Home page of professor Raine Kortet

      Dr. Raine Kortet, Professor.

      Current address:
      Department of Environmental & Biological Sciences,
      University of Eastern Finland,
      P.O. Box 111, FI-80101 Joensuu, Finland.
      Phone: +358 500 957749.
      Fax: +358 13 318 039.
      E-mail (# to be removed): raine.kortet#@uef.fi

      Research group:

      Citation metrics:

Personal information

Name: Raine Kalevi Kortet.
Sex: Male.
Date and place of birth: September 10, 1975, Raahe, Finland.
Languages: Finnish, English, Swedish.

Education & Qualifications


Ph.D., University of Jyväskylä, 29.1. 2003 (Limnology and Hydrobiology "excellent"). Thesis dissertation (Parasitism, Reproduction and Sexual Selection of Roach, Rutilus rutilus L.) awarded with the grade "approved with honours", the most outstanding dissertations, only approx. 5% of the total.

MSc, University of Jyväskylä, 13.4. 2000 (Hydrobiology Laudatur "good", Ecology Cumlaude Approbatur "good", Biology Approbatur "excellent", Statistics Approbatur "good", Computer Sciences some courses, Environmental Sciences some courses, Chemistry some courses). 2002 Education Approbatur "good".


Adjunct Professor (‘Dosentti’), University of Oulu, 1.7. 2009-. Subject: Animal Ecology.

Adjunct Professor (‘Dosentti’), University of Helsinki, 4.4. 2007-. Subject: Ecology and Evolutionary Biology. Teaching skills “excellent” evaluated by the faculty 29.1. 2007.

Adjunct Professor ('Dosentti'), University of Jyväskylä, 1.1. 2005-. Subject: Hydrobiology and Limnology: Ecological Immunology / Ecological Parasitology. Teaching skills "good" evaluated by the faculty 25.8. 2004.

Areas of specialization

Aquatic ecology, evolutionary ecology, behavioral ecology, ecological parasitology and ecological immunology: empirical and experimental research using mainly fishes, crayfishes, insects and arachnids.

Main Research Interests

I am located at the Joensuu campus of the University of Eastern Finland, where I am leading the research group of Aquatic Ecology and Behavioural Ecology. My team consists of several senior researchers and post-docs, as well as PhD and MSc students. The research group collaborates actively with many Finnish (e.g. Universities of Helsinki, Oulu, Jyväskylä and Turku) and international (e.g. USA, Canada, Germany, UK, Norway) research partners.

I am personally interested in various topics, including life-history trade-offs and the interplay between sexual selection and natural selection (i.e. predation and parasitism). Recently, I have been involved with the research on animal personalities. I am also interested in applied aspects of aquatic ecology and parasitology (e.g. fish and crayfish diseases as a potential tool to monitor health of the aquatic ecosystems).

My work can be categorised into following areas:

- Life-history trade-offs, sexual selection and parasitism in teleosts (e.g. sexual ornamentation, spawning behaviour, intrasexual competition and sperm competition).

- Seasonal changes in immunological parameters and hormone levels.

- Animal personalities and behavioural syndromes in general.

- Predator-prey interactions in fishes and in insects (e.g. indirect cues and anti-predatory responses).

- Survival of the hathery-reared fish releases in wild environment (finding out ways to improve the survival).

- Behavioural and immunological ecology of crayfishes including crayfish plague and other disease-related problems.

- Papillomatosis (epidermic hyperplasia, a skin disease of fishes), and its potential use as a indicator-tool of ecosystem health.

- Studies related to sexual selection for indirect (i.e. pheromone) cues.

- 'Ecological parasitology' and 'ecological immunology' in invertebrates. I have been working with e.g. snails, beetles, field crickets and wolf spiders.

- The distribution, invasion and impacts of the deer ked (Lipoptena cervi (L.) on the host species.

See my publications listed by the topics.

Research & working experience

University of Eastern Finland, Department of Environmental & Biological Sciences:
Professor 1.1.2016 - .
University of Western Ontario, Department of Biology:
Visiting Professor 1.5.2016 - 28.2.2017.
University of Eastern Finland, Department of Biology:
Professor 1.1.2010 - 31.12.2015 (head of the department: 1.1.2014-31.12.2015).
University of Joensuu, Faculty of Biosciences:
Professor 1.1.2009 - 31.12.2009 (on leave of absence: 1.1.2009-30.6.2009).
University of Oulu, Department of Biology:
University researcher 1.8.2007-30.6.2009,
Assistant Professor 1.10.2006-30.6.2009 (on leave of absence: 1.10.2006-28.2.2007 & 1.8.2007-30.6.2009).
University of Helsinki, Department of Biological & Environmental Sciences:
University researcher 1.8.2005-28.2.2007.
University of California, Davis, Neurobiology, Physiology & Behavior:
Postdoctoral researcher 2.8.2003-29.7.2005.
University of Jyväskylä, Department of Biological and Environmental Science:
Postdoctoral researcher 1.2.2003-31.7.2003,
Researcher (PhD-student) 14.4.2000-31.1.2003,
Research assistant 1.1.1999-13.4.2000,
Conference assistant 1.9.1999-13.9.1999.
Raahe Steel, Research Center:
Researcher (study of biology of Lake Kuljunlahti roach) 3.2.1999-20.12.1999.
Raahe Steel, Coking plant, Dry-quenching plant:
Factory worker (three months per year) 1995, 1996, 1997, 1998.
Finnish Defence Forces, Sodankylä Artillery Unit:
Fire observer, gunner (national service) 22.8.1994-18.4.1995.
Raahen seudun metsänhoitoyhdistys:
Forestry worker (one month) 1994.

