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RESEARCH ARTICLE

Advancing fish early life-history research: daily age determination of walleye pollock (Gadus chalcogrammus) with Fourier transform–near-infrared spectroscopy

Esther D. Goldstein https://orcid.org/0000-0002-0499-7288 A * , Mary E. Matta A , Charles D. Waters https://orcid.org/0000-0003-4606-3202 B , Heather K. Fulton-Bennett https://orcid.org/0000-0002-5938-9714 B , Brenna C. Hsieh A , Craig R. Kastelle https://orcid.org/0000-0002-6681-1602 A , Johanna J. Vollenweider B , Jakub T. Sliwinski C and Thomas E. Helser A
+ Author Affiliations
- Author Affiliations

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

B Auke Bay Laboratories, Alaska Fisheries Science Center, NMFS, NOAA, 17109 Point Lena Loop Road, Juneau, AK 99801, USA.

C Trace Lab, University of Washington School of Oceanography, 1501 NE Boat Street, Seattle, WA 98105, USA.

* Correspondence to: esther.goldstein@noaa.gov

Handling Editor: Haseeb Randhawa

Marine and Freshwater Research 76, MF25053 https://doi.org/10.1071/MF25053
Submitted: 7 March 2025  Accepted: 28 May 2025  Published: 25 June 2025

© 2025 The Author(s) (or their employer(s)). Published by CSIRO Publishing

Abstract

Context

Fish daily age information provides insight into growth, phenology and recruitment but is rarely incorporated into the decision-making process for management because of labor-intensive data collection.

Aims

We tested Fourier transform–near-infrared (FT-NIR) spectroscopy as a rapid method (~1.0 v. ~60 min per otolith for microscopic methods) to determine daily age of walleye pollock (pollock; Gadus chalcogrammus), which supports an important fishery.

Methods

We reared two annual cohorts of pollock over 3 years to accrue a comprehensive dataset for calibration models.

Key results

FT-NIR spectroscopy provided an improvement over predicting daily age from otolith weight, fish weight or fish length. A calibration model developed using microscopically aged fish had a root mean square error (RMSE) from cross validation (CV) of 12.2 days. Models developed from the full dataset with ages assigned from cohort hatch dates had a test dataset RMSE of 35.4–40.7 days and a CV RMSE of 29.4–36.9 days.

Conclusions

Spectra were affected by size-at-age, which could affect model efficacy because growth and size are influenced by environmental variability. This can be addressed with robust calibration datasets, model testing and model updating.

Implications

FT-NIR spectroscopy age prediction is likely to be sufficient to capture population-scale shifts in hatch dates of pollock.

Keywords: daily age, Fourier transform–near-infrared spectroscopy, Gadus chalcogrammus, Gulf of Alaska, LA-ICP-MS, otolith, otolith microchemistry, walleye pollock.

References

Almeida LZ, Laurel BJ, Thalmann HL, Miller JA (2024) Warmer, earlier, faster: cumulative effects of Gulf of Alaska heatwaves on the early life history of Pacific cod. Elementa: Science of the Anthropocene 12, 00050.
| Crossref | Google Scholar |

Arrington MB, Helser TE, Benson IM, Essington TE, Matta ME, Punt AE (2022) Rapid age estimation of longnose skate (Raja rhina) vertebrae using near-infrared spectroscopy. Marine and Freshwwater Research 73, 71-80.
| Crossref | Google Scholar |

Atkinson D (1994) Temperature and organism size—a biological law for ectotherms? Advances in Ecological Research 25, 1-58.
| Google Scholar |

Bailey KM, Macklin SA, Reed RK, Brodeur RD, Ingraham WJ, Piatt JF, Shima M, Francis RC, Anderson PJ, Royer TC, Hollowed AB, Somerton DA, Wooster WS (1995) ENSO events in the northern Gulf of Alaska, and effects on selected marine fisheries. Vol. 36, pp. 78–96. (California Cooperative Oceanic Fisheries Investigations) Available at http://pubs.er.usgs.gov/publication/2002017

Barnett BK, Benson IM, Helser TE, Lowerre-Barieri SK, Menendez HS (2024) Investigating the use of FT-NIR spectroscopy to age gag grouper (Mycteroperca microlepis), a protogynous hermaphroditic species. In ‘Proceedings of the Fourth Research Workshop on the Rapid Estimation of Fish Age Using Fourier Transform Near Infrared Spectroscopy’, 3–7 April 2023, Seattle, WA, USA. (Eds ME Matta, TE Helser) AFSC Processed Report 2024-01, pp. 99–126. (Alaska Fisheries Science Center, NOAA, National Marine Fisheries Service: Seattle, WA, USA) Available at https://repository.library.noaa.gov/view/noaa/56377

