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Manuscript Number: FISH9546

Title: Current status of the pink cusk eel (Genypterus blacodes) industry
and its potential to diversify aquaculture in Chile

Article Type: Review article

Keywords: Genypterus blacodes; overfishing; management measure; Chilean

aquaculture.

Corresponding Author: Professor Jennie Marianne Risopatron, MSc

Corresponding Author's Institution: La Frontera University

First Author: Kelly Dumorné, PhD

Order of Authors: Kelly Dumorné, PhD; Osvaldo Merino, PhD; Elías

Figueroa, PhD; Iván Valdebenito; Jorge G. Farías; Jennie Marianne
Risopatron, MSc

Manuscript Region of Origin: CHILE

Abstract: Chile is a country with abundant aquatic resources that has

allowed fishery and aquaculture to develop in the last decades. Pink cusk
eel (Genypterus blacodes) is an important ocean species commercialized in
Chile and other countries such as Australia and New Zealand. This species
constitute a very important resource in artisanal and industrial fishing.
However, in recent decades the volume of landings has fallen considerably
due to overexploitation. Although G. blacodes possesses an attractive
commercial farming potential, the reproductive aspects associated with
this species have not been addressed in detail. Nevertheless, in the
current context, where the diversification of aquaculture through the
production of new species is an important issue for the sector,
cultivation of G. blacodes offers a real alternative for Chilean
aquaculture, presenting excellent medium and long-term expectations not
only for satisfying domestic demand, but also for increasing exports.
Currently, its production is totally dependent on the harvest of
broodstocks or gametes from wild populations. The potential to farm G.
blacodes will open interesting business prospects for Chile in the coming
years. This review summarizes the current status of G. blacodes and its
potential for aquaculture, focusing on Chile where it is a widespread and
traditionally important species.
*Cover Letter

Temuco, 09th October 2019

Dear Dr.
G. A. Rose
Editors-in-Chief
Fisheries Research

We wish to submit an original review entitled "Current status of the pink cusk eel
(Genypterus blacodes) industry and its potential to diversify aquaculture in Chile" for
consideration by Reviews in Fisheries Science & Aquaculture. We confirm that this work is
original and has not been published elsewhere, nor is it currently under consideration for
publication elsewhere.
This review provides the current status of G. blacodes and its potential for aquaculture. We
believe that this manuscript is appropriate for publication by Fisheries Research because
your Journal is essential reading for scientists and people linked to fishing management and
aquaculture.

Sincerely,

MSc. Jennie Risopatrón G.

Directora
Centro de Excelencia en Biotecnología de la Reproducción (BIOREN-CEBIOR)
Facultas de Medicina
Universidad de La Frontera
Temuco-Chile
*Highlights (for review)

Highligths
 The potential to farm G. blacodes will open interesting business prospects for
Chile in the coming years.

 The cultivation of G. blacodes offers a real alternative for Chilean aquaculture.

 Strategies to protect the reproductive potential of G. blacodes are to reassess

capture size and average size at sexual maturity.

*Manuscript including abstract
Click here to view linked References

1 Current status of the pink cusk eel (Genypterus blacodes) industry and its potential
2 to diversify aquaculture in Chile
3
4 Kelly Dumornéa,b, Osvaldo Merinoa, Elías Figueroac, Iván Valdebenitod, Jorge G.
5 Faríasa,e, and Jennie Risopatróna,f*
6
a
7 Center of Excellence of Biotechnology in Reproduction (BIOREN-CEBIOR), Faculty of

8 Medicine,University of La Frontera, Temuco, Chile.

b
9 Vice-Rectory for Research and Postgraduate Studies, Graduate Academic Direction,

10 University of La Frontera, Temuco, Chile.

c
11 Nucleus of Research in Food Production, Department of Biological and Chemical

12 Sciences, Faculty of Natural Resources, Catholic University of Temuco, Temuco, Chile.

d
13 Faculty of Natural Resources, Catholic University of Temuco, Temuco, Chile.
e
14 Chemical Engineering Department, Faculty of Engineering and Science, University of

