Rapp. P.-v. Réun. Cons. int. Explor. M er, 187: 108-114. 1987 Exceptional Plankton Blooms Conclusions of discussions: Convener’s report M iles Parker Parker, Miles. 1987. Exceptional Plankton Blooms. Conclusions of discussions: Con vener’s report. - Rapp. P.-v. Réun. Cons. int. Explor. Mer, 187: 108—114. This report draws on the papers presented to the meeting, the records of the session rapporteurs, and the considerations of discussion groups that met on the two evenings of the meeting to draw some general conclusions related to the terms of reference of the special meeting. It reflects as accurately as possible the consensus prevailing among the active participants. Miles Parker: Ministry o f Agriculture, Fisheries and Food, Directorate o f Fisheries Research, Fisheries Laboratory, Remembrance Avenue, Burnham-on-Crouch, Essex CM08HA, UK. T h e incidence o f excep tion al blo o m s There are very few long time-series data on exceptional bloom incidence. In assessing such data as are available, distinction needs to be made between oceanic and open shelf waters (A 6 ‘) on the one hand and on the other, coastal and estuarine waters where terrestrial influences may be more marked. Since the mid-1960s, the dinoflagellate G yrodinium aureolum has occurred in E uropean coastal and shelf waters and has given rise to exceptional blooms in many areas and caused fish kills (A9, B7, B8, D2, D8, D9). The causes for its recent upsurge are unknown but it has, however, been suggested (A9) that it is a recently introduced species to E uropean waters; introduced spe cies may thrive excessively in the first years following their introduction. Apart from the special case of G. aureolum, it is more difficult to point to trends in inshore waters of the Atlantic. There has been a southward extension over a long time-scale of Paralytic Shellfish Poisoning (PSP) problems associated with both low and high biomasses of Gonyaulax tamarensis on the east coast of Canada and the U SA , apparently for natural climatic reasons. Fluctuations in the degree of toxicity of shellfish in the Bay of Fundy have been correlated with long time-scale oceanographic and meteorological changes and with the 18-year lunar cycle on tidal mixing activity (C2). In Europe, exhaustive studies failed to identify trends in blooms in the Oslofjord, though changes in species composition have been noted (D l). On the east coast of the United Kingdom, PSP toxin has been recorded each year since 1968, but no trends in toxicity have been observed; shellfish toxicity is known to have occurred in this area earlier in the century. In the Germ an Bight, where blooms of Ceratium spp. and other species have caused concern recently (B6), it is also known that such events have occurred at intervals over the last century. It is worth comparing results from inshore areas o u t side the North Atlantic, such as the Seto Inland Sea of Japan where clear trends in bloom incidence have been associated with trends in organic pollution (B13). Exceptional blooms must be seen against the general background of the normal cycle of phytoplankton bio mass. D ata from the Continuous Plankton Recorder (CPR) (A6) for the Northeast Atlantic suggest a down ward trend in the incidence of periods of higher than normal biomass of diatoms and Ceratium spp. in the last 20 years with an increase since the mid-1970s of the more generalized “Phytoplankton Colour” index. These trends may be superimposed on general downward trends in zoo- and phytoplankton abundance. In summary, there is little evidence in Atlantic waters for any rising trend in bloom incidence, though the data base is very small. Changes in species composition have been observed, though the reason for these changes are unclear. Causal factors Physical factors 'References to meeting papers in this text are given as the programme num ber cited in the Annex to the Introduction (e.g., A l , C3, etc.). 108 Recent research has notably improved the understand ing of the importance of physical factors in bloom in- cidence (A l). The widespread extent of observed changes (A6) in the plankton points to the important role of long-term and widescale climatic and ocean ographic changes, though how these changes might be related to bloom incidence is not well understood. On a smaller scale, areas of high biomass or produc tion are associated with boundaries (pycnoclines and fronts) and upwelling areas (A7). This is also true of inshore areas where the effect of river inputs, in addi tion to tidal forces and other coastal influences on the stability of the coastal waters, may be an important factor, especially in exceptional blooms of diatoms early in the year (B15). Later in the year, as nutrient levels decline, dinoflagellate populations, which are a normal component of the summer plankton, are noticeably high at boundary systems and the incidence of summer storms followed by calm weather is related to the in cidence of exceptional blooms. Interaction of weather (C2) (especially wind) with the sea and sea-sediments are also im portant factors, for example, with respect to resuspension in the water co lumn of resting stages (cysts) of some dinoflagellates from the sea bed (C6). Coastal transport and smallerscale transport processes (e.g., convergences) also have a role to play in ensuring that