Beć KB, Huck CW (2019) Breakthrough potential in near-infrared spectroscopy: spectra simulation. A review of recent developments. Frontiers in Chemsitry 7, 48.
| Crossref | Google Scholar |

Benson IM, Helser TE, Marchetti G, Barnett BK (2023) The future of fish age estimation: deep machine learning coupled with Fourier transform near-infrared spectroscopy of otoliths. Canadian Journal of Fisheries and Aquatic Sciences 80, 1482-1494.
| Crossref | Google Scholar |

Benson IM, Helser TE, Barnett BK (2024) Fourier transform near infrared spectroscopy of otoliths coupled with deep learning improves age prediction for long-lived northern rockfish. Fisheries Research 278, 107116.
| Crossref | Google Scholar |

Bobelyn E, Serban A-S, Nicu M, Lammertyn J, Nicolai BM, Saeys W (2010) Postharvest quality of apple predicted by NIR-spectroscopy: study of the effect of biological variability on spectra and model performance. Postharvest Biology and Technology 55, 133-143.
| Crossref | Google Scholar |

Bowker AH (1948) A test for symmetry in contingency tables. Journal of the American Statistical Association 43, 572-574.
| Crossref | Google Scholar |

Brown AL, Busby MS, Mier KL (2001) Walleye pollock Theragra chalcogramma during transformation from the larval to juvenile stage: otolith and osteological development. Marine Biology 139, 845-851.
| Crossref | Google Scholar |

Campana SE (1992) Measurement and interpretation of the microstructure of fish otoliths. In ‘Otolith microstructure examination and analysis’. (Eds DK Stevenson, SE Campana) Canadian Special Publication of Fisheries and Aquatic Sciences 117, pp. 59–71. (Canadian Special Publication of Fisheries and Aquatic Sciences: Ottawa, ON, Canada) Available at https://publications.gc.ca/collections/collection_2016/mpo-dfo/Fs41-31-117-eng.pdf

Campana SE (2001) Accuracy, precision and quality control in age determination, including a review of the use and abuse of age validation methods. Journal of Fish Biology 59, 197-242.
| Crossref | Google Scholar |

Campana SE, Jones CM (1992) Analysis of otolith microstructure data. In ‘Otolith microstructure examination and analysis’. (Eds DK Stevenson, SE Campana) Canadian Special Publication of Fisheries and Aquatic Sciences 117, pp. 73–100. (Canadian Special Publication of Fisheries and Aquatic Sciences: Ottawa, ON, Canada) Available at https://publications.gc.ca/collections/collection_2016/mpo-dfo/Fs41-31-117-eng.pdf

Chang WYB (1982) A statistical method for evaluating the reproducibility of age determination. Canadian Journal of Fisheries and Aquatic Sciences 39, 1208-1210.
| Crossref | Google Scholar |

Chang M-Y, Geffen AJ (2013) Taxonomic and geographic influences on fish otolith microchemistry. Fish and Fisheries 14, 458-492.
| Crossref | Google Scholar |

Couture JJ, Singh A, Rubert-Nason KF, Serbin SP, Lindroth RL, Townsend PA (2016) Spectroscopic determination of ecologically relevant plant secondary metabolites. Methods in Ecology and Evolution 7, 1402-1412.
| Crossref | Google Scholar |

Dahl K, O’Malley J., Barnett B, Kline B, Widdrington J (2024) Otolith morphometry and Fourier transform near-infrared (FT-NIR) spectroscopy as tools to discriminate archived otoliths of newly detected cryptic species, Etelis carbunculus and Etelis boweni. Fisheries Research 272, 106927.
| Crossref | Google Scholar |

Dougherty AB (2008) Daily and sub-daily otolith increments of larval and juvenile walleye pollock, Theragra chalcogramma (Pallas), as validated by alizarin complexone experiments. Fisheries Research 90, 271-278.
| Crossref | Google Scholar |

Dougherty AB, Bailey KM, Mier KL (2007) Interannual differences in growth and hatch date distributions of age-0 year walleye pollock Theragra chalcogramma (Pallas) sampled from the Shumagin Islands region of the Gulf of Alaska, 1985–2001. Journal of Fish Biology 71, 763-780.
| Crossref | Google Scholar |

