Arild Folkvord




Research groups


The focus of my research is on growth and survival mechanisms in the early life of fishes. The research is experimentally oriented, and covers aspects of marine juvenile production and process oriented recruitment studies. Recently I have focused on experimental studies on herring from different populations and their adaptations to native environments. 

Size-selective mortality in cultured fish is often due to aggression and cannibalism. A large effort has been made to elucidate factors promoting cannibalism in captive cod (e.g. hunger level, size, size differences etc.). Size- and temperature dependent growth in juvenile flatfish and cod has also been studied with the aim of determining the growth potential under culture conditions. The growth process is very dynamic during the early life stages, and controlled laboratory experiments are carried out to calibrate methods used for growth characterization (e.g. RNA:DNA analysis, otolith microstructure analysis, fatty acid analysis).

The age of field caught fish larvae is often difficult to assess. Fortunately, growth increments are usually formed on a daily basis in the ear stones (otoliths) of fish larvae. It is thus possible to estimate the larval age in days in a similar manner as one determines the age of older fish by counting yearly zones in the otoliths and scales. The growth pattern in the otolith also reflects the previous growth history of the larva, and can be used to determine where, and under what conditions, the larva grew up. The otolith has therefore often been referred to as the "black box" in fish. An important part of the ageing research is related to the accuracy and precision of the age estimation. Knowledge in this field is of great importance in process studies and fisheries management.

I currently hold an adjunct position at Institute of Marine Research.


BIO102 Oranismal biology 2 (autumn)

BIO​​​​​299 Research practice in biology (spring/autumn) 

BIO325 Ocean science (autumn)

BIO340 Special topics in Fisheries and Marine Biology (spring/autumn) (on demand)


Master's Programme in Biology - Fisheries Biology and Management (thesis supervision)

Master's Programme in Biology - Aquaculture (thesis supervision)

Master's Programme in Biology - Marine Biology (thesis supervision)


Previously tought courses:

BIO213 Marine Ecology

BIO308 Early life history of fish (now only as special topic)

BIO300A Biological Data Analysis and Research Design (now Academic writing)


Esamples of previously suggested Master thesis topics

Selective mortality of herring larvae 

Inferring cod feeding history from otolith isotopic composition


Other topics are available on request and after discussion with supervisor 

A list of previous Master and Phd students are provided here 

Academic article
Academic lecture
Masters thesis
Doctoral dissertation
Short communication
Popular scientific lecture
Academic chapter/article/Conference paper
Programme participation

See a complete overview of publications in Cristin.


Link to Google scholar


Original scientific publications in peer-review journals:

A126. Berg F, Shirajee S, Folkvord A, Aanestad Godiksen J, Skaret G, Slotte A (2021) Early life growth is affecting timing of spawning in the semelparous Barents Sea capelin (Mallotus villosus). Progress in Oceanography:102614,

A125. Allan BJM, Ray JL, Tiedemann M, Komyakova V, Vikebø F, Skaar KS, Stiasny MH, Folkvord A, Nash RDM, Stenevik EK, Kjesbu OS (2021) Quantitative molecular detection of larval Atlantic herring (Clupea harengus) in stomach contents of Atlantic mackerel (Scomber scombrus) marks regions of predation pressure. Sci Rep 11:5095,

A124. Stenevik EK, Nash RDM, Folkvord A (2021) Development stage distribution as a proxy for feeding success and growth for first feeding Norwegian spring spawning herring larvae. Journal of Fish Biology 98:1112-1119,

A123. Han F, Jamsandekar M, Pettersson ME, Su L, Fuentes-Pardo AP, Davis BW, Bekkevold D, Berg F, Casini M, Dahle G, Farrell ED, Folkvord A, Andersson L (2020) Ecological adaptation in Atlantic herring is associated with large shifts in allele frequencies at hundreds of loci. eLife 9:e61076,

A122. Rafati N, Chen J, Herpin A, Pettersson ME, Han F, Feng C, Wallerman O, Rubin C-J, Péron S, Cocco A, Larsson M, Trötschel C, Poetsch A, Korsching K, Bönigk W, Körschen HG, Berg F, Folkvord A, Kaupp UB, Schartl M, Andersson L (2020) Reconstruction of the birth of a male sex chromosome present in Atlantic herring. Proceedings of the National Academy of Sciences 117:24359-24368,

A122. Folkvord A, Lakso E, Laupsa M, Meier S, Musialak LA, Sundby S (2020) Swimbladder filling in herring larvae: effects of food oil on the water surface. Marine Biology Research 16:446-457,

A121. Otterå H, Johansen T, Folkvord A, Dahle G, Solvang Bingh MK, Westgård JI, Glover KA (2020) The pantophysin gene and its relationship with survival in early life stages of Atlantic cod. Royal Society Open Science 7:191983,

A120. Blanco E, Reglero P, Ortega A, Folkvord A, de la Gándara F, Hernández de Rojas A, Moyano M (2020) First estimates of metabolic rate in Atlantic bluefin tuna larvae. Journal of Fish Biology 97:1296-1305,

A119. Tonheim S, Slotte A, Andersson L, Folkvord A, Berg F (2020) Comparison of otolith microstructure of herring larvae and sibling adults reared under identical early life conditions. Frontiers in Marine Science 7:1-13,