the organisms reach the correct environment and are not dispersed from it (B l, A4, A5). Biological factors The role of biological factors is clearly important. Full understanding of exceptional bloom events depends on a thorough knowledge of the life cycles of the organisms concerned (B l); such information is often lacking. Among other relevant biological factors, reduced grazing pressure, such as occurs particularly in response to inhibitory properties of some dinoflagellates, may be im portant to the development of large populations of algae. The role of vertical migration (A2) in ensuring both effective nutrient and light utilization and concen tration in surface waters aided by physical mechanisms may be a key to the development of surface “red wa te r” . Some bloom species appear to be well adapted to a changeable light environment. N utrients The role of nutrient enrichment is less clear (B l). Cer tainly, where nutrient levels are high, production will occur somewhere, though nutrient-rich coastal waters are often highly turbid and well mixed, which reduces the opportunity for exceptional growth (A2). Similarly, it is clear that in many cases it is the flux of nutrients rather than their concentration which is important to the development of algal populations, though relatively low fluxes may be sufficient to sustain already existing large populations. Some long-term data sets on inorga nic nitrogen and phosphorous nutrient concentrations are available, but data on fluxes of nutrients are far more limited and the roles of inputs or of recycling through grazers, bacteria, or the sediments are poorly understood. Current understanding suggests that the physical structure of the water and its relationship to the distribution and availability of nutrients is a major fac tor in bloom development (C l). A n th ro p o g en ic factors - hypernutrification and eutrophication In relation to discussion of the effects of anthropogenic influences on the incidence of exceptional blooms, it is also necessary here to define the terms “hypernutrifica tion” and “eutrophication” . Hypernutrification involves substantial and measur able increases in concentrations of dissolved nutrients (which may include inorganic and organic compounds of nitrogen, phosphorous, silicon, iron, and other trace metals and vitamins). Current practice is generally to use measurements of inorganic nitrogen and phospho rous because of their relative ease of measurement. It should be noted, however, that laboratory investiga tions of phytoplankton have shown that a wide range of substances can control growth, including not only sub stances which can be considered as nutrients, but also substances such as toxic metals and materials which control their availability to phytoplankton (B13). Hypernutrification can in theory lead to increased phytoplankton biomass and productivity in regions and at times in which phytoplankton growth is limited by nutrient availability. In freshwater ecosystems, this is referred to as eutrophication and, by analogy, this term has also come to be used in the marine context. If high nutrient levels affect algal populations, they might do so in three ways: (a) by directly increasing overall annual biomass, perhaps by extension of the growth season, (b) by leading to sporadic exceptional outbursts of growth, and/or (c) by changing species composition. The latter effect would be particularly hard to distinguish from natural changes in species com position which occur all the time, for example, such as those noted in CPR data throughout the eastern North Atlantic (A6). As stated earlier, there is little evidence to suggest that actual increases in exceptional bloom incidence (other than G. aureolum) have oc curred in Atlantic waters. Equally, it is hard to find any evidence for an overall measurable increase in biomass in these waters, though the data available are very lim ited. If such effects were to occur, they would be most likely to happen in areas where the spatial and temporal scales of enrichment were such as to allow time for algal growth, i.e., in enclosed embayments or estuaries or in coastal areas affected by major river inputs. Even in these areas it might be difficult to separate nutrient 109 effects from purely physical ones, (e.g., the effect of freshwater input on stability). Such areas might include the New York Bight, the G erm an Bight and southern North Sea, the Kattegat (B7, B8), parts of the Baltic (B4, B5), and areas such as the Oslofjord (D l). While nutrient inputs compared with the available ‘pool’ are undoubtedly high in these areas, it is still not easy to point to clear examples of eutrophication. The availabil ity of phosphate in summer may allow increased pro duction by N-fixing blue-green algae in the Baltic, and the high level of production of Phaeocystis (D5, DIO) in the southeastern North Sea may point to a case of eutrophication. Equally, if elevated nutrient concentra tions occur, exceptional meteorological and hydrographic conditions may permit outbursts of growth (B6). The case mentioned earlier of the Seto Inland Sea (B13) raises a further question in relation to eutrophica tion and especially the potential role of anthropogenic influences. T here are many field and laboratory data that point to the significance on the