Papers in international journals and books

(If not linked, ask for PDF by email!)

2021 & in press

YARIPOUR, S., KEKÄLÄINEN, J., HUUSKONEN, H., JANHUNEN, M. & KORTET, R. 2021: Ultra-acute exposure to cadmium does not impair whitefish sperm motility. Journal of Fish Biology, in press.

HATANPÄÄ, A., HUUSKONEN, H., JANHUNEN, M., KORTET, R. & PIIRONEN, J. 2021: Spawning season movements of transported landlocked Atlantic salmon in a newly restored river habitat. Canadian Journal of Fisheries and Aquatic Sciences, 78: 184-192.

ERONEN, A., KEKÄLÄINEN, J., PIIRONEN, J., HYVÄRINEN, P., HUUSKONEN, H., JANHUNEN, M., YARIPOUR, S. & KORTET, R. 2021: Sperm motility and offspring pre- and post-hatching survival in hybridization crosses among a landlocked and two anadromous Atlantic salmon populations: implications for conservation. Canadian Journal of Fisheries and Aquatic Sciences, 78: 483-492.


YARIPOUR, S., KEKÄLÄINEN, J., HYVÄRINEN, P., KAUNISTO, S., PIIRONEN, J., VAINIKKA, A., KOLJONEN, M-L., KOSKINIEMI, J. & KORTET, R. 2020: Does enriched rearing during early life affect sperm quality or skin colouration in the adult brown trout? Aquaculture, 529: 735648.

KYNKÄÄNNIEMI, KORTET, R. & LAAKSONEN, S. 2020: Range expansion and reproduction of the ectoparasitic deer ked (Lipoptena cervi) in its novel host, the Arctic reindeer (Rangifer tarandus tarandus) in Finland. Parasitology Research, 119: 3113-3117.

HUUSKONEN, H., SUBIRON I FOLGUERA, J., KORTET, R., AKKANEN, J., VAINIKKA, A., JANHUNEN, M., KEKÄLÄINEN, J. 2020: Do whitefish (Coregonus lavaretus) larvae show adaptive variation in the avoidance of microplastic ingestion? Environmental Pollution, 262: 114353.

HATANPÄÄ, A., HUUSKONEN, H., KEKÄLÄINEN, J., KORTET, R., HYVÄRINEN, P., VITELLETTI, M.L. & PIIRONEN, J. 2020: Early winter foraging success, swimming performance and morphology of juvenile landlocked Atlantic salmon reared under semi-wild and hatchery conditions. Canadian Journal of Fisheries and Aquatic Sciences, 77: 770-778.

ADAIR, K.L., BOST, A., BUENO, E., KAUNISTO, S., KORTET, R., PETERS-SCHULZE, G., MARTINSON, V.G. & DOUGLAS, A.E. 2020: Host determinants of among-species variation in microbiome composition in drosophilid flies. The ISME Journal, 14: 217-229.


ÅGREN, A., VAINIKKA, A., JANHUNEN, M., HYVÄRINEN, P., PIIRONEN, J. & KORTET, R. 2019: Experimental crossbreeding reveals strain-specific variation in mortality, growth and personality in the brown trout (Salmo trutta). Scientific Reports, 9: 2771. (PDF)

MAKKONEN, J., KOKKO, H., GÖKMEN, G., WARD, J., UMEK, J., KORTET, R., PETRUSEK, A. & JUSSILA, J. 2019: The signal crayfish (Pacifastacus leniusculus) in Lake Tahoe (USA) hosts multiple Aphanomyces species. Journal of Invertebrate Pathology, 166: 107218.

SOININEN, N., BELINSKIJ, A., SIMILÄ, J. & KORTET, R. 2019: Too important to fail? Evaluating legal adaptive capacity for increasing coastal and marine aquaculture production in EU-Finland. Marine Policy, 110: 103498.

KORTET, R., LAAKKONEN, M.V.M, TIKKANEN, J., VAINIKKA, A. & HIRVONEN, H. 2019: Size-dependent stress response in juvenile Arctic charr (Salvelinus alpinus) under prolonged predator conditioning. Aquaculture Research, 50: 1482-1490. (PDF)

HUUSKONEN, H., PIIRONEN, J., SYVÄRANTA, J., ERONEN, R., BIASI, C., KIISKINEN, P., KORTET, R. & VAINIKKA, A. 2019: Diet and movements of pikeperch (Sander lucioperca) in a large oligotrophic lake with an exceptionally high pikeperch yield. Ecology of Freshwater Fish, 28: 533-543.

TRAKIMAS, G., KRAMS, R., KRAMA, T., KORTET, R., HAQUE, S., LUOTO, S., EICHLER INWOOD, S., BUTLER, D.M., JOERS, P., HAWLENA, D., RANTALA, M.J., ELFERTS, D., CONTRERAS-GARDUÑO, J. & KRAMS, I. 2019: Ecological stoichiometry: a link between developmental speed and physiological stress in an omnivorous insect. Frontiers in Behavioral Neuroscience, 13:42. (PDF)

PETRÓCZKI, K., VAINIKKA, A. & KORTET, R. 2019: Does acoustic environment modify boldness and related life-history traits in field cricket nymphs? Acta Ethologica, 22: 105-112. (PDF)


VÄÄTÄINEN, R., HUUSKONEN, H., HYVÄRINEN, P., KEKÄLÄINEN, J., KORTET, R., ARNEDO, M.T. & VAINIKKA, A. 2018: Do metabolic traits, vulnerability to angling, or capture method explain boldness variation in Eurasian perch? Physiological and Biochemical Zoology, 91: 1115-1128. (PDF)

