Recent articles, open access:

Cervin, L., T. Harkonen and K.C. Harding. 2020. Multiple stressors and data deficient populations; a comparative life-history approach sheds new light on the extinction risk of the highly vulnerable Baltic harbour porpoises (Phocoena phocoena). Environment International 144, 106076. https://doi.org/10.1016/j.envint.2020.106076

Silva, W.T.A.F., E. Bottagisio, T. Härkönen, A. Galatius, M.T. Olsen and K.C. Harding. 2021. Risk for overexploiting a seemingly stable seal population: influence of multiple stressors and hunting.  Ecosphere. https://doi.org/10.1002/ecs2.3343

Silva, W.T.A.F., K.C. Harding, G.M. Marques, C. Sonne, R. Dietz, T. Härkönen and J.-P. Desforges. 2020. Life cycle bioenergetics of the gray seal (Halichoerus grypus) in the Baltic Sea: population response to environmental stress Environmental International. Oct 7;145:106145. https://doi:10.1016/j.envint.2020.106145.

Full list:

1. Harding K.C. and Härkönen T.J. (1995). Estimating mean age at sexual maturity in the crabeater seal (Lobodon carcinophagus). Can. J. Fish. Aquat. Sci. 52: 2347-2352.

2. Härkönen T., Harding K.C. and Lunneryd S.G. (1999). Age- and sex-specific behaviour in harbour seals (Phoca vitulina) leads to biased estimates of vital population parameters.   J. Appl. Ecol. 36:825-841.

3. Harding, KC. and Härkönen, T. (1999). Development in the Baltic grey seal (Halichoerus grypus) and ringed seal (Phoca hispida) populations during the 20th century. Ambio 28:619-627.

4. Sih, A., Ziemba, R. and Harding, K.C. (2000). New insights on how temporal variation in predation risk shapes prey behavior. TREE 15: 3-4.

5. Härkönen T. and Harding K.C. (2001). Spatial structure of harbour seal populations and the implications thereof. Canadian Journal of Zoology. 79: 2115-2127.

6. Härkönen, T., Harding, K.C. and Heide-Jørgenssen M.-P. (2002) Rates of increase in age structured populations, a lesson from the European harbour seal. Canadian Journal of Zoology. 80: 1498-1510.

7. Harding K.C. and J.M. McNamara. (2002). A unifying framework for metapopulation dynamics. American Naturalist 160:173-185.

8. Harding, K.C., Härkönen, T. and H. Caswell. (2002). The 2002 European seal plague: Epidemiology and population consequences. Ecology Letters.  5: 727-732.

9. Harding K.C., Härkönen, T. and J. Pineda. (2003). Estimating quasi-extinction risk of European harbour seals.: a reply to Lonergan and Harwood. Ecology Letters 6: 894-897.

10. McNamara J.M. and K.C. Harding. (2004). Measurement error and estimates of population extinction risk. Ecology Letters. 7: 16-20.

11. Harding, K.C. , M. Fujiwara, T. Härkönen and Y. Axberg. (2005). Mass dependent energetics and survival in harbour seal pups. Functional Ecology. 19: 129-135.

12. Härkönen, T, K.C. Harding, S. Goodman, and K. Johannesson (2005) Colonization history of the Baltic harbor seals: integrating archaeological, behavioral, and genetic data, T. Marine Mammal Science 21: 695-716.

13. Harding, K.C., J Hansen and S. Goodman. (2005) Acquired immunity and stochasticity in epidemic intervals impede the evolution of host disease resistance. American Naturalist. 166: 722-730.

14. T. Härkönen, R. Dietz, P. Reijnders, J. Teilmann, P. Thompson, K.C. Harding,  A. Hall, S. Brasseur, U. Siebert, S. Goodman, P. Jepson, T. Dau Rasmussen. (2006) The 1988 and 2002 phocine distemper virus epidemics in European harbour seals. Diseases of Aquatic Organisms 68(2): 115-130.

15. Hansén B.J.L. and K.C. Harding. (2006) On the potential impact of harbour seal predation on the cod population in the eastern North Sea. J Sea Res.

16. Houston, A.I., Stephens, P.A., Boyd, I.L., Harding, K.C. & McNamara, J.M. (2007) Capital or income breeding? A theoretical model of female reproductive strategies.  Behavioral Ecology 18, 241-250.

17. Leidenberger, S., K.C. Harding, T. Härkönen. (2007) Phocid seals, seal lice and heartworms: a terrestrial host-parasite system conveyed to the marine environment. Dis Aquat Org. 77:235-253.

18. Harding, K. C., Härkönen, T., Helander, B. and Karlsson, O. (2007). Status of Baltic grey seals: Population assessment and risk analysis. Nammco Scientific Publications. s. 33-56. No. 81986

19. Härkönen T., K.C. Harding, T. Dau Rasmussen, J. Teilmann, R. Dietz. (2007) Age- and Sex-specific Mortality Patterns in an Emerging Wildlife Epidemic: the Phocine Distemper in European Harbour Seals. PLoS ONE 2(9): e887.

20. Stephens P.A., C. Carbone, I.L. Boyd, J.M. McNamara, K.C. Harding and A.I. Houston.. (2008) The scaling of diving time budgets: insights from an optimality approach. American Naturalist 171: 305-314.

21. Harkonen T., Bäcklin, B.M., Barrett, T., Bergman, A., Corteyn, M., Dietz, R., Harding, K.C., Malmsten, J., Roos, A. and J. Teilmann. (2008). Mass mortality in harbour seals and harbour porpoises caused by an unknown pathogen. Vet. Rec. 162: 555-556.

22. Harkonen, T., M. Jüssi, I. Jüssi, M. Verevkin, E. Helle, R. Sagitov, K.C. Harding. (2008) Seasonal activity budget of adult Baltic ringed seals (Phoca hispida botnica). PLoS ONE 3(4): e2006.doi:10.1371/journal.pone.0002006.

23. Roy, M., Harding K.C. and R.D. Holt. 2008. Generalizing Levins metapopulation model in explicit space: models of intermediate complexity. J. Theoretical Biology. 255 152-161.

24. Jagers and Harding . 2009. Viability of small populations haunted by random catastrophes. Math Pop Studies. 16 (3) 177-198.

25. Harkonen, T. and K.C. Harding. 2010. Predicting recurrent PDV epizootics in European harbour seals (Phoca vitulina). NAMMCO Scientific Publications: 8   Pages: 275-284.

26. Svensson C.J., A. Eriksson, T. Härkönen and K.C. Harding (2011). Detecting Density Dependence in Recovering Seal Populations Ambio 40:52-59.

27. M. P. Heide-Jørgensen, K.L. Laidre,  R.G. Hansen, M.L. Burt, D.L. Borchers, J. Hansén, K.C. Harding, M. Rasmussen, R. Dietz, and J. Teilmann. (2012) Rate of increase and current abundance of humpback whales in West Greenland. Journal of Cetacean Research and Management 12: 1 - 14

28. Leidenberger S., K.C. Harding and P.R. Jonsson. (2012) Ecology and distribution of the isopods Idotea spp. in the Baltic Sea: key species in a changing environment. J. Crustacean Biol. 32: 359-381.

29. KC Harding, Begon M, B. A. Eriksson and Wennberg. (2012) Increased migration in host-pathogen metapopulations can cause host extinction. J. Theor. Biol. 298: 1-7.

30. Sundqvist, L., Harkonen, T., Svensson, C.J. and K.C. Harding. 2012. Linking Climate Trends to Population Dynamics in the Baltic Ringed Seal: Impacts of Historical and Future Winter Temperatures. Ambio 41: 865-871.

31. Harkonen, T., K.C. Harding, S. Wilson, M. Baimukanov, L. Dmitrieva, C. J. Svensson, S. J. Goodman. 2012. Collapse of a Marine Mammal Species Driven by Human Impacts. Plos ONE 7:1-9 

32. Garnier, R., S. Gandon, K.C. Harding, T. Boulinier 2014. Length of intervals between epidemics: evaluating the influence of maternal transfer of immunity. Ecology and Evolution, 4: 568-575. DOI: 10.1002/ece3.955

33. Stephens, P.A., A. I. Houston, K.C. Harding, L. Boyd and J.M McNamara. (2014) Capital and income breeding in pinnipeds: the role of food supply. Ecology. http://www.esajournals.org/doi/abs/10.1890/13-1434.1

34. Singh, A., K.C. Harding, HRV., Reddy, A. Godhe. (2014) An assessment of Dinophysis blooms in the coastal Arabian Sea. Harmful Algae. 34: 29-35.

35. Arcalis-Planas, A.,  S. Sveegaard, O. Karlsson, K.C. Harding, A. Wåhlin, T. Harkonen and J. Teilmann (2015). Limited use of sea ice by the Ross seal (Ommatophoca rossii), in Amundsen Sea, Antarctica, using telemetry and remote sensing data. Polar Biology: 38: 445-461.

36. Garde, E. S. H. Hansen, S. Ditlevsen, K. B. Tvermosegaard, J. Hansen, K. C. Harding and M.P. Heide-Jørgensen. 2015. Life history parameters of narwhals (Monodon monoceros) from Greenland. Journal of Mammalogy 96(4).