one hand of vitamin ( D l l ) and growth factors in determining species com po sition and on the other, to the role of natural organic chelating agents in controlling the toxicity and nutri tional availability of metals on algae. Recent research suggests that the metal-complexing compounds in sew age effluent may be chemically similar to naturally oc curring humic and fulvic acids in natural waters. Since dinoflagellate growth is often stimulated by the addition of soil extracts or similar enrichments containing humic acids, it is possible that pollution of coastal areas could enhance bloom development due to input of organic rather than to inorganic compounds. The Seto Inland Sea experience suggests that bloom incidence there was at least partially related to organic inputs as well as to nutrient inputs, but there are few data available on either the natural levels of such substances or the levels of inputs and fluxes in Atlantic waters. In summary, on the basis of data on exceptional blooms, there is little evidence for the existence of large-scale eutrophic effects in North Atlantic waters. However, such effects are more likely to be observed through study of annual phytoplankton biomass as a whole, rather than exceptional bloom incidence. Suit able data sets are not available for most areas; in partic ular, biological data and long time-series data sets are lacking. A ttention should be addressed especially to the areas identified above and to other estuarine areas re ceiving large natural or anthropogenic inputs of nutri ents, w hether organic or inorganic. Further research is needed on the role played by organic enrichment. Predictability o f “e x c e p tio n a l” even ts Understanding of the oceanography of bloom incidence is now sufficient to identify areas in which there is a high probability of bloom occurrence, either on the basis of 110 long time-series data o r a knowledge of the physical structure present (A l, A6). Hydrographic or physical oceanographic variables and phenom ena associated with blooms and related events, e.g., hypoxia, have been modelled in a preliminary m anner in certain re gions. These models are capable of predicting condi tions which can lead to bloom circumstances and events. Relatively simple biological models (B l), incorporating data particularly on grazing rates and the behaviour and physiology of bloom-forming species and the dominant nutrient flux, can also be used to predict the probability of bloom events in given areas. Prediction of the group of species (diatom, dinoflagellates, flagellates) likely to dominate the bloom is feasible on the basis of the d e gree of turbulence/stability of the water column, though it is more difficult to make predictions as to particular species dominance since information on the life histo ries and specific growth rates of many species is lacking. However, these models may be extended to prediction of harmful effects either through estimation of the ef fects of biomass on oxygen consumption or through the relationships which are beginning to be established be tween toxicity and the nutrient status of the cell. Such predictability is purely probabalistic since the actual concentrations or values of physical, chemical, and biological factors that lead to a particular bloom cannot be predicted absolutely through the application of such models. Probabalistic prediction, however, is probably adequate for overall environmental manage ment purposes and should be used at this stage as a means of directing resources to the longer-term mon itoring of areas theoretically likely to have a higher incidence of blooms. O n the other hand, mariculture and shellfish fishery managers require prediction of ac tual events over much shorter time and smaller spatial scales; this can only come from a system of monitoring at the areas to be protected (C l). Predictive capabilities could be im proved by: (a) an improved knowledge of the biology and life his tories of blooming species; (b) collection of long time-series data on phytoplank ton selected species composition and biomass, in areas identified above. One source of historical data would be the recent (100 years) sediment rec ord of cysts and pigments which could be obtained from areas of high natural sediment accumulation and low bioturbation (C6). Satellite and aircraft remote sensing may provide useful support for field programmes, especially in synopticity, but their use in gathering long time-series data needs further investigation. A utom ated methods for the con tinuous collection of biological data also need further development. M a n a g e m e n t o f public health problem s related to e xc ep tion al b loom even ts Seafood toxicity associated with exceptional blooms (in the sense defined in the paragraph on “Nutrients” above) has a long history in the N orth Atlantic area (C l). Most reports have related to Paralytic Shellfish Poisoning (PSP) (C5, B9). More recently, Diarrhetic Shellfish Poisoning (DSP) has been reported in E u ropean waters (C3, C4, C5, D9). However, the defini tion and diagnosis of this syndrome are also relatively recent and there are earlier records which may be attrib utable to DSP. PSP and DSP result from the consumption of shellfish which have accumulated toxins produced and contained in certain