Dougherty A, Bailey K, Vance T, Cheng W (2012) Underlying causes of habitat-associated differences in size of age-0 walleye pollock (Theragra chalcogramma) in the Gulf of Alaska. Marine Biology 159, 1733-1744.
| Crossref | Google Scholar |

Doyle MJ, Mier KL (2015) Early life history pelagic exposure profiles of selected commercially important fish species in the Gulf of Alaska. Deep-Sea Research – II. Topical Studies in Oceanography 132, 162-193.
| Crossref | Google Scholar |

Duffy-Anderson JT, Barbeaux SJ, Farley E, Heintz R, Horne JK, Parker-Stetter SL, Petrik C, Siddon EC, Smart TI (2016) The critical first year of life of walleye pollock (Gadus chalcogrammus) in the eastern Bering Sea: implications for recruitment and future research. Deep-Sea Research – II. Topical Studies in Oceanography 134, 283-301.
| Crossref | Google Scholar |

Evans GT, Hoenig JM (1998) Testing and viewing symmetry in contingency tables, with application to readers of fish ages. Biometrics 54, 620-629.
| Crossref | Google Scholar |

Farrés M, Platikanov S, Tsakovski S, Tauler R (2015) Comparison of the variable importance in projection (VIP) and of the selectivity ratio (SR) methods for variable selection and interpretation. Journal of Chemometrics 29, 528-536.
| Crossref | Google Scholar |

Fuller K (2021) Exploring effects of sample storage preparation and tissue type on Fourier transform–near infrared spectroscopy (FT-NIRS) ageing across fish taxa. PhD thesis, University of South Carolina, Columbia, SC, USA.

Goldstein ED, Helser TE, Vollenweider JJ, Sreenivasan A, Sewall FF (2021) Rapid and reliable assessment of fish physiological condition for fisheries research and management using Fourier transform near-infrared spectroscopy. Frontiers in Marine Science 8, 690934.
| Crossref | Google Scholar |

Haenlein M, Kaplan AM (2004) A beginner’s guide to partial least squares analysis. Understanding Statistics 3, 283-297.
| Crossref | Google Scholar |

Heintz RA, Siddon EC, Farley EV Jr, Napp JM (2013) Correlation between recruitment and fall condition of age-0 pollock (Theragra chalcogramma) from the eastern Bering Sea under varying climate conditions. Deep-Sea Research – II. Topical Studies in Oceanography 94, 150-156.
| Crossref | Google Scholar |

Helser TE, Benson I, Erickson J, Healy J, Kastelle C, Short JA (2019) A transformative approach to ageing fish otoliths using Fourier transform near infrared spectroscopy: a case study of eastern Bering Sea walleye pollock (Gadus chalcogrammus). Canadian Journal of Fisheries and Aquatic Sciences 76, 780-789.
| Crossref | Google Scholar |

Houde ED (2016) Recruitment variability. In ‘Fish reproductive biology’. (Eds T Jakobsen, MJ Fogarty, BA Megrey, E Moksness) pp. 98–187. (Wiley–Blackwell) doi:10.1002/9781118752739.ch3

Izzo C, Reis-Santos P, Gillanders BM (2018) Otolith chemistry does not just reflect environmental conditions: a meta-analytic evaluation. Fish and Fisheries 19, 441-454.
| Crossref | Google Scholar |

Johnson DW, Grorud-Colvert K, Sponaugle S, Semmens BX (2014) Phenotypic variation and selective mortality as major drivers of recruitment variability in fishes. Ecology Letters 17, 743-755.
| Crossref | Google Scholar | PubMed |

Jones CM (2013) Growth and mortality of pre and post-settlement age-0 red snapper, Lutjanus campechanus (Poey 1860), in the Gulf of Mexico. PhD thesis, University of South Alabama, AL, USA.