A118. Seljestad GW, Quintela M, Faust W, Halvorsen KT, Besnier F, Jansson E, Dahle G, Knutsen H, André C, Folkvord A, Glover KA (2020) Genetic analysis identifies geographically distinct groups of ballan wrasse, a cleaner fish, and divergence between sympatric phenotypes. Ecology and Evolution 10:6120-6135,

A117. Berg F, Andersson L, Folkvord A (2020) Respiration rates of herring larvae at different salinities, and effects of previous environmental history. Marine Ecology Progress Series 650:141-152,

A116. Berg F, Østgaard HD, Slotte A, Andersson L, Folkvord A (2021) A combination of genetic and phenotypic characterization of spring- and autumn-spawning herring suggests gene flow between populations. ICES Journal of Marine Science 78:694-703,

A115. Irgens C, Folkvord A, Otterå H, Kjesbu OS (2020) Otolith growth and zone formation during first maturity and spawning of Atlantic cod (Gadus morhua). Canadian Journal of Fisheries and Aquatic Sciences 77:113-123,

A114. Godiksen JA, Ming TS, Folkvord A, Grønkjær P (2019) Effects of temperature on tissue–diet isotopic spacing of nitrogen and carbon in otolith organic matter. Marine and Freshwater Research 72:1757–1767,

A113. Pettersson ME, Rochus CM, Han F, Chen JF, Hill J, Wallerman O, Fan GY, Hong XN, Xu QW, Zhang H, Liu SS, Liu X, Haggerty L, Hunt T, Martin FJ, Flicek P, Bunikis I, Folkvord A, Andersson L (2019) A chromosome-level assembly of the Atlantic herring genome-detection of a supergene and other signals of selection. Genome Res 29:1919-1928,

A112. Hill J, Enbody ED, Pettersson M, Sprehn CG, Bekkevold D, Folkvord A, Kleinau G, Scheerer P, Andersson L (2019) Recurrent convergent evolution at amino acid residue 261 in fish rhodopsin. Proceedings of the National Academy of Sciences 116:18473-18478,

A111. Blanco E, Reglero P, Hernández de Rojas A, Ortega A, de la Gandara F, Folkvord A (2019) The effect of nutritional condition on the growth to post-flexion of Atlantic bluefin tuna and Atlantic bonito larvae. Journal of Experimental Marine Biology and Ecology,

A110. Slotte A, Husebø Å, Berg F, Stenevik EK, Folkvord A, Fossum P, Mosegaard H, Vikebø F, Nash RDM (2019) Earlier hatching and slower growth, a key to survival in the early life history of Norwegian spring spawning herring. Marine Ecology Progress Series 617-618:25-39,

A109. Berg F, Slotte A, Andersson L, Folkvord A (2019) Genetic origin and salinity history influence reproductive success of Atlantic herring. Marine Ecology Progress Series 617-618:81-94,

A108.   Blanco E, P Reglero, A Ortega, F de la Gandara, A Folkvord, (2018). Size-selective mortality during the piscivorous stage of laboratory-reared Atlantic bluefin tuna larvae as inferred from otolith microstructure analysis. Journal of Experimental Marine Biology and Ecology. 59, 36-43.

A107.  Salvanes AGV, Christiansen H, Seivåg ML, Taha Y, Henseler C, Kjesbu OS, Folkvord A, Utne Palm AC, Currie B, Ekau W, Van der Plas A, Gibbons MJ (2018) Variation in growth, morphology and reproduction of the bearded goby (Sufflogobius bibarbatus) in hypoxic waters of northern Benguela. J Mar Syst 188:81-97.

A106.  Reglero P, Blanco E, Alemany F, Ferrá C, Alvarez-Berastegui D, Ortega A, de la Gándara F, Aparicio A, Folkvord A (2018) Vertical distribution of Atlantic bluefin tuna (Thunnus thynnus) and bonito (Sarda sarda) larvae is related to temperature preference. Mar Ecol Prog Ser 594:231-243,

A105. Berg F, Almeland OW, Skadal J, Slotte A, Andersson L, Folkvord A (2018) Genetic factors have a major effect on growth, number of vertebrae and otolith shape in Atlantic herring (Clupea harengus). PLoS One 13:e0190995,

A104. Folkvord A, Koedijk R, Grahl-Nielsen O, Meier S, Olsen BR, Blom G, Otterlei E, Imsland A (2018) You are what you eat? Differences in lipid composition of cod larvae reared on natural zooplankton and enriched rotifers. Aquaculture Nutrition 24:224-235,

A103. Feng C, Petterson M, Lamichhaney S, Rubin CJ, Rafati N, Casini M, Folkvord A, Andersson L (2017) Moderate nucleotide diversity in the Atlantic herring is associated with a low mutation rate. eLife:1-14,

A102. Berg F, Husebø Å, Godiksen JA, Slotte A, Folkvord A (2017) Spawning time of Atlantic herring (Clupea harengus) populations with a restricted area reflects their otolith growth at the larval stage. Fisheries Research 194:68-75,

A101. Irgens C, Kjesbu OS, Folkvord A (2017) Ontogenetic development of otolith shape during settlement of juvenile Barents Sea cod (Gadus morhua). ICES Journal of Marine Science 74:2389–2397,

