Deep-sea Seamount Fisheries: a review of global status and future prospects/Pesquerias de Aguas Profundas Realizadas en Montes Submarinos: Revision Global de su Estado y Perspectivas Futuras.
Seamounts are prominent features of the seafloor throughout the oceans of the world. Their numbers are unknown, but recent studies based largely on satellite altimetry have estimated that there could be tens of thousands of large seamounts (Wessel, 2001; Kitchingman & Lai, 2004; Hillier & Watts, 2007). Their distribution is widespread throughout the world's oceans, especially in the Pacific (Fig. 1).
Seamounts support a large number and wide diversity of fish species. A total of almost 800 species have been recorded from seamounts (Froese & Sampang, 2004; Morato et al., 2006; Morato & Clark, 2007). Most of these also occur widely on the continental shelf and slope habitat, but seamounts can be an important habitat for commercially valuable species that may form dense aggregations for spawning or feeding (Clark, 2001) and on which a number of large-scale fisheries have developed in the deep-sea.
[FIGURE 1 OMITTED]
Many of these fisheries, however, have not been sustainably fished or managed. A number have shown a 'boom and bust' pattern, with catches rapidly developing and declining within a decade (e.g. Uchida & Tagami, 1984; Clark, 2001; Vinnichenko, 2002a). These patterns have raised concerns over whether such fisheries can be sustainable (e.g. Roberts, 2002; Gianni, 2004; Stone et al., 2004).
This paper briefly describes deep-sea seamount fish and trawl fisheries, reviews aspects of their sustainability, discusses what is required for effective management of both fisheries and the seamount habitat, and considers prospects for large-scale commercial exploitation of fish resources on seamounts. The study is restricted to demersal finfish, although seamount fisheries also occur for surface and midwater fish, crustaceans, and squids (see Rogers, 1994 and papers in Pitcher et al., 2007).
Description of seamount fishes
The definition of "deep sea" varies between organizations and countries. The FAO criterion is beyond the continental shelf/slope break, typically occurring at about 200 m. However, this depth-limit means a large number of shallow-water species are included where their depth distribution extends beyond 200 m. Hence, in this paper, the focus is on a more ecological definition that recognizes fish with low productivity relative to inshore continental shelf species, which are typically deeper than 400-500 m (FAO, 2008) (Table 1). It is not possible to generalize about biological characteristics, as deepwater seamount species span a wide range of productivity values (e.g. rubyfish are relatively short-lived and fast growing compared with orange roughy), but it is widely recognized that these deep-sea species are less productive and more vulnerable to fishing pressure than shelf species (e.g. Gordon, 2005; Japp & Wilkinson, 2007; Sissenwine & Mace, 2007).
Deep-sea trawl fisheries occur on seamounts for a number of species. These include alfonsino (Beryx splendens), black cardinalfish (Epigonus telescopus), orange roughy (Hoplostethus atlanticus), southern boarfish (Pseudopentaceros richardsoni), macrourid rattails (primarily roundnose grenadier Coryphaenoides rupestris), oreos (several species of the family Oreosomatidae, including the smooth oreo Pseudocyttus maculatus and the black oreo Allocyttus niger), and toothfish (Patagonian toothfish Dissostichus eleginoides and Antarctic toothfish D. mawsoni) (Clark et al., 2007). Other fisheries occur over seamounts, such as those for pelagic species (mainly tunas), near-bottom fishing for mackerels, and target species for smaller scale line fisheries (e.g. black scabbardfish Aphanopus carbo) (FAO, 2004; da Silva & Pinho, 2007).
Many of the main commercial seamount species are widespread, especially through the Atlantic, Indian, and South Pacific oceans (Table 2). A number of Southern Hemisphere species are found in the North Atlantic, but do not extend into the North Pacific (e.g. orange roughy, oreos). Some species are more localised to the North Atlantic (e.g. roundnose grenadier, blue ling, Sebastes mentella, S. marinus), and sablefish occur only in the North Pacific.
Seamount fisheries have taken place, or currently occur, on a large number of seamounts throughout the worlds' oceans. These are prominent in the Pacific ocean, but also in the southern Indian ocean, the Mid-Atlantic Ridge in the north Atlantic, and off the African coast in the south Atlantic (Fig. 2).