Kimura D, Anderl DM, Goetz BJ (2007) Seasonal marginal growth on otoliths of seven Alaska groundfish species support the existence of annual patterns. Alaska Fishery Research Bulletin 12, 243-251.
| Google Scholar |

Kucheryavskiy S (2020) mdatools - R package for chemometrics. Chemometrics and Intelligent Laboratory Systems 198, 103937.
| Crossref | Google Scholar |

Leahy SM, Jerry DR, Wedding BBC, Robins JB, Wright CL, Sadekov A, Boyle S, Jones DB, Williams SM, Grauf S, Pavich L, McLennan M, Sellin MJ, Goldsbury JA, Saunders RJ (2023) Multimethod approach to advance provenance determination of fish in stocked systems. Canadian Journal Fisheries Aquatic Sciences 80, 1410-1424.
| Crossref | Google Scholar |

Levine M, Jones DT, McGowan DW (2024) Results of the acoustic-trawl survey of walleye pollock (Gadus chalcogrammus) in the Gulf of Alaska, June–July 2021 (DY2021-04). AFSC Processed Report 2024-07. (Alaska Fisheries Science Center, National Marine Fisheries Service, National Oceanic and Atmospheric Administration: Seattle, WA, USA) Available at https://repository.library.noaa.gov/view/noaa/61451/noaa_61451_DS1.pdf

Li B, Shertzer KW, Lynch PD, Ianelli JN, Legault CM, Williams EH, Methot RD, Brooks EN, Deroba JJ, Berger AM, Sagarese SR, Brodziak JKT, Taylor IG, Karp MA, Wetzel CR, Supernaw M (2021) A comparison of 4 primary age-structured stock assessment models used in the United States. Fishery Bulletin 119, 149-167.
| Crossref | Google Scholar |

Litzow MA, Bailey KM, Prahl FG, Heintz R (2006) Climate regime shifts and reorganization of fish communities: the essential fatty acid limitation hypothesis. Marine Ecology Progress Series 315, 1-11.
| Crossref | Google Scholar |

Litzow MA, Ciannelli L, Puerta P, Wettstein JJ, Rykaczewski RR, Opiekun M (2018) Non-stationary climate–salmon relationships in the Gulf of Alaska. Proceedings of the Royal Society B: Biological Sciences 285, 20181855.
| Crossref | Google Scholar |

Matta ME, Helser TE (Eds) (2024) ‘Proceedings of the Fourth Research Workshop on the Rapid Estimation of Fish Age Using Fourier Transform Near Infrared Spectroscopy’, 3–7 April 2023, Seattle, WA, USA. AFSC Processed Report 2024-01. (Alaska Fisheries Science Center, NOAA, National Marine Fisheries Service: Seattle, WA, USA) Available at https://repository.library.noaa.gov/view/noaa/56377

Matta ME, Kimura DK (Eds) (2012) Age determination manual of the Alaska Fisheries Science Center Age and Growth Program. NOAA Professional Paper NMFS 13, United States Department of Commerce.

Mayerhöfer TG, Pahlow S, Popp J (2020) The Bouguer–Beer–Lambert law: shining light on the obscure. ChemPhysChem 21, 2029-2046.
| Crossref | Google Scholar | PubMed |

Mazur MM, Wilson MT, Dougherty AB, Buchheister A, Beauchamp DA (2007) Temperature and prey quality effects on growth of juvenile walleye pollock Theragra chalcogramma (Pallas): a spatially explicit bioenergetics approach. Journal of Fish Biology 70, 816-836.
| Crossref | Google Scholar |

McBride RS (2015) Diagnosis of paired age agreement: a simulation of accuracy and precision effects. ICES Journal of Marine Science 72, 2149-2167.
| Crossref | Google Scholar |

Monnahan CC (2024) Toward good practices for Bayesian data-rich fisheries stock assessments using a modern statistical workflow. Fisheries Research 275, 107024.
| Crossref | Google Scholar |

Monnahan CC, Adams GD, Ferriss BE, Shotwell SK, McKelvey DR, McGowan DW (2023) Assessment of the walleye pollock stock in the Gulf of Alaska. North Pacific Fishery Management Council, Anchorage, AK, USA.

Muhling BA, Brodie S, Smith JA, Tommasi D, Gaitan CF, Hazen EL, Jacox MG, Auth TB, Brodeur RD (2020) Predictability of species distributions deteriorates under novel environmental conditions in the California Current System. Frontiers in Marine Science 7, 589.
| Crossref | Google Scholar |

Nishimura A, Yamada J (1988) Geographical differences in early growth of walleye pollock Theragra chalcogramma, estimated by back-calculation of otolith daily growth increments. Marine Biology 97, 459-465.
| Crossref | Google Scholar |

Oke KB, Mueter F, Litzow MA (2022) Warming leads to opposite patterns in weight-at-age for young versus old age classes of Bering Sea walleye pollock. Canadian Journal of Fisheries and Aquatic Sciences 79, 1655-1666.
| Crossref | Google Scholar |