A100. Blanco E, Reglero P, Ortega A, Gándara Fdl, Fiksen Ø, Folkvord A (2017) The effects of light, darkness and intermittent feeding on the growth and survival of reared Atlantic bonito and Atlantic bluefin tuna larvae. Aquaculture 479:333-339,

A99. Martinez Barrio A, Lamichhaney S, Fan G, Rafati N, Pettersson M, Zhang H, Dainat J, Ekman D, Höppner M, Jern P, Martin M, Nystedt B, Liu X, Chen W, Liang X, Shi C, Fu Y, Ma K, Zhan X, Feng C, Gustafson U, Rubin C-J, Sällman Almén M, Blass M, Casini M, Folkvord A, Laikre L, Ryman N, Ming-Yuen Lee S, Xu X, Andersson L (2016) The genetic basis for ecological adaptation of the Atlantic herring revealed by genome sequencing. eLife 5:e12081,

A98. Folkvord A, Vollset KW, Catalán IA (2015) Differences in growth and survival between cod and herring early stages co-reared at variable prey concentrations. Journal of Fish Biology 87:1176-1190,

A97. Folkvord A, Gundersen G, Albretsen J, Asplin L, Kaartvedt S, Giske J (2016) Impact of hatch-date on early life growth and survival of Mueller's pearlside (Maurolicus muelleri) larvae, and life history consequences. Canadian Journal of Fisheries and Aquatic Sciences 73:163-176,

A96. Maneja RH, Frommel AY, Browman HI, Geffen AJ, Folkvord A, Piatkowski U, Durif CMF, Bjelland R, Skiftesvik AB, Clemmesen C (2015) The swimming kinematics and foraging behavior of larval Atlantic herring (Clupea harengus L.), are unaffected by elevated pCO2. Journal of Experimental Marine Biology and Ecology 466:42–48,

A95. Reglero P, Blanco E, Ortega A, Fiksen Ø, Gándara F, Seoka M, Viguri FJ, Folkvord A (2015) Prey selectivity in piscivorous bluefin tuna larvae reared in the laboratory. Journal of Plankton Research 37:2-5,

A94. Jung K-M, Folkvord A, Kjesbu OS, Sundby S (2014) Experimental parameterisation of principal physics in buoyancy variations of marine teleost eggs. PLoS One 9:e104089,

A93. Frommel AY, Maneja RH, Lowe DM, Pascoe C, Geffen AJ, Folkvord A, Piatkowski U, Clemmesen C (2014) Organ damage in Atlantic herring larvae as a result of ocean acidification. Ecological Applications 24:1131-1143,

A92. Reglero P, Ortega A, Blanco E, Fiksen Ø, Viguri FJ, Gándara F, Seoka M, Folkvord A (2014) Size-related differences in growth and survival in piscivorous fish larvae fed different prey types. Aquaculture 433:94-101,

A91. Folkvord A, Jørgensen C, Korsbrekke K, Nash RDM, Nilsen T, Skjæraasen JE (2014) Trade-offs between growth and reproduction in wild Atlantic cod. Canadian Journal of Fisheries and Aquatic Sciences 71:1106-1112,

A90. Imsland AK, Ólafsson K, Skírnisdóttir S, Gunnarsson S, Oddgeirsson M, van Damme S, Helyar SJ, Skadal J, Folkvord A (2014) Life history of turbot in Icelandic waters: intra- and inter-population genetic diversity and otolith tracking of environmental temperatures. Fisheries Research 155:185-193,

A89. Vollset KW, Catalán IA, Fiksen Ø, Folkvord A (2013) Effect of food deprivation on distribution of larval and early juvenile cod in experimental vertical temperature and light gradients. Marine Ecology Progress Series 475:191-201,

A88. Ritchie K, Folkvord A, Hufthammer AK (2013) Oxygen isotope ratios in cod otoliths used to reveal seasonality of fishing at Late Mesolithic sites in Denmark. Archaeofauna 22:95-104,

A87. Maneja RH, Frommel AY, Geffen AJ, Folkvord A, Piatkowski U, Chang MY, Clemmesen C (2013) Effects of ocean acidification on the calcification of otoliths of larval Atlantic cod, Gadus morhua L. Marine Ecology Progress Series 477:251–258,,

A86. Hansen TW, Folkvord A, Grøtan E, Sæle Ø (2013) Genetic ontogeny of pancreatic enzymes in Labrus bergylta larvae and the effect of feed type on enzyme activities and gene expression. Comp Biochem Physiol B Biochem Mol Biol 164:176-184,

A85. Karlson S, Michalsen K, Folkvord A (2013) Age determination of Atlantic halibut (Hippoglossus hippoglossus L.) along the coast of Norway - status, and future potentials. ICES Journal of Marine Science 70:50-55,

A84. Maneja RH, Frommel AY, Browman HI, Skiftesvik AB, Clemmesen C, Geffen AJ, Folkvord A, Piatkowski U, Durif C, Bjelland R (2013) The swimming kinematics of larval Atlantic cod, Gadus morhua L., are resilient to elevated seawater pCO2. Marine Biology 160:1963-1972,

A83. Jung K-M, Folkvord A, Kjesbu OS, Agnalt A-L, Thorsen A, Sundby S (2012) Egg buoyancy variability in local populations of Atlantic cod (Gadus morhua). Marine Biology 159:1969-1980,