The largest seamount trawl fisheries have occurred in the Pacific ocean (Koslow, 2007; Clark et al., 2007). In the 1960s to 1980s, large-scale fisheries for pelagic armourhead and alfonsino took place on the Hawaiian and Emperor seamount chains in the north Pacific. In total, about 800,000 ton of pelagic armour-head and about 80,000 ton of alfonsino were taken. In the southwest Pacific, fisheries for orange roughy, oreos, and alfonsino were large and continue to be locally important. Orange roughy has also been the target of fisheries on seamounts off the central coast of Chile in the southeastern Pacific, on the Mid-Atlantic Ridge in the north Atlantic, off the west coast of southern Africa, and in the southwest Indian ocean. Roundnose grenadier was an important fishery for the Soviet Union in the North Atlantic, where catches were over 200,000 tonnes. Smaller fisheries for alfonsino, mackerel, and cardinalfish have occurred on various seamounts in the mid-Atlantic, southeast Pacific, and off the coast of north Africa. In the southern ocean, fisheries for toothfish, notothenids, and icefish can occur on seamounts as well as on slope and bank areas. Most of these seamounts are fished with bottom trawls, but several are also subjected to midwater trawl and longline fisheries.
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In total, the international catch of the main commercial demersal fish species on seamounts by distant-water fishing fleets is estimated to be over 2.15 million tonnes of fish since 1968 (Table 3) (Clark et al., 2007). Hence, seamount fisheries have contributed only a very small proportion of the total global wild-fish catch, which averaged about 60 million tonnes per year over the same period (Csirke, 2005).
South American fisheries
Most large volume deepwater fisheries have occurred in other regions, but several fisheries have developed off South America on seamounts. Off Brazil, a major programme of exploratory trawling between 2000 and 2006 covered a large part of the coast, although seamounts were of secondary importance in this work, which focused on the upper slope for finfish and shrimps (Perez et al., 2009b). Seamounts of the Fernando de Noronha and Vitoria-Trinidade chains have been exploited, although mainly with longlines (Hazin et al., 1998, Martins et al, 2005) and only limited trawling for groupers (Epinephelus spp.) (Perez et al. 2009a). During the 1980s, trawlers from the then Soviet Union fished on the Vitoria-Trinidade and Martin Vaz seamount chains and developed a fishery for alfonsino on the Rio Grande Rise (Clark et al., 2007).
In the southeastern Pacific, Soviet trawling occurred on the Nazca and Salas y Gomez Ridges for mackerel (Trachurus murphyi) and redbaits (Emmelichthys spp.), mainly in the 1970s but sporadic exploratory fishing also occurred in the 1980s (Clark et al., 2007; Galvez, 2009). Off Chile, a commercial fishery for orange roughy and alfonsino developed on seamounts around the Juan Fernandez Archipelago in 1998 (Labbe & Arana, 2001, Guerrero & Arana, 2009). Orange roughy catches peaked at 1,870 ton in 2001, but declined thereafter (Paya et al., 2005). This fishery was closed except for a research quota in 2006 following decreasing catches and stock size estimates (Niklitschek et al., 2005, Sissenwine & Mace, 2007).
SUSTAINABILITY OF DEEPWATER SEAMOUNT FISHERIES
Deep-sea seamount fisheries, even within areas of national jurisdiction, have typically not maintained high catch levels over time. There are many examples of 'boom and bust' fisheries that developed and declined rapidly, sometimes within a few years or a decade (e.g. Uchida & Tagami, 1984; Koslow et al., 2000; Clark, 2001). Orange roughy is commonly cited as an example of this, as few of its fisheries in the world have proven to be sustainable (e.g. Branch, 2001; Clark, 2001; Lack et al, 2003; Sissenwine & Mace, 2007).
