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Advances in the aquatic sciences
RESEARCH ARTICLE (Open Access)

Validation of age determination using otoliths of the European anchovy (Engraulis encrasicolus L.) in the Bay of Biscay

A. Uriarte A D , I. Rico A , B. Villamor B , E. Duhamel C , C. Dueñas B , N. Aldanondo A and U. Cotano A
+ Author Affiliations
- Author Affiliations

A AZTI Tecnalia, Marine Research Division, Herrera Kaia Portualdea z/g, E-20110 Pasaia, Spain.

B Instituto Español de Oceanografía (IEO), Promontorio de San Martín s/n, E-39080 Santander, Cantabria, Spain.

C Institut Français de Recherche et d’Exploitation de la Mer (IFREMER), Lorient station, 8 rue François Toullec F-56100, Lorient, France.

D Corresponding author. Email: auriarte@azti.es

Marine and Freshwater Research 67(7) 951-966 https://doi.org/10.1071/MF15092
Submitted: 3 March 2015  Accepted: 30 January 2016   Published: 28 April 2016

Journal Compilation © CSIRO Publishing 2016 Open Access CC BY-NC-ND

Abstract

Validation of the age determination procedure using otoliths of European anchovy in the Bay of Biscay was achieved by monitoring very strong year-classes in successive spring catches and surveys, as well as the seasonal occurrence of edge types. Historical corroboration of the ageing method was obtained by cross-correlation between successive age groups by year-classes in catches and surveys (1987–2013). Summary annual growth in length is also presented. Yearly annuli consist of a hyaline zone (either single or composite) and a wide opaque zone, disrupted occasionally by some typical checks (mainly at age-0 and age-1 at peak spawning time). Age determination, given a date of capture, requires knowledge of the typical annual growth pattern of otoliths, their seasonal edge formation by ages and the most typical checks. Most opaque growth occurs in summer and is minimal (translucent) in winter. Opaque zone formation begins earlier in younger fish (in spring), and this helps distinguish age-1 from age-2+.

Additional keywords: age estimation, checks, Engraulidae, otolith formation.


References

Aguayo, H. M. (1976). Edad y crecimiento de la anchoveta (Engraulis ringens) del norte de Chile (Arica-Iquique). Investigaciones Pesqueras 23, 1–25.

Aldanondo, N., Cotano, U., Etxebeste, E., Irigoien, X., Álvarez, P., Martínez de Murguía, A., and Herrero, D. L. (2008). Validation of daily increments deposition in the otoliths of European anchovy larvae (Engraulis encrasicolus L.) reared under different temperature conditions. Fisheries Research 93, 257–264.
Validation of daily increments deposition in the otoliths of European anchovy larvae (Engraulis encrasicolus L.) reared under different temperature conditions.Crossref | GoogleScholarGoogle Scholar |

Aldanondo, N., Cotano, U., and Etxebeste, E. (2011). Growth of young-of-the-year European anchovy (Engraulis encrasicolus L.) in the Bay of Biscay. Scientia Marina 75, 227–235.

Aldanondo, N., Cotano, U., Álvarez, P., and Uriarte, A. (2016). Validation of the first annual increment deposition in the otoliths of European anchovy in the Bay of Biscay based on otolith microstructure analysis. Marine and Freshwater Research 67, 943–950.
Validation of the first annual increment deposition in the otoliths of European anchovy in the Bay of Biscay based on otolith microstructure analysis.Crossref | GoogleScholarGoogle Scholar |

Anon. (1989). Report of the Working Group on the Assessment of Pelagic Stocks in Divisions VIIIc and IXa and Horse Mackerel. ICES CM 1989. International Council for the Exploration of the Sea (ICES), Copenhagen.

Anon. (1991). Report of the Working Group on the Assessment of the Stocks of Sardine, Horse Mackerel and Anchovy. ICES CM 1991. International Council for the Exploration of the Sea (ICES), Copenhagen.

Anon. (1992). Report of the Working Group on the Assessment of the Stocks of Mackerel Horse Mackerel, Sardine and Anchovy. ICES CM 1992. International Council for the Exploration of the Sea (ICES), Copenhagen.

