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Advances in the aquatic sciences
RESEARCH ARTICLE

Stock structure of Pacific cod (Gadus macrocephalus) around the Korean Peninsula: an otolith microchemical perspective

Kali R. Stone https://orcid.org/0000-0002-4853-9035 A C , Craig R. Kastelle https://orcid.org/0000-0002-6681-1602 A , Irina M. Benson A , Thomas E. Helser A , Jonathan A. Short A and Sukyung Kang B
+ Author Affiliations
- Author Affiliations

A Resource Ecology and Fisheries Management Division, Alaska Fisheries Science Center, National Marine Fisheries Service, National Oceanic and Atmospheric Administration, 7600 Sand Point Way NE, Seattle, WA 98115, USA.

B Fisheries Resources Management Division, National Fisheries Research and Development Institute, 216 Gijanghaean-ro, Gijang-eup, Gijang-gun, Busan 46083, South Korea.

C Corresponding author. Email: kali.stone@noaa.gov

Marine and Freshwater Research 72(6) 774-786 https://doi.org/10.1071/MF20223
Submitted: 16 July 2020  Accepted: 24 September 2020   Published: 14 December 2020

Abstract

Sustainable management of fishery resources is predicated on a foundational understanding of the biogeography of fish stocks and the delineation of stocks into appropriate management units. Despite notable fluctuations in the catch of commercially valuable Pacific cod (Gadus macrocephalus) in South Korea, relatively little is known regarding the stock structure and migratory trends of this species in the region. Here, otolith microchemistry was used to evaluate the stock structure and ontogenetic migratory trends of Pacific cod from five spawning grounds around the Korean Peninsula. Statistically significant between-region discrimination was evident and resolved Pacific cod around the Korean Peninsula into two distinct stocks. Specimens were classified to region of capture using quadratic discriminant analysis of age-0 and capture elemental signatures with overall accuracies of 71.12 and 79.1% respectively, lending support to the notion that Pacific cod demonstrate natal philopatry. Analysis of the elemental signatures over the first 2 years of life exhibited clear trends indicative of shifts in habitat use, suggesting that trace elements function well as indicators of ontogenetic migration. Our work provides information that is complementary to other direct and indirect methods of monitoring migratory trends and delineating stocks, which are integral components of effective species and ecosystem management plans.

Keywords: fishery ecology, LA-ICP-MS, laser ablation inductively coupled plasma mass spectrometry, natal philopatry, otolith chemistry, otolith microchemistry, spawning site fidelity.


References

Alaska Fisheries Science Center (2020). New study shows pacific cod eggs are highly vulnerable to changes in bottom temperature. In NOAA Fisheries News. Available at https://www.fisheries.noaa.gov/feature-story/new-study-shows-pacific-cod-eggs-are-highly-vulnerable-changes-bottom-temperature [Verified 15 May 2020].

Bailon, M. X., David, A., Park, Y., Kim, E., and Hong, Y. (2018). Total mercury, methyl mercury, and heavy metal concentrations in Hyeongsan River and its tributaries in Pohang city, South Korea. Environmental Monitoring and Assessment 190, 274.
Total mercury, methyl mercury, and heavy metal concentrations in Hyeongsan River and its tributaries in Pohang city, South Korea.Crossref | GoogleScholarGoogle Scholar | 29644448PubMed |

Benson, I. M., Kastelle, C. R., Helser, T. E., Short, J. A., and Anderl, D. M. (2019). Age interpretation in eulachon (Thaleichthys pacificus) as suggested by otolith microchemical signatures. Environmental Biology of Fishes 102, 629–643.
Age interpretation in eulachon (Thaleichthys pacificus) as suggested by otolith microchemical signatures.Crossref | GoogleScholarGoogle Scholar |

Campana, S. E. (2005). Otolith elemental composition as a natural marker of fish stocks. In ‘Stock Identification Methods’. (Eds S. X. Cadrin, K. D. Friedland, and J. R. Waldman.) Chapter 12, pp. 227–245. (Academic Press.) https://doi.org/10.1016/B978-012154351-8/50013-7

Campana, S. E., and Thorrold, S. R. (2001). Otoliths, increments, and elements: keys to a comprehensive understanding of fish populations? Canadian Journal of Fisheries and Aquatic Sciences 58, 30–38.
Otoliths, increments, and elements: keys to a comprehensive understanding of fish populations?Crossref | GoogleScholarGoogle Scholar |

