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Publications (send me an e-mail for reprints)

Behringer, D., Karvonen, A., Bojko, J. (2018). Parasite avoidance behaviours in aquatic environments. Philosophical Transactions of the Royal Society B, in press.


Karvonen, A., Lindström, K. (2018). Spatiotemporal and gender-specific parasitism in two species of gobiid fish. Ecology and Evolution, in press.


Klemme, I., Karvonen, A. (2018). Shoaling with infected conspecifics does not improve resistance to trematode infection. Ethology 124, 170-176.


Karvonen, A., Faltýnková, A., Mah Choo, J., Valtonen, E.T. (2017). Infection, specificity and host manipulation of Australapatemon sp. (Trematoda, Strigeidae) in two sympatric species of leeches (Hirudinea). Parasitology 144, 1346-1355.

Klemme, I., Karvonen, A. (2017). Vertebrate defence against parasites: interactions between avoidance, resistance and tolerance. Ecology and Evolution 7, 561-571.


Klemme, I., Karvonen, A. (2016). Learned parasite avoidance is driven by host personality and resistance to infection in a fish-trematode interaction. Proceedings of the Royal Society B 283, 20161148.

Klemme, I., Kortet, R., Karvonen, A. (2016). Parasite infection in a central sensory organ of fish does not affect host personality. Behavioral Ecology 27, 1533-1537.

Klemme, I., Louhi, K.-R., Karvonen, A. (2016). Host infection history modifies co-infection success of multiple parasite genotypes. Journal of Animal Ecology 85, 591-597.

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.

Louhi, K.-R., Sundberg, L.-R., Jokela, J., Karvonen, A. (2015). Interactions among bacterial strains and fluke genotypes shape virulence of co-infection. Proceedings of the Royal Society B 282, 20152097.  

Seppälä, O., Louhi, K.-R., Karvonen, A., Rellstab, C., Jokela, J. (2015). Relative reproductive success of co-infecting parasite genotypes under intensified within-host competition. Infection, Genetics and Evolution 36, 450-455.

Karvonen, A., Lucek, K., Marques, D.A., Seehausen, O. (2015). Divergent macroparasite infections in parapatric Swiss lake-stream pairs of threespine stickleback (Gasterosteus aculeatus). PLoS ONE 10(6): e0130579.

Seppälä, O., Karvonen, A., Kuosa, M., Haataja, M., Jokela, J. (2013). Are sick individuals weak competitors? Competitive ability of snails parasitized by a gigantism-inducing trematode. PLoS ONE 8(10): e79366.

Rellstab, C., Karvonen, A., Louhi, K.-R., Jokela, J. (2013). Genotype-specific vs. cross-reactive host immunity against a macroparasite. PLoS ONE 8(10): e78427.

Louhi, K.-R., Karvonen, A., Rellstab, C., Jokela, J. (2013). Genotypic and phenotypic variation in transmission traits of a complex life cycle parasite. Ecology and Evolution 3, 2116-2127.


Karvonen, A., Kristjánsson, B.K., Skúlason, S., Lanki, M., Rellstab, C., Jokela, J. (2013). Water temperature, not fish morph, determines parasite infections of sympatric Icelandic threespine sticklebacks (Gasterosteus aculeatus). Ecology and Evolution 3, 1507-1517.

Louhi, K.-R., Karvonen, A., Rellstab, C., Louhi, R., Jokela, J. (2013). Prevalence of infection as a predictor of multiple genotype infection frequency in parasites with multiple-host life cycle. Journal of Animal Ecology 82, 191-200.

Karvonen, A., Lundsgaard-Hansen, B., Jokela, J., Seehausen, O. (2013). Differentiation in parasitism among ecotypes of whitefish segregating along depth gradients. Oikos 122, 122-128.

Seppälä, O., Karvonen, A., Valtonen E.T. (2012). Behavioral mechanisms underlying ‘specific’ host manipulation by a trophically transmitted parasite. Evolutionary Ecology Research 14, 73-81.

Seppälä, O., Karvonen, A., Rellstab, C., Louhi, K.-R., Jokela, J. (2012). Reciprocal interaction matrix reveals complex genetic and dose-dependent specificity among coinfecting parasites. American Naturalist 180, 306-315.

Karvonen, A., Seehausen, O. (2012). The role of parasitism in adaptive radiations – when might parasites promote and when might they constrain ecological speciation? International Journal of Ecology, Vol. 2012, Article ID 208169.

Karvonen, A., Rellstab, C., Louhi, K.-R., Jokela, J. (2012). Synchronous attack is advantageous - mixed genotype infections lead to higher infection success in trematode parasites. Proceedings of the Royal Society B 279, 171-176.

Karvonen, A. (2012). Diplostomum spathaceum and Related Species. In: Fish Parasites: Pathobiology and Protection (Eds. Woo, P.T.K. and Buchmann, K.). pp. 260-269. CAB International.