A82. Ferreira AS, Stenevik EK, Vollset KW, Korneliussen R, Folkvord A (2012) Vertical migration of Norwegian spring-spawning herring larvae in relation to predator and prey distribution. Marine Biology Research 8:605-614,

A81. Meyer S, Caldarone EM, Chicharo MA, Clemmesen C, Faria AM, Faulk C, Folkvord A, Holt GJ, Høie H, Kanstinger P, Malzahn A, Moran D, Petereit C, Støttrup JG, Peck MA (2012) On the edge of death: Impacts of temperature, species and body size on RNA-DNA ratios of starving marine fish larvae. Journal of Marine Systems 93:11-24,

A80. Frommel AY, Maneja RH, Lowe DM, Malzahn A, Geffen A, Folkvord A, Piatkowski U, Reusch T, Clemmesen C (2012) Severe tissue damage in Atlantic cod larvae under increasing ocean acidification. Nature Climate Change 2:42-46,

A79. Koedijk R, Imsland AK, Folkvord A, Stefansson SO, Jonassen TM, Foss A (2012) Larval rearing environment influences the physiological adaptation in juvenile Atlantic cod, Gadus morhua. Aquaculture International 20:467–479,

A78. Catalán IA, Vollset KW, Morales-Nin B, Folkvord A (2011) The effect of temperature gradients and stomach fullness on the vertical distribution of larval herring in experimental columns. Journal of Experimental Marine Biology and Ecology 404:26-32,

A77. Geffen AJ, Høie H, Folkvord A, Hufthammer AK, Andersson C, Ninnemann U, Pedersen RB, Nedreaas K (2011) High-latitude climate variability and its effect on fisheries resources as revealed by fossil cod otoliths. ICES Journal of Marine Science 68:1081-1089,

A76. Vollset KW, Folkvord A, Browman HI (2011) Foraging behaviour of larval cod (Gadus morhua) at low light intensities. Marine Biology 158:1125-1133,

A75. Imsland AK, Koedijk R, Stefansson SO, Foss A, Hjorleifsdottir S, Hreggvidsson GO, Otterlei E, Folkvord A (2011) A retrospective approach to fractionize variation in body mass of Atlantic cod Gadus morhua. Journal of Fish Biology 78:251-264,

A74. Meier S, Morton HC, Nyhammer G, Grøsvik BE, Makhotin V, Geffen A, Boitsov S, Kvestad KA, Bohne-Kjersem A, Goksøyr A, Folkvord A, Klungsøyr J, Svardal A (2010) Development of Atlantic cod (Gadus morhua) exposed to produced water during early life stages. Effects on embryos, larvae, and juvenile fish. Marine Environmental Research 70:383-394,

A73. Folkvord A, Koedijk R, Lokøy V, Imsland AK (2010) Timing and selectivity of mortality in reared Atlantic cod revealed by otolith analysis. Environmental Biology of Fishes 89:513-519,

A72. Seljeset O, Vollset K, Folkvord A, Geffen AJ (2010) The role of prey concentration and size range in the growth and survival of larval cod. Marine Biology Research 6:251-262,

A71. Koedijk RM, le Francois NR, Blier PU, Foss A, Folkvord A, Ditlacadet D, Lamarre SG, Stefansson SO, Imsland AK (2010) Ontogenetic effects of diet during early development on growth performance, myosin mRNA expression and metabolic enzyme activity in Atlantic cod juveniles reared at different salinities. Comparative Biochemistry and Physiology A 156:102-109,

A70. Catalán IA, Folkvord A, Palomera I, Quilez-Badia G, Kallianoti F, Tselepides A, Kallianotis A (2010) Growth and feeding patterns of European anchovy (Engraulis encrasicolus) early life stages in the Aegean Sea (NE Mediterranean). Estuarine, Coastal and Shelf Science 86:299-312,

A69. Koedijk RM, Folkvord A, Foss A, Pittman K, Stefansson SO, Handeland S, Imsland AK (2010) The influence of first-feeding diet on the Atlantic cod Gadus morhua phenotype: survival, development and long-term consequences for growth. Journal of Fish Biology 77:1-19,

A68. Hufthammer AK, Høie H, Folkvord A, Geffen AJ, Andersson CA, Ninnemann US (2010) Seasonality of human site occupation based on stable oxygen isotope ratios of cod otoliths. Journal of Archaeological Science 37:78-83,

A67.   Bat NK, CT Vinh, A Folkvord, A Johannessen, K Tsuchiya, S Segawa, 2009. Age and  growth of mitre squid Photololigo chinensis in the Tonkin Gulf of Vietnam based on statolith microstructure. La mer. 47, 57-65.

A66.   Vollset KW, Ø Fiksen, A Folkvord, 2009. Vertical distribution of larval cod (Gadus morhua) in experimental temperature gradients. Journal of Experimental Marine Biology and Ecology. 379, 16-32.

A65.   Folkvord A, Ø Fiksen, H Høie, A Johannessen, E Otterlei, KW Vollset, 2009. What can size distributions within cohorts tell us about ecological processes in fish larvae? Scientia Marina. 74, 119-130.

A64.   Husebø Å, EK Stenevik, A Slotte, P Fossum, A Salthaug, F Vikebø, S Aanes, A Folkvord, 2009. Effects of hatching time on year-class strength in Norwegian spring spawning herring (Clupea harengus). ICES Journal of Marine Science. 66, 1710-1717.

