Professor Lotta Kvarnemo

Contact details

E-mail: lotta.kvarnemo(at)

My background

  • University of Gothenburg, Sweden; Zoology: Fish Ecology, Professor 2005-onwards (1 Sep 2012-28 Feb 2014: leave of absence)
  • Åbo Akademi University, Finland; Biological Sciences: Environmental Biology, Senior lecturer 2012-2014
  • Stockholm University, Sweden; Zoology, Assistant Professor (Forskarassistent) 2000-2005 (5 yrs)
  • Stockholm University, Sweden; Zoology, Associate Professor (Docent) 2000
  • University of Jyväskylä, Finland; Biology, Post Doc 1999-2000 (6 months)
  • Stockholm University, Sweden; Zoology, Repatriation 1998-1999 (NFR 1 yr)
  • University of Western Australia, Australia; Zoology, Post Doc 1996-1998 (NFR 2 yrs)
  • Uppsala University, Sweden; Zoology, PhD 1996
  • Åbo Akademi University, Finland; Biology, MSc 1990

Research Interests

The main focus of my research is sexual selection, mainly in animals with male care, investigated from a behavioural and evolutionary ecological perspective. I am particularly interested in questions related to the evolution of parental care , mating systems, sperm competiton, operational sex ratio, filial cannibalism, etc.

Below I describe my main interests in a little more detail, and what is done together with whom. Abbreviations used: ÅAU=Åbo Akademi University; GU=University of Gothenburg; SU=Stockholm University; UWA=University of Wester Australia; UU=Uppsala University.

Male care

When the male provides care to the offspring, this behaviour is tightly linked to natural selection, via offspring survival, but also sexual selection, e.g. via competition for mates, mate choice, potential reproductive rate, paternity. Male care is more common than female care or biparental care among fishes. Thus, fish are good for studying these questions.

Together with former PhD students/post docs Ola Svensson and Maria Norevik Andén (previously Lissåker), SU/GU, and Master students Therese Stenlund, Karin Olsson and Johannes Björk, GU, I have explored questions about sneaking, male care and filial cannibalism in the marine fish sand goby, Pomatoschistus minutus. I am also collaborating with Colette St Mary, Univ of Florida, Elisabet Forsgren, NINA, Norway, Kai Lindström, ÅAU, and Erica Leder, Univ of Turku, Finland, Ola Svensson, and Jon Havenhand, GU, and other researchers on various comparative projects on sand, two-spotted and painted gobies.

Together with Adam G. Jones, Texas A&M Univ, and Glenn I. Moore, Museum of Western Australia, I have studied sexual selection and the genetic mating system of the Western Australian seahorse, Hippocampus subelongatus. With Lucy Woodall, NHM, UK, Sara Lourie, McGill, Canada, and Erica Leder , University of Turku, Finland, now added to this collaborative network, our questions have broadened further, to also include taxonomy, population genetics and genetic sex markers.

Ongoing projects also include post-mating sexual selection in the pipefish Syngnathus typhle, done together with Ingrid Ahnesjö, UU, Adam Jones, Kenyon Mobley, Charlyn Partridge, Texas A&M Univ, and my current & former PhD students Gry Sagebakken and Inês Braga Gonçalves, GU. Master students Josefin Sundin and Emma Martinsson, GU, have also been involved in this work.

Operational Sex Ratio (OSR)

OSR is the ratio of males to females ready to mate in a population at a given time. It is an important predictor of mating competition, and thus also sexual selection.

During my own PhD, I studied the influence of temperature, food and nest site availability on OSR, potential reproductive rates and mating competition. Using sand gobies, I investigated dynamics in animal sex-roles and how variation in sexual selection can be modulated by the environment. This is a theme I have recently returned to, especially in the context of environmental change, within a Linnnaeus project, called CeMEB (centre for marine evolutionary biology), funded by the Swedish research councils VR & Formas.