Rellstab, C., Louhi, K.-R., Karvonen, A., Jokela, J. (2011). Analysis of trematode parasite communities in fish eye lenses by pyrosequencing of naturally pooled DNA. Infection, Genetics and Evolution 11, 1276-1286.

Faltýnková, A., Karvonen, A., Valtonen, E.T. (2011). Establishment and interspecific associations in two species of Ichthyocotylurus (Trematoda) parasites in perch (Perca fluviatilis). Parasites & Vectors 4, 85.


Seppälä, O., Karvonen, A., Haataja, M., Kuosa, M., Jokela, J. (2011). Food makes you a target: disentangling genetic, physiological and behavioral effects determining susceptibility to infection. Evolution 65, 1367-1375.

Seppälä, O., Karvonen, A., Valtonen, E.T. (2011). Eye fluke-induced cataracts in natural fish populations: is there potential for host manipulation? Parasitology 138, 209-214.

Karvonen, A., Rintamäki, P., Jokela, J., Valtonen, E.T. (2010). Increasing water temperature and disease risks in aquatic systems: climate change increases the risk of some, but not all, diseases. International Journal for Parasitology 40, 1483-1488.

Louhi, K.-R., Karvonen, A., Rellstab, C., Jokela, J. (2010). Is the population genetic structure of complex life cycle parasites determined by the geographic range of the most motile host? Infection, Genetics and Evolution 10, 1271-1277.

Karvonen, A., Halonen, H., Seppälä, O. (2010). Priming of host resistance to protect cultured rainbow trout against eye flukes and parasite-induced cataracts. Journal of Fish Biology 76, 1508-1515.

Faltýnková, A., Karvonen, A., Valtonen, E.T. (2009). Being successful in the world of narrow opportunities: transmission patterns of the trematode Ichthyocotylurus pileatus. Parasitology 136, 1375-1382.

Aldhoun, J.A., Faltýnková, A, Karvonen, A., Horák, P. (2009). Schistosomes in the North: a unique finding from a prosobranch snail using molecular tools. Parasitology International 58, 314-317.

Karvonen, A., Valtonen, E.T. (2009). Between-population similarity in intestinal parasite community structure of pike (Esox lucius) - effects of distance and historical connections. Journal of Parasitology 95, 505-511.

Karvonen, A., Seppälä, O., Valtonen, E.T. (2009). Host immunisation shapes interspecific associations in trematode parasites. Journal of Animal Ecology 78, 945-952.

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.


Seppälä, O., Karvonen, A., Valtonen, E.T., Jokela, J. (2009). Interactions among co-infecting parasite species: a mechanism maintaining genetic variation in parasites? Proceedings of the Royal Society B 276, 691-697.

Faltýnková, A., Valtonen, E.T., Karvonen, A. (2008). Spatial and temporal structure of the trematode component community in Valvata macrostoma (Gastropoda, Prosobranchia). Parasitology 135, 1691-1699.

Karvonen, A., Seppälä, O. (2008). Eye fluke infection and lens size reduction in fish: a quantitative analysis. Diseases of Aquatic Organisms 80, 21-26.

Karvonen, A., Seppälä, O. (2008). Effect of eye fluke infection on the growth of whitefish (Coregonus lavaretus) - an experimental approach. Aquaculture 279, 6-10.

Seppälä, O., Liljeroos, K., Karvonen, A., Jokela, J. (2008). Host condition as a constraint for parasite reproduction. Oikos 117, 749-753.

Seppälä, O., Karvonen, A., Valtonen E.T. (2008). Shoaling behaviour of fish under parasitism and predation risk. Animal Behaviour 75, 145-150.

Santos, M.J., Karvonen, A., Pedro, J.C., Faltýnková, A., Seppälä, O., Valtonen, E.T. (2007). Qualitative and quantitative behavioral traits in a community of furcocercarial trematodes – tools for species separation? Journal of Parasitology 93, 1319-1323.

Seppälä, O., Karvonen, A., Valtonen E.T. (2007). Phenotypic variation in infectivity of Diplostomum spathaceum cercariae within a population. Journal of Parasitology 93, 1244-1246.

Markkula, S.E., Karvonen, A., Salo, H., Valtonen, E.T., Jokinen, E.I. (2007). Ultraviolet B irradiation affects resistance of rainbow trout (Oncorhynchus mykiss) against bacterium Yersinia ruckeri and trematode Diplostomum spathaceum. Photochemistry and Photobiology 83, 1263-1269.

Karvonen, A., Bagge, A.M., Valtonen, E.T. (2007). Interspecific and intraspecific interactions in the monogenean communities of fish: a question of study scale? Parasitology 134, 1237-1242.


Seppälä, O., Karvonen, A., Valtonen, E.T. (2006). Host manipulation by parasites and risk of non-host predation: is manipulation costly in an eye fluke-fish interaction? Evolutionary Ecology Research 8, 871-879.

Karvonen, A., Terho, P., Seppälä, O., Jokela, J., Valtonen, E.T. (2006). Ecological divergence of closely related Diplostomum (Trematoda) parasites. Parasitology 133, 229-235.

Hakalahti, T., Karvonen, A., Valtonen, E.T. (2006). Climate warming and disease risks in temperate regions – Argulus coregoni and Diplostomum spathaceum as case studies. Journal of Helminthology 80, 93-98.

Karvonen, A., Savolainen, M., Seppälä, O., Valtonen, E.T. (2006). Dynamics of Diplostomum spathaceum infection snail hosts at a fish farm. Parasitology Research 99, 341-345.

Seppälä, O., Karvonen, A., Valtonen, E.T. (2006). Susceptibility of eye fluke infected fish to predation by bird hosts. Parasitology 132, 575-579.

Karvonen, A., Cheng, G.-H., Seppälä, O., Valtonen, E.T. (2006). Intestinal distribution and fecundity of two species of Diplostomum parasites in definitive hosts. Parasitology 132, 357-362.

Karvonen, A., Cheng, G.-H., Valtonen, E.T. (2005). Within-lake dynamics in the similarity of parasite assemblages of perch (Perca fluviatilis). Parasitology 131, 817-823.

Bagge, A.M., Sasal, P., Valtonen, E.T., Karvonen, A. (2005). Infracommunity level aggregation in the monogenean communities of crucian carp (Carassius carassius). Parasitology 131, 367-372.

Karvonen, A., Bagge, A.M., Valtonen, E.T. (2005). Parasite assemblages of crucian carp (Carassius carassius) – is depauperate composition explained by lack of parasite exchange, extreme environmental conditions or host unsuitability? Parasitology 131, 273-278.

Seppälä, O., Karvonen, A., Valtonen, E.T. (2005). Impaired crypsis of fish infected with a trophically transmitted parasite. Animal Behaviour 70, 895-900.


Seppälä, O., Karvonen, A., Valtonen, E.T. (2005). Manipulation of fish host by eye flukes in relation to cataract formation and parasite infectivity. Animal Behaviour 70, 889-894.

Karvonen, A., Paukku, S., Seppälä, O., Valtonen, E.T. (2005). Resistance against eye flukes: naïve versus previously infected fish. Parasitology Research 95, 55-59.

Karvonen, A., Seppälä, O., Valtonen, E.T. (2004). Eye fluke-induced cataract formation in fish: quantitative analysis using an opthalmological microscope. Parasitology 129, 473-478.

Rintamäki-Kinnunen, P., Karvonen, A, Anttila, P., Valtonen, E.T. (2004). Diplostomum spathaceum metacercarial infection and colour change in salmonid fish. Parasitology Research 93, 577-581.

Seppälä, O., Karvonen, A., Valtonen, E.T. (2004). Parasite induced change in host behaviour and susceptibility to predation in an eye fluke - fish interaction. Animal Behaviour 68, 257-263.

Karvonen, A., Valtonen, E.T. (2004). Helminth assemblages of whitefish (Coregonus lavaretus) in interconnected lakes: similarity as a function of species specific parasites and geographical separation. Journal of Parasitology, 90 (3), 471-476.

Karvonen, A., Seppälä, O., Valtonen, E.T. (2004). Parasite resistance and avoidance behaviour in preventing eye fluke infections in fish. Parasitology 129, 159-164.

Karvonen, A., Kirsi, S., Hudson, P.J., Valtonen, E.T. (2004). Patterns of cercarial production from Diplostomum spathaceum: terminal investment or bet hedging? Parasitology 129, 87-92.

Karvonen, A., Hudson, P.J., Seppälä, O., Valtonen, E.T. (2004). Transmission dynamics of a trematode parasite: exposure, acquired resistance and parasite aggregation. Parasitology Research 92, 183-188.

Karvonen A., Paukku, S., Valtonen, E.T., Hudson, P.J. (2003). Transmission, infectivity and survival of Diplostomum spathaceum cercariae. Parasitology 127, 217-224.


Other publications:

Karvonen, A., Hakalahti, T., Seppälä, O., Valtonen, E.T. (2005). Sustainable production of healthy fish – tackling parasitic threats with knowledge on their ecology. In: Jalkanen, A. & Nygren, P. (eds.), Sustainable use of renewable natural resources - from principles to practices. University of Helsinki Department of Forest Ecology Publications 34, 77-88.

Pulkkinen, K., Karvonen, A, Valtonen E.T. (2004). Lapamato – harvinainen muttei hävinnyt. Duodecim 120, 1912. (in Finnish)

Karvonen, A. (2004). Transmission of Diplostomum spathaceum between intermediate hosts. PhD-Thesis. Jyväskylä Studies in Biological and Environmental Science 139. University of Jyväskylä. 40 p.


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