A63.   Folkvord A, H Høie, A Johannessen, T Solbakken, 2009. Effects of prey concentration, light regime and parental origin on growth and survival of herring larvae under controlled experimental conditions. ICES Journal of Marine Science. 66, 1702-1709.

A62.   Vollset KW, O Seljeset, Ø Fiksen, A Folkvord, 2009. A common garden experiment with larval Northeast Arctic and Norwegian coastal cod cohorts in replicated mesocosms. Deep-Sea Research II. 56, 1984-1991.

A61.   Busch KET, A Folkvord, H Otterå, WF Hutchinson, T Svåsand, 2009. Effects of female spawning experience and larval size on feeding and growth of cod larvae Gadus morhua L. reared in mesocosms. Marine Biology Research. 5, 286-296.

A60.   Urtizberea A, Ø Fiksen, A Folkvord, X Irigoien, 2008. Modelling growth of larval anchovies including diel feeding patterns, temperature and body size. Journal of Plankton Research. 30, 1369-1383.

A59.   Li L, H Høie, AJ Geffen, E Heegard, J Skadal, A Folkvord, 2008. Back-calculation of previous fish size using individually tagged and marked Atlantic cod (Gadus morhua). Canadian Journal of Fisheries and Aquatic Sciences. 65, 2496-2508.

A58.   Høie H, A Folkvord, H Mosegaard, L Li, LA Clausen, B Nordberg, AJ Geffen, 2008. Restricted fish feeding reduces cod otolith opacity. Journal of Applied Ichthyology. 24, 138-143.

A57.   Imsland AK, A Foss, R Koedijk, A Folkvord, SO Stefansson, TM Jonassen, 2007. Persistent growth effects of temperature and photoperiod in Atlantic cod Gadus morhua. Journal of Fish Biology. 71, 1371-1382.

A56.   González-Quirós R, I Munuera , A Folkvord, 2007. Cell cycle analysis of brain cells as a growth index in larval cod at different feeding conditions and temperatures. Scientia Marina. 71, 485-497.

A55.   Tian T, Ø Fiksen, A Folkvord, 2007. Estimating larval fish growth under size-dependent mortality: a numerical analysis of bias. Canadian Journal of Fisheries and Aquatic Sciences. 64, 554-562.

A54.   Imsland AK, A Foss, T Alvseike, A Folkvord, SO Stefansson, TM Jonassen, 2007. Interaction between temperature and photoperiod on growth and feeding of Atlantic cod (Gadus morhua): possible secondary effects. Canadian Journal of Fisheries and Aquatic Sciences. 64, 239-248.

A53.   Kristiansen T, Ø Fiksen, A Folkvord, 2007. Modelling feeding, growth, and habitat selection in larval Atlantic cod (Gadus morhua): observations and model predictions in a macrocosm environment. Canadian Journal of Fisheries and Aquatic Sciences. 64, 136-151.

A52.   Bang A, P Grønkjær, A Folkvord, 2007. Possible fitness costs of high and low standard metabolic rates in larval herring Clupea harengus, as determined by otolith microstructure. Marine Ecology Progress Series. 331, 233-242.

A51.   Meisfjord J, F Midtøy, A Folkvord, 2006. Validation of daily increment deposition in otoliths of juvenile Limnothrissa miodon (Clupeidae). Journal of Fish Biology. 69, 1845-1848.

A50.   Imsland AK, A Foss, R Koedijk, A Folkvord, SO Stefansson, TM Jonassen, 2006. Short- and long-term differences in growth, feed conversion efficiency and deformities in juvenile Atlantic cod (Gadus morhua) startfed on rotifers and zooplankton. Aquaculture Research. 37, 1015-1027.

A49.   Caldarone EM, CM Clemmesen, E Berdalet, TJ Miller, A Folkvord, GJ Holt, MP Olivar, IM Suthers, 2006. Intercalibration of four spectrofluorometric protocols for measuring RNA/DNA ratios in larval and juvenile fish. Limnology and Oceanography: Methods. 4, 153-163.

A48.   Simonsen CS, P Munk, A Folkvord, SA Pedersen, 2006. Feeding ecology of Greenland halibut and sandeel larvae off West Greenland. Marine Biology. 149, 937-952.

A47.   Imsland AK, A Foss, A Folkvord, SO Stefansson, TM Jonassen, 2005. The interrelation between temperature regimes and fish size in juvenile Atlantic cod (Gadus morhua): effects on growth and feed conversion efficiency. Fish Physiology and Biochemistry. 31, 347-361.

A46.   Imsland AK, A Foss, A Folkvord, SO Stefansson, TM Jonassen, 2005. Genotypic response to photoperiod treatment in Atlantic cod (Gadus morhua). Aquaculture. 250, 525-532.

A45.   Høie H, A Folkvord, 2006. Estimating the timing of growth rings in Atlantic cod otoliths using stable oxygen isotopes. Journal of Fish Biology. 68, 826-837.

A44.   Imsland AK, A Foss, G Nævdal, T Johansen, A Folkvord, SO Stefansson, TM Jonassen, 2004. Variations in growth in haemoglobin genotypes of Atlantic cod. Fish Physiology and Biochemistry. 30, 47-55.

A43.   Folkvord A, 2005. Comparison of size-at-age of larval Atlantic cod (Gadus morhua) from different populations based on size- and temperature-dependent growth models. Canadian Journal of Fisheries and Aquatic Sciences. 62, 1037-1052.

