a higher size at recruitment to the fishery (at 45–50 cm), although these must take account of the availability of the bass population to the various fisheries (Pawson et al., 2005). The selectivity values for bass suggest that the fishery in the North Sea, for example, is not taking full advantage of the increased productivity of the population and implies that further increases in minimum landing size (MLS) will not necessarily lead to increased catches because bass older than 8 years are less catchable in trawls and nets. Uniquely, for a European marine fish species, sea bass are especially important to inshore artisanal fishers and recreational anglers. The potential effects of increasing the MLS for bass in UK coastal waters have been simulated using selectivity patterns for métiers in the inshore fishery and the underlying stock structure derived from the models presented here (M. T. Smith, pers. comm.), and they have indicated the magnitude and timeframe of costs to the commercial fishery through an initial reduction in catches, as well as potential socio-economic benefits to recreational anglers of an increase in the abundance of medium-sized and large fish (Drew Associates, 2004). Continued development and application of these models will have an important role in monitoring and assessing bass stocks and fishery trends, and in evaluating the likelihood of success of implementing new management measures.

We are especially grateful to Sarah Walmsley (Cefas) for updating and checking the assessment data used in these models of the UK sea bass fishery. The study was funded by the UK's Department for the Environment, Food and Rural Affairs (Defra).

Top Introduction Material and methods Results Discussion References


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