15 La Frontera, Temuco, Chile.

f
16 Department of Basic Sciences, Faculty of Medicine, University of La Frontera,

17 Temuco, Chile

18

19

20

*
21 Correspondence: Jennie Risopatrón, Center of Excellence of Biotechnology in

22 Reproduction (CEBIOR-BIOREN), Faculty of Medicine, University of La Frontera,

23 Temuco, Chile, Avenida Francisco Salazar 01145, Box 54D. Temuco 4811230, Chile.

24 Email: jennie.risopatron@ufrontera.cl

25
26
27
28
1
29 ABSTRACT
30
31 Chile is a country with abundant aquatic resources that has allowed fishery and

32 aquaculture to develop in the last decades. Pink cusk eel (Genypterus blacodes) is an

33 important ocean species commercialized in Chile and other countries such as Australia

34 and New Zealand. This species constitute a very important resource in artisanal and

35 industrial fishing. However, in recent decades the volume of landings has fallen

36 considerably due to overexploitation. Although G. blacodes possesses an attractive

37 commercial farming potential, the reproductive aspects associated with this species have

38 not been addressed in detail. Nevertheless, in the current context, where the

39 diversification of aquaculture through the production of new species is an important

40 issue for the sector, cultivation of G. blacodes offers a real alternative for Chilean

41 aquaculture, presenting excellent medium and long-term expectations not only for

42 satisfying domestic demand, but also for increasing exports. Currently, its production is

43 totally dependent on the harvest of broodstocks or gametes from wild populations. The

44 potential to farm G. blacodes will open interesting business prospects for Chile in the

45 coming years. This review summarizes the current status of G. blacodes and its

46 potential for aquaculture, focusing on Chile where it is a widespread and traditionally

47 important species.

48

49 Keywords: Genypterus blacodes, overfishing, management measure, Chilean

50 aquaculture
51

52

53

54

55

2
56 1. Introduction
57
58 The aquaculture industry generates revenue by the application of different

59 technologies to the production of aquatic species in high demand for human

60 consumption (Ovando-Solis, 2013). The sustainability of world fisheries is an important

61 issue, especially in terms of current trends in fish abundance and the prospects for stock,

62 food security and biodiversity conservation (Worm et al., 2009; Youn et al., 2014;

63 McClanahan et al., 2015). In recent years there has been an increase in aquaculture

64 production; it currently provides half of all the fish destined for human consumption,

65 with the foreseeable future growth of the fisheries sector deriving mainly from

66 aquaculture (FAO, 2016).

67 During recent years, aquaculture production has increased considerably due to

68 growth of world population (Aksnes and Browman, 2016; Lahnsteiner et al., 2009).

69 This increase in aquaculture production has been driven in part by growing concern

70 about the state of fish stocks and the need to provide information for policy and decision

71 makers globally. Since each fish stock is typically unique, and experimental approaches

72 cannot be used to predict their response to fishing, modelling and simulation are the

73 most frequently used methods in fisheries science (Jarić et al., 2012), play a major role

74 in providing management advice. Fish play a key role in the world’s food supply, and

75 consumption has grown steadily during the last five decades (FAO, 2014; WHO, 2016).

76 Aquaculture production has also increased, due to demand from consumers and

77 the depletion/restriction of wild capture fisheries; this industry, now accounting for 50%

78 of all fish consumed globally, is expected to become the main source of fish and

79 seafood by 2030 (Henchion et al., 2017). During recent years, the percentage of

80 overexploited fish stocks has increased (FAO, 2010). Chile offers great potential for the

81 development of freshwater and marine aquaculture (FAO, 2010, 2014) (Table 2).
3
 82 It is among the ten most important aquaculture producers in the world, with

83 abundant aquatic resources that have allowed aquaculture, especially of Atlantic salmon

84 (Salmo salar), to develop strongly in recent decades. Chile has made significant strides

85 in increasing the volume and the value of both capture fisheries and aquaculture

86 production (Bórquez and Hernández, 2009). Genypterus blacodes is a benthic predator

87 found at depths from 22 to 1000 m, distributed in both in the Pacific and Atlantic waters

88 of South America (Díaz et al., 2012); populations also exist in the waters of New

89 Zealand and southern Australia (Baker et al., 2014; Contreras-Reyes et al., 2018)

90 (Figure 1A). A specie with potential to farm and will open interesting business

91 prospects for Chile in the coming years.