A42.   Folkvord A, A Johannessen, E Moksness, 2004. Temperature dependent otolith growth in Norwegian spring spawning herring (Clupea harengus) larvae. Sarsia. 89, 297-310.

A41.   Skajaa K, A Fernö, A Folkvord, 2004. Ontogenetic- and condition-related effects of starvation on responsiveness in herring larvae (Clupea harengus L.) during repeated attacks by a model predator. Journal of Experimental Marine Biology and Ecology. 312, 253-269.

A40.   Høie H, CA Dahl, A Folkvord, Ø Karlsen, 2004. Precision and accuracy of stable isotope signals in otoliths of pen reared cod (Gadus morhua L.) when sampled with high resolution micromill. Marine Biology. 144, 1039-1049.

A39.   Høie H, E Otterlei, A Folkvord, 2004. Temperature-dependent fractionation of stable oxygen isotopes in otoliths of juvenile cod (Gadus morhua L). ICES Journal of Marine Science. 61, 243-251.

A38.   Fox CJ, A Folkvord, AJ Geffen, 2003. Otolith micro-increment formation in herring Clupea harengus larvae in relation to growth rate. Marine Ecology Progress Series. 264, 83-94.

A37.   Skajaa K, A Fernö, A Folkvord, 2003. Swimming, feeding and predator avoidance in cod larvae (Gadus morhua L.): trade-offs between hunger and predation risk. in: Browman HI, AB Skiftesvik (Eds.), The Big Fish Bang: Proceedings of the 26th Annual Larval Fish Conference. Institute of Marine Research, Bergen, Norway, pp. 105-121.

A36.   Høie H, A Folkvord, E Otterlei, 2003. Effect of somatic and otolith growth rate on stable isotopic composition of early juvenile cod (Gadus morhua L) otoliths. Journal of Experimental Marine Biology and Ecology. 289, 41-58.

A35.   Tytler P, CJ Fox, A Folkvord, 2002. Glycoconjugates in the otolithic membrane of herring larvae: a possible framework for encoding the life history recorder in fishes. Journal of Fish Biology. 61, 39-49.

A34.   Otterlei E, A Folkvord, G Nyhammer, 2002. Temperature dependent otolith growth of larval and early juvenile Atlantic cod (Gadus morhua). ICES Journal of Marine Science. 59, 851-860.

A33.   Johannessen A, G Blom, A Folkvord, 2000. Differences in growth pattern between spring and autumn spawned herring (Clupea harengus L.) larvae. Sarsia. 85, 461-466.

A32.   Folkvord A, G Blom, A Johannessen, E Moksness, 2000. Growth dependent age estimation in herring (Clupea harengus L.) larvae. Fisheries Research. 46, 91-103.

A31.   Otterlei E, G Nyhammer, A Folkvord, SO Stefansson, 1999. Temperature- and size-dependent growth of larval and early juvenile Atlantic cod (Gadus morhua): a comparative study of Norwegian coastal cod and northeast Arctic cod. Canadian Journal of Fisheries and Aquatic Sciences. 56, 2099-2111.

A30.   Fiksen Ø, A Folkvord, 1999. Modelling growth and ingestion processes in herring Clupea harengus larvae. Marine Ecology Progress Series. 184, 273-289.

A29.   Høie H, A Folkvord, A Johannessen, 1999. The influence of different  parental combinations and incubation temperature on the RNA and DNA content of herring (Clupea harengus L.) larvae at hatching: a pilot study. Journal of Fish Biology. 55, 110-118.

A28.   Høie H, A Folkvord, A Johannessen, 1999. Maternal, paternal and temperature effects on otolith size of young herring (Clupea harengus L.) larvae. Journal of Experimental Marine Biology and Ecology. 234, 167-184.

A27.   Suneetha K-B, A Folkvord, A Johannessen, 1999. Responsiveness of selected condition measures of herring, Clupea harengus, larvae to starvation in relation to ontogeny and temperature. Environmental Biology of Fishes. 54, 191-204.

A26.   Imsland AK, T Nilsen, A Folkvord, 1998. Stochastic simulation of size variation in turbot: possible causes analysed with an individual-based model. Journal of Fish Biology. 53, 237-258.

A25.   Sunde LM, A Imsland, A Folkvord, SO Stefansson, 1998. Effects of size grading on growth and survival of juvenile turbot at two temperatures. Aquaculture International. 6, 19-32.

A24.   Blom G, A Folkvord, A Johannessen, P Fossum, 1997. Interannual variations in condition indices of larval Norwegian spring-spawning herring. Journal of Fish Biology. 51, 370-384.

A23.   Folkvord A, K Rukan, A Johannessen, E Moksness, 1997. Early life history of herring larvae in contrasting feeding environments determined by otolith microstructure analysis. Journal of Fish Biology. 51, 250-263.

A22.   Blom G, A Folkvord, 1997. A snapshot of cannibalism in 0-group Atlantic cod (Gadus morhua) in a marine pond. Journal of Applied Ichthyology-Zeitschrift Fur Angewandte Ichthyologie. 13, 177-181.

A21.   Imsland AK, A Folkvord, ÓDB Jónsdóttir, SO Stefansson, 1997. Effects of exposure to extended photoperiods during the first winter on long-term growth and age at first maturity in turbot (Scophthalmus maximus). Aquaculture. 159, 125-141.