dinoflagellates. PSP is associated with the species variously referred to as Gonyaulax tamarensis or excavata, while there is some uncertainty about the causative organisms of DSP; D inophysis acuminata ap pears to be the most likely candidate, though certain Prorocentrum spp. have also been brought into question in the past. It is important that these taxonomic and biological issues be cleared up for monitoring and man agement purposes; in particular, D. acuminata needs to be cultured so that its toxicity can be assessed. A t the time of writing, it is clear that PSP is a recur rent problem in some areas of the coastal regions of New Brunswick, Canada (C2, D4) and New England, and the USA (C l) in the Northwest Atlantic, and N or way ( D l), United Kingdom, Spain (C5), and Portugal (B9) in the Northeast Atlantic. DSP has been reported primarily in the N ortheast Atlantic, namely in the N eth erlands (C3), France (C4), Ireland (D9), and the Iber ian Peninsula (C5). Monitoring is an im portant tool for management (C l). Traditionally, monitoring has concentrated on the incidence of toxic shellfish. Different assays for PSP and DSP are available (C3, C4); both are bioassays and, while far from satisfactory, are adequate to provide basic management information on degrees of toxicity. However, there is an urgent and continuing need to update assay methods, to reduce their costs and to make them simpler and quicker to apply. Recent experience with DSP suggests that toxicity may occur quite sud denly and that monitoring of phytoplankton in addition to toxicity, may give valuable forewarning. Unfortunately, monitoring programmes in many ar eas are non-uniform. Since information on shellfish tox icity is often relevant outside the immediate area af fected, there is a need for uniformity with respect to: (a) standardization of criteria for monitoring station selection; (b) intercalibration of assay methods between laborato ries; (c) collection of associated data sets with respect to meteorology, hydrography, and species composi tion of phytoplankton present at the time of collec tion; (d) data presentation and analysis, possibly using agreed com puter formats. In this respect, a coordinated archiving of data would be valuable to improve access and assessment. Faced with toxicity problems, resource managers have options including blanket closure, seasonal clo sure, and closure in response to monitoring results (C l). Safe application of the latter requires an aggressive monitoring programme but has cost/benefit advantages in that it allows maximum resource utilization. The greatest negative impact is witnessed when a monitoring programme is used solely to protect public health (C l, C3, D9). The seafood industry can suffer near-collapse when public alarm is raised over “toxic shellfish” . This can be avoided when the sensitivity of the issue is acknowledged and the public is made aware of the risks, causes/effects, etc., by the prompt issue of balanced factual announcements by the controlling au thorities. Negative “halo” effects include: (a) decline in marketability of seafoods in unaffected regions; (b) decline in marketability of unaffected species within affected regions; (c) decline in marketability of seafoods beyond the time scale of toxicity. At a national and international level, there is a need to review and agree upon accepted quarantine and safety levels. There is also a need for the establishment of rapid communication networks to report on monitoring data and management action. M a n a g e m e n t o f e x c ep tio n a l-b lo o m related problem s in mariculture I n tro d u c tio n Fish kills and sub-lethal effects in mariculture have been particularly associated with the spread of Gyrodinium aureolum in European waters (B7, C5, C6, D2, D8, D13), though several other species have been impli cated, particularly an unidentified ‘flagellate X ’ (B8, D2, D8, D12) and, in 1984, the PSP-producing species G onyaulax excavata (D14). So far, these problems have been limited to the eastern North Atlantic, though G. aureolum has recently bloomed in Canadian waters. Wild, caged, and pond fish, shellfish and marine in vertebrate mortality, or reduced growth due to excep tional blooms have occurred in many E uropean coun tries and have caused substantial economic losses, espe cially in Ireland (D8, D9, D12), Scotland (D2), the Faroes (D14), and Norway (D l). Ill The time is now ripe for a synthesis to be made of the experience of these problems in recent years, so that advice may be formulated for mariculture managers. The following paragraphs indicate the main headings under which such advice should be formulated. There is a need both for fish farmers to be aware of the factors listed below and for central government expertise to be available to provide specialist and site-specific advice, and to train fish farmers in basic techniques. Licence conditions for fish farms would include provisions con cerning the availability of basic equipment and the training of personnel. Predictability - site selection and m onitoring [t is now possible to identify a series of factors which allow risk assessment of sites otherwise potentially suit able for mariculture ( B l, C l , D2). These should be professionally examined before production commences; in general, the fish farmer should