92 In Chile, adults exhibit demersal behaviour and are usually found at depths

93 between 45 and 350 m (Nyegaard et al., 2004). Other species of Genypterus are found

94 around South Africa (G. capensis) and South America (G. chilensis). In Chile G.

95 blacodes populations are divided into northern and southern stocks, each with different

96 characteristics (Baker et al., 2014; Contreras-Reyes et al., 2018). According to existing

97 records, the pink cusk-eel fishery is carried on in Chilean waters from Coquimbo to

98 Cape Horn (41º28'-57º00'S) (Wiff et al., 2007) (Figure 1B). The reproductive activity of

99 the species occurs mainly during the summer in Patagonian coastal waters, from 41ºS to

100 45ºS (Wiff et al., 2011).

101 G. blacodes is one of two species of the genus Genypterus traditionally exploited

102 in the Argentine Sea and in Chile (Cohen et al., 2015); it is also one the most important

103 species for Chile in terms of intentionality of fishing (Wiff and Quiroz, 2010). Annual

104 catches of G. blacodes in Chile range from 45-50 thousand tons (Chong et al., 2014).

105 During recent decades, fish farmers around the world have been attracted by G.

106 blacodes farming, especially in Chile, Australia, Argentina and New Zealand.

4
107 There is a current management plan to repopulate the environment with this

108 species. There are no international databases recording the production and sales of

109 farmed G. blacodes. Although aquaculture protocols for the management of broodstock

110 males are well established, there is still a significant lack of data and statistics that

111 would appropriately depict the current state of G. blacodes aquaculture. Commercial

112 capture of G. blacodes represents a significant source of income for Chilean fisheries,

113 however, the population has declined to such a degree that major concerns have been

114 raised over its long-term survival (Chong et al., 2014; Wiff et al., 2006). It has very

115 great potential for aquaculture in Chile, due to the exceptional quality and high

116 commercial value of its flesh (Chong et al., 2014). There is limited information in the

117 literature on the functional aspects of the reproductive biology of G. blacodes (Freijo et

118 al., 2009).

119 Studies of G. blacodes in other parts of the world have suggested the existence

120 of multiple stocks in smaller areas than the distribution presented by this species in

121 Chile (Wiff et al., 2011). In Chile, sexual maturation of G. blacodes bred in captivity

122 has not been yet reported; however, Dumorné et al. (2018a) have reported a fertilizing

123 capacity rate (> 70%) using intratesticular sperm from wild males in ova from captive

124 female breedstock. This study showed that the use of intratesticular spermatozoa for in

125 vitro fertilization is a key tool for breeding this species in captivity and hence would

126 allow its introduction in the aquaculture industry. Sexual maturation can be influenced

127 by a number of factors including age, size, geographical distribution, season, nutritional

128 status and fishing pressure (Smith et al., 2005).

129

130

5
131 There are numerous publications of studies of G. blacodes fisheries, covering

132 areas such as: taxonomy, stomach contents, age and growth parameters (Chong and

133 Aguayo, 1990); macroscopic and microscopic structure of the ovary in samples from the

134 Atlantic range, description of spawning stages from Argentinean waters

135 (Machinandiarena et al., 1998); regional morphometric variations in New Zealand

136 (Colman, 1995); instantaneous rate of natural mortality (Ojeda et al., 1986); population

137 structure (Canales-Aguirre et al., 2010a), growth (Wiff et al., 2007), reproduction