Anon. (1993). Report of the Working Group on the Assessment of the Stocks of Mackerel Horse Mackerel, Sardine and Anchovy. ICES Document CM 1993. International Council for the Exploration of the Sea (ICES), Copenhagen.

Arneri, E., Carpi, P., Donato, F., and Santojanni, A. (2011). Growth in small pelagic fishes and its implications in their population dynamics. Biologia Marina Mediterranea 18, 106–113.

Astudillo, A. (1986). The Anchovy in the Bay of Biscay: Recent Data on the Fishery. ICES CM 1986/H:45. International Council for the Exploration of the Sea (ICES), Copenhagen.

Basilone, G., Guisande, C., Patti, B., Mazzola, S., Cuttitta, A., Bonanno, A., and Kallianiotis, A. (2004). Linking habitat conditions and growth in the European anchovy (Engraulis encrasicolus). Fisheries Research 68, 9–19.
Linking habitat conditions and growth in the European anchovy (Engraulis encrasicolus).Crossref | GoogleScholarGoogle Scholar |

Beckman, D. W., and Wilson, C. A. (1995). Seasonal timing of opaque zone formation in fish otoliths. In ‘Recent Developments in Fish Otolith Research’. (Eds D. H. Secor, J. M. Dean and S. E. Campana.) pp. 27–43. (University of South Carolina Press: Columbia, SC.)

Bellido, J. M., Pierce, G. J., Romero, J. L., and Millan, M. (2000). Use of frequency analysis methods to estimate growth of anchovy (Engraulis encrasicolus L. 1758) in the Gulf of Cádiz (SW, Spain). Fisheries Research 48, 107–115.
Use of frequency analysis methods to estimate growth of anchovy (Engraulis encrasicolus L. 1758) in the Gulf of Cádiz (SW, Spain).Crossref | GoogleScholarGoogle Scholar |

Blaszczyk, P. (1999). Length and age composition and growth of anchovy Engraulis encrasicolus (L. 1758) caught in the East Schelde, The Netherlands, in 1987. Folia Universitatis Agriculturae Stetinensis 192, 5–11.

Boyra, G., Martinez, U., Cotano, U., Santos, M., Irigoien, X., and Uriarte, A. (2013). Acoustic surveys for juvenile anchovy in the Bay of Biscay: abundance estimate as an indicator of the next year’s recruitment and spatial distribution patterns. ICES Journal of Marine Science 70, 1354–1368.
Acoustic surveys for juvenile anchovy in the Bay of Biscay: abundance estimate as an indicator of the next year’s recruitment and spatial distribution patterns.Crossref | GoogleScholarGoogle Scholar |

Brandhorst, W., Castello, J. P., Cousseau, M. B., and Capezzani, D. A. (1974). Evaluation of anchovy resources (Engraulis anchoita) offshore Argentina and Uruguay. VIII. Spawning, growth and population structure. Physis (Rio de Janeiro, Brazil) 33, 37–58.

Campana, S. E. (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.
Accuracy, precision and quality control in age determination, including a review of the use and abuse of age validation methods.Crossref | GoogleScholarGoogle Scholar |

Campana, S. E., and Neilson, J. D. (1985). Microstructure of fish otoliths. Canadian Journal of Fisheries and Aquatic Sciences 42, 1014–1032.
Microstructure of fish otoliths.Crossref | GoogleScholarGoogle Scholar |

Canales, T. M., and Leal, E. (2009). Parámetros de historia de vida de la anchoveta Engraulis ringens Jenyns, 1842, en la zona centro norte de Chile. Revista de Biología Marina y Oceanografía 44, 173–179.

Cappo, M., Eden, P., Newman, S. J., and Robertson, S. (2000). A new approach to validation of periodicity and timing of opaque zone formation in the otoliths of eleven species of Lutjanus from the central Great Barrier Reef. Fishery Bulletin 98, 474–488.

Casselman, J. M. (1987). Determination of age and growth. In ‘The Biology of Fish Growth’. (Eds A. H. Weatherley and H. S. Gill.) pp. 209–242. (Academic Press: London.)