Cho, Y.-K., and Kim, K. (2000). Branching mechanism of the Tsushima Current in the Korea Strait. Journal of Physical Oceanography 30, 2788–2797.
Branching mechanism of the Tsushima Current in the Korea Strait.Crossref | GoogleScholarGoogle Scholar |

Choi, Y. M. (2013). Pacific cod (Gadus macrocephalus). In ‘Top 100 Items of Fisheries Products for Practical Research and Communication Promotion with Fishermen’. (Ed. Y. H. Jung.) Report number PR-2013-FR-007, pp. 182–204. (National Institute of Fisheries Science: Busan, South Korea.) [In Korean].

Chung, S., Kim, S., and Kang, S. (2013). Ecological relationship between environmental factors and Pacific cod (Gadus macrocephalus) catch in the southern East/Japan Sea. Animal Cells and Systems 17, 374–382.
Ecological relationship between environmental factors and Pacific cod (Gadus macrocephalus) catch in the southern East/Japan Sea.Crossref | GoogleScholarGoogle Scholar |

Cohen, J. (1960). A coefficient of agreement for nominal scales. Educational and Psychological Measurement 20, 37–46.
A coefficient of agreement for nominal scales.Crossref | GoogleScholarGoogle Scholar |

Collins, S. M., Bickford, N., McIntyre, P. B., Coulon, A., Ulseth, A. J., Taphorn, D. C., and Flecker, A. S. (2013). Population structure of a neotropical migratory fish: contrasting perspectives from genetics and otolith microchemistry. Transactions of the American Fisheries Society 142, 1192–1201.
Population structure of a neotropical migratory fish: contrasting perspectives from genetics and otolith microchemistry.Crossref | GoogleScholarGoogle Scholar |

Cunningham, K. M., Canino, M. F., Spies, I. B., and Hauser, L. (2009). Genetic isolation by distance and localized fjord population structure in Pacific cod (Gadus macrocephalus): limited effective dispersal in the northeastern Pacific Ocean. Canadian Journal of Fisheries and Aquatic Sciences 66, 153–166.
Genetic isolation by distance and localized fjord population structure in Pacific cod (Gadus macrocephalus): limited effective dispersal in the northeastern Pacific Ocean.Crossref | GoogleScholarGoogle Scholar |

de Mitcheson, Y. S. (2016). Mainstreaming fish spawning aggregations into fishery management calls for a precautionary approach. Bioscience 66, 295–306.
Mainstreaming fish spawning aggregations into fishery management calls for a precautionary approach.Crossref | GoogleScholarGoogle Scholar |

Elsdon, T. S., Wells, B. K., Campana, S. E., Gillanders, B. M., Jones, C. M., Limburg, K. E., Secor, D. H., Thorrold, S. R., and Walther, B. D. (2008). Otolith chemistry to describe movements and life-history parameters of fishes: hypotheses, assumptions, limitations and inferences. Oceanography and Marine Biology - an Annual Review 46, 297–330.
Otolith chemistry to describe movements and life-history parameters of fishes: hypotheses, assumptions, limitations and inferences.Crossref | GoogleScholarGoogle Scholar |

Feyrer, F., Hobbs, J., Baerwald, M., Sommer, T., Yin, Q.-Z., Clark, K., May, B., and Bennett, W. (2007). Otolith microchemistry provides information complementary to microsatellite DNA for a migratory fish. Transactions of the American Fisheries Society 136, 469–476.
Otolith microchemistry provides information complementary to microsatellite DNA for a migratory fish.Crossref | GoogleScholarGoogle Scholar |

Fisher, M. (2018). Korean stock structure and transoceanic divergence in Pacific cod (Gadus macrocephalus). M.Sc. Thesis, University of Washington, Seattle, WA, USA.

Gillanders, B. M. (2002). Temporal and spatial variability in elemental composition of otoliths: implications for determining stock identity and connectivity of populations. Canadian Journal of Fisheries and Aquatic Sciences 59, 669–679.
Temporal and spatial variability in elemental composition of otoliths: implications for determining stock identity and connectivity of populations.Crossref | GoogleScholarGoogle Scholar |

Gustafson, R. G., Lenarz, W. H., McCain, B. B., Schmitt, C. C., Grant, W. S., Builder, T. L., and Methot, R. D. (2000). Status review of Pacific hake, Pacific cod, and walleye pollock from Puget Sound, Washington. NOAA Technical Memorandum NMFS-NWFSC-44, US Department of Commerce, National Oceanic and Atmospheric Administration, National Marine Fisheries Service, Northwest Fisheries Science Center, Seattle, WA, USA.