New Zealand fisheries have dominated global catches (Fig. 3), although note that this figure includes continental slope habitats as well as seamounts. The New Zealand fishery is really the only one that has persisted over time and much of this is due to fishing grounds in a restricted area of the Chatham Rise, to the east of the main New Zealand islands. Seamounts account for between 40 and 60% of the catch in New Zealand (Clark & O'Driscoll, 2003), even though serial depletion has occurred in some areas (Clark, 1999; Clark et al., 2000). The Australian fishery was very large for a number of years between 1989 and 1993, when high catch rates of spawning fish on St Helens seamount were regularly made, but the stocks were rapidly depleted and quotas were progressively reduced (Lack et al., 2003; Bax et al., 2005; Koslow, 2007). The St Helens fishery is now closed completely. A similar situation occurred in Namibia and Chile where, despite extensive research and precautionary management objectives, catches have not been sustained, and fisheries are now very small or just by catch. The southwest Indian ocean is another area where large catches were taken for a short time, made worse by a total lack of any management on the High Seas, which saw an uncontrolled increase in effort in the early 2000s (from five vessels in 1999 to over 35 in 2000) and a sharp drop in catches and catch rates (FAO, 2002; Japp & James, 2005). Sissenwine & Mace (2007) described these catch histories as following two patterns: in the first, small stocks were fished down rapidly in only a few years before effective management could be implemented and, in the second, larger stocks were initially overestimated by research, and this was often coupled by non-conservative management practises and "fishing-down" phases, which lead to excessive depletion.
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Orange roughy is one of the least productive commercial species, but is not necessarily an extreme example. Fisheries for other deep-sea species have also shown low resilience to large catches, such as the pelagic armourhead fishery off Hawaii in the 1970s, alfonsino in the north Atlantic, roundnose grenadier on the Mid-Atlantic Ridge, and deepwater notothenids in the southern ocean (Clark et al., 2007). These fisheries have sometimes maintained catches by moving to new grounds or by switching to other species as the target species biomass has declined (e.g. increase in alfonsino catches as pelagic armourhead was over-fished on the Hawaiian seamounts (Clark et al., 2007)). However, even relatively shallow species (e.g. pink maomao, Caprodon longimanus) can be rapidly depleted over seamounts, evidenced by a short-lived fishery on the Lord Howe Rise, where Japanese catch rates in 1976 decreased from 1.7 to 0.2 ton h-1 over one year with a catch of not much more than 1000 ton (Sasaki, 1986). Sissenwine & Mace (2007) listed 44 deep-sea (> 200 m) area species combinations, 27 of which included stocks classed as overexploited or depleted. No stocks were identified as recovering.
Once overexploited, few deep-sea fisheries have shown signs of recovery. There are situations in which fishing success for orange roughy has improved with a reduction in effort levels, and fishers have reported increased catches of alfonsino and pelagic armourhead in some areas when the seamounts or fishing grounds have not been fished for a period. However, this may in part be related to fewer disturbances of aggregations with reduced trawling than an increase in stock size (Clark & Tracey, 1991). Orange roughy stocks in many areas generally continued to decline even when the catch has been reduced to levels thought by scientists to be sustainable. Some analyses have suggested depletion has not been excessive (Hilborn et al., 2006) but uncertainty about stock assessments and, in particular, assumptions about recruitment are critical in such evaluations. Irregular recruitment levels may be a key factor for the recovery of deep-sea species (Clark, 2001; Dunn, 2007).
The example of orange roughy fisheries, in particular, has given strong insights into three generic aspects that contribute to the lack of sustainability of deep-sea seamount fisheries:
Deep species often exhibit high longevity (up to 100 years for several species, e.g. redfish, orange roughy), late maturation (sometimes > 20 years before becoming mature, e.g. orange roughy, oreos), slow growth, low fecundity (e.g. deepwater sharks, orange roughy), intermittent recruitment (occurs with most species, but with long-lived species there could be decades between good year classes), and spawning may not occur every year (Morato et al., 2006; Morato & Clark, 2007). Intermittent spawning behaviour and the migration of aggregations to spawning grounds ("intermittent aggregation") have also been proposed to explain why some decreases in stock size appear to have been greater than expected based solely on the fisheries catch or why the levels of depletion have varied between different fishing grounds (e.g. Butterworth & Brandao, 2005). These types of fish generally have low rates of natural mortality and low production rates, meaning recovery is slow. Table 1 indicates that several of the major deepwater commercial fish species have a population doubling time ("resilience") of 14 years or greater (e.g. cardinalfish, orange roughy, redfish) and high vulnerability indices of 70 to 80. Their biology is not evolved to cope with high levels of natural predation and so they are more vulnerable to overfishing than shallow water shelf species.