Cendrero, O., Cort, J. L., and Cárdenas, E. (1981). Revisión de algunos datos sobre la biología de la anchoa, Engraulis encrasicholus (L.) del Mar Cantábrico. Boletín del Instituto Español de Oceanografía 311, 118–123.

Cermeño, P., Uriarte, A., De Munguía, A. M., and Morales-Nin, B. (2003). Validation of daily increment formation in otoliths of juvenile and adult European anchovy. Journal of Fish Biology 62, 679–691.
Validation of daily increment formation in otoliths of juvenile and adult European anchovy.Crossref | GoogleScholarGoogle Scholar |

Cermeño, P., Uriarte, A., Morales-Nin, B., Cotano, U., and Álvarez, P. (2008). Setting up interpretation criteria for ageing juvenile European anchovy otoliths. Scientia Marina 72, 733–742.

Collins, R. A., and Spratt, J. D. (1969). Age determination of northern anchovies, Engraulis mordax, from otoliths. In ‘The Northern Anchovy (Engraulis mordax) and its Fishery 1965–1968’. (Ed. J. D. Messersmith.) Department of Fish and Game, Fish Bulletin 147, pp. 39–55. (State of California, The Resources Agency.)

Cort, J. L., Cendrero, O., and Iribar, X. (1976). La anchoa, Engraulis encrasicholus (L.), del Cantábrico. Resultados de las campañas de 1974, 1975 y 1976. Boletín del Instituto Español de Oceanografía 220, 1–34.

Cubillos, L. A., Arcos, D. F., Bucarey, D. A., and Canales, M. T. (2001). Seasonal growth of small pelagic fish off Talcahuano, Chile (37°S, 73°W): a consequence of their reproductive strategy to seasonal upwelling? Aquatic Resources 14, 115–124.
Seasonal growth of small pelagic fish off Talcahuano, Chile (37°S, 73°W): a consequence of their reproductive strategy to seasonal upwelling?Crossref | GoogleScholarGoogle Scholar |

Erkoyuncu, I., and Ozdamar, E. (1989). Estimation of the age, size and sex composition and growth parameters of anchovy (Engraulis encrasicholus L.) in the Black sea. Fisheries Research 7, 241–247.
Estimation of the age, size and sex composition and growth parameters of anchovy (Engraulis encrasicholus L.) in the Black sea.Crossref | GoogleScholarGoogle Scholar |

Gonzalez-Salas, C., and Lenfant, P. (2007). Interannual variability and intraannual stability of the otolith shape in European anchovy Engraulis encrasicolus (L.) in the Bay of Biscay. Journal of Fish Biology 70, 35–49.
Interannual variability and intraannual stability of the otolith shape in European anchovy Engraulis encrasicolus (L.) in the Bay of Biscay.Crossref | GoogleScholarGoogle Scholar |

Guerault, D., and Avrilla, J. (1973). L’anchois du golfe de Gascogne, captures de 1972, données biologiques et biometriques. ICES CM 1973/J:11. International Council for the Exploration of the Sea (ICES), Copenhagen.

Guerault, D., and Avrilla, J. (1974). L’anchois du golfe de Gascogne, taille, âge et croissance. ICES CM 1974/J:17. International Council for the Exploration of the Sea (ICES), Copenhagen.

He, W., Li, Z., Liu, J., Li, Y., Murphy, B. R., and Xie, S. (2008). Validation of a method of estimating age, modelling growth, and describing the age composition of Coilia mystus from the Yangtze Estuary, China. ICES Journal of Marine Science 65, 1655–1661.
Validation of a method of estimating age, modelling growth, and describing the age composition of Coilia mystus from the Yangtze Estuary, China.Crossref | GoogleScholarGoogle Scholar |

Hernández, C., Villamor, B., Abaunza, P., and Landa, J. (2009). Age and growth of European anchovy (Engraulis encrasicolus) in the Bay of Biscay (NE Atlantic), 1994–2008. In ‘Report of the Workshop on Age Reading of European Anchovy (WKARA)’, 9–13 November 2009, Sicily, Italy. ICES CM 2009/ACOM:43. (International Council for the Exploration of the Sea (ICES): Copenhagen.) Available at http://www.ices.dk/sites/pub/Publication%20Reports/Expert%20Group%20Report/acom/2009/WKARA/WKARA%20Report%202009.pdf [Verified 2 January 2016].