37. Lehnert, K., Weirup L., Harding K.C., Härkönen T., Karlsson O. and J. Teilmann. 2017. Antarctic seals: Molecular biomarkers as indicators for diet, pollutant exposure and health effects. Science of the Total Environment 599-600: 1693-1704.

38. Kauhala K., Bäcklin B.M., J. Raitaniemi and K.C. Harding. 2017. The effect of prey quality and ice conditions on the nutritional status of Baltic grey seals of different age groups. Mammal Research 62:351-362.

39. Pertoldi C., L.F. Jensen, A.K.O. Alstrup, O.L. Munk, T.B. Pedersen, C. Sonne, R. Dietz, T. Daugaard-Petersen, H.E. Kortegaard, M. Tange-Olsen, K.C. Harding and T. Hammer Jensen. (2018). Prevalence of skull pathologies in European harbour seals (Phoca vitulina) during 1981-2014. Mammal Research, 63: 55-63.

40. Saremi S., M. Isaksson and K.C. Harding. 2018. Bio accumulation of radioactive Caesium in marine mammals in the Baltic Sea – reconstruction of a historical time series. Sci. Tot. Env. 631: 7-12. https://doi.org/10.1016/j.scitotenv.2018.02.282

41. Harding, K.C., Salmon, M. Teilmann, J., Dietz D. and T. Harkonen. (2018). Population wide decline in somatic growth in harbour seals – early signs of density dependence. Frontiers in Ecology and Evolution. https://doi.org/10.3389/fevo.2018.00059

42. Iben Stokholm, Tero Härkonen, Karin C. Harding, Ursula Siebert, Kristina Lehnert, Rune Dietz, Jonas Teilmann, Anders Galatius, Linnea Worsøe Havmøller, Emma Carroll, Ailsa Hall, and Morten Tange Olsen 2019. Phylogenomic insights to the origin and spread of Phocine Distemper Virus in European harbour seals in 1988 and 2002. Diseases of Aquatic Organisms. DOI: 10.3354/dao03328

43. E. K. Brault, P.L. Koch, D.P. Costa, M.D. McCarthy, L.A. Hückstädt, K. Goetz, K.W. McMahon, M.E. Goebel, O. Karlsson, J. Teilmann, T. Härkönen and K.C. Harding. 2019. Trophic position and foraging ecology of Ross, Weddell, and crabeater seals revealed by compound-specific isotope analysis. Marine Ecology Progress Series DOI: 10.3354/meps12856

44. Näpflin, K. , Becks, L., Bensch, S., Ellis, V.A., Hafer, N., Harding, K. C., Lindén, S., O’Connor, E., Olsen, M.T., Roved, J., Sackton, TB, Shultz, AJ, Venkatakrishnan, V., Videvall, E., Westerdahl, H., Winternitz, JC and Edwards, SV. (2019) Genomics of co-evolutionary interactions between hosts and pathogens: challenges and opportunities across scales. PeerJ. 10.7717/peerj.8013

45.  Sonne, C., U. Siebert, K. Gonnsen, J.-P. Desforges, I. Eulaers, S. Persson, A. Roos, B.-M. Bäcklin, K. Kauhala, M. Olsen, K.C. Harding, G. Treu, A. Galatius, E. Andersen-Ranberg, S. Gross, J. Lakemeyer, K. Lehnert, S.S. Lam, P. Wan-Xi and R. Dietz. (2020) Health effects from contaminant exposure in Baltic Sea birds and marine mammals: A review.  Environment International.

46. Cervin, L., T. Harkonen and K.C. Harding. 2020. Multiple stressors and data deficient populations; a comparative life-history approach sheds new light on the extinction risk of the highly vulnerable Baltic harbour porpoises (Phocoena phocoena). Environment International 144, 106076. https://doi.org/10.1016/j.envint.2020.106076

47. Silva, W.T.A.F., E. Bottagisio, T. Härkönen, A. Galatius, M.T. Olsen and K.C. Harding. (2021). Risk for overexploiting a seemingly stable seal population: influence of multiple stressors and hunting.  Ecosphere. https://doi.org/10.1002/ecs2.3343

48. Silva, W.T.A.F., K.C. Harding, G.M. Marques, C. Sonne, R. Dietz, T. Härkönen and J.-P. Desforges. (2020). Life cycle bioenergetics of the gray seal (Halichoerus grypus) in the Baltic Sea: population response to environmental stress Environmental International. Oct 7;145:106145. doi: 10.1016/j.envint.2020.106145.