gather as much in formation as possible about the local, physical, chem ical, and biological environment. A distinction needs to be made between risks of exceptional blooms occurring in situ close to cages or of the transport of exceptional blooms from offshore areas to cages. Key factors in site risk assessment include: (a) water exchange. High exchange rates will reduce risks of in situ blooms, but may allow transport inshore of offshore blooms; (b) the presence of offshore frontal or upwelling sys tems, which may be seed areas for offshore blooms. (In respect of the first two items, the ideal site is fully mixed and communicates with a fully mixed sea area); (c) accumulation of soft sediments, which may indicate poor circulation; (d) nutrient status of local waters; (e) recorded or surveyed phytoplankton community, including data on PSP or DSP incidence; (f) cyst occurrence in local sediments (C6). Inevitably, there will be some conflict between these factors in individual circumstances. A risk assessment for Scottish waters based on local application of these factors has been carried out by the Scottish Marine Biological Association (D2). Especially in identified high-risk areas, it is important that fish farmers monitor the environment of their cages and any local areas such as fronts which could give rise to problems. There is a need for training of fish farmers in basic techniques, including dinoflagellate identifica tion; courses in the latter have been held in Ireland, Norway, and the U nited Kingdom. (It is important that the cost of such courses should not militate against attendance.) Monitoring methods must be as simple as possible. 112 Offshore events may be monitored using local fisher men to collect observations of discoloured water and water samples. Surface w ater samples may be adequate, though the use of techniques such as the Lund tube to obtain integrated water column samples, is preferred. Initial analysis should concentrate on presence or ab sence of key species; more detailed identification and counting should be limited to high biomass samples. Near-cage monitoring can be simply carried out with a basic set of tools including oxygen and temperature probe, secchi disc, a cheap microscope, and suitable preservatives for algal samples (D8). All fish farmers should regularly make records of tem perature and oxy gen profiles and secchi depth. The latter serves as a support to phytoplankton sampling by indicating changes in water clarity which occur, inter alia, as a result of bloom development. In high-risk areas, farm ers must seek and have available specialist advice and will need to carry out more extensive monitoring. Some techniques for bioassay of w ater quality (e.g ., the oyster embryo bioassay) may be useful in such circumstances (D3), and research to develop other bioassays using sensitive species (e.g., Arenicola) o r bio-chemical m eth ods, is needed. Prediction o f specific events Detailed analysis of w eather patterns, associated with blooms in both offshore and inshore areas, may allow the development of a w eather warning-system similar to that used in terrestrial agriculture for Potato Blight infestation. Small-scale in situ events are inherently more difficult to predict, though statistical records of incidence, w eather patterns, and nutrient conditions will all be relevant. For effective short time-scale prediction, data are needed in real time, both in terms of rapid work-up of samples and of good centralized communication net works which should be government based. Site m anagem ent The importance of good cage husbandry was stressed. While it is not very likely that cages can significantly affect their local environment by inorganic and organic nutrient input, every effort should be made to reduce cage impact (e.g., by avoiding wasteful over-feeding) (D8). Rotation of cage sites allows the seabed beneath cages to recover from the effects of rain-out. Limitation on size of farms according to site conditions is impor tant. General fish health is important in so far as diseased or parasitized fish will more easily succumb to the ef fects of blooms. Conditions suitable for disease or parasitization may also be suitable for in situ bloom forma tion, so health may also be a monitoring tool. In high-risk sites, shore-based installations with al ternative shallow or deep intake pipes are a sensible option. Where young fish are expected to feed on plankton, care should be taken on the timing of stocking in rela tion to the time of development of toxic blooms. In some areas, adequate growth may be attained be tween early winter and the onset of the bloom season. In this strategy, trout are the most appropriate fish to use. M an agem ent options in b lo o m events Cage movement, especially in in situ blooms may be a useful option, but is likely to be impracticable in large farms or in response to large-scale blooms. Feeding should be stopped during blooms as it only increases the stress on the fish (D8). Early slaughtering of fish may allow some recovery of stock value (D8). For this approach to be effective, the species of fish should be m arketable as small young fish (i.e. trout, not salmon) and freezer or processing facil ities should be availabe to avoid sudden m arket gluts. Biological control does not yet