138 (Baker et al., 2014), mortality (Wiff et al., 2011), food consumption (Wiff et al., 2015),

139 Genetic and morphological (Smith and Paulin, 2010), quality and morphology gamete

140 (Dumorné et al., 2018a, b). It is important to promote further research into G. blacodes,

141 particularly in view of its great productive potential for Chile. G. blacodes farming

142 presents excellent medium and long-term prospects, not only for satisfying domestic

143 demand but also for increasing exports. Biotechnological reproductive management

144 needs to be consolidated for reproductive processes, gamete management and in vitro

145 embryo production, with a view to promoting and developing a sustainable, competitive

146 industry. The aim of this review is to provide new background on G. blacodes that

147 allows to develop strategy to conserve and repopulate the environment of this species

148 for establishing a sustainable export trade for Chile.

149
150 2. Management and capture of Genypterus blacodes
151
152 Capture fisheries and aquaculture supplied the world with about 148.5 million

153 tons of fish for human consumption in 2010, providing an apparent per capita supply of

154 18.4 kg-1 (FAO 2010). The first step in successful fisheries management is to clearly

155 define and prioritize management objectives (Beddington et al., 2007).

6
156 Markets in the form of individual transferable quotas (ITQs), and the growing

157 global integration of fish markets, will increasingly provide the organizing and

158 regulatory principles for many fisheries (Squires et al., 1995). Chile lands around 2.5

159 million tons of fish every year, the eighth largest catch of any country in the world

160 (FAO, 2014). New fishery legislation has been passed in Chile since the early 1990s.

161 The Fishery and Aquaculture Law (FAL) of 1991 reformed the division of the right to

162 fish between the artisanal and industrial subsectors, introduced sea-zoning strategies,

163 and established a differential individual transferable quota (ITQ) system. The FAL

164 established different fishery-management regimes according to the degree of species

165 exploitation. For fully exploited species such as G. blacodes, vessels must be registered

166 for specific fisheries and management is based on total allowable catches, determined

167 by the government on the basis of stock assessments and quota allocations to

168 individuals and registered vessels (Gómez-Lobo et al., 2007; Castilla, 2010;).

169 The first definition of stock for G. blacodes was made by Chong and Aguayo

170 (1990) based on a morphometric analysis of otoliths. Microsatellites are currently one of

171 the most popular and most used genetic markers in molecular ecology and in the

172 fisheries industry (O’Connell and Wright, 1997). These markers are especially useful

173 for evaluating patterns of population genetic variability, and the micro evolutionary

174 mechanisms behind these patterns (Barbará et al., 2007). Studies based on

175 morphometric analysis of G. blacodes populations have been carried out in Australian

176 (Ward and Reilly, 2001) and New Zealand waters (Colman, 1995; Smith and Francis,

177 1982). Canales-Aguirre et al. (2010b) used microsatellites to determine genetic

178 variability and population in zones where G. blacodes is being fully exploited in

179 Chilean waters. During the 1980s the fleet began targeting G. blacodes directly, and

180 high levels of capture have been recorded since the early 1990s.

7
181 The population attributes of G. blacodes, such as its low resilience to

182 exploitation, slow growth, medium longevity and sedentary life-style, facilitate the

183 study of life history trade-offs (Wiff et al., 2007). According to indicators studied by

184 Céspedes et al. (2011), the size structures of industrial capture of G. blacodes in Chile

185 between 2009 and 2010 showed unimodal distributions, with a greater presence of

186 juveniles. New Zealand has determined the existence of at least three G. blacodes stocks

187 through several techniques including allozymes (Smith and Francis, 1982),

188 morphometry (Colman, 1995), vital parameters and size structure (Horn, 1993). The

189 maximum length of G. blacodes in Chile is 160 cm, while the average length at capture

190 is 80-90 cm (Baker et al., 2014). The capture of a significant proportion of juveniles

191 (under 90 cm length) is a significant feature of the G. blacodes fishery, considering that

192 the age at first maturity is 6 years (82 cm) (Wiff et al., 2011).

193 Studies using allozyme and microsatellite techniques to look at the stock

194 structure of G. blacodes inhabiting the Australian coast have found no statistical

195 evidence to reject or accept the hypothesis of a single stock in the distribution. It is

196 therefore difficult to determine whether the existence of genetic variations is local

197 (Canales-Aguirre et al., 2010a). Landings of G. blacodes have been decreasing

198 considerably in Chile (Figure 2). However, the fishery has been on the increase since

199 2014, with a steep increase during the last two years. In Chile, according to Contreras et

200 al. (2014), the stock of G. blacodes is at 20% spawning biomass. The artisanal fishery

201 contributed 60% of national landings in 2013, followed by the industrial fleet with 40%.

202 In recent years, given the high levels of recorded catches, the stock is in a state of

203 overexploitation; it remains at risk of overfishing due to high fishing mortality rates

204 (Wiff et al., 2011). In the current context of stock depletion, demand for G. blacodes is

205 increasing and further upward movement of prices is observed.

8
206 With increased marketing efforts and considering population growth, the

207 demand for this product could increase tremendously (Janko, 2014). Types of fishing,

208 fishing methods, socio-economic factors, lack of facilities and infrastructure, ineffective

209 administration, lack of expertise and lack of scientific data are major challenges for the

210 fishery management and conservation of G. blacodes. Thus installing effective

211 management with regular follow-up of stakeholders, and encouraging the development

212 of aquaculture, are very important for sustaining resources and satisfying demand.

213
214 3. Reproductive aspects of Genypterus blacodes
215
216 Studies on the reproductive status of G. blacodes along the coast of Chile are

217 relatively sparse and consider the austral zone as a single closed population. They

218 include studies on fecundity and maturity (Chong, 1993; Paredes and Bravo, 2005),

219 however, key aspects of the reproduction of this species, and how these may differ

220 between management zones and over time, remain largely unknown (Baker et al.,

221 2014). G. blacodes presents low fertility and a high frequency of spawning events

222 (Cordo et al., 2002). It has been established that G. blacodes is of a partial spawning

223 type, with a pattern of indeterminate oocyte development. Since it has relatively low

224 levels of fecundity (approximately 500.000 oocytes) and matures at a larger size than,

225 for example, the two Chilean hake species, Merluccius gayi and Merluccius australis, it

226 may be more susceptible to overexploitation than hake (Paredes and Bravo, 2005).

227 During the spawning season, G. blacodes moves to the deepest layers of its habitat to

228 reproduce, as deep as a thousand meters (Freijo et al., 2009). The eggs of Genypterus

229 spp. resemble tiny balloons of very transparent glass and fluctuate between the surface

230 and a depth of 500 metres; the tiny organisms are born in sheet form, called

231 leptocephali.

9
232 While studies on the reproduction of this species are improving, questions

233 remain about many aspects of its life history and reproduction stages (George-

234 Nascimento and Muñoz, 1997). The culture of G. blacodes in Chile is in the initial

235 stage, and is mainly oriented towards the capture of wild juveniles. The production of

236 embryos and larvae requires research to obtain essential, detailed reproductive

237 information. Before the species can be cultivated, the reproduction cycle must be

238 controlled to ensure continuous production of viable gametes and embryos. The

239 production of embryos of marine fish has received great attention in recent years,

240 because efficient production of embryos with a high probability of survival allows the

241 production of viable larvae and consequently of viable offspring. However, achieving

242 constant production of embryos is complicated, mainly due to the risk of high mortality

243 rates. Gamete quality is essential in species introduced into aquaculture (Valdebenito et

244 al., 2013; Bobe, 2015)

245 As a research group, our efforts have focused on improving the quality of the

246 intratesticular spermatozoa of G. blacodes, through research focused on aspects that can

247 improve their quality and efficacy for in vitro fertilization. Investigations into G.