As a post-doc in Leigh W. Simmons' group at UWA, I focused on how OSR interacts with variation in mate quality in affecting mate choice, in two species of Australian bush crickets. I have also addressed theoretical issues related to OSR and mating competition, together with Leigh Simmons, Ingrid Ahnesjö and Sami Merilaita, and tested some of the assumptions together with Andreas Nyman and Ola Svensson.

Sperm Competition

When multiple males mate with the same female there is a risk of sperm competition over the fertilisation of the female's eggs.

Together with Leigh Simmons, I have investigated how males decrease their ejaculate expenditure with a perceived increase in sperm competition intensity, in the Australian bushcricket Kawanaphila nartee. In collaboration with Kai Lindström, ÅAU, and Adam Jones, Texas A&M University, USA, I have investigated sexual selection in relation to sneaker success in the sand goby.

I have also focused on sperm competition in relation to the evolution of male care. In a phylogenetic study on fish with Malin Ah-King and Birgitta Tullberg, SU, we have shown that high paternity is not an necessary prerequisite for the evolution of male care, and in a theoretical model I have explored a previously unappreciated benefit of providing care, namely that aspects of male care behaviours may help prevent or reduce sperm competition. Together with Malin Ah-King, Helena Elofsson, and others, we have also investigated the mode of fertilization in Nerophis ophidion, a pipefish with male brood care.

Research group

Previous PhD students:

  • Gry Sagebakken (2006-2012): Paternal care and brood reduction in a pipefish.Animal Ecology, Dept of Biological and Environmental Sciences, GU
  • Ines Goncalves (2006-2011): Egg size evolution in relation to oxygen conditions and paternal care in pipefishes and gobies. Animal Ecology, Dept of Zoology, GU
  • Maria Prager (2005-2010): Phylogeny and ecology of plumage signal diversity in African widowbirds and bishops. Animal Ecology, Dept of Zoology, GU.
  • Maria Lissåker (2002-2007): Parental care and filial cannibalism in the sand goby. Animal Ecology, Dept of Zoology, SU.
  • Helena Strömberg (2003-2004): Internal bioerosion of coral reefs. Animal Ecology, Dept of Zoology, SU (started 1999).
  • Ola Svensson (2000-2004): Sperm competition in gobiid fishes. Animal Ecology, Dept of Zoology, SU.

Host of visiting researchers, post docs, etc.

  • Topi Lehtonen: Post doc: 2013-2015 (2 yrs)
  • Gry Sagebakken: Project researcher: 2013 (4 months)
  • Miguel Correirz: Project researcher: 2013 (5 months)
  • Ola Svensson: Research assistant position: 2010-2015 (4 yrs); Post doc: 2007-2010 (2 yrs)
  • Maria Norevik Andrén (formerly Lissåker): Post doc 2012 (6 months)
  • Lucy Woodall: Post doc 2011 (2 months)
  • Isabelle Côté: Guest professor 2011 (3 months)
  • Takashi Takegaki: Visting researcher 2009 (7 months)
  • Colette St. Mary: Visting researcher 2006 and 2007 (3 months)

Previous exam students:

  • Emma Martinsson (2013): Behaviour of brooding male pipefish in different oxygen conditions (GU, 15 hp)
  • Sara Sundström (2012): 'Utveckling av pedagogiska skyltar på Borås djurpark' (Application work in Animal Ecology, GU, 15 hp).
  • Johannes Björk (2011): 'Male preference for nest sites in relation to their own body size: an experimental investigation'. (GU, 60 hp).
  • Karin Olsson (2008): 'An ecological niche? A study on nest material preference in sand gobies (Pomatoschistus minutus)' (GU, 15 hp).
  • Josefin Sundin (2008): 'Does male condition affect female choice in the sex-role reversed pipefish Syngnathus typhle?' (GU, 30 hp).
  • Karin Olsson (2007): 'Relative cost of courtship behaviour in nest-building sand gobies.' (GU, 30 hp).
  • Therese Stenlund (2006): 'Effects on parental care in sand goby, Pomatoschistus minutus, caused by an egg predator and different oxygen levels' (GU, 30 hp).
  • Anna Singer (2003): 'Does nest site availability influence sneaking rate in the sand goby Pomatoschistus minutus' (SU, 30 p).
  • Andreas Nyman (2002): 'Intensitet i parningskonkurrens hos sandstubb i relation till äggtäckningsgrad i boet' (SU, 20 p).
  • Maria Lissåker (2001): 'Effects of a low oxygen environment on parental effort and filial cannibalism in the male sand goby, Pomatoschistus minutus' (SU, 20 p).
  • Glenn I Moore (1997-1999): 'Mating patterns and sexual selection in the Western Australian Seahorse Hippocampus subelongatus' (Master of Science, UWA).
  • Ola Svensson (1995): 'Parental behaviour depending on paternity assurance and food availability in the common goby Pomatoschistus microps' (UU, 20 p).
  • Anna Karlsson (1993): 'Why do female sand gobies (Pomatoschistus minutus) prefer males with eggs in their nests?' (UU, 20p).