KEKÄLÄINEN, J., OSKOEI, P., JANHUNEN, M., KOSKINEN, H., KORTET, R. & HUUSKONEN, H. 2018: Sperm pre-fertilization thermal environment shapes offspring phenotype and performance. Journal of Experimental Biology, 221: jeb181412. (PDF)

ALIORAVAINEN, N., HYVÄRINEN, P., KORTET, R., HÄRKÖNEN, L. & VAINIKKA, A. 2018: Survival of crossbred brown trout under experimental pike predation and stocking in the wild. Boreal Environment Research, 23: 267-281. (PDF)

FERGUSON, L.V., KORTET, R. & SINCLAIR, B.J. 2018: Eco-immunology in the cold: the role of immunity in shaping the overwintering survival of ectotherms. Journal of Experimental Biology, 221: jeb163873. (PDF)

YARIPOUR, S., ZAMANPOORE, M. & KORTET, R. 2018: Fin morphology variation in Aphanius farsicus in two local populations. Aquaculture Reports, 11: 38-41. (PDF)

KRAMS, I.A., TRAKIMAS, G., KECKO, S., ELFERTS, D., KRAMS, R., LUOTO, S., RANTALA, M.J., MÄND, R., KUUSIK, A., KEKÄLÄINEN, J., JÕERS, P., KORTET, R. & KRAMA, T. 2018: Linking organismal growth, coping styles, stress reactivity, and metabolism via responses against a selective serotonin reuptake inhibitor in an insect. Scientific Reports, 8: 8599. (PDF)

MAKKONEN, J., JUSSILA, J., PANTELEIT, J., KELLER, N.S., SCHRIMPHF, A., THEISSINGER, K., KORTET, R., MARTIN-TORRIJOS, L., SANDOVAL-SIERRA, J.V., DIEGUEZ-URIBEONDO, J. & KOKKO, H. 2018: MtDNA allows the sensitive detection and haplotyping of the crayfish plague disease agent Aphanomyces astaci showing clues about its origin and migration. Parasitology, 145: 1210-1218. (PDF)


KORTET, R., LAUTALA, T., KEKÄLÄINEN, J., TASKINEN, J. & HIRVONEN, H. 2017: Maternal effects in vulnerability to eye-parasites and correlations between behavior and parasitism in juvenile Arctic charr. Ecology and Evolution, 7: 8780-8787. (PDF)

KRAMS, I.A., NIEMELÄ, P.T., TRAKIMAS, G., KRAMS, R., BURGHARD, G.M., KRAMA, T., KUUSIK, A., MÄND, M., RANTALA, M.J., MÄND, R., KEKÄLÄINEN, J., SIRKKA, I., LUOTO, S. & KORTET, R. 2017: Metabolic rate associates with, but does not generate, covariation between behaviours in western stutter-trilling crickets, Gryllus integer. Proceedings of the Royal Society B, 284: 20162481. (PDF)


MAKKONEN, J., VESTERBACKA, A., MARTIN, F., JUSSILA, J., DIEGUEZ-URIBEONDO, J., KORTET, R. & KOKKO, H. 2016: Mitochondrial genomes and comparative genomics of Aphanomyces astaci and Aphanomyces invadans. Scientific Reports, 6: 36089. (PDF)

DIRIENZO, N., NIEMELÄ, P.T., HEDRICK, A.V. & KORTET, R. 2016: Adult bacterial exposure increases behavioral variation and drives higher repeatability in field crickets. Behavioral Ecology and Sociobiology, 70: 1941-1947. (PDF)

KARVONEN, A., AALTO-ARANEDA, M., VIRTALA, A-M., KORTET, R., KOSKI, P. & HYVÄRINEN, P. 2016: Enriched rearing environment and wild genetic background can enhance survival and disease resistance of salmonid fishes during parasite epidemics. Journal of Applied Ecology, 53: 213-221. doi: 10.1111/1365-2664.12568 (PDF)

GRUBER, C., TULONEN, J., KORTET, R. & HIRVONEN, H. 2016: Resource availability and predation risk influence contest behavior and dominance hierarchies in crayfish. Behavioral Ecology and Sociobiology, 70: 1305-1317. (PDF)

KLEMME, I., KORTET, R. & KARVONEN, A. 2016: Parasite infection in a central sensory organ of a fish does not affect host personality. Behavioral Ecology, 27: 1533-1538. (PDF)

MANDIMA, J., KORTET, R. & SARVALA, J. 2016: Limnothrissa miodon (Boulenger, 1906) in Lake Kariba: daily ration and population food consumption estimates, and potential application to predict the fish stock biomass from prey abundance. Hydrobiologia, 780: 99-111. (PDF)

KAUNISTO, S., RAUNISMAA, I., KORTET, R. & YLÖNEN, H. 2016: Summer time predation on the obligatory off-host stage of an invasive ectoparasite. Parasitology, 143: 1960–1973. (PDF)

LAI, Y.-T., KEKÄLÄINEN, J. & KORTET, R. 2016: Infestation with the parasitic nematode Philometra ovata does not impair behavioral sexual competitiveness or odor attractiveness of the male European minnow (Phoxinus phoxinus). Acta Ethologica, 19: 103-11. (PDF)

LAI, Y.-T., KEKÄLÄINEN, J. & KORTET, R. 2016: Does parasitic infection by the body cavity dwelling Philometra ovata (Nematoda) impair swimming performance of male European minnow (Phoxinus phoxinus)? Marine & Freshwater Behaviour & Physiology, 49: 47-61. (PDF)


JUSSILA J., RUOKONEN, T., SYVÄRANTA, J., KOKKO, H., VAINIKKA, A., MAKKONEN, J. & KORTET, R. 2015: It takes time to see the menu from the body: an experiment on stable isotope composition in freshwater crayfishes. Knowledge and Management of Aquatic Ecosystems, 416: 25. doi: 10.1051/kmae/2015021. (PDF)