248 blacodes sperm physiology were reported in Dumorné et al. (2018 a, b). The authors

249 evaluated some aspects of the sperm biology of G. blacodes, in particular assessing the

250 effects of pH, temperature and osmolality on the motility of G. blacodes intratesticular

251 spermatozoa, as well as quality parameters such as mitochondrial membrane potential,

252 fertilization rate and plasma membrane integrity. The results of these studies allow a

253 baseline to be established for further research, and protocols for artificial reproduction

254 of this species to be developed and optimized. The information gathered in the present

255 research will be useful for developing the biotechnology of G. blacodes (Dumorné et

256 al., 2018 a, b).

10
257 Considering the importance of fish reproduction in captivity and in the wild, it is

258 advisable to extend research into cellular aspects, thus providing useful information for

259 the development of means of cultivation and other inputs used in the management of

260 gametes of this species.

261
262 4. Destination markets for Chilean Genypterus blacodes

263
264 Aquaculture in Chile is oriented mainly towards the production of fish; it is

265 carried on mostly in coastal marine environments, and to a lesser degree in freshwater

266 environments, almost exclusively in just two of the country’s administrative regions.

267 The first efforts at commercial aquaculture in Chile were carried out in the period 1921-

268 1973, through plans and initiatives based on extensive and semi-intensive systems

269 (Basulto, 2003). However, large-scale commercial aquaculture began in the 1980s,

270 consistent with a national economic policy that embraced private activity and opening

271 to international trade, and as a response to the increasing overexploitation of local

272 fishery stocks of native species destined for international markets. Aquaculture

273 developed quickly, geared towards foreign trade and based on the cultivation of species

274 of high commercial value for which the cultivation technologies were already known.

275 The greatest technological development has been associated with fish-farming, followed

276 by the cultivation of scallops and oysters. In general, cultured mussels and algae are

277 small- and medium-scale crops, with lower levels of investment and technology. The

278 remarkable development of the salmon industry since the 1980s has been based on the

279 appropriate and advantageous natural conditions that the country offers, lower

280 production costs, the global macroeconomic environment, and the public and private

281 support given to the production stage (Agosín, 1999).

11
282 Market access and appropriate technology are necessary conditions for

283 aquaculture to develop, providing good quality food, business earnings, and

284 employment for marginalized people (Belton et al., 2012; Béné et al., 2016). The

285 aquaculture and wild capture of G. blacodes are entirely controlled in Chile, although

286 live specimens are traded internationally in some countries. Demand for G. blacodes is

287 growing and markets are expanding, and Chile, together with Argentina and New

288 Zealand, is already established as an important exporter. G. blacodes catches in Chile

289 are severely regulated by quotas, which are lower than for other species, as a result of a

290 management system which has poor conservation parameters (Wiff et al., 2011; Chong

291 et al., 2014).

292 The global market for G. blacodes is around 45.000 tons per year. The main

293 export market for G. blacodes from Chile is Spain, which receives 72% of frozen

294 products and 100% of fresh chilled fish; the next largest markets are United States,

295 Brazil, Russia and Portugal (SUBPESCA, 2013). Table 1 shows main Chilean export

296 markets of G. blacodes (Chong et al., 2014). The export price in Chile varies between

297 US$3.50 per kilo for fresh chilled produce and US$4.20 per kilo for frozen format,

298 positioning it as a very attractive business. Artisanal sector catches are processed mainly

299 in fresh chilled format, while the industrial sector is essentially processed for frozen fish

300 (SUBPESCA, 2013). In Chile, the marketing chain for G. blacodes from the artisanal

301 sector is quite simple: the fish are distributed either to wholesalers nationwide who

302 serve their regions, or to processing plants that are responsible for processing the fish

303 mainly in fresh chilled format for export. After capture, G. blacodes is either processed

304 immediately or kept alive, requiring a reliable, consistent supply of crabs for feed.