Ono K, Licandeo R, Muradian ML, Cunningham CJ, Anderson SC, Hurtado-Ferro F, Johnson KF, McGilliard CR, Monnahan CC, Szuwalski CS, Vero JL, Vert-Pre KA, Whitten AR, Punt AE (2015) The importance of length and age composition data in statistical age-structured models for marine species. ICES Journal of Marine Science 72, 31-43.
| Crossref | Google Scholar |

Panfili J, de Pontual H, Troadec H, Wright PJ (2002) ‘Manual of fish sclerochronology.’ (Ifremer-IRD: Brest, France)

Passerotti MS, Jones CM, Swanson CE, Quattro JM (2020a) Fourier-transform near infrared spectroscopy (FT-NIRS) rapidly and non-destructively predicts daily age and growth in otoliths of juvenile red snapper Lutjanus campechanus (Poey, 1860). Fisheries Research 223, 105439.
| Crossref | Google Scholar |

Passerotti MS, Helser TE, Benson IM, Barnett BK, Ballenger JC, Bubley WJ, Reichert MJM, Quattro JM (2020b) Age estimation of red snapper (Lutjanus campechanus) using FT-NIR spectroscopy: feasibility of application to production ageing for management. ICES Journal of Marine Science 77, 2144-2156.
| Crossref | Google Scholar |

Passerotti MS, Reichert MJM, Robertory BA, Marsh Z, Stefik M, Quattro JM (2022) Physicochemical mechanisms of FT-NIRS age prediction in fish otoliths. Marine and Freshwater Research 73, 846-865.
| Crossref | Google Scholar |

Paton C, Hellstrom J, Paul B, Woodhead J, Hergt J (2011) Iolite: freeware for the visualisation and processing of mass spectrometric data. Journal of Analytical Atomic Spectrometry 26, 2508-2518.
| Crossref | Google Scholar |

Pomerantsev AL, Rodionova OY (2014) Concept and role of extreme objects in PCA/SIMCA. Journal of Chemometrics 28, 429-438.
| Crossref | Google Scholar |

Rankin TL, Sponaugle S (2011) Temperature influences selective mortality during the early life stages of a coral reef fish. PLoS ONE 6, e16814.
| Crossref | Google Scholar | PubMed |

Rinnan Å, van den Berg F, Engelsen SB (2009) Review of the most common pre-processing techniques for near-infrared spectra. TrAC Trends in Analytical Chemistry 28, 1201-1222.
| Crossref | Google Scholar |

Rodionova OY, Pomerantsev AL (2020) Detection of outliers in projection-based modeling. Analytical Chemistry 92, 2656-2664.
| Crossref | Google Scholar | PubMed |

Rogers LA, Dougherty AB (2019) Effects of climate and demography on reproductive phenology of a harvested marine fish population. Global Change Biology 25, 708-720.
| Crossref | Google Scholar | PubMed |

Rogers LA, Monnahan CC, Williams K, Jones DT, Dorn MW (2024) Climate-driven changes in the timing of spawning and the availability of walleye pollock (Gadus chalcogrammus) to assessment surveys in the Gulf of Alaska. ICES Journal of Marine Science 82, fsae005.
| Crossref | Google Scholar |

Saas E, Dorval E, Porzio DL, Schwartzkopf BD (2024) Development of methods for collecting spectral data from Fourier transform near-infrared spectroscopy to age three small coastal pelagic species (Pacific sardine, Pacific mackerel, northern anchovy) in the northeast Pacific Ocean. NOAA Technical Memorandum, NOAA-TM-NMFS-SWFSC-694, United States Department of Commerce.

Searcy SP, Sponaugle S (2001) Selective mortality during the larval–juvenile transition in two coral reef fishes. Ecology 82, 2452-2470.
| Crossref | Google Scholar |

Searcy SP, Eggleston DB, Hare JA (2007) Is growth a reliable indicator of habitat quality and essential fish habitat for a juvenile estuarine fish? Canadian Journal of Fisheries and Aquatic Sciences 64, 681-691.
| Crossref | Google Scholar |

Shima M, Hollowed AB, VanBlaricom GR (2002) Changes over time in the spatial distribution of walleye pollock (Theragra chalcogramma) in the Gulf of Alaska, 1984–1996. Fishery Bulletin 100, 307-323.
| Google Scholar |