DIRIENZO, N., NIEMELÄ, P.T., SKOG, A., VAINIKKA, A. & KORTET, R. 2015: Juvenile pathogen exposure affects the presence of personality in adult field crickets. Frontiers in Ecology and Evolution, 3(36): 1-10. doi: 10.3389/fevo.2015.00036. (PDF)

KORTET, R., SIRKKA, I., LAI, Y.-T., VAINIKKA, A. & KEKÄLÄINEN, J. 2015: Personality differences in two minnow populations that differ in their parasitism and predation risk. Frontiers in Ecology and Evolution, 3(9): 1-8. doi: 10.3389/fevo.2015.00009. (PDF)

JUSSILA, J., VREZEC, A., MAKKONEN, J., KORTET, R. & KOKKO, H. 2015: Invasive crayfish and their invasive diseases in Europe; focus on virulence evolution of the crayfish plague. In: Biological Invasions in Changing Ecosystems: Vectors, Ecological Impacts, Management and Predictions, Canning-Clode, J., ed. De Gruyter Open. (PDF)

LAI, Y.-T., TASKINEN, J., KEKÄLÄINEN, J. & KORTET, R. 2015: Infection ecology of Philometra ovata (Nematoda: Philometridae) in a wild European minnow (Phoxinus phoxinus) population in Finland. Parasitology, 142: 926-937. (PDF)

KORHONEN, E., VERA, C.P., PULLIAINEN, A.T., SIRONEN, T., AALTONEN, K., KORTET, R., HÄRKÖNEN, L., HÄRKÖNEN, S., PAAKKONEN, T., YLÖNEN, H. & VAPALAHTI, O. 2015: Molecular detection of Bartonella spp. in deer ked pupae, adult keds and moose blood in Finland. Epidemiology and Infection, 143(3):578-85. doi: 10.1017/S0950268814001411. (PDF)

KAUNISTO, S., HÄRKÖNEN, L., RANTALA, M.J. & KORTET, R. 2015: Early life temperature modifies adult encapsulation response in an invasive ectoparasite. Parasitology, 142: 1290-1296. (PDF)

KAUNISTO, S., YLÖNEN, H. & KORTET, R. 2015: Passive sinking into the snow as possible survival strategy during the off-host stage in an insect ectoparasite. Folia Parasitologica, 62: 038. (PDF)


KORTET, R., VAINIKKA, A., JANHUNEN, M., PIIRONEN, J. & HYVÄRINEN, P. 2014: Behavioral variation shows heritability in juvenile brown trout Salmo trutta. Behavioral Ecology and Sociobiology, 68: 927-934. (PDF)

HUUSKONEN, H., SUHONEN, A., GRUBER, C., VAINIKKA, A., HIRVONEN, H. & KORTET, R. 2014: Metabolic rate in the signal crayfish (Pasifastacus leniusculus) is temporally consistent and elevated at molting. Marine and Freshwater Behaviour and Physiology, 47: 205-209. (PDF)

GRUBER, C., VAINIKKA, A., HIRVONEN, H., RANTALA, M.J. & KORTET, R. 2014: Endogenous seasonal variation in the encapsulation response of the noble crayfish (Astacus astacus). Annales Zoologici Fennici, 51: 433-444. (PDF)

KEKÄLÄINEN, J., LAI, Y.-T., VAINIKKA, A., SIRKKA, I. & KORTET, R. 2014: Do brain parasites alter host personality? – Experimental study in minnows. Behavioral Ecology and Sociobiology, 68: 197-204. (PDF)

MAKKONEN, J., KOKKO, H., VAINIKKA, A., KORTET, R. & JUSSILA, J. 2014: Dose-dependent mortality of the noble crayfish (Astacus astacus) to different strains of the crayfish plague (Aphanomyces astaci). Journal of Invertebrate Pathology, 115: 86-91. (PDF)

AYDIN, H., KOKKO, H., MAKKONEN, J., KORTET, R., KUKKONEN, H, & JUSSILA, J. 2014: The signal crayfish is vulnerable to both the As and PsI-isolates of the crayfish plague. Knowledge and Management of Aquatic Ecosystems, 413: 3. DOI: 10.1051/kmae/2014004. (PDF)

GRUBER, C., KORTET, R., VAINIKKA, A., HYVÄRINEN, P., RANTALA, M.J., PIKKARAINEN, A., JUSSILA, J., MAKKONEN, J., KOKKO, H. & HIRVONEN, H. 2014: Variation in resistance to the invasive crayfish plague and immune defence in the native noble crayfish. Annales Zoologici Fennici, 51(4): 371-389. (PDF)

JUSSILA, J., MAKKONEN, J., VAINIKKA, A., KORTET, R. & KOKKO, H. 2014: Crayfish plague dilemma: how to be a courteous killer? Boreal Environment Research, 19: 235-244. (PDF)

KYNKÄÄNNIEMI, S-M., TIKKAKOSKI, M., KORTET, R., HÄRKÖNEN, L., KAITALA, A., PAAKKONEN, T., MUSTONEN, A-M., NIEMINEN, P., HÄRKÖNEN, S., YLÖNEN, H. & LAAKSONEN, S. 2014: Acute impacts of the deer ked (Lipoptena cervi) infestation on reindeer (Rangifer tarandus tarandus) behaviour. Parasitology Research, 113: 1489-1497. (PDF)


LAI, Y.-T., KEKÄLÄINEN, J. & KORTET, R. 2013: Male ornamentation in the European minnow (Phoxinus phoxinus) signals swimming performance. Ethology, 119: 1077-1085. (PDF)