List of publications

  • Kvarnemo, C (2014). Sexual selection. Oxford Bibliographies Online: Evolutionary Biology. doi: 10.1093/9780199941728-0029

  • Sundin, J., Sagebakken, G. & Kvarnemo, C. (2013). Female mate choice is not affected by mate condition in a fish with male care. Acta Ethologica 16: 189-194. doi:10.1007/s10211-013-0149-1

  • Kvarnemo, C & Simmons, LW. (2013). Polyandry as a mediator of sexual selectionbefore and after mating. Philosophical Transctions of the Royal Society B 368: 20120042.

  • Takegaki, T, Svensson, O & Kvarnemo, C (2012). Socially induced tactic change in 2 types of sand goby sneaker males. Behavioral Ecology 23: 742-750. doi:10.1093/beheco/ars022

  • Björk, JR & C Kvarnemo (2012). Mechanisms behind size-assortative nest choice by sand goby males in absence of intrasexual competition. Animal Behaviour 83: 55-62. doi:10.1016/j.anbehav.2011.09.033

  • Kvarnemo, C (2011). Stressed singers and symmetric lovers. Behavioral Ecology 22: 11-12. doi: 10.1093/beheco/arq082

  • Kvarnemo, C, Mobley, KB, Partridge, C, Jones, AG & I Ahnesjö (2011). Evidence of paternal nutrient provisioning to embryos in the pipefish Syngnathus typhle. Journal of Fish Biology 78: 1725-1737. doi:10.1111/j.1095-8649.2011.02989.x

  • Mobley, KB, Kvarnemo, C, Ahnesjö, I, Partridge, C, Berglund, A & AG Jones (2011). The effect of maternal body size on embryo survivorship in the broods of pregnant male pipefish. Behavioral Ecology and Sociobiology. 65: 1169-1177. doi:10.1007/s00265-010-1129-3

  • Braga Goncalves, I, Ahnesjö, I & C Kvarnemo (2011). The relationship between female body size and egg size in pipefishes. Journal of Fish Biology 78: 1847-1854. doi:10.1111/j.1095-8649.2011.02984.x

  • ärvi-Laturi, M, Lindström, K, Kvarnemo, C & O Svensson (2011). Alternative mating tactics under high risk of egg predation. Journal of Zoology 2011: 1-7. doi:10.1111/j.1469-7998.2011.00788.x

  • Sagebakken, G, Ahnesjö I & I Braga Gonçalves & C Kvarnemo (2011). Multiply-mated males show higher embryo survival in a paternally caring fish. Behavioral Ecology 22: 625-629. doi:10.1093/beheco/arr023

  • Kvarnemo, C (2010). Parental care. In: Evolutionary Behavioral Ecology (Eds D F Westneat & C W Fox). Oxford University Press, New York.