Gwak, W. S., and Nakayama, K. (2011). Genetic variation and population structure of the Pacific cod Gadus macrocephalus in Korean waters revealed by mtDNA and msDNA markers. Fisheries Science 77, 945–952.
Genetic variation and population structure of the Pacific cod Gadus macrocephalus in Korean waters revealed by mtDNA and msDNA markers.Crossref | GoogleScholarGoogle Scholar |

Hinckley, S., Stockhausen, W. T., Coyle, K. O., Laurel, B. J., Gibson, G. A., Parada, C., Hermann, A. J., Doyle, M. J., Hurst, T. P., Punt, A. E., and Ladd, C. (2019). Connectivity between spawning and nursery areas for Pacific cod (Gadus macrocephalus) in the Gulf of Alaska. Deep-sea Research – II. Topical Studies in Oceanography 165, 113–126.
Connectivity between spawning and nursery areas for Pacific cod (Gadus macrocephalus) in the Gulf of Alaska.Crossref | GoogleScholarGoogle Scholar |

Hurst, T., Laurel, B., and Ciannelli, L. (2010). Ontogenetic patterns and temperature-dependent growth rates in early life stages of Pacific cod (Gadus macrocephalus). Fishery Bulletin 108, 382–392.

Hurst, T. P., Cooper, D. W., Duffy-Anderson, J. T., and Farley, E. V. (2014). Contrasting coastal and shelf nursery habitats of Pacific cod in the southeastern Bering Sea. ICES Journal of Marine Science 72, 515–527.
Contrasting coastal and shelf nursery habitats of Pacific cod in the southeastern Bering Sea.Crossref | GoogleScholarGoogle Scholar |

Izzo, C., Reis-Santos, P., and Gillanders, B. M. (2018). Otolith chemistry does not just reflect environmental conditions: A meta-analytic evaluation. 19, 441–454.
Otolith chemistry does not just reflect environmental conditions: A meta-analytic evaluation.Crossref | GoogleScholarGoogle Scholar |

Johnston, C. G., and Anderl, D. M. (2012). Pacific cod (Gadus macrocephalus). In ‘Age Determination Manual of the Alaska Fisheries Science Center Age and Growth Program’. (Eds M. E. Matta and D. K. Kimura.) NOAA Professional Paper NMFS 13, pp. 25–30. (US Department of Commerce, National Marine Fisheries Service: Seattle, WA, USA.)

Kastelle, C., Helser, T., McKay, J., Johnston, C., Anderl, D., Matta, M., and Nichol, D. (2017). Age validation of Pacific cod (Gadus macrocephalus) using high-resolution stable oxygen isotope (δ18O) chronologies in otoliths. Fisheries Research 185, 43–45.
Age validation of Pacific cod (Gadus macrocephalus) using high-resolution stable oxygen isotope (δ18O) chronologies in otoliths.Crossref | GoogleScholarGoogle Scholar |

Kent, A. J. R., and Ungerer, C. A. A. (2006). Analysis of light lithophile elements (Li, Be, B) by laser ablation ICP-MS: comparison between magnetic sector and quadrupole ICP-MS. The American Mineralogist 91, 1401–1411.
Analysis of light lithophile elements (Li, Be, B) by laser ablation ICP-MS: comparison between magnetic sector and quadrupole ICP-MS.Crossref | GoogleScholarGoogle Scholar |

Kerr, L. A., and Campana, S. E. (2014). Chapter eleven – chemical composition of fish hard parts as a natural marker of fish stocks. In ‘Stock Identification Methods: Applications in Fishery Science’. (Eds S. X. Cadrin, L. A. Kerr, and S. Mariani.) pp. 205–234. (Academic Press: San Diego, CA, USA.) https://doi.org/10.1016/B978-0-12-397003-9.00011-4

Kim, K.-R., Cho, Y.-K., Kang, D.-J., and Ki, J.-H. (2005). The origin of the Tsushima Current based on oxygen isotope measurement. Geophysical Research Letters 32, L03602.
The origin of the Tsushima Current based on oxygen isotope measurement.Crossref | GoogleScholarGoogle Scholar |

Kim, M.-J., An, H. S., and Choi, K.-H. (2010). Genetic characteristics of Pacific cod populations in Korea based on microsatellite markers. Fisheries Science 76, 595–603.
Genetic characteristics of Pacific cod populations in Korea based on microsatellite markers.Crossref | GoogleScholarGoogle Scholar |

Laikre, L., Palm, S., and Ryman, N. (2005). Genetic population structure of fishes: implications for coastal zone management. Ambio 34, 111–119.
Genetic population structure of fishes: implications for coastal zone management.Crossref | GoogleScholarGoogle Scholar | 15865307PubMed |

Laurel, B. J., Hurst, T. P., Copeman, L. A., and Davis, M. W. (2008). The role of temperature on the growth and survival of early and late hatching Pacific cod larvae (Gadus macrocephalus). Journal of Plankton Research 30, 1051–1060.
The role of temperature on the growth and survival of early and late hatching Pacific cod larvae (Gadus macrocephalus).Crossref | GoogleScholarGoogle Scholar |

Laurel, B. J., Ryer, C. H., Knoth, B., and Stoner, A. W. (2009). Temporal and ontogenetic shifts in habitat use of juvenile Pacific cod (Gadus macrocephalus). Journal of Experimental Marine Biology and Ecology 377, 28–35.
Temporal and ontogenetic shifts in habitat use of juvenile Pacific cod (Gadus macrocephalus).Crossref | GoogleScholarGoogle Scholar |

Laurel, B. J., Copeman, L. A., and Parrish, C. C. J. M. B. (2012). Role of temperature on lipid/fatty acid composition in Pacific cod (Gadus macrocephalus) eggs and unfed larvae. Marine Biology 159, 2025–2034.
Role of temperature on lipid/fatty acid composition in Pacific cod (Gadus macrocephalus) eggs and unfed larvae.Crossref | GoogleScholarGoogle Scholar |

Laurel, B. J., Spencer, M., Iseri, P., and Copeman, L. A. J. P. B. (2016). Temperature-dependent growth and behavior of juvenile Arctic cod (Boreogadus saida) and co-occurring North Pacific gadids. Polar Biology 39, 1127–1135.
Temperature-dependent growth and behavior of juvenile Arctic cod (Boreogadus saida) and co-occurring North Pacific gadids.Crossref | GoogleScholarGoogle Scholar |

Lee, S.-G., and Rahimi Midani, A. (2014). National comprehensive approaches for rebuilding fisheries in South Korea. Marine Policy 45, 156–162.
National comprehensive approaches for rebuilding fisheries in South Korea.Crossref | GoogleScholarGoogle Scholar |

Lee, M., Bae, W., Chung, J., Jung, H.-S., and Shim, H. (2008). Seasonal and spatial characteristics of seawater and sediment at Youngil Bay, southeast coast of Korea. Marine Pollution Bulletin 57, 325–334.
Seasonal and spatial characteristics of seawater and sediment at Youngil Bay, southeast coast of Korea.Crossref | GoogleScholarGoogle Scholar | 18514230PubMed |

Lee, K., Cha, H. K., Kim, Y. H., Lee, J. Y., and Jeong, S. G. (2016). Comparison of biological characteristics of East Sea and Yellow Sea cod (Gadus macrocephalus). Korean Journal of Fisheries and Aquatic Sciences 49, 499–508.
Comparison of biological characteristics of East Sea and Yellow Sea cod (Gadus macrocephalus).Crossref | GoogleScholarGoogle Scholar |

Lewandoski, S. A., Bishop, M. A., and McKinzie, M. K. (2018). Evaluating Pacific cod migratory behavior and site fidelity in a fjord environment using acoustic telemetry. Canadian Journal of Fisheries and Aquatic Sciences 75, 2084–2095.
Evaluating Pacific cod migratory behavior and site fidelity in a fjord environment using acoustic telemetry.Crossref | GoogleScholarGoogle Scholar |

Low, L.-L. (2014). International affairs and research collaboration: NOAA–Korea joint project agreement. (NOAA Fisheries, Alaska Fisheries Science Center: Seattle, WA, USA.) Available at https://archive.fisheries.noaa.gov/afsc/Quarterly/jas2014/divrptsREFM10.htm [Verified 1 January 2020].

Matta, M. E., Miller, J. A., Short, J. A., Helser, T. E., Hurst, T. P., Rand, K. M., and Ormseth, O. A. (2019). Spatial and temporal variation in otolith elemental signatures of age-0 Pacific cod (Gadus macrocephalus) in the Gulf of Alaska. Deep-sea Research – II. Topical Studies in Oceanography 165, 268–279.
Spatial and temporal variation in otolith elemental signatures of age-0 Pacific cod (Gadus macrocephalus) in the Gulf of Alaska.Crossref | GoogleScholarGoogle Scholar |

McGarigal, K., Cushman, S., and Stafford, S. (2000). ‘Multivariate Statistics for Wildlife and Ecology Research.’ (Springer: New York, NY, USA.)

Miller, J. A. (2007). Scales of variation in otolith elemental chemistry of juvenile staghorn sculpin (Leptocottus armatus) in three Pacific Northwest estuaries. Marine Biology 151, 483–494.
Scales of variation in otolith elemental chemistry of juvenile staghorn sculpin (Leptocottus armatus) in three Pacific Northwest estuaries.Crossref | GoogleScholarGoogle Scholar |

Miller, J. A., DiMaria, R. A., and Hurst, T. P. (2016). Patterns of larval source distribution and mixing in early life stages of Pacific cod (Gadus macrocephalus) in the southeastern Bering Sea. Deep-sea Research – II. Topical Studies in Oceanography 134, 270–282.
Patterns of larval source distribution and mixing in early life stages of Pacific cod (Gadus macrocephalus) in the southeastern Bering Sea.Crossref | GoogleScholarGoogle Scholar |

Nichol, D. G., Kotwicki, S., and Zimmermann, M. (2013). Diel vertical migration of adult Pacific cod Gadus macrocephalus in Alaska. Journal of Fish Biology 83, 170–189.
Diel vertical migration of adult Pacific cod Gadus macrocephalus in Alaska.Crossref | GoogleScholarGoogle Scholar | 23808699PubMed |

Ra, K., Kim, E.-S., Kim, K.-T., Kim, J.-K., Lee, J.-M., and Choi, J.-Y. (2013). Assessment of heavy metal contamination and its ecological risk in the surface sediments along the coast of Korea. Journal of Coastal Research 65, 105–110.
Assessment of heavy metal contamination and its ecological risk in the surface sediments along the coast of Korea.Crossref | GoogleScholarGoogle Scholar |

Rand, K. M., Munro, P., Neidetcher, S. K., and Nichol, D. G. (2014). Observations of seasonal movement from a single tag release group of Pacific cod in the eastern Bering Sea. Marine and Coastal Fisheries 6, 287–296.
Observations of seasonal movement from a single tag release group of Pacific cod in the eastern Bering Sea.Crossref | GoogleScholarGoogle Scholar |

Song, N., Liu, M., Yanagimoto, T., Sakurai, Y., Han, Z.-Q., and Gao, T.-X. (2016). Restricted gene flow for Gadus macrocephalus from Yellow Sea based on microsatellite markers: geographic block of Tsushima Current. International Journal of Molecular Sciences 17, 467.
Restricted gene flow for Gadus macrocephalus from Yellow Sea based on microsatellite markers: geographic block of Tsushima Current.Crossref | GoogleScholarGoogle Scholar | 27043534PubMed |

Spies, I., and Punt, A. E. (2015). The utility of genetics in marine fisheries management: a simulation study based on Pacific cod off Alaska. Canadian Journal of Fisheries and Aquatic Sciences 72, 1415–1432.
The utility of genetics in marine fisheries management: a simulation study based on Pacific cod off Alaska.Crossref | GoogleScholarGoogle Scholar |

Stoner, A. W., Laurel, B. J., and Hurst, T. P. (2008). Using a baited camera to assess relative abundance of juvenile Pacific cod: field and laboratory trials. Journal of Experimental Marine Biology and Ecology 354, 202–211.
Using a baited camera to assess relative abundance of juvenile Pacific cod: field and laboratory trials.Crossref | GoogleScholarGoogle Scholar |

Stroganov, A., and Orlov, A. (2012). Special characteristics of the formation of population structure in Pacific cod. In ‘Advances in Zoology Research’. (Ed. O. P. Jenkins.) pp. 169–185. (NOVA Science Publishers: New York, NY, USA.)