NIEMELÄ, P.T., DINGEMANSE, N., ALIORAVAINEN, N., VAINIKKA, A. & KORTET, R. 2013: Personality pace-of-life hypothesis: testing genetic associations among personality and life-history. Behavioral Ecology, 24: 935-941. (PDF)

JUSSILA, J., KOKKO, H., KORTET, R. & MAKKONEN, J. 2013: Aphanomyces astaci PsI-genotype isolates from different Finnish signal crayfish stocks show variation in their virulence but still kill fast. Knowledge and Management of Aquatic Ecosystems, 411: 10. http://dx.doi.org/10.1051/kmae/2013077. (PDF)

NORDEIDE, J.T., KEKÄLÄINEN, J., JANHUNEN, M. & KORTET, R. 2013: Female ornaments revisited – are they correlated with offspring quality? Journal of Animal Ecology, 82: 26-38. (PDF)

TASKINEN, J., SUNDBERG, L-R., & KORTET, R. 2013: Sex and sexual ornamentation associated with survival of the cyprinid fish, Rutilus rutilus, under disease stress. Ecological Parasitology & Immunology, 2: ID 235652. doi:10.4303/epi/235652. (PDF)

NIEMELÄ, P.T., VAINIKKA, A., FORSMAN, J.T., LOUKOLA, O.J. & KORTET, R. 2013: How does variation in the environment and individual cognition explain the existence of consistent behavioural differences? Ecology & Evolution, 3(2): 457-464. (PDF)

PÖLKKI, M., KORTET, R., HEDRICK A. & RANTALA, M.J. 2013: Dominance is not always an honest signal of male quality but females may be able to detect the dishonesty. Biology Letters, 9: 20121002. doi:10.1098/rsbl.2012.1027. (PDF)

KEKÄLÄINEN, J., FIGENSCHOU, L., JANHUNEN, M., KORTET, R., PEUHKURI, N. & RUDOLFSEN, G. 2013: Hatchery selection may depress the number of motile sperm but intensify selection for their swimming velocity in the Arctic charr. Aquaculture International, 21: 405-411. (PDF)

FIGENSCHOU, L., FOLSTAD, I., RUDOLFSEN, G., HANSSEN, S.A., KORTET, R., SKAU, P.A., KILLIE, J.E., OSKAM, I. & STRAND, H. 2013: The relative effect of parasites and social status on sperm traits in Arctic charr. Behavioral Ecology, 24: 497-504. (PDF)


MAKKONEN, J., JUSSILA, J., KORTET, R., VAINIKKA, A. & KOKKO, H. 2012: Differing virulence of Aphanomyces astaci isolates and elevated resistance of noble crayfish Astacus astacus against crayfish plague. Diseases of Aquatic Organisms, 102: 129-136. (PDF)

LAI, Y.-T., KEKÄLÄINEN, J., TASKINEN, J. & KORTET, R. 2012: Non-invasive diagnosis for Philometra ovata (Nematoda) infection in the common minnow Phoxinus phoxinus. Parasitology Research, 111: 2411-2418. (PDF)

VAINIKKA, A., KOSKIMÄKI, J., NIEMELÄ, P.T. & KORTET, R. 2012: Composition of the Eurasian perch (Perca fluviatilis) catches in ice fishing: Does capture order predict body size? Fisheries Research, 115: 24-30. (PDF)

KORTET, R., NIEMELÄ, P., VAINIKKA, A. & LAAKSO, J. 2012: Females prefer bold males; an analysis of boldness, mate choice and bacterial resistance in the field cricket Gryllus integer. Ecological Parasitology and Immunology, 1: ID 235580. doi:10.4303/epi/235580. (PDF)

NIEMELÄ, P., VAINIKKA, A., HEDRICK, A.V. & KORTET, R. 2012: Integrating behavior with life history: boldness of the field cricket, Gryllus integer, during ontogeny. Functional Ecology, 26: 450-456. (PDF)

HEDRICK, A.V. & KORTET, R. 2012: Effects of body size on selectivity for mating cues in different sensory modalities. Biological Journal of the Linnean Society, 105: 160-168. (PDF)

CÓRDOBA-AGUILAR, A., RUIZ-SILVA, D., GONZÁLEZ-TOKMAN, A., CONTRERAS-GARDUÑO, J., PERETTI, A., MORENO-GARCÍA, M.A., RANTALA, M.J., KOSKIMÄKI, J., KORTET, R. & SUHONEN, J. 2012: No firm evidence of immunological costs of insect oviposition and copulation: a test with dragonflies (Zygoptera). Odonatologica, 41(1): 7-15 (PDF)

NIEMELÄ, P., VAINIKKA, A., LAHDENPERÄ, S. & KORTET, R. 2012: Nymphal density, behavioral development, and life history in a field cricket. Behavioral Ecology and Sociobiology, 66: 645-652. (PDF)

HEDRICK, A.V. & KORTET, R. 2012: Sex differences in the repeatability of boldness over metamorphosis. Behavioral Ecology and Sociobiology, 66: 407-412. (PDF)

KAUNISTO, S., KORTET, R., HÄRKÖNEN, S., KAITALA, A., LAAKSONEN, S. & YLÖNEN, H. 2012: Do small mammals prey upon an invasive ectoparasite of cervids? Canadian Journal of Zoology, 90: 1044-1050. (PDF)

KAUNISTO, S., KORTET, R., KOSKIMÄKI, J., HÄRKÖNEN, S., KAITALA, A., VÄLIMÄKI, P., LAAKSONEN, S., HÄRKÖNEN, L. & YLÖNEN, H. 2012: Avian predation on a parasitic fly of cervids during winter: can host-related cues increase the predation risk? Biological Journal of the Linnean Society, 106: 275-286. (PDF)