Many species aggregate on seamounts or ridge peaks because of depth or oceanographic conditions (e.g. Koslow, 1997). Such aggregations will be more vulnerable to overfishing and rapid depletion than those in which species are more dispersed on shelf or slope habitats. When aggregations are formed for spawning, the effects may be greater because of the possible disruption of spawning processes and reduced reproductive success (although this has rarely been documented). Target trawling on seamounts is often localized, and the density of tows per seamount area can be high (e.g. O'Driscoll & Clark, 2005). Heavy bottom trawl gear is used to tow on the rough, hard bottom, which is often characteristic of seamounts, and the invertebrate fauna, often dominated by large, slow-growing, sessile organisms, are especially vulnerable to damage by fishing gear (e.g. Clark & Koslow, 2007). Fishing grounds often occur offshore and so are carried out by large powerful vessels with the ability to work large gear, catch and process large amounts of fish, and stay at sea for long periods.
Economic considerations can also be important. The market value of some of the deepwater species is high, which creates an incentive for commercial operators to target the species (Japp & Wilkinson, 2007). Good catches will offset the relatively high operating costs of vessels offshore and create pressure to continue fishing as stocks decline or if there are high costs associated with industry funding of research and management (e.g. Francis & Clark, 2005). Research and management limitations Francis & Clark (2005) described issues affecting the sustainability of orange roughy based on the New Zealand experience. As well as a lack of knowledge of the biological characteristics and processes (mentioned above), they emphasised how standard stock assessment techniques were often difficult to apply in the deep-sea environment and given the aggregating nature of the species. Punt (2005) and Sissenwine & Mace (2007) also discussed difficulties for estimating the stock size of orange roughy and how uncertain such stock assessments are likely to be. This uncertainty in the science has, at times, led to subsequent management responses being too slow or insufficient (Bax et al., 2005; Francis & Clark, 2005). Precautionary management is required and the standard target reference levels and management concepts (e.g. MSY, fishing down practices) applied in several deep-sea fishing countries (e.g. New Zealand) have proven risky and insufficiently conservative (Sissenwine & Mace, 2007).
Management of seamount fisheries
The "ecosystem approach" to fisheries management is now widely advocated and applied in deep-sea fisheries (FAO, 2003). However, the inherent restrictions on obtaining sufficient stock assessment or benthic habitat data (compared with nearshore shelf/slope fisheries) mean that management regimes typically operate at a low level of knowledge, and management action must occur in a highly precautionary manner. There is an increasing body of literature on data, reporting requirements, and appropriate management actions to help ensure the sustainability of deep-sea fisheries (Francis & Clark, 2003; FAO, 2007, 2008; Sissenwine & Mace, 2007; Probert et al., 2007; Morato & Pitcher, 2008; Rogers et al, 2008).
Various management actions now include closed seamounts, fishing method or gear restrictions, depth limits, individual seamount catch quotas, bycatch quotas, and habitat exclusion areas (e.g. hydrothermal vents) (Probert et al., 2007). Closed areas are a common method to protect the habitat, but can be problematic because they can exclude fishing from productive grounds. Recent initiatives by some fishing consortia (e.g. southwest Indian Ocean Fisher's Association, New Zealand Deepwater Group) have instigated "Benthic Protected Areas (BPAs)" which the fishers voluntarily avoid to prevent seafloor damage by bottom trawling. Industrial "buy-in" to environmental protection is a positive step as long as science is involved to help design the BPA distribution and size so they are representative and effective. Typically, the fine spatial scale needed to research and manage sea mount stocks is problematic. Serial depletion of fish stocks can occur quickly (Clark, 1999), yet catch rates can be maintained even though biomass is being reduced (Clark, 2001; Lack et al., 2003; Francis & Clark, 2005; Sissenwine & Mace, 2007). There is no easy answer to setting appropriate precautionary catch limits for new seamount fisheries, although restricting effort to only a few vessels (e.g. Namibian orange roughy fisheries; Boyer et al., 2001) and imposing limits on the catch from a single seamount or feature (e.g. New Zealand; Rogers et al., 2008) can help reduce the risk of rapid over-exploitation. An analysis of seamount catch over time indicates that the initial orange roughy biomass on a single seamount feature may generally be only a few thousand tonnes (Clark et al., 2001).