A20.   Bergstad OA, A Folkvord, 1997. Dispersal of tagged juvenile turbot Scophthalmus maximus on the Norwegian Skagerrak coast. Fisheries Research. 29, 211-215.

A19.   Imsland AK, A Folkvord, GL Grung, SO Stefansson, GL Taranger, 1997. Sexual dimorphism in growth and maturation of turbot, Scophthalmus maximus (Rafinesque, 1810). Aquaculture Research. 28, 101-114.

A18.   Folkvord A, L Ystanes, A Johannessen, E Moksness, 1996. RNA:DNA ratios and growth of herring (Clupea harengus L.) larvae reared in mesocosms. Marine Biology. 126, 591-602.

A17.   Imsland AK, LM Sunde, A Folkvord, SO Stefansson, 1996. The interaction of temperature and fish size on growth of juvenile turbot. Journal of Fish Biology. 49, 926-940.

A16.   Folkvord A, IM Haaland, O Grahl-Nielsen, A Johannessen, 1996. Fatty acid composition of herring (Clupea harengus L.) larvae off western Norway. Sarsia. 8, 299-305.

A15.   Stenevik EK, P Fossum, A Johannessen, A Folkvord, 1996. Identification of Norwegian spring spawning herring (Clupea harengus L.) larvae from spawning grounds off western Norway applying otolith microstructure analysis. Sarsia. 80, 285-292.

A14.   Hallaråker H, A Folkvord, SO Stefansson, 1995. Growth of juvenile halibut (Hippoglossus hippoglossus L.) related to temperature, day length and feeding regime. Netherlands Journal of Sea Research. 34, 139-147.

A13.   Imsland AK, A Folkvord, SO Stefansson, 1995. Growth, oxygen consumption and activity of juvenile turbot (Scophthalmus maximus) reared under different temperatures and photo periods. Netherlands Journal of Sea Research. 34, 149-159.

A12.   Folkvord A, E Moksness, 1995. RNA/DNA ratios and growth of herring larvae. Marine Ecology Progress Series. 121, 311-312.

A11.   Folkvord A, V Øiestad, PG Kvenseth, 1994. Growth patterns of three cohorts of Atlantic cod larvae (Gadus morhua L.) studied in a macrocosm. ICES Journal of Marine Science. 51, 325-336.

A10.   Folkvord A, G Blom, O Dragesund, A Johannessen, O Nakken, G Nævdal, 1994. A conceptual framework for enhancing and stabilizing recruitment of marine fish stocks. Aquaculture and Fisheries Management. 25, 245-258.

A09.   Folkvord A, 1993. Prey recognition in stomachs of cannibalistic juvenile cod (Gadus morhua L.). Sarsia. 78, 97-100.

A08.   Otterå H, A Folkvord, 1993. Allometric growth in juvenile cod (Gadus morhua L.) and possible effects on cannibalism. Journal of Fish Biology. 43, 643-645.

A07.   Folkvord A, H Otterå, 1993. Effects of initial size distribution, day length and feeding frequency on growth, survival and cannibalism in juvenile Atlantic cod (Gadus morhua L.). Aquaculture. 114, 243-260.

A06.   Nævdal G, A Folkvord, E Otterlei, S Thorkildsen, 1992. Growth rate related to genotype of 0-group cod at three environmental temperatures. Sarsia. 77, 71-73.

A05.   Booman CI, A Folkvord, JR Hunter, 1991. Responsiveness of starved northern anchovy Engraulis mordax larvae to predatory attacks by adult anchovy. Fishery Bulletin U.S. 89, 707-711.

A04.   Folkvord A, 1991. Growth, survival and cannibalism of cod juveniles (Gadus morhua L.): effects of feed type, starvation and fish size. Aquaculture. 97, 41-59.

A03.   Øiestad V, T Pedersen, A Folkvord, Å Bjordal, PG Kvenseth, 1987. Automatic feeding and harvesting of juvenile Atlantic cod (Gadus morhua L.) in a pond. Modelling Identification and Control. 8, 39-46.

A02.   Folkvord A, JR Hunter, 1986. Size specific vulnerability of northern anchovy (Engraulis mordax) larvae to predation by fishes. Fishery Bulletin U.S. 84, 859-869.

A01.   Øiestad V, PG Kvenseth, A Folkvord, 1985. Mass production of Atlantic cod juveniles (Gadus morhua) in a Norwegian saltwater pond. Transactions of the American Fisheries Society. 114, 590-595.


Other scientific publications with original content:


B19.   Sundby S, P Fossum, AD Sandvik, FB Vikebø, A Aglen, L Buhl-Mortensen, A Folkvord, K Bakkeplass, P Buhl-Mortensen, M Johannessen, M Jørgensen, T Kristiansen, CS Landa, MS Myksvoll, RDM Nash, 2013. KunnskapsInnhenting Barentshavet – LOfoten – Vesterålen (KILO). Fisken og Havet, 1-188.

B18.   Kristiansen T, Ø Fiksen, A Folkvord, 2005. Larval cod in ponds and models:the role of habitat selection. Globec International Newsletter. 11, 22.