Hoedt, F. E. (1990). Growth of the tropical anchovy, Stolephorus nelsoni, in Northern Australia. In ‘Tuna Baitfish in the Indo-Pacific Region: Proceedings of a Workshop’, 11–13 December 1989, Honiara, Solomon Islands. (Eds S. J. M. Blaber and J. W. Copland.) pp. 147–149. (Australian Centre for International Agricultural Research: Canberra.)

Hoedt, F. E. (1992). Age and growth of a large tropical anchovy, Thryssa hamiltoni (Gray): a comparison of ageing techniques. Australian Journal of Marine and Freshwater Research 43, 953–971.
Age and growth of a large tropical anchovy, Thryssa hamiltoni (Gray): a comparison of ageing techniques.Crossref | GoogleScholarGoogle Scholar |

Holden, M. J., and Raitt, D. F. S. (1974). Manual of Fisheries Science. Part 2. Methods of Resource Investigations and Their Application. FAO Fisheries Technical Paper number 115. FAO, Rome.

Hsu, C. C., and Tzeng, W. N. (2009). Validation of annular deposition in scales and otoliths of flathead mullet Mugil cephalus. Zoological Studies 48, 640–648.

Ibaibarriaga, L., Fernandez, C., Uriarte, A., and Roel, B. A. (2008). A two-stage biomass dynamic model for the Bay of Biscay anchovy: a Bayesian approach. ICES Journal of Marine Science 65, 191–205.
A two-stage biomass dynamic model for the Bay of Biscay anchovy: a Bayesian approach.Crossref | GoogleScholarGoogle Scholar |

Ibaibarriaga, L., Fernandez, C., and Uriarte, A. (2011). Gaining information from commercial catch for a Bayesian two-stage biomass dynamic model: application to Bay of Biscay anchovy. ICES Journal of Marine Science 68, 1435–1446.
Gaining information from commercial catch for a Bayesian two-stage biomass dynamic model: application to Bay of Biscay anchovy.Crossref | GoogleScholarGoogle Scholar |

Ibaibarriaga, L., Uriarte, A., Laconcha, U., Bernal, M., Santos, M., Chiflet, M., and Irigoien, X. (2013). Modelling the spatio-temporal distribution of age-1 Bay of Biscay anchovy (Engraulis encrasicolus) at spawning time. Scientia Marina 77, 461–472.
Modelling the spatio-temporal distribution of age-1 Bay of Biscay anchovy (Engraulis encrasicolus) at spawning time.Crossref | GoogleScholarGoogle Scholar |

ICES (2008a). Report of the Workshop on Age Reading of Baltic Herring (WKARBH), 9–13 June 2008, Riga, Latvia. ICES CM 2008/ACOM:36. (International Council for the Exploration of the Sea (ICES): Copenhagen.)

ICES (2008b). Report of the Workshop on Age Reading on Baltic Sprat (WKARBS), 17–20 March 2008, Klaipeda, Lithuania. ICES CM 2008/ACOM:37. (International Council for the Exploration of the Sea (ICES): Copenhagen.)

ICES (2009). Report of the Workshop on Age Reading of European Anchovy (WKARA), 9–13 November 2009, Sicily, Italy. ICES CM 2009/ACOM:43. (International Council for the Exploration of the Sea (ICES): Copenhagen.)

ICES (2011). Report of the Workshop on Age Reading of European Atlantic Sardine (WKARAS), 14–18 February 2011, Lisbon, Portugal. ICES CM 2011/ACOM:42. (International Council for the Exploration of the Sea (ICES): Copenhagen.)

ICES (2014). Report of the Working Group on Southern Horse Mackerel, Anchovy and Sardine (WGHANSA), 20–25 June 2014, Copenhagen, Denmark. ICES CM 2014/ACOM:16. (International Council for the Exploration of the Sea (ICES): Copenhagen.)

ICSEAF (1983). Otolith Interpretation Guide. Number 1 Hake. International Commission for the Southeast Atlantic fisheries, Madrid.

Irigoien, X., Fiksen, Ø., Cotano, U., Uriarte, A., Alvarez, P., Arrizabalaga, H., Boyra, G., Santos, M., Sagarminaga, Y., Otheguy, P., Etxebeste, E., Zarauz, L., Artetxe, I., and Motos, L. (2007). Could Biscay Bay anchovy recruit through a spatial loophole? Progress in Oceanography 74, 132–148.
Could Biscay Bay anchovy recruit through a spatial loophole?Crossref | GoogleScholarGoogle Scholar |

Iversen, S. A., Zhu, D., Johannessen, A., and Toresen, R. (1993). Stock size, distribution and biology of anchovy in the Yellow Sea and East China Sea. Fisheries Research 16, 147–163.
Stock size, distribution and biology of anchovy in the Yellow Sea and East China Sea.Crossref | GoogleScholarGoogle Scholar |

Jensen, A. C. (1965). A standard terminology and notation for otolith readers. ICNAF Research Bulletin 2, 5–7.

Junquera, S. (1986). Pêche de l’anchois (Engraulis encrasicholus) dans le golfe de Gascogne et sur le littoral atlantique de Galice depuis 1920. Variations quantitatives. Revue des Travaux de l’Institut des Pêches Maritimes 48, 133–142.

Karacam, H., and Düzgünes, E. (1990). Age, growth and meat yield of the European anchovy (Engraulis encrasicolus, L. 1758) in the Black Sea. Fisheries Research 9, 181–186.
Age, growth and meat yield of the European anchovy (Engraulis encrasicolus, L. 1758) in the Black Sea.Crossref | GoogleScholarGoogle Scholar |

Khemiri, S., Gaamour, A., Meunier, F. J., and Zylberger, L. (2007). Age and growth of Engraulis encrasicolus (Clupeiforme: Engraulidae) in the Tunisian waters. Cahiers de Biologie Marine 48, 259–269.

Machias, A., Somarakis, S., Drakopoulos, P., Magoulas, A., and Koutsikopoulos, C. (2000). Evaluation of the Southern Greek Anchovy Stocks. Project 97–0048, Final Report. Institute of Marine Biology, Crete.

Mallicote, D. L., and Parrish, R. H. (1981). Seasonal growth patterns of California stocks of northern anchovy, Engraulis mordax, pacific mackerel, Scomber japonicus, and jack mackerel, Trachurus symmetrlcus. California Cooperative Oceanic Fisheries Investigations Report 22, 69–81.

Massé, J. (1996). Acoustic observations in the Bay of Biscay: schooling, vertical distribution, species assemblages and behaviour. Scientia Marina 60, 227–234.

Massé, J., Duhamel, E., Petitgas, P., Doray, M., and Huret, M. (2016). Spring acoustic surveys: Pelgas survey. In ‘Pelagic Surveys Series for Sardine and Anchovy in ICES Areas VIII and IX (WGACEGG): Towards an Ecosystem Approach’. (Eds J. Masse, A. Uriarte, M. M. Angelico and P. Carrera.) ICES Cooperative Research Report 332. (International Council for the Exploration of the Sea (ICES): Copenhagen.)

Melo, Y. C. (1984). Ages studies on anchovy Engraulis capensis Gichrist off south west Africa. South African Journal of Marine Science 2, 19–31.
Ages studies on anchovy Engraulis capensis Gichrist off south west Africa.Crossref | GoogleScholarGoogle Scholar |

Morales-Nin, B. (1989). Age and growth of the southern stock of Peruvian anchoveta based on otolith microstructures and length frequency analysis. In ‘ICLARM Conference Proceedings 18. The Peruvian Upwelling System: Dynamics and Interactions’. (Eds D. Pauly, P. Muck, J. Mendo, and I. Tsukayama.) pp. 179–188. (Instituto del Mar del Perú (IMARPE): Callao, Peru; Deutsche Gesellschaft für Technische Zusammenarbeit (GTZ), GmbH; Eschborn, Germany; and International Center for Living Aquatic Resources Management: Manila, Philippines.)

Morales-Nin, B., and Panfili, J. (2002). Age estimation. In ‘Manual of Fish Sclerochronology’. (Eds J. Panfili, H. Troadec, H. de Pontual and P. J. Wright.) pp. 91–98. (IFREMER and IRD: Brest, France).

Morales-Nin, B., and Pertierra, J. P. (1990). Growth rates of the anchovy Engraulis encrasicolus and the sardine Sardina pilchardus in the northwestern Mediterranean Sea. Marine Biology 107, 349–356.
Growth rates of the anchovy Engraulis encrasicolus and the sardine Sardina pilchardus in the northwestern Mediterranean Sea.Crossref | GoogleScholarGoogle Scholar |

Motos, L. (1996). Reproductive biology and fecundity of the Bay of Biscay anchovy population (Engraulis encrasicolus L.). Scientia Marina 60, 195–207.

Motos, L., Uriarte, A., and Valencia, V. (1996). The spawning environment of the Bay of Biscay anchovy (Engraulis encrasicolus L.). Scientia Marina 60, 117–140.

Motos, L., Uriarte, A., Prouzet, P., Santos, M., Alvarez, P., and Sagarminaga, Y. (2005). Assessing the Bay of Biscay anchovy population by DEPM: a review 1989–2001. In ‘Report of the SPACC Meeting on Small Pelagic Fish Spawning Habitat Dynamics and the Daily Egg Production Method (DEPM)’. (Eds L. R. Castro, P. Freón, C. D. van der Lingen and A. Uriarte.) GLOBEC Report 22. pp. 88–90. (GLOBEC International Project Office: Plymouth, UK.)

Navaz J.M. and Lozano F. (1966). Donnes sur la taille, l’âge et la croissance de l’anchois (Engraulis encrasicholus L.) de la côte Basque Espagnole. ICES CM 1966/J:6. International Council for the Exploration of the Sea (ICES), Copenhagen.

Newberger, T. A., and Houde, E. D. (1995). Population biology of bay anchovy Anchoa mitchilli in the mid Cheasapeake Bay. Marine Ecology Progress Series 116, 25–37.
Population biology of bay anchovy Anchoa mitchilli in the mid Cheasapeake Bay.Crossref | GoogleScholarGoogle Scholar |

Palomares, M. L., Muck, P., Mendo, J., Chuman, E., Gomez, O., and Pauly, D. (1987). Growth of the Peruvian anchoveta (Engraulis ringens), 1953 to 1982. In ‘The Peruvian Anchoveta and its Upwelling Ecosystem: Three Decades of Change’. (Eds D. Pauly and I. Tsukayama.) ICLARM Studies and Reviews. 15, pp. 117–141. (Instituto del Mar del Peru (IMARPE): Callao, Peru; Deutsche Gesellschaft für Technische Zusammenarbeit (GTZ), GmbH: Eschborn, Germany; and International Center for Living Aquatic Resources Management: Manila, Philippines.).

Panfili, J., Troadec, H., de Pontual, H., and Wright, P. J. (Eds) (2002). ‘Manual of Fish Sclerochronology.’ (IFREMER and IRD: Brest, France.)

Pannella, G. (1971). Fish otoliths: daily growth layers and periodical patterns. Science 173, 1124–1127.
Fish otoliths: daily growth layers and periodical patterns.Crossref | GoogleScholarGoogle Scholar |

Pauly, D., and Munro, J. L. (1984). Once more on the comparison of growth in fish and invertebrates. Fishbyte 2, 21.

Pecquerie, L., Petitgas, P., and Kooijman, S. A. L. M. (2009). Modeling fish growth and reproduction in the context of the dynamic energy budget theory to predict environmental impact on anchovy spawning duration. Journal of Sea Research 62, 93–105.
Modeling fish growth and reproduction in the context of the dynamic energy budget theory to predict environmental impact on anchovy spawning duration.Crossref | GoogleScholarGoogle Scholar |

Pecquerie, L., Fablet, T., de Pontual, H., Bonhommeau, S., Alunno-Bruscia, M., Petitgas, P., and Kooijman, S. A. L. M. (2012). Reconstructing individual food and growth histories from biogenic carbonates. Marine Ecology Progress Series 447, 151–164.
Reconstructing individual food and growth histories from biogenic carbonates.Crossref | GoogleScholarGoogle Scholar |

Pertierra, J. P. (1987). Crecimiento del boquerón (Engraulis encrasicolus, L. 1758) (Pisces, Engraulidae) de la costa catalana (Mediterráneo noroccidental). Investigaciones Pesqueras 51, 263–275.

Petitgas, P., and Grellier, P. (2003). Size selective processes for anchovy in Biscay, 2000–2002: recruitment, adult survival and spawning. ICES CM, N:07. International Council for the Exploration of the Sea (ICES), Copenhagen.

Petitgas, P., Uriarte, A., Nogueira, E., Massé, J., and Cotano, U. (2010). Bay of Biscay anchovy. In ‘Life Cycle Spatial Patterns of Small Pelagic Fish in the Northeast Atlantic’. (Ed. P. Petitgas.) ICES Cooperative Research Report number 306, pp. 40–44. (International Council for the Exploration of the Sea (ICES): Copenhagen.)

Petitgas, P., Alheit, J., Peck, M. A., Raab, K., Irigoien, X., Huret, M., van der Kooij, J., Pohlmann, T., Wagner, C., Zarraonaindia, I., and Dickey-Collas, M. (2012). Anchovy population expansion in the North Sea. Marine Ecology Progress Series 444, 1–13.
Anchovy population expansion in the North Sea.Crossref | GoogleScholarGoogle Scholar |

Ricker, W. E. (1975). Computation and Interpretation of Biological Statistics of Fish Populations. Bulletin 191, Fisheries Research Board of Canada, Ottawa, ON.

Santiago, J., and Arrizabalaga, H. (2005). An integrated growth study for North Atlantic albacore (Thunnus alalunga Bonn. 1788). ICES Journal of Marine Science 62, 740–749.
An integrated growth study for North Atlantic albacore (Thunnus alalunga Bonn. 1788).Crossref | GoogleScholarGoogle Scholar |

Santos, M., Uriarte, A., Boyra, G., and Ibaibarriaga, L. (2016). Anchovy DEPM surveys 2003–2012 in the Bay of Biscay (subarea VIII) BIOMAN. In ‘Pelagic Surveys Series for Sardine and Anchovy in ICES Areas VIII and IX (WGACEGG): Towards an Ecosystem Approach’. (Eds J. Massé, A. Uriarte, M. M. Angelico and P. Carrera.) ICES Cooperative Research Report 332. (International Council for the Exploration of the Sea (ICES): Copenhagen.)

Sinovčić, G. (2000). Anchovy, Engraulis encrasicolus (Linnaeus, 1758): biology, population dynamics and fisheries case study. Acta Adriatica 41, 3–53.

Spratt, J. D. (1975). Growth rate of northern anchovy Engraulis Mordax, in Southern California waters calculated from otoliths. California Fish and Game 61, 116–126.

Swan, S. C., and Gordon, J. D. M. (2001). A review of age estimation in macrourid fishes with new data on age validation of juveniles. Fisheries Research 51, 177–195.
A review of age estimation in macrourid fishes with new data on age validation of juveniles.Crossref | GoogleScholarGoogle Scholar |

Taylor, C. C. (1962). Growth equations with metabolic parameters. Journal du Conseil International pour l’Exploration de la Mer 27, 270–286.
Growth equations with metabolic parameters.Crossref | GoogleScholarGoogle Scholar |

Thomas, R. M. (1983). Back-calculation of time of hyaline ring formation in the otoliths of the pilchard off South West Africa. South African Journal of Marine Science 1, 3–18.
Back-calculation of time of hyaline ring formation in the otoliths of the pilchard off South West Africa.Crossref | GoogleScholarGoogle Scholar |

Thomas, R. M. (1984). A method of age determination for the South West African pilchard Sardinops ocellata. South African Journal of Marine Science 2, 63–70.
A method of age determination for the South West African pilchard Sardinops ocellata.Crossref | GoogleScholarGoogle Scholar |

Thomas, R. M. (1985). Growth rate of the pilchard off south west Africa, 1971–1983. Investigational report 128, Sea Fisheries Research Institute, South Africa.

Tiroba, G., Rawlinson, N. J. F., Nichols, P. V., and Leqata, J. L. (1990). Length–frequency Analysis of Major Baitfish Species in Solomon Islands. In ‘Tuna Baitfish in the Indo-Pacific Region: Proceedings of a Workshop’, 11–13 December 1989, Honiara, Solomon Islands. (Eds S. J. M. Blaber and J. W. Copland.) pp. 114–133. (Australian Centre for International Agricultural Research: Canberra.)

Uriarte, A. (2015). Consistency of relative changes in biomass and proportions at age 1 in anchovy surveys. Preliminary results. In ‘ICES 2015. First Interim Report of the Working Group on Acoustic and Egg Surveys for Sardine and Anchovy in ICES Areas VII, VIII and IX (WGACEGG)’, 17–21 November 2014, Vigo, Spain. ICES CM 2014/SSGESST:21, pp. 88–91. (International Council for the Exploration of the Sea (ICES). Copenhagen.)

Uriarte, A., and Astudillo, A. (1987). The anchovy in the Bay of Biscay: new data and analysis of the fishery, 1974–1987. ICES CM 1987/H:20. International Council for the Exploration of the Sea (ICES), Copenhagen.

Uriarte, A., Prouzet, P., and Villamor, B. (1996). Bay of Biscay and Ibero Atlantic anchovy populations and their fisheries. Scientia Marina 60, 237–255.

Uriarte, A., Sagarminaga, Y., Scalabrin, C., Valencia, V., Cermeño, P., De Miguel, E, Gómez Sánchez, J.A., Jiménez, M. (2001). Ecology of Anchovy Juveniles in the Bay of Biscay 4 Months after Peak Spawning: Do They Form Part of the Plankton? ICES CM/W:20. International Council for the Exploration of the Sea (ICES), Copenhagen.

Uriarte, A., Blanco, M., Cendrero, O., Grellier, P., Millán, M., Morais, A., and Rico, I. (2002). Report of the Workshop on Anchovy Otoliths from Subarea VIII and Division IXa. Working Document to the ICES Working Group on the Assessment of Mackerel, Horse Mackerel, Sardine and Anchovy. Available at http://www.ices.dk/community/Documents/PGCCDBS/ANCHOVY%20OTOLITH%20WORKSHOP_2002.pdf [Verified 28 February 2015].

Vaz, S., Petitgas, P., Beillois, P., and Masse, J. (2002). Time and Spatial Variations of Anchovy Biometric Parameters in the Bay of Biscay from 1983 to 2002. ICES Document CM 2002/O:27. International Council for the Exploration of the Sea (ICES), Copenhagen.

Waldron, M. E. (1994). Validation of annuli of South African anchovy Engraulis capensis, using daily otoliths increments. ICES Journal of Marine Science 51, 233–234.
Validation of annuli of South African anchovy Engraulis capensis, using daily otoliths increments.Crossref | GoogleScholarGoogle Scholar |

Waldron, M. E., and Kerstan, M. (2001). Age validation in horse mackerel (Trachurus trachurus) otoliths. ICES Journal of Marine Science 58, 806–813.
Age validation in horse mackerel (Trachurus trachurus) otoliths.Crossref | GoogleScholarGoogle Scholar |

Williams, T., and Bedford, B. C. (1974). The use of otoliths for age determination. In ‘The Ageing of Fish’. (Ed. T. B. Bagenal.) pp 114–123. (Unwin Brothers: London, UK.)

Xiao, Y. (1994). Von Bertalanffy growth models with variability in, and correlation between, K and L∞. Canadian Journal of Fisheries and Aquatic Sciences 51, 1585–1590.
Von Bertalanffy growth models with variability in, and correlation between, K and L.Crossref | GoogleScholarGoogle Scholar |