  • Kvarnemo, C, Svensson, O & W Manson (2010). Investment in testes, sperm duct glands and lipid reserves differs between male morphs but not between early and late breeding season in Pomatoschistus minutus. Journal of Fish Biology 76: 1609-1625. doi:10.1111/j.1095-8649.2010.02587.x

  • Braga Goncalves, I, Mobley, KB, Ahnesjö, I, Sagebakken, G, Jones AG & C Kvarnemo (2010). Reproductive compensation in broad-nosed pipefish females. Proceedings of the Royal Society B 277: 1581-1589. doi: 10.1098/rspb.2009.2290

  • Sagebakken, G, Ahnesjö, I, Mobley, KB, Braga Goncalves I & C Kvarnemo (2010). Brooding fathers, not siblings, take up nutrients from embryos. Proceedings of the Royal Society B 277: 971-977. doi: 10.1098/rspb.2009.1767

  • Kvarnemo, C (2009). Sex hos djur. In: Djuren och människan - Om den moderna biologin och dess världsbild. (Ed LJ Erkell). Studentlitteratur, Lund.

  • Kvarnemo, C, Lindenfors, P, Ah-King, M & I Ahnesjö (2009). Gender perspectives on the development of sexual selection theory. ISBE newsletter 21: 11-12.

  • Olsson, K H, Kvarnemo, C & O Svensson (2009). Relative costs of courtship behaviours in nest-building sand gobies. Animal Behaviour 77: 541-546. doi: 10.1016/j.anbehav.2008.10.021

  • Partridge, C, Ahnesjö, I, Kvarnemo, C, Mobley, KB, Berglund, A & AG Jones (2009). The effect of perceived female parasite load on postcopulatory male choice in a sex-role-reversed pipefish. Behavioral Ecology and Sociobiology 63: 345-454. doi: 10.1007/s00265-008-0668-3

  • Ahnesjö, I, Forsgren, E & C Kvarnemo (2008).+ Variation in sexual selection in fishes. In: Fish Behaviour (Eds Magnhagen, C, Braithwaite, VA, Forsgren, E & BG Kapoor). Enfield, NH: Science Publishers Inc.

  • Kvarnemo, C, Moore, GI & AG Jones (2007). Sexually selected females in the monogamous Western Australian seahorse Hippocampus subelongatus. Proceedings of the Royal Society London Series B 274: 521-525. doi:10.1098/rspb.2006.3753

  • Svensson, O & C Kvarnemo (2007). Parasitic spawning in sand gobies: an experimental assessment of nest-opening size, sneaker male cues, paternity and filial cannibalism. Behavioral Ecology 18: 410-419. doi:10.1093/beheco/arl098

  • Kvarnemo C (2006). Evolution and maintenance of male care: Is increased paternity a neglected benefit of care? Behavioral Ecology 17: 144-148. doi:10.1093/beheco/ari097

  • Kvarnemo, C & S Merilaita (2006)+ Mating distribution and its temporal dynamics affect operational sex ratio: a simulation study. Biological Journal of the Linnean Society 89: 551-559.

  • Lissåker, M & C Kvarnemo (2006). Ventilation or nest defence - parental trade-offs in a fish with male care. Behavioral Ecology and Sociobiology 60: 864-873. doi:10.1007/s00265-006-0230-0

  • Simmons, L W & C Kvarnemo (2006). Costs of breeding and their effects on the direction of sexual selection. Proceedings of the Royal Society London Series B 273: 465-470. doi:10.1098/rspb.2005.3309

  • Nyman, A, Kvarnemo, C and O Svensson (2006). The capacity for additional matings does not affect male mating competition in the sand goby. Animal Behaviour 71: 865-870. doi:10.1016/j.anbehav.2005.06.020

  • Singer, A, Kvarnemo C, Lindström, K & O Svensson (2006). Genetic mating patterns studied in pools with manipulated nest site availability and two populations of Pomatoschistus minutus. Journal of Evolutionary Biology 19: 1641-1650. doi:10.1111/j.1420-9101.2006.01114.x

  • Wedell, N, Kvarnemo, C, Lessells, CM & T Tregenza (2006). Sexual conflict and life histories. Review paper for Animal Behaviour 71: 999-1011. doi:10.1016/j.anbehav.2005.06.023