VAINIKKA, A., RANTALA, M.J., NIEMELÄ, P., HIRVONEN, H. & KORTET, R. 2011: Boldness as a consistent personality trait in the noble crayfish, Astacus astacus. Acta Ethologica, 14: 17-25. (PDF)

JUSSILA, J., MAKKONEN, J., VAINIKKA, A., KORTET, R. & KOKKO, H. 2011: Latent crayfish plague (Aphanomyces astaci) infection in a robust wild noble crayfish (Astacus astacus) population. Aquaculture, 321: 17-20. (PDF)

HUUSKONEN, H., KEKÄLÄINEN, J., PANDA, B., SHIKANO, T. & KORTET, R. 2011: Embryonic survival and larval predator-avoidance ability in mutually ornamented whitefish. Biological Journal of the Linnean Society, 103: 593-601. (PDF)

PAAKKONEN, T., MUSTONEN, A-M., KÄKELÄ, R., KILJANDER, T., KYNKÄÄNNIEMI, S-M., LAAKSONEN, S., SOLISMAA, M., KORTET, R., PUUKKA, K., SAARELA, S., HÄRKÖNEN, L., KAITALA, A., YLÖNEN, H. & NIEMINEN, P. 2011: Experimental infection of the deer ked (Lipoptena cervi) has no negative effects on the physiology of the captive reindeer (Rangifer tarandus tarandus). Veterinary Parasitology, 179: 180-188. (PDF)

JANHUNEN, M., KEKÄLÄINEN, J., KORTET, R., HYVÄRINEN, P. & PIIRONEN, J. 2011: No evidence for an indirect benefit from female mate preference in Arctic charr, Salvelinus alpinus, but female ornamentation decreases offspring viability. Biological Journal of the Linnean Society, 103: 602-611. (PDF)

VÄLIMÄKI, P., KAITALA, A., MADSLIEN, K., HÄRKÖNEN, L., VÁRKONYI, G., HEIKKILÄ, J., JAAKOLA, M., YLÖNEN, H., KORTET, R. & YTREHUS, B. 2011: Geographical variation in host use of a blood-feeding ectoparasitic fly: implications for population invasiveness. Oecologia, 166: 985-995. (PDF)


KORTET, R., HEDRICK, A.V. & VAINIKKA, A. 2010: Parasitism, predation and the evolution of animal personalities. Ecology Letters, 13: 1449-1458. (PDF)

KEKÄLÄINEN, J., HUUSKONEN, H., TUOMAALA, M. & KORTET, R. 2010: Both male and female sexual ornaments reflect offspring performance in a fish. Evolution, 64: 3149-3157. (PDF)

KEKÄLÄINEN, J., RUDOLFSEN, G., JANHUNEN, M., FIGENSCHOU, L., PEUHKURI, N., TAMPER, N. & KORTET, R. 2010: Genetic and potential non-genetic benefits increase offspring fitness of polyandrous females in non-resource based mating system. BMC Evolutionary Biology, 10: 20. (PDF)

VÄLIMÄKI, P., MADSLIEN, K., MALMSTEN, J., HÄRKÖNEN, L., HÄRKÖNEN, S., KAITALA, A., KORTET, R., LAAKSONEN, S., MEHL, R., REDFORD, L., YLÖNEN, H. & YTREHUS, B. 2010: Fennoscandian distribution of an important parasite of cervids, the deer ked (Lipoptena cervi), revisited. Parasitology Research, 107: 117-125. (PDF)

HÄRKÖNEN, L., HÄRKÖNEN, S., KAITALA, A., KAUNISTO, S., KORTET, R., LAAKSONEN, S. & YLÖNEN, H. 2010: Predicting range expansion of an ectoparasite – the effect of spring and summer temperatures on deer ked (Lipoptena cervi, Diptera: Hippoboscidae) performance along a latitudinal gradient. Ecography, 33: 906-912. (PDF)

LAAKSONEN, S., PUSENIUS, J., KUMPULA, J., VENÄLÄINEN, A., KORTET, R., OKSANEN, A. & HOBERG, E. 2010: Climate change promotes the emergence of serious disease outbreaks for Filarioid nematodes. EcoHealth, 7: 7-13. (PDF)

KYNKÄÄNNIEMI, S-M., KORTET, R., HÄRKÖNEN, L., KAITALA, A., PAAKKONEN, T., MUSTONEN, A-M., NIEMINEN, P., HÄRKÖNEN, S., YLÖNEN, H. & LAAKSONEN, S. 2010: Threat of an invasive parasitic fly, the deer ked (Lipoptena cervi) to the reindeer (Rangifer tarandus tarandus): experimental infection and treatment. Annales Zoologici Fennici, 47: 28-36. (PDF)

KORTET, R., HÄRKÖNEN, L., HOKKANEN, P., HÄRKÖNEN, S., KAITALA, A., KAUNISTO, S., LAAKSONEN, S., KEKÄLÄINEN, J. & YLÖNEN, H. 2010: Experiments on the ectoparasitic deer ked that often attacks humans; preferences for body parts, colour and temperature. Bulletin of Entomological Research, 100: 279-285. (PDF)


JANHUNEN, M., RUDOLFSEN, G., KEKÄLÄINEN, J., FIGENSCHOU, L., PEUHKURI, N. & KORTET, R. 2009: Spawning coloration and sperm quality in a large lake population of Arctic charr (Salmonidae: Salvelinus alpinus L.). Biological Journal of the Linnean Society, 98: 794-802. (PDF)

VOUTILAINEN, A., VALDEZ, H., KARVONEN, A., KORTET, R., KUUKKA, H., PEUHKURI, N., PIIRONEN, J. & TASKINEN, J. 2009: Infectivity of trematode eye flukes in farmed salmonid fish - effects of parasite and host origins. Aquaculture, 293: 108-112. (PDF)

VOUTILAINEN, A., VAN OOIK, T., PUURTINEN, M., KORTET, R. & TASKINEN, J. 2009: Relationship between prevalence of trematode parasite Diplostomum sp. and population density of its snail host Lymnaea stagnalis in lakes and ponds in Finland. Aquatic Ecology, 43: 351-357. (PDF)

KETOLA, T., KORTET, R. & KOTIAHO, J.S. 2009: Endurance in exercise is associated with courtship call rate in decorated crickets, Gryllodes sigillatus. Evolutionary Ecology Research, 11: 1131-1139. (PDF)

KAITALA, A., KORTET, R., HÄRKÖNEN, S., LAAKSONEN, S., HÄRKÖNEN, L., KAUNISTO, S. & YLÖNEN, H. 2009: Deer ked; an ectoparasite of moose, in Finland: a brief review of its biology and invasion. Alces, 45: 85-88.

LAAKSONEN, S., SOLISMAA, M., KORTET, R., KUUSELA, J. & OKSANEN, A. 2009: Vectors and transmission dynamics for Setaria tundra (Filarioidea; Onchocercidae), a parasite of reindeer in Finland. Parasites and Vectors, 2: 3. (PDF)

KORKEA-AHO, T.L., VAINIKKA, A., KORTET, R. & TASKINEN, J. 2009: Factors affecting between-lake variation in the occurrence of epidermal papillomatosis in roach, Rutilus rutilus (L.). Journal of Fish Diseases, 32: 263-270. (PDF)

VAINIKKA, A., TASKINEN, J., LÖYTYNOJA, K., JOKINEN, E.I. & KORTET, R. 2009: Measured immunocompetence relates to the proportion of dead parasites in a wild roach population. Functional Ecology, 23: 187-195. (PDF)

KAUNISTO, S., KORTET, R., HÄRKÖNEN, L., HÄRKÖNEN, S., YLÖNEN, H. & LAAKSONEN, S. 2009: New bedding site examination-based method to analyse deer ked (Lipoptena cervi) infection in cervids. Parasitology Research, 104: 919-925. (PDF)

LAAKSONEN, S., SOLISMAA, M., ORRO, T., KUUSELA, J, SAARI, S., KORTET, R., NIKANDER, S., OKSANEN, A. & SUKURA, A. 2009: Setaria Tundra microfilariae in reindeer and other cervids in Finland. Parasitology Research, 104: 257-265. (PDF)


KAITALA, A., KORTET, R., HÄRKÖNEN, S., LAAKSONEN, S., & YLÖNEN, H. 2008: Invasion of the moose ectoparasite Lipoptena cervi in Finland. In: Proceedings of the VIth International Moose Symposium, ed. Baskin L.M., Yakutsk, pp. 37-39. ISBN 978-5-471-00431-3.

KORTET, R. & VAINIKKA, A. 2008: Seasonality of innate immunity; evolutionary aspects and latest updates. In: New Research on Innate Immunity, eds. Durand M. & Morel C.V. Nova Science Publishers: NY, pp. 13-45. ISBN 978-1-60456-549-2. (PDF)


YLÖNEN, H., KORTET, R., MYNTTI, J. & VAINIKKA, A. 2007: Predator odour recognition and antipredatory response in fish: Does the prey know the predator diel rhythm? Acta Oecologica, 31: 1-7. (PDF)

KORTET, R. & HEDRICK, A. 2007: A behavioural syndrome in the field cricket Gryllus integer: intrasexual aggression is correlated with activity in a novel environment. Biological Journal of the Linnean Society, 91: 475-482. (PDF)

KETOLA, T., KORTET, R. & KOTIAHO, J.S. 2007: Testing theories of sexual selection in decorated crickets (Gryllodes sigillatus). Evolutionary Ecology Research, 9: 869-885. (PDF)

KORTET, R., RANTALA, M.J. & HEDRICK, A. 2007: Boldness in anti-predation behavior and immune defence in field crickets. Evolutionary Ecology Research, 9: 185-197. (PDF)


AHTIAINEN, J.J., ALATALO, R., KORTET, R. & RANTALA, M.J. 2006: Immune function, dominance and mating success in drumming male wolf spiders Hygrolycosa rubrofasciata. Behavioral Ecology and Sociobiology, 60: 826-832. (PDF)

HEDRICK, A.V. & KORTET, R. 2006: Hiding behaviour in two cricket populations that differ in predation pressure. Animal Behaviour, 72: 1111-1118. (PDF)

VÄÄNÄNEN, S., KORTET, R. & RANTALA, M.J. 2006: Dominance and immune function in the F1 generation of wild caught field crickets. Behaviour, 143: 701-712. (PDF)

RANTALA, M.J., ERIKSSON, P.C.J., VAINIKKA, A. & KORTET, R. 2006: Male steroid hormones and female preference for male body odor. Evolution and Human Behavior, 27: 259-269. (PDF)


VAINIKKA, A., KORTET, R., PAUKKU, S., RANTALA, M.J. & PIRHONEN, J. 2005: What do male tench, Tinca tinca (L.), advertise with morphological ornaments? Acta Ethologica, 8: 70-78. (PDF)

VAINIKKA, A., JOKINEN, E.I., KORTET, R., PAUKKU, S., PIRHONEN, J., RANTALA, M.J. & TASKINEN, J. 2005: Effects of testosterone and ß-glucan on immune functions in tench. Journal of Fish Biology, 66: 348-361. (PDF)