Effects of bottom trawling on the wider benthic community and habitat also need to be considered (e.g. Dayton et al., 1995; Hall, 1999; Clark & Koslow, 2007). The deep-sea fish community includes species that have low productivity and can be vulnerable to the effects of fishing, even if the fishery targets a different species. Seamounts can host endemic species or species with a very restricted geographical distribution (Rogers, 1994; Richer de Forges et al., 2000), as well as habitat-forming fauna such as deep-sea corals and sponges that are regarded as indicators of "vulnerable marine ecosystems" (FAO, 2008). This has been part of the justification for calls from NGOs in recent years for a moratorium on bottom trawling on the High Seas. Management, therefore, needs to balance exploitation and conservation, both of fisheries and seamount habitats (Probert et al., 2007). A mixture of protected and open seamounts is one strategy that appears to be successful off New Zealand (Brodie & Clark, 2003).
Seamount fisheries for deep-sea species in the future are likely to be small volume, high value fisheries. The track record of the deeper species such as orange roughy and oreos indicates that large catches will only last a few years, and highly precautionary catch limits may be needed if they are to be sustainable. More productive seamount species such as alfonsino, scabbardfish, grenadier, and armourheads are more resilient to heavy fishing and have been classified as less vulnerable (Gordon, 2005), but stocks appear to be variable and are still unlikely to withstand high catch levels for more than a few years. Where estimates have been made, the biomass of many fish stocks on seamounts is relatively low and does not exceed several hundreds of thousands of tonnes for even the most abundant species (e.g. Sasaki, 1986; Vinnichenko, 1998, 2002b). An analysis of New Zealand seamount fisheries for orange roughy suggests that few seamounts can support long-term catches of more than a few hundred tonnes per year (Clark et al., 2001).
Japp & Wilkinson (2007) and Sissenwine & Mace (2007) have both summarized "deep-sea" catches as reported in FAO statistics up to 2005. The catch trends for the main seamount species or family groupings typically show a decline after an initial rapid increase as the fisheries developed. Although FAO statistics cover very large regions of the world and do not distinguish seamounts from slope fisheries, these patterns are consistent with those estimated from specific seamount data by Clark et al. (2007).
Clark et al. (2007) summarized the global status of the large offshore seamount fisheries. Their conclusions were that intense fishery pressure on the seamounts of the Corner Rise, north Azores area of the Mid-Atlantic Ridge, and the Vavilov, Walvis, southwest Indian, Emperor and Hawaiian ridges has resulted in the depression of many fish stocks, with no increase of catch in these areas likely within the next few years. Relatively new fishing grounds such as seamounts on the Norfolk, Lord Howe, Louisville, and south Tasman Rise have been targeted for deepwater species such as orange roughy and are also fully exploited. Some areas of the northern Mid-Atlantic Ridge and some offshore seamounts located in Antarctic waters and the central oceanic regions may have some potential for further exploitation. However, the volume of deep-sea species is likely to be small and short-lived if fisheries are not managed carefully. Such management will also need, in the future, to take account of balancing fisheries management with conservation of the benthic environment.
This paper is based on a presentation at the Deep Sea fisheries special session at the XII COLACMAR in Brazil, April 2007. My thanks to the Convenor of that session (Dr. Angel Perez) and the conference organizers for funding to attend the conference. The contents of the paper draw on a wide range of research over the last decade, but particularly a NIWA project on Seamounts: their importance to fisheries and marine ecosystems (FRST contract no. CO1X0508). Two anonymous journal referees made numerous useful comments to improve the manuscript.