  • Ah-King, M, Elofsson, H, Kvarnemo, C, Rosenqvist, G & A Berglund (2006). Why is there no sperm competition in a pipefish with externally brooding males? Insights from sperm activation and morphology. Journal of Fish Biology 68: 958-962. doi:10.1111/j.1095-8649.2006.00971.x

  • Strömberg, H & Kvarnemo, C (2005). The effects of territorial damselfish on cryptic bioeroding organisms on dead Acropora formosa. Journal of Experimental Marine Biology and Ecology 327: 91-102. doi:10.1016/j.jembe.2005.06.012

  • Svensson, O & C Kvarnemo (2005). The importance of sperm competition risk and nest appearance for male behaviour and female choice in Pomatoschistus minutus. Behavioral Ecology 16: 1042-1048. doi:10.1093/beheco/ari085

  • Ah-King, M, Kvarnemo, C & BS Tullberg (2005). The influence of territoriality and mating system on the evolution of male care: a phylogenetic study on fish. Journal of Evolutionary Biology 18: 371-382. doi:10.1111/j.1420-9101.2004.00823.x

  • Kvarnemo, C & LW Simmons (2004). Testes investment and spawning mode in pipefishes and seahorses (Syngnathidae). Biological Journal of the Linnean Society 83: 369-376.

  • Svensson, O, Nyman, A & C Kvarnemo (2004). Costly courtship or dishonest display? Intensely displaying sand goby (Pomatoschistus minutus) males have lower lipid content. Journal of Fish Biology 64: 1425-1429.

  • Kvarnemo, C, Simmons, LW, Ah-King, M & H Elofsson (2003). The myth of the sperm cloud - or why is testes size not affected by spawning mode in seahorses and pipefishes. Abstract of the Fisheries Society of the British Isles Symposium, Norwich, UK. Journal of Fish Biology, Suppl A, 63: 235.

  • Svensson, O & C Kvarnemo (2003). Sexually selected nest building - Pomatoschistus minutus males build smaller nest-openings in the presence of sneaker males. Journal of Evolutionary Biology 16: 896-902.

  • Lissåker, M, Kvarnemo, C & O Svensson (2003). Effects of a low oxygen environment on parental effort and filial cannibalism in the male sand goby, Pomatoschistus minutus. Behavioral Ecology 14: 374-381.

  • Jones, AG, Moore, GI, Kvarnemo, C, Walker, D & JC Avise (2003). Sympatric speciation as a consequence of male pregnancy in seahorses Proceedings of the National Academy of Sciences. 100: 6598-6603. doi:10.1073/pnas.1131969100

  • Kvarnemo, C & I Ahnesjö (2002).+ Operational sex ratios and mating competition. In: Sex Ratios: Concepts and Research Methods, ed ICW Hardy. Cambridge University Press, Cambridge pp 366-382.

  • Svensson, O & C Kvarnemo (2001). Nest quality and sperm competition in sand gobies, Pomatoschistus minutus. Abstract of the XXVII International Ethological Conference, Tübingen, Germany Ethology, Suppl 36, 271.

  • Ahnesjö, I, C Kvarnemo & S Merilaita (2001).+ Using potential reproductive rates to predict mating competition. Behavioral Ecology 12: 397-401.

  • Jones, AG, Walker, D, Kvarnemo, C, Lindström, K & JC Avise (2001). How cuckoldry can decrease the opportunity for sexual selection: data and theory from a genetic parentage analysis of the sand goby, Pomatoschistus minutus. Proceedings of the National Academy of Sciences 98:9151-9156. doi:10.1073/pnas.171310198

  • ones, AG, Walker, D, Lindström, K, Kvarnemo, C & JC Avise (2001). Surprising similarity of sneaking rates and genetic mating patterns in two populations of sand goby experiencing disparate sexual selection regimes. Molecular Ecology 10: 461-469.