49. Dietz, R, Sonne, C, Jenssen, BM, Das, K, de Wit, CA, Harding, K.C., Siebert, U, Olsen MT. 2021. The Baltic Sea: an ecosystem with multiple stressors. Environmental International. 147(6409):106324. DOI: 10.1016/j.envint.2020.106324

*50. Jonasson, J.,  T. Harkonen, S. Edwards, L. Sundqvist and K.C. Harding. 2022. A unifying framework for estimating generation time in age-structured populations: implications for phylogenetics and conservation biology. American Naturalist. https://doi.org/10.1086/719667

*51. Lohmann, R., M. Serageldin, E. Brault, R. Dickhut, K.C. Harding, T. Harkonen, O. Karlsson, K. Lehnert and J. Teilmann. 2021. Bioaccumulation of PCBs, OCPs and PBDEs in marine mammals from West Antarctica. Frontiers in Marine Science https://doi.org/10.3389/fmars.2021.768715

52. Liu X., Schjøtt SR, Granquist SM, Rosing-Asvid A, Dietz R, Teilmann J, Galatius A, Cammen K, O´Corry-Crowe G, Harding K, Härkönen T, Hall A, Carroll EL, Kobayashi Y, Hammill M, Stenson G, Frie AK, Lydersen C, Kovacs KM, Andersen LW, Hofmann J, Goodman SJ, Vieira FG, Heller R, Moltke I, Olsen MT (2022) Origin and expansion of the world’s most widespread pinniped: range-wide population genomics of the harbour seal (Phoca vitulina). Molecular Ecology, https://doi.org/10.1111/mec.16365

*53. Infantes, E., D. Carroll, W.T.A.F., Silva, T. Harkonen, S.V. Edwards and K. C. Harding (2022). An automated work-flow for pinniped surveys: A new tool for monitoring population dynamics. Front. Ecol. Evol., Vol:10.  DOI 10.3389/fevo.2022.905309

54. Mauritsson, K., Desforges, JP. & Harding, K.C. (2022) Maternal Transfer and Long-Term Population Effects of PCBs in Baltic Grey Seals Using a New Toxicokinetic–Toxicodynamic Population Model. Arch Environ Contam Toxicol. https://doi.org/10.1007/s00244-022-00962-3

55. Johannesson, K., Leder, E. H., André, C., Dupont, S., Eriksson, S. P., Harding, K., Havenhand, J. N., Jahnke, M., Jonsson, P. R., Kvarnemo, C., Pavia, H., Rafajlović, M., Rödström, E. M., Thorndyke, M., & Blomberg, A. (2023). Ten years of marine evolutionary biology - challenges and achievements of a multidisciplinary research initiative. Evolutionary Applications. https://doi.org/10.1111/eva.13389

56. Carroll, D, M. Ahola, A. Carlsson, M. Sköld and K.C. Harding. (2024). 120-years of ecological monitoring data shows that the risk of overhunting is increased by environmental degradation for an isolated marine mammal population: the Baltic grey seal.  Journal of Animal Ecology 93, 525-539. https://doi.org/10.1111/1365-2656.14065

57. Carroll D., Infantes E., Pagan E. V., and K.C. Harding (2024). Approaching a population level assessment of body size in pinnipeds using drones, an early warning of environmental degradation. Remote Sensing in Ecology and Conservation. https://doi.org/10.1002/rse2.413

58. Carroll, D.,  M.P. Ahola, A.M. Carlsson, A. Galatius, K.T. Nilssen, T. Härkönen, K.C. Harding (2025). Declining harbour seal abundance in the Kattegat-Skagerrak and Danish Straits. PlosOne https://doi.org/10.1371/journal.pone.0326933

59. Carroll D., Harvey-Carroll J., Härkönen T. and K.C. Harding (2025). Harbor Seals (Phoca vitulina) Sighted 34 Years After Their Inclusion in a Capture-Mark-Recapture Study. Marine Mammal Science. https://onlinelibrary-wiley-com.ezproxy.ub.gu.se/doi/10.1111/mms.70073?af=R

60. Kappa F., T. Härkönen, C. Freitas, D. Carroll, K.C. Harding (2025) Shifting harbour seal (Phoca vitulina) diet may reflect ecosystem changes in Skagerrak. Marine Mammal Science https://doi.org/10.1111/mms.70069

61. Ulrika Ervander, Willian Talvane Arestides Ferreira Da Silva, Karin Hårding, Micael Jonsson. 2025. Status and future prospect of deregistered woodland key habitats in Northwestern Sweden, Plos One - 2025 

BOOK CHAPTERS

1. Harding K.C., J.M. McNamara and R.D. Holt (2006) Understanding invasions in patchy habitats through metapopulation theory. Pages 371-403 In: Cadotte MW, McMahon, Fukami T, eds. Conceptual ecology and invasions biology:reciprocal approaches to nature. Springer.

2. Harding, K.C. Härkönen, T., B. Helander and O. Karlsson (2007). Status of Baltic grey seals: Population assessment and risk analysis. NAMMCO Scientific Publications, 6: 33-56.