305

12
306 Live storage includes maintenance in water tanks near processing facilities and dry

307 transport to the destination market; it enables the industry to control the transport and

308 processing time of live G. blacodes. Processing includes a series of operations from

309 slaughter to the final product. Live storage technology allows G. blacodes to be

310 transported to overseas markets in good condition (Figure 3). Marketing and promotion

311 efforts are required to increase Chile’s market share and positioning, however it will be

312 of the utmost importance to ensure optimal handling from catch to destination market in

313 order to secure the supply of consistent, high-quality products.

314
315 5. Culture of native and introduced fish in Chile
316
317 The development of industrial aquaculture in Chile and the world, with a profile

318 of rapid technological development and a wide knowledge of the species cultivated,

319 requires adequate control of reproductive processes (Zohar and Mylonas, 2001). In

320 Chile, the principal species farmed are Atlantic salmon (Salmo salar), coho or Pacific

321 salmon (Oncorhynchus kisutch), rainbow trout (Oncorhynchus mykiss) and king

322 salmon (Oncorhynchus tschawytscha) (Tabla 2). Their production accounted for 57% of

323 all fisheries exports and 93% of aquaculture exports in 2004. Turbot (Psetta maxima),

324 cultivated in clay ponds, contributed 0.2% of the volume exported in 2004. Semi-

325 intensive aquaculture allows greater yield from cultivation areas, better growth and

326 larger production, whereas intensive aquaculture systems generate the largest returns in

327 export sales. Salmon is the single most important type of fish cultured under intensive

328 systems: breeding and smolt rearing are carried out in on-shore facilities, while

329 fattening takes place in floating pens in the sea. There are approximately 200 salmon

330 farms in Chile, most of them highly intensive in terms of capital and technology

331 (SENAPESCA 2013).

13
332 Successful intensive production of G. blacodes depends on high-quality,

333 sexually mature males and females. Sexual maturation and reproductive behaviour are

334 controlled by an interaction of endogenous and exogenous forces (Baggerman, 1980;

335 Baroiller and Jalabert, 1989). Cultivation of G. blacodes requires control of the

336 reproductive cycle and broodstock management systems; an alternative is to rear

337 broodstock in recirculating aquaculture systems adapted to their maturation cycles

338 (Migaud, 2006; Philipsen, 2008). Many studies have found broodstock nutrition to have

339 an effect on the reproductive quality of farmed fish (Henrotte et al., 2010). In G.

340 blacodes, as in most commercially important farmed finfish, hormone stimulation is

341 frequently applied to synchronize ovulation in females and/or stimulate semen release in

342 males. One strategy to protect the reproductive potential of this species is to evaluate

343 gamete quality, broodstock management and larval culture.

344 The main challenge for repopulating the environment with G. blacodes will be to

345 optimize and improve production in captivity to reduce mortality rates and maximize

346 survival of the larvae, as well as improving in vivo storage conditions. To produce G.

347 blacodes larvae with good yields, knowledge of the nutritional requirements of the

348 species particularly in the first larval stages is essential. It is known that the nutrition of

349 the broodstock has a direct impact on the quality of the larvae (Valdebenito et al., 2013).

350 Studies of artificial reproduction methods are still needed, with emphasis on broodstock

351 reproductive physiology to define the optimal individuals for stable mass production of

352 G. blacodes larvae and fry.

353
354 6. Concluding remarks

355 Recent investments in new technologies, fishing vessels, processing plants and

356 skilled human resources have made the Chilean fishing and aquaculture industries

357 highly competitive in the global context.

14
358 As aquaculture continues to grow and expand worldwide, it offers a promising

359 field for Chile, with great potential for expansion and development. The best

360 development prospects up to 2030 are in diversification into native species. Chile is

361 making concerted efforts to manage its fisheries in an appropriate, sustainable manner,

362 however, future developments in fisheries and aquaculture will require an increased

363 emphasis on sustainability. G. blacodes occupies a good position in the aquaculture

364 industry and market, and can become an important tool for promoting sustainable

365 development. This sector is in an expansion phase in Chile, with unsatisfied demand;

366 however, overexploitation has caused a significant impact on the biomass of G.