  • Kvarnemo, C & E Forsgren (2000). The influence of potential reproductive rates and variation in mate quality on male and female choosiness in the sand goby. Behavioral Ecology and Sociobiology 48: 378-384. doi: 10.1007/s002650000246

  • Kvarnemo, C, Moore, GI, Jones, AG, Nelson, WS & JC Avise (2000). Monogamous pair bonds and mate switching in the Western Australian seahorse Hippocampus subelongatus. Journal of Evolutionary Biology 13: 882-888.

  • Kvarnemo, C & LW Simmons (1999). Variance in female quality, OSR and male mate choice in a bushcricket. Behavioral Ecology and Sociobiology 45: 245-252.

  • Kvarnemo, C (1998). Temperature modulates competitive behaviour: why do sand goby males fight more in warmer water. Ethology Ecology & Evolution 10: 105-114.

  • Kvarnemo, C (1998). Why male cannibalism won't cause a female biased OSR - a comment on Smith & Wootton's paper. Reviews in Fish Biology and Fisheries 8: 93-98.

  • Kvarnemo, C & LW Simmons (1998). Male potential reproductive rate influences mate choice in a bushcricket. Animal Behaviour 55: 1499-1506.

  • Kvarnemo, C, Svensson, O & E Forsgren (1998). Parental behaviour in relation to food availability in the common goby. Animal Behaviour 56: 1285-1290.

  • Jones, AG, Kvarnemo, C, Moore, GI, Simmons, LW & JC Avise (1998). Microsatellite evidence for monogamy and sex-biased recombination in the Western Australian seahorse Hippocampus angustus. Molecular Ecology 7: 1497-1506.

  • Svensson, O, Magnhagen, C, Forsgren E & C Kvarnemo (1998). Parental behaviour in relation to the occurrence of sneaking in the common goby. Animal Behaviour 56: 175-179.

  • Kvarnemo C (1997). Food affects the potential reproductive rates of sand goby females, but not of males. Behavioral Ecology 8: 605-611.

  • Simmons, LW & C Kvarnemo (1997). Ejaculate expenditure by male bushcrickets decreases with sperm competition intensity. Proceedings of the Royal Society London Series B 264: 1203-1208.

  • Kvarnemo C (1996). Temperature affects operational sex ratio and intensity of male-male competition - an experimental study on sand gobies. Behavioral Ecology 7: 208-212.

  • Kvarnemo, C & I Ahnesjö (1996).+ The dynamics of operational sex ratios and competition for mates. Trends in Ecology & Evolution 11: 404-408.

  • Forsgren, E, Kvarnemo, C & K Lindström (1996).+ Differences in sexual selection pressures between two populations of sand gobies: the importance of nest availability. Evolution 50: 646-654.

  • Forsgren, E, Karlsson, A & C Kvarnemo (1996). Female sand gobies choose males whose nests contain eggs, and thereby gain direct benefits. Behavioral Ecology and Sociobiology 39: 91-96.

  • Kvarnemo, C (1996). Hos sandstubben tar pappan hand om barnen. Flora & Fauna, Nr 5-6: 43-47 (reviewed popular scientific publication)

  • Kvarnemo, C (1995). Size-assortative nest choice in the absence of competition in males of the sand goby, Pomatoschistus minutus. Environmental Biology of Fishes 43: 233-239.

  • Kvarnemo, C, Forsgren, E & C Magnhagen (1995). Effects of sex ratio on intra- and intersexual behaviour in sand gobies. Animal Behaviour 50: 1455-1461.

  • Kvarnemo, C (1994). Temperature differentially affects male and female reproductive rates in the sand goby: consequences for operational sex ratio. Proceedings of the Royal Society London Series B 256: 151-156. doi:10.1098/rspb.1994.0063

  • Magnhagen, C & L Kvarnemo (1989). Big is better: the importance of size for reproductive success in male Pomatoschistus minutus (Pisces, Gobiidae). Journal of Fish Biology 35: 755-763.

According to ISI Web of Science these publications have been cited 1762 times (Jan 2014). Mean: 32.0 citations/publication; 70.5 citations/year; H-index: 24. + means input was shared equally among the authors.