Sturrock, A. M., Trueman, C. N., Darnaude, A. M., and Hunter, E. (2012). Can otolith elemental chemistry retrospectively track migrations in fully marine fishes? Journal of Fish Biology 81, 766–795.
Can otolith elemental chemistry retrospectively track migrations in fully marine fishes?Crossref | GoogleScholarGoogle Scholar | 22803735PubMed |

Sturrock, A. M., Trueman, C. N., Cooper, M. J., and Hunter, E. (2014). Physiological influences can outweigh environmental signals in otolith microchemistry research. Marine Ecology Progress Series 500, 245–264.
Physiological influences can outweigh environmental signals in otolith microchemistry research.Crossref | GoogleScholarGoogle Scholar |

Suda, A., Nagata, N., Sato, A., Narimatsu, Y., Nadiatul, H. H., and Kawata, M. (2017). Genetic variation and local differences in Pacific cod Gadus macrocephalus around Japan. Journal of Fish Biology 90, 61–79.
Genetic variation and local differences in Pacific cod Gadus macrocephalus around Japan.Crossref | GoogleScholarGoogle Scholar | 27723107PubMed |

Svedäng, H., Stål, J., Sterner, T., and Cardinale, M. (2010). Consequences of subpopulation structure on fisheries management: cod (Gadus morhua) in the Kattegat and Öresund (North Sea). Reviews in Fisheries Science 18, 139–150.
Consequences of subpopulation structure on fisheries management: cod (Gadus morhua) in the Kattegat and Öresund (North Sea).Crossref | GoogleScholarGoogle Scholar |

Tanner, S. E., Reis-Santos, P., and Cabral, H. N. (2016). Otolith chemistry in stock delineation: a brief overview, current challenges and future prospects. Fisheries Research 173, 206–213.
Otolith chemistry in stock delineation: a brief overview, current challenges and future prospects.Crossref | GoogleScholarGoogle Scholar |

Thomas, O. R. B., and Swearer, S. E. (2019). Otolith biochemistry – a review. Reviews in Fisheries Science & Aquaculture 27, 458–489.
Otolith biochemistry – a review.Crossref | GoogleScholarGoogle Scholar |

Vareda, J. P., Valente, A. J. M., and Durães, L. (2019). Assessment of heavy metal pollution from anthropogenic activities and remediation strategies: a review. Journal of Environmental Management 246, 101–118.
Assessment of heavy metal pollution from anthropogenic activities and remediation strategies: a review.Crossref | GoogleScholarGoogle Scholar | 31176176PubMed |

Weidman, C. R., and Millner, R. (2000). High-resolution stable isotope records from North Atlantic cod. Fisheries Research 46, 327–342.
High-resolution stable isotope records from North Atlantic cod.Crossref | GoogleScholarGoogle Scholar |

Wilson, M. T., Dougherty, A., Matta, M. E., Mier, K. L., and Miller, J. A. (2018). Otolith chemistry of juvenile walleye pollock Gadus chalcogrammus in relation to regional hydrography: evidence of spatially split cohorts. Marine Ecology Progress Series 588, 163–178.
Otolith chemistry of juvenile walleye pollock Gadus chalcogrammus in relation to regional hydrography: evidence of spatially split cohorts.Crossref | GoogleScholarGoogle Scholar |

Wu, R., Lin, J., and Li, B. (2016). Spatial and temporal variability of sea surface temperature in eastern marginal seas of China. Advances in Meteorology 2016, 1–9.
Spatial and temporal variability of sea surface temperature in eastern marginal seas of China.Crossref | GoogleScholarGoogle Scholar |

Ying, Y., Chen, Y., Lin, L., and Gao, T. (2011). Risks of ignoring fish population spatial structure in fisheries management. Canadian Journal of Fisheries and Aquatic Sciences 68, 2101–2120.
Risks of ignoring fish population spatial structure in fisheries management.Crossref | GoogleScholarGoogle Scholar |

Zhang, C. I. (1984). Pacific cod of South Korean waters. International North Pacific Fisheries Commision 42, 116–129.