appear to be feasible and the use of algicides should be strongly opposed. In both cases, the wrong species may be attacked exacer bating blooms; algicides may be toxic to fish. More experimental research is needed on: (a) deep cages (penetrating the thermocline and allow ing fish access to sub-thermocline water); (b) cage lowering (for the same purpose). Observations that forced vertical circulation of water through cages (using pumped air ventouri systems) re lieves the symptoms of G. aureolum effects require corroboration and further experiment (D8). There is no good theoretical basis for these observations as yet. Appendix Proposals and recom m en dation s o f the m eeting There is an urgent need to improve spatial and temporal coverage of phytoplankton data. In particular: (a) Existing data sets should be assembled and re-in vestigated, particularly to compile time series. (b) Cross-calibration of CPR and other instruments (e.g., fluorimeters) for recording plankton data is needed. (c) Examination of recent sediments from non-bioturbated accumulating areas to establish time series of phytoplankton activity from cyst and pitment records should be carried out (perhaps in conjunction with cur rent ICES sediment pollution studies). (d) The application of remote sensing to these prob lems should be examined in detail, including the use of existing aircraft surveillance operations and aircraft of opportunity. ICES should put pressure on the E u ropean Space Agency to provide an Ocean Colour In strument as soon as possible. With respect to questions of eutrophication: (a) A ttention should be focussed not on bloom inci dence but on the overall patterns of annual biomass. (b) A ttention should be focussed on coastal and estuarine areas in which terrestrial inputs are significant in relation to the “pool” of nutrients in the sea (in partic ular to the New York Bight, the Southern and G erm an Bights of the North Sea, the Kattegat and areas of the Baltic Sea, and areas such as the Oslofjord, and the more enclosed estuaries). (c) Research is needed on the role of organic sub stances in phytoplankton production, including organic nutrients, vitamins, and natural chelating agents, espe cially where they may be components of sewage and other organic wastes. Research is needed on the biology of species which cause exceptional blooms: (a) Knowledge of life cycles, in particular of resting stages, is lacking in many species. (b) Taxonomic and related confusion needs to be cleared up, in particular with respect to Gonyaulax tamarensis! excavata, ‘flagellate X ’, and the causative o r ganisms of DSP; ICES is urged to press ahead with publication of its leaflet series on Plankton Identifica tion, which will include phytoplankton species. (c) More information is needed on growth rates of ex ceptional bloom-forming species. With respect to exceptional blooms posing public health risks: (a) D evelopment of improved rapid toxin assays and cross-calibration of techniques is needed. (b) Monitoring of phytoplankton as well as toxicity should be carried out especially in DSP-affected areas. (c) National action is needed in some countries to har monize monitoring programmes and to ensure rapid information flow to resource managers. Calm and prompt public statements are essential to limit negative impact on commercial fisheries, especially molluscan shellfisheries. (d) International action is necessary to ensure rapid flow of information on toxic events and to review and harmonize safety and quarantine levels. With respect to problems posed by exceptional blooms for mariculture, it is proposed that the preliminary statement in the section on ‘M anagement of excep tional-bloom-related problems in mariculture’ should 113 be worked on further by a new ICES Working G roup on the M anagement of Problems Posed by Exceptional Blooms for Mariculture with the following terms of reference: (a) to establish means to collect and exchange informa tion on the incidence of problems due to excep tional blooms on mariculture operations; (b) to consider means of improving the predictability of bloom events on time - and space - scales relevant to fish farmers, including analysis of weather p at terns in relation to bloom incidence, (c) to consider proposals for research into management techniques for overcoming the effects of excep tional blooms, (d) to prepare advice for M em ber G overnments and for fish farmers on site selection, monitoring and prediction, and management options for bloom events. 114 Member States of ICES should be encouraged to con tinue to submit data on the incidence of exceptional blooms and related events to A nnales Biologiques for publication in the section dealing with this issue, com piled by D r J. P. Mommaerts. Selected papers from this Special Meeting should be published in the Rapports et Procès-Verbaux series, un der the editorship of D r M. M. Parker and D r P. Tett. A c k n o w le d g e m e n t s I wish to thank the participants in the evening discussion groups, and especially Paul Tett and Kathy Richardson, for valuable discussion and assistance in preparing this report. My thanks also to the ICES Secretariat for mak ing it possible to produce the first draft of this report during the Statutory Meeting.
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