B17.   Belchier M, C Clemmesen, D Cortés, A Doan, A Folkvord, A García, AJ Geffen, H Høie, A Johannessen, E Moksness, Hd Pontual, T Rámirez, D Schnack, B Sveinsbø, 2004. Recruitment studies: Manual of tools for recruitment studies. ICES Techniques in Marine Environmental Science, 1-35.

B16.   Salvanes AGV, A Aksnes, A Folkvord, M Hordnes, H Høie, F Midtøy, J Nejstgaard, E Otterlei, JE Skjæraasen, T Sørnes, AC Utne-Palm, 2003. Environmental influence on fish stocks. IFM Rapport, 1-21.

B15.   Høie H, A Folkvord, A Johannessen, 2000. A multivariate analysis of condition of herring larvae from different environmental conditions, ICES Council Meeting 2000/R:04, pp. 1-10.

B13.   Folkvord A, E Otterlei, A Svardal, 1999. Alkylerte fenolers hormonelle virkning på torsk - generasjonseffekter. Vekst, overlevelse og kjønnsdifferensiering. IFM Rapport, 1-11.

B12.   Fiksen Ø, A Folkvord, 1999. Maternal effects and the benefit of yolk supply in cod larvae in different enfironments - a simulation model, ICES Council Meeting 1999/Y:21, pp. 1-6.

B12.   Fiksen Ø, E Otterlei, A Folkvord, 2000. Experiments and models as reciprocal tools to understand environmental links in recruitment dynamics, ICES Council Meeting 2000/R:02, pp. 1-8.

B11.   Folkvord A, G Blom, JV Helvik, BI Holmqvist, A Johannessen, E Otterlei, SO Stefansson, 1998. Teori- og eksperimentbasert studie av rekruttering hos marin fisk - Sluttrapport fra del I av strategisk program. IFM Rapport, 1-46.

B10.   Folkvord A, A Johannessen, 1998. Otolitt mikrostruktur, presisjon, nøyaktighet og anvendelse. IFM Rapport, 1-19.

B09.   Moksness E, K Rukan, L Ystanes, A Folkvord, A Johannessen, 1995. Comparison of somatic and otolith growth in North Sea herring (Clupea harengus L.) larvae; evaluation of growth dynamics in mesocosms. University of South Carolina Press, Columbia.

B08.   Johannessen A, G Blom, A Folkvord, H Svendsen, 1995. The effect of local wind on the distribution of Norwegian spring spawning herring (Clupea harengus L.) larvae. Elsevier, Amsterdam.

B07.   Folkvord A, 1993. Experimental studies of growth and mortality of larval and juvenile cod (Gadus morhua L.) with special emphasis on cannibalism. University of Bergen, pp. 153.

B06.   Folkvord A, H Otterå, 1991. Effects of size distribution, feeding and light regime on growth and survival of cod juveniles in tanks. European Aquaculture Society, Ghent, Belgium.

B05.   Folkvord A, 1989. Growth and cannibalism of cod fry (Gadus morhua) in intensive systems. European Aquaculture Society, Bredene, Belgium.

B04.   Øiestad V, A Folkvord, PG Kvenseth, 1985. Growth-patterns of Atlantic cod larvae (Gadus morhua L.) from first feeding to metamorphosis studied in a mesocosm, ICES Council Meeting 1985/Mini-Symposium/No. 9.

B03.   Folkvord A, PG Kvenseth, T Pedersen, V Øiestad, 1985. Mass-production of juvenile cod (Gadus morhua L.) in a pond: results and new approaches in 1985, ICES Council Meeting 1985/F:63.

B02.   Folkvord A, 1985. Size specific vulnerability of northern anchovy Engraulis mordax larvae to predation by fishes. University of California, San Diego, pp. 96.

B01.   Øiestad V, PG Kvenseth, A Folkvord, 1983. Mass-production of cod fry in a pond in Western Norway. with additional feeding post-metamorphosis, ICES Council Meeting 1983/F:5, pp. 10.


Overview papers, reviews, book chapters, synopsis etc.


C06.   Mosegaard H, A Folkvord, PJ Wright, 2002. V. Some uses of individual age data. B. Ecological applications. Ifremer-IRD coedition, Brest, France.

C05.   Folkvord A, H Mosegaard, 2002. V. Some uses of individual age data. A. Age and growth analysis. Ifremer-IRD coedition, Brest, France.

C04.   Wright PJ, J Panfili, A Folkvord, H Mosegaard, J Meunier, 2002. IV. Validation methods. A. Direct validation. Ifremer-IRD coedition, Brest, France.

C03.   Butler JL, A Folkvord, 2000. Section 2: Estimation of fish age and growth. Fisheries Research. 46, 51-52.

C02.   Folkvord A, 1997. Ontogeny of cannibalism in larval and juvenile fish with special emphasis on Atlantic cod, Gadus morhua L. Chapman and Hall Ltd., London.

C01.   Folkvord A, H Botnen, PG Kvenseth, 1991. Erfaringer med oppdrett av torskeyngel i poller i Sør-Norge. Institute of Marine Research, Bergen, Norway.


Participant and WP leader of RCN project HypOnFjordFish 2020-2024

Leader of RCN GENSINC project, 2016-2020

Leader of RCN project EMBRC Norway 2016-2020

Participant in IMR project RECNOR 2017-2020



M.Sc. University of San Diego, USA 1985

Dr. scient. Universitetet i Bergen 1993

Universitetspedagogikk 1994, 2016