367 blacodes. The development of cultivation techniques is considered a good solution to

368 overexploitation. One of the strategies to protect the reproductive potential of this

369 species is to reassess capture size and average size at sexual maturity.

370 Undoubtedly, future economic growth should be based on incorporating added

371 value to the extracted fish, since the physical production of the species has not exceeded

372 catch volumes in recent years. Currently, there is little information on the reproductive

373 biology of G. blacodes, although it is thought to be easy to rear, having a low

374 metabolism compared to other fishes. The main challenge for G. blacodes will be to

375 optimise production in captivity and reduce the mortality rate by improving live storage

376 conditions from harvest to destination markets. Both the cultivation and capture of G.

377 blacodes require more attention in Chile, as landings of the species are dwindling. The

378 existing technologies for egg collection and rearing of larvae are leading to

379 improvements in G. blacodes farming, and progress has also been made in the

380 replacement of wild seed stock by fingerlings from production facilities.

381

15
382 The lack of data and statistics makes it difficult to assess all the shifts and trends

383 in G. blacodes aquaculture. Fisheries and aquaculture represent important industrial

384 sectors of the Chilean economy. The development and implementation of a particular

385 fishery or aquaculture management system have important environmental, economic

386 and social repercussions, consequently, major modifications are needed in the

387 productive practices of Chilean fisheries, including the extraction and processing of

388 caught fish as well as the development and expansion of aquaculture. Diversification

389 into culture systems for more native species could help to consolidate Chilean

390 aquaculture systems in economic, social and ecological terms.

391 Acknowledgements

392 This review paper was supported by the Fund for the Commission of Scientific and

393 Technological Development CONICYT, FONDECYT/POSTDOCTORAL (No.

394 3180765) FONDECYT/REGULAR (No. 1180387) Chile and Universidad de La

395 Frontera/ POSTDOCTORAL (KD).

396

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589

590

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606 Table legend

607 Table 1. Main Chilean export markets of Genypterus blacodes.

608

609 Table 2. Main fish cultured and traded in Chile.

610

611 Figure legends

612 Figure 1. A) Gographic distribution of Genypterus blacodes in South America and

613 southern Australia and around New Zealand (www.aquamaps.org); B) Gographic

614 distribution of Genypterus blacodes from North and South Area in Chile (Chong et al.,

615 2014 modified).

616

617 Figure 2. Total capture of Genypterus blacodes in Chile. (Chong et al., 2014 modified).

618

619 Figure 3. Illustration of a typical supply chain of Genypterus blacodes from catch to

620 market.

621

25
Table

Table 1. Main Chilean export markets of Genypterus blacodes.

Volume/tonnes
Year Frozen Live
2009 1.512 25
2010 1.195 36
2011 1.215 43
2012 823 27
2013 780 10
2014 263 4
Source: Secretary of Fisheries, Chile (Chong et al., 2014 modified).
Table

Table 2. Main fish cultured and traded in Chile.

Production system Traded
Species Cultured/wild-caught Fresh Frozen
Salmo Salar +/- + +
Genypterus blacodes -/+ + +
Genypterus capensis -/+ + +
Genypterus chilensis -/+ + +
Eleginops maclovinus +/+ + +
Merluccius gayi gayi -/+ + +
Merluccius australis -/+ + +
Micromesistius australis -/+ + +
Macruronus magellanicus -/+ + +
Brama australis -/+ + +
Dissostichus eleginoides +/+ + +
Seriolela porosa -/+ + +
Seriolella caerulea -/+ + +
Oncorhynchus mykiss +/+ + +
Oncorhynchus kisutch +/+ + +
Source: Secretary of Fisheries, Chile (www.subpesca.cl).
Figure
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*Declaration of Interest Statement

Declaration of interests

x The authors declare that they have no known competing financial interests or personal relationships
that could have appeared to influence the work reported in this paper.

☐The authors declare the following financial interests/personal relationships which may be considered
as potential competing interests: