Prof John Harwood


room:  (The Observatory)
The Observatory
Buchanan Gardens
University of St Andrews
St Andrews
KY16 9LZ

General Information

Population and Conservation Biology : Effects of individual variation and spatial structure on the population dynamics, genetics and epidemiology of vertebrates, particularly marine mammals. Interactions between anthropogenic activities and ecosystem function.

Research groups:


NERC Sea Mammal Research Unit


Recent Publications

Donovan, CR, Harris, CM, Milazzo, L, Harwood, J, Marshall, L & Williams, R 2017, 'A simulation approach to assessing environmental risk of sound exposure to marine mammals' Ecology and Evolution, vol 7, no. 7, pp. 2101-2111. DOI: 10.1002/ece3.2699
Tyack, PL, Bailey, H, Crocker, D, Estes, JE, Francis, CD, Harwood, J, Schwacke, L, Thomas, LJ & Wartzok, D 2016, Approaches to Understanding the Cumulative Effects of Stressors on Marine Mammals. National Academies Press, Washington DC. National Academies Press, Washington DC. DOI: 10.17226/23479
Fleishman, E, Costa, DP, Harwood, J, Kraus, S, Moretti, D, New, LF, Schick, RS, Schwarz, LK, Simmons, SE, Thomas, L & Wells, RS 2016, 'Monitoring population-level responses of marine mammals to human activities' Marine Mammal Science, vol 32, no. 3, pp. 1004-1021. DOI: 10.1111/mms.12310
Pirotta, E, Harwood, J, Thompson, P, New, L, Cheney, B, Arso Civil, M, Hammond, PS, Donovan, CR & Lusseau, D 2015, 'Predicting the effects of human developments on individual dolphins to understand potential long-term population consequences' Proceedings of the Royal Society B: Biological Sciences, vol 282, no. 1818, 20152109. DOI: 10.1098/rspb.2015.2109
King, SL, Schick, RS, Donovan, CR, Booth, CG, Burgman, M, Thomas, L & Harwood, J 2015, 'An interim framework for assessing the population consequences of disturbance' Methods in Ecology and Evolution, vol 6, no. 10, pp. 1150-1158. DOI: 10.1111/2041-210X.12411
Wiff, R, Barrientos, MA, Milessi, AC, Quiroz, JC & Harwood, J 2015, 'Modelling production per unit of food consumed in fish populations' Journal of Theoretical Biology, vol 365, pp. 67-75. DOI: 10.1016/j.jtbi.2014.10.004
Moretti, D, Thomas, L, Marques, T, Harwood, J & Dilley, A 2014, 'Errutum: A risk function for behavioral disruption of blainville's beaked whales mesoplodon densirostris from mid-frequency active sonar (PLoS ONE (2014) 9:12 (e85064) doi:10.1371/journal.pone.0085064)' PLoS One, vol 9, no. 12. DOI: 10.1371/journal.pone.0116555
Pirotta, E, New, L, Harwood, J & Lusseau, D 2014, 'Activities, motivations and disturbance: an agent-based model of bottlenose dolphin behavioral dynamics and interactions with tourism in Doubtful Sound, New Zealand' Ecological Modelling, vol 282, pp. 44-58. DOI: 10.1016/j.ecolmodel.2014.03.009
New, LF, Clark, J, Costa, D, Fleishman, E, Hindell, M, Klanjšček, T, Lusseau, D, Kraus, S, McMahon, C, Robinson, P, Schick, RS, Schwartz, L, Simmons, S, Thomas, L, Tyack, PL & Harwood, J 2014, 'Using short-term measures of behaviour to estimate long-term fitness of southern elephant seals.' Marine Ecology Progress Series, vol 496, pp. 99-108. DOI: 10.3354/meps10547
Moretti, D, Thomas, L, Marques, TA, Harwood, J, Dilley, A, Neales, B, Shaffer, J, Mccarthy, E, New, LF, Jarvis, S & Morrissey, R 2014, 'A risk function for behavioral disruption of Blainville’s beaked whales (Mesoplodon densirostris) from mid-frequency active sonar' PLoS One, vol 9, no. 1, e85064. DOI: 10.1371/journal.pone.0085064
Smout, SC, Rindorf, A, Hammond, PS, Harwood, J & Matthiopoulos, J 2014, 'Modelling prey consumption and switching by UK grey seals' ICES Journal of Marine Science, vol 71, no. 1, pp. 81-89. DOI: 10.1093/icesjms/fst109
Harris, DV, Matias, L, Thomas, L, Harwood, J & Geissler, W 2013, 'Applying distance sampling to fin whale calls recorded by single seismic instruments in the northeast Atlantic.' Journal of the Acoustical Society of America, vol 134, no. 5, 3522. DOI: 10.1121/1.4821207
Russell, DJ, McConnell, BJ, Thompson, D, Duck, CD, Morris, C, Harwood, J & Matthiopoulos, J 2013, 'Uncovering the links between foraging and breeding regions in a highly mobile mammal' Journal of Applied Ecology, vol 50, no. 2, pp. 499-509. DOI: 10.1111/1365-2664.12048
Schick, RS, New, L, Thomas, L, Costa, D, Hindell, M, McMahon, C, Robinson, P, Simmons, S, Thums, M, Harwood, J & Clark, J 2013, 'Estimating resource acquisition and at-sea body condition of a marine predator' Journal of Animal Ecology, vol 82, no. 6, pp. 1300-1315. DOI: 10.1111/1365-2656.12102
New, LF, Harwood, J, Thomas, L, Donovan, CR, Clark, J, Hastie, GD, Thompson, P, Cheney, B, Scott Hayward, LAS & Lusseau, D 2013, 'Modelling the biological significance of behavioural change in coastal bottlenose dolphins in response to disturbance' Functional Ecology, vol 27, pp. 314-322. DOI: 10.1111/1365-2435.12052
Cammen, K, Hoffman, JI, Knapp, LA, Harwood, J & Amos, W 2011, 'Geographic variation of the major histocompatibility complex in Eastern Atlantic grey seals (Halichoerus grypus).' Molecular Ecology, vol 20, no. 4, pp. 740-752. DOI: DOI: 10.1111/j.1365-294X.2010.04975.x
Smout, SC, Asseburg, C, Matthiopoulos, J, Fernández, C, Redpath, S, Thirgood, S & Harwood, J 2010, 'The functional response of a generalist predator' PLoS One, vol 5, no. 5, e10761. DOI: 10.1371/journal.pone.0010761
Lonergan, M, Hall, A, Thompson, H, Thompson, PM, Pomeroy, P & Harwood, J 2010, 'Comparison of the 1988 and 2002 phocine distemper epizootics in British harbour seal Phoca vitulina populations' Diseases of Aquatic Organisms, vol 88, no. 3, pp. 183-188. DOI: 10.3354/dao02153
Reynolds, TJ, King, R, Harwood, J, Frederiksen, M, Harris, MP & Wanless, S 2009, 'Integrated Data Analysis in the Presence of Emigration and Mark Loss' Journal of Agricultural, Biological and Environmental Statistics, vol 14, no. 4, pp. 411-431. DOI: 10.1198/jabes.2009.08008
Mollett, A, Schofield, C, Miller, I, Harwood, J, Harris, CM & Donovan, CR 2009, Environmental Risk Management Capability: Advice on Minimising the Impact of Both Sonar and Seismic Offshore Operations on Marine Mammals. in Offshore Europe Oil and Gas Conference and Exhibition : proceedings : 8-11 September 2009, Aberdeen Exhibition and Conference Center UK, Aberdeen UK. Society of Petroleum Engineers, pp. 1-8, 2009 SPE Offshore Europe Oil and Gas Conference and Exhibition, Aberdeen, United Kingdom, 8 September. DOI: 10.2118/124301-MS
Cabrelli, A, Harwood, J, Matthiopoulos, J, New, LF & Thomas, L 2009, A critical review of the literature on population modelling. CREEM Technical Report, no. 2009-2, University of St Andrews.
Hall, AJ & Harwood, J 2009, Mass Mortalities. in WF Perrin, B Wursig & JGM Thewissen (eds), Encyclopedia of Marine Mammals. Academic Press Inc., pp. 709-711.
Zimmer, WMX, Harwood, J, Tyack, PL, Johnson, M & Madsen, PT 2008, 'Passive acoustic detection of deep-diving beaked whales' Journal of the Acoustical Society of America, vol 124, no. 5, pp. 2823-2832. DOI: 10.1121/1.2988277
Harris, CM, Travis, JMJ & Harwood, J 2008, 'Evaluating the influence of epidemiological parameters and host ecology on the spread of phocine distemper virus through populations of harbour seals' PLoS One, vol 3, no. 7, e2710. DOI: 10.1371/journal.pone.0002710
Matthiopoulos, J, Smout, SC, Winship, A, Thompson, D, Boyd, IL & Harwood, J 2008, 'Getting beneath the surface of marine mammal - fisheries competition' Mammal Review, vol 38, no. 2-3, pp. 167-188. DOI: 10.1111/j.1365-2907.2008.00123.x