Shotwell SK, Pirtle JL, Watson JT, Deary AL, Doyle MJ, Barbeaux SJ, Dorn MW, Gibson GA, Goldstein ED, Hanselman DH, Hermann AJ, Hulson PJF, Laurel BJ, Moss JH, Ormseth OA, Robinson D, Rogers LA, Rooper CN, Spies I, Strasburger WW (2022) Synthesizing integrated ecosystem research to create informed stock-specific indicators for next generation stock assessments. Deep-Sea Research – II. Topical Studies in Oceanography 198, 105070.
| Crossref | Google Scholar |

Siddon EC, Kristiansen T, Mueter FJ, Holsman KK, Heintz RA, Farley EV (2013) Spatial match–mismatch between juvenile fish and prey provides a mechanism for recruitment variability across contrasting climate conditions in the eastern Bering Sea. PLoS ONE 8, e84526.
| Crossref | Google Scholar | PubMed |

Sogard SM (1997) Size-selective mortality in the juvenile stage of teleost fishes: a review. Bulletin of Marine Science 60, 1129-1157.
| Google Scholar |

Stahl JP, Kruse GH (2008) Spatial and temporal variability in size at maturity of walleye pollock in the eastern Bering Sea. Transactions of the American Fisheries Society 137, 1543-1557.
| Crossref | Google Scholar |

Stauffer G (2004) NOAA protocols for groundfish bottom trawl surveys of the Nation’s fishery resources. NOAA Technical Memorandum, NMFS-F/SPO-65, United States Department of Commerce.

Stevenson DK, Campana SE (Eds) (1992) Otolith microstructure examination and analysis. Canadian Special Publication of Fisheries and Aquatic Sciences 117. (Department of Fisheries and Oceans: Ottawa, ON, Canada) Available at https://publications.gc.ca/collections/collection_2016/mpo-dfo/Fs41-31-117-eng.pdf

Summerfelt RC, Hall GE (Eds) (1987) ‘Age and growth of fish.’ (Iowa State University Press: Ames, IA, USA)

Szuwalski CS, Hollowed AB (2016) Climate change and non-stationary population processes in fisheries management. ICES Journal of Marine Science 73, 1297-1305.
| Crossref | Google Scholar |

TenBrink T, Neidetcher S, Arrington M, Benson I, Conrath C, Helser T (2022) Fourier transform near infrared spectroscopy as a tool to predict spawning status in Alaskan fishes with variable reproductive strategies. Journal of Near Infrared Spectroscopy 30, 179-188.
| Crossref | Google Scholar |

Townsend H, Harvey CJ, deReynier Y, Davis D, Zador SG, Gaichas S, Meijerman M, Hazen EL, Kaplan IC (2019) Progress on implementing ecosystem-based fisheries management in the United States through the use of ecosystem models and analysis. Frontiers in Marince Science 6, 641.
| Crossref | Google Scholar |

von Szalay PG, Raring NW (2018) Data Report: 2017 Gulf of Alaska bottom trawl survey, NOAA Technical Memorandum NMFS-AFSC-374. US Department of Commerce.

Wedding BB, Forrest AJ, Wright CL, Grauf S, Exley P, Poole SE (2014) A novel method for the age estimation of Saddletail snapper (Lutjanus malabaricus) using Fourier transform-near infrared (FT-NIR) spectroscopy. Marine and Freshwwater Research 65, 894-900.
| Crossref | Google Scholar |

Williams BC, Kruse GH, Dorn MW (2016) Interannual and spatial variability in maturity of walleye pollock Gadus chalcogrammus and implications for spawning stock biomass estimates in the Gulf of Alaska. PLoS ONE 11, e0164797.
| Crossref | Google Scholar | PubMed |

Wilson MT, Mier KL, Jump CM (2013) Effect of region on the food-related benefits to age-0 walleye pollock (Theragra chalcogramma) in association with midwater habitat characteristics in the Gulf of Alaska. ICES Journal of Marine Science 70, 1396-1407.
| Crossref | Google Scholar |

Wood SN (2017) ‘Generalized additive models: an introduction with R’, 2nd edn. (CRC Press: Boca Raton, FL, USA)

Zador SG, Holsman KK, Aydin KY, Gaichas SK (2017) Ecosystem considerations in Alaska: the value of qualitative assessments. ICES Journal of Marine Science 74, 421-430.
| Crossref | Google Scholar |