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Received: 15 August 2008; Accepted: 9 March 2009
Malcolm R. Clark
NIWA, Private Bag 14-901, Wellington, New Zealand
Corresponding author: Malcolm R. Clark (firstname.lastname@example.org)
Table 1. Bathymetric distribution and biological characteristics of commercial fish species on seamounts. Depth is the main commercial range. Data on maximum age, resilience (population doubling time), and intrinsic extinction vulnerability, the latter based on Cheung et al. (2005) and consisting of a relative measure ranging from 1 (very low) to 100 (very high), are from Fishbase. Tabla 1. Distribucion batimetrica y caracteristicas biologicas de los peces comerciales capturados en montes submarinos. La profundidad se refiere a rangos en que se efectuan las pescas comerciales: edad maxima, productividad relativa (lapso en que se duplica la poblacion) y vulnerabilidad intrinseca de extincion. De acuerdo a Cheung et al. (2005), una medida relativa varia de 1 (muy bajo) a 100 (muy alto), datos tomados de Fishbase. Species Scientific name Main depth range (m) Alfonsino Beryx splendens 300-600 Cardinalfish Epigonus telescopus 500-800 Rubyfish Plagiogeneion rubiginosum 250-450 Blue ling Molva dypterygia 250-500 Black scabbardfish Aphanopus carbo 600-800 Sablefish Anoplopoma fimbria 500-1,000 Pink maomao Caprodon spp. 300-450 Southern boarfish Pseudopentaceros richardsoni 600-900 Pelagic armourhead Pseudopentaceros wheeleri 250-600 Orange roughy Hoplostethus atlanticus 600-1,200 Oreos Pseudocyttus maculatus, 600-1,200 Allocyttus niger Bluenose Hyperoglyphe antarctica 300-700 Redfish Sebastes spp. (S. marinus, 400-800 S. mentella, S. proriger) Roundnose grenadier Coryphaenoides rupestris 800-1,000 Toothfish Dissostichus spp. 500-1,500 Notothenid cods Notothenia spp. 200-600 Species Maximum Resilience Intrinsic age (years) extinction (years) vulnerability Alfonsino 25 5-14 65 Cardinalfish 100 14 70 Rubyfish 10 2-4 49 Blue ling 20 5-14 73 Black scabbardfish 30 5-14 63 Sablefish 115 >14 76 Pink maomao 2-4 50 Southern boarfish 2-4 42 Pelagic armourhead 11 7 57 Orange roughy 150 >14 74 Oreos 100 >14 72 Bluenose 15 5-14 74 Redfish 75 >14 75 Roundnose grenadier 54 5-14 78 Toothfish 20-30 5-14 86 Notothenid cods 15-20 5-14 58 Table 2. Geographical distribution of commercial fish species (+ indicates occurrence in that ocean). Tabla 2. Distribucion geografica de las especies comerciales (+ indica presencia en un oceano particular). Species North South North South Atlantic Atlantic Pacific Pacific Alfonsino + + + + Cardinalfish + + + Rubyfish + + Blue ling + Black scabbardfish + Sablefish + Pink maomao + + Southern boarfish + + Pelagic armourhead + Orange roughy + + + Oreos + + Bluenose + + Redfish + + Roundnose grenadier + Toothfish + + Notothenid cods + + Species Indian Southern Ocean Ocean Alfonsino + Cardinalfish + Rubyfish + Blue ling Black scabbardfish Sablefish Pink maomao Southern boarfish + Pelagic armourhead Orange roughy + Oreos + + Bluenose + Redfish Roundnose grenadier Toothfish + + Notothenid cods + + Table 3. Historic catches of the main fish species from seamounts, major fishing periods, and main gear types used in seamount fisheries (derived from data in Clark et al., 2007). Tabla 3. Capturas historicas de las especies mas relevantes capturadas en montes submarinos, periodo de pesca mas importante y principal tipo de arte de pesca empleado en la pesqueria (datos tomados de Clark et al., 2007). Common name Total historical Main catch (ton) fishery years Alfonsino 166950 1978-present Orange roughy 419100 1978-present Oreos 145150 1970-present Cardinalfish 52100 1978-present Redfish 54450 1996-present Southern boarfish 9600 1982-present Pelagic armourhead 800000 1968-1982 Mackerel species 148200 1970-1995 Roundnose grenadier 217000 1974-present Blue ling 10000 1979-80 Scabbard fish 75000 1973-2002 Sablefish 1400 1995-present Bluenose 2500 1990-present Rubyfish 1500 1995-present Pink maomao 2000 1972-1976 Notothenid cods 36250 1974-1991 Toothfish 12250 1990-present Total 2153470 Common name Gear type Alfonsino Bottom and midwater trawl, some longline Orange roughy Bottom trawl Oreos Bottom trawl Cardinalfish Bottom (and midwater trawl) Redfish Bottom and midwater trawl Southern boarfish Bottom trawl Pelagic armourhead Bottom and midwater trawl Mackerel species (Bottom) and midwater trawl Roundnose grenadier Bottom, and midwater trawl Blue ling Bottom trawl Scabbard fish Bottom and midwater trawl Sablefish (Bottom trawl), longline Bluenose Bottom and midwater trawl Rubyfish Bottom and midwater trawl Pink maomao Bottom and midwater trawl Notothenid cods Bottom trawl Toothfish Bottom trawl, longline Total
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|Author:||Clark, Malcolm R.|
|Publication:||Latin American Journal of Aquatic Research|
|Date:||Nov 1, 2009|
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