VAINIKKA, A., JOKELAINEN, T., KORTET, R. & YLÖNEN, H. 2005: Predation risk allocation or direct vigilance response in the predator interaction between perch (Perca fluviatilis L.) and pike (Esox lucius L.)? Ecology of Freshwater Fish, 14: 225-232. (PDF)

KORTET, R. & HEDRICK, A. 2005: The scent of dominance: female field crickets use odour to predict the outcome of male competition. Behavioral Ecology and Sociobiology, 59: 77-83. (PDF)

LUONG, L.T., HEDRICK, A.V. & KORTET, R. 2005: Prevalence and intensity of Cephalobium microbivorum (Nematoda: Diplogasterida) infection in three species of Gryllus field crickets. Parasitology Research, 97: 336-339. (PDF)

AHTIAINEN, J.J, RANTALA, M.J., ALATALO, R.V. & KORTET, R. 2005: A trade-off between investment in sexual signalling and immune function in a natural population of the drumming wolf spider Hygrolycosa rubrofasciata. Journal of Evolutionary Biology, 18: 985-991. (PDF)


KORTET, R., VAINIKKA, A., RANTALA, M.J. & TASKINEN, J. 2004: Sperm quality, secondary sexual characters and parasitism in roach (Rutilus rutilus). Biological Journal of the Linnean Society, 81: 111-117. (PDF)

KORTET, R. & TASKINEN, J. 2004: Parasitism, condition and number of front head breeding tubercles in roach (Rutilus rutilus L.). Ecology of Freshwater Fish, 13: 119-124. (PDF)

KORTET, R. & HEDRICK, A. 2004: Detection of the spider predator, Hololena nedra by naïve juvenile field crickets (Gryllus integer) using indirect cues. Behaviour, 141: 1189-1196. (PDF)

VAINIKKA, A., JOKINEN, E.I., KORTET, R. & TASKINEN, J. 2004: Gender- and season-dependent relationships between testosterone, oestradiol and immune functions in wild roach. Journal of Fish Biology, 64: 227-240. (PDF)

RANTALA, M.J & KORTET, R. 2004: Male dominance and immunocompetence in the field cricket (Gryllus bimaculatus). Behavioral Ecology, 15: 187-191. (PDF)

AHTIAINEN, J.J., ALATALO, R.V., KORTET, R. & RANTALA, M.J. 2004: Sexual advertisement and immune function in an arachnid species (Lycosideae). Behavioral Ecology, 15: 602-606. (PDF)

KORTET, R., VAINIKKA, A., RANTALA, M.J., MYNTTI, J. & TASKINEN, J. 2004: In vitro embryo survival and early viability of larvae in relation to male sexual ornaments and parasite resistance in roach, Rutilus rutilus L. Journal of Evolutionary Biology, 17: 1337-1344. (PDF)

KORTET, R., TASKINEN, J., VAINIKKA, A. & YLÖNEN, H. 2004: Breeding tubercles, papillomatosis and dominance behavior of male roach (Rutilus rutilus) during the breeding period. Ethology, 110: 591-601. (PDF)

VAINIKKA, A., KORTET, R. & TASKINEN, J. 2004: Epizootic cutaneous papillomatosis, cortisol and ornamentation during and after breeding in roach Rutilus rutilus. Diseases of Aquatic Organisms, 60: 189-195. (PDF)


KORTET, R., TASKINEN, J., SINISALO, T. & JOKINEN, I. 2003: Breeding -related seasonal changes in immunocompetence, health state and condition of the cyprinid fish, Rutilus rutilus, L. Biological Journal of the Linnean Society, 78: 117-127. (PDF)

KORTET, R., VAINIKKA, A., RANTALA, M.J., JOKINEN, I. & TASKINEN, J. 2003: Sexual ornamentation, androgens and papillomatosis in male roach (Rutilus rutilus). Evolutionary Ecology Research, 5: 411-419. (PDF)

RANTALA, M.J. & KORTET, R. 2003: Courtship song and immune function in the field cricket Gryllus bimaculatus. Biological Journal of the Linnean Society, 79: 503-510. (PDF)

RANTALA, M.J., KORTET, R., KOTIAHO, J.S., VAINIKKA, A. & SUHONEN, J. 2003: Condition dependence of pheromones and immune function in the grain beetle Tenebrio molitor. Functional Ecology, 17: 534-540. (PDF)

KORTET, R., VAINIKKA, A. & TASKINEN, J. 2003: Effect of epidermal papillomatosis on survival of the freshwater fish Rutilus rutilus. Diseases of Aquatic Organisms, 57: 163-165. (PDF)

RANTALA, M.J., VAINIKKA, A. & KORTET, R. 2003: The role of juvenile hormone in immune function and pheromone production trade-offs: a test of the immunocompetence handicap principle. Proceedings of the Royal Society of London B, 270: 2257-2261. (PDF)


KORTET, R., TASKINEN, J. & VAINIKKA, A. 2002: Epizootic cutaneous papillomatosis in roach Rutilus rutilus: sex and size dependence, seasonal occurrence and between population differences. Diseases of Aquatic Organisms, 52(3): 185-190. (PDF)

TASKINEN, J. & KORTET, R. 2002: Dead and alive parasites: Sexual ornaments signal resistance in the male fish, Rutilus rutilus. Evolutionary Ecology Research, 4: 919-929. (PDF)

RANTALA, M.J, JOKINEN, I., KORTET, R., VAINIKKA, A. & SUHONEN, J. 2002: Do pheromones reveal male immunocompetence? Proceedings of the Royal Society of London B, 269: 1681-1685. (PDF)

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Last modified: April 28, 2021.