Robinson, RA, Harwood, J & and 18 others, N 2008, 'Travelling through a warming world: climate change and migratory species' Endangered Species Research. DOI: 10.3354/esr00162
Harris, CM, Matthiopoulos, J & Harwood, J 2007, 'Influence of the physical environment and conspecific aggression on the spatial arrangement of breeding grey seals' Ecological Informatics, vol 2, no. 4, pp. 308-317. DOI: 10.1016/j.ecoinf.2007.09.001
Matthiopoulos, J, Graham, K, Smout, SC, Asseburg, C, Redpath, S, Thirgood, S, Hudson, P & Harwood, J 2007, 'Sensitivity to assumptions in models of generalist predation on a cyclic prey' Ecology, vol 88, no. 10, pp. 2576-2586. DOI: 10.1890/06-0483.1
Buckland, ST, Newman, KB, Fernandez, C, Thomas, L & Harwood, J 2007, 'Embedding population dynamics models in inference.' Statistical Science, vol 22, no. 1, pp. 44-58. DOI: 10.1214/088342306000000673
Harwood, J 2007, 'Is there a role for ecologists in an ecosystem approach to the management of marine resources?' Aquatic Conservation: Marine and Freshwater Ecosystems, vol 17, no. 1, pp. 1-4. DOI: 10.1002/aqc.812
Buckland, ST, Newman, KB, Fernandez, C, Thomas, L & Harwood, J 2006, 'Defining matrix population models and fitting them to data' pp. 15-27.
Harrison, P, Buckland, ST, Thomas, L, Harris, R, Pomeroy, P & Harwood, J 2006, 'Incorporating movement into models of grey seal population dynamics' Journal of Animal Ecology, vol 75, no. 3, pp. 634-645. DOI: 10.1111/j.1365-2656.2006.01084.x
Asseburg, C, Harwood, J, Matthiopoulos, J & Smout, SC 2006, The functional response of generalist predators, and their implications for the monitoring of marine ecosystems. in IL Boyd, S Wanless & CJ Camphuysen (eds), Top predators in marine ecosystems: their role in monitoring and management . Cambridge University Press, pp. 262-274.
Matthiopoulos, J, Harwood, J & Thomas, L 2005, 'Metapopulation consequences of site fidelity for colonially breeding mammals and birds' Journal of Animal Ecology, vol 74, no. 4, pp. 716-727. DOI: 10.1111/j.1365-2656.2005.00970.x
Thomas, L, Buckland, ST, Newman, KB & Harwood, J 2005, 'A unified framework for modelling wildlife population dynamics' Australian and New Zealand Journal of Statistics, vol 47, no. 1, pp. 19-34. DOI: 10.1111/j.1467-842x.2005.00369.x
Everard, K, Harwood, J, Matthiopoulos, J, McConnell, BJ, Sharples, R, Harris, CM & Warnes, CZ 2005, Defining biologically relevant management units for seal populations in the UK. Unknown Publisher.
Gaggiotti, OE, Brooks, SP, Amos, W & Harwood, J 2004, 'Combining demographic, environmental and genetic data to test hypotheses about colonisation events in metapopulations' Molecular Ecology, vol 13, no. 4, pp. 811-825. DOI: 10.1046/j.1365-294X.2003.02028.x
Harwood, J & McLaren, IA Modelling interactions between marine mammals and fisheries: model structure, assumptions and data requirements..
Harwood, J & Stokes, K 2003, 'Coping with uncertainty in ecological advice: lessons from fisheries' Trends in Ecology and Evolution, vol 18, no. 12, pp. 617-622. DOI: 10.1016/j.tree.2003.08.001
Lonergan, M & Harwood, J 2003, 'The potential effects of repeated outbreaks of phocine distemper among harbour seals: a response to Harding et al.(2002)' Ecology Letters, vol 6, no. 10, pp. 889-893. DOI: 10.1046/j.1461-0248.2003.00510.x
Matthiopoulos, J, Harwood, J & Thomas, L 2003, Modelling the Short-Term Movements of Grey Seals between Breeding Colonies and their Long-Term Population Consequences. Unknown Publisher.
Matthiopoulos, J & Harwood, J 2003, Estimating Annual Pup Production in Grey Seal Breeding Colonies. Unknown Publisher.
Harwood, J 2002, 'Mitigating the effects of acoustic disturbance in the oceans' Aquatic Conservation: Marine and Freshwater Ecosystems, vol 12, pp. 485-488. DOI: 10.1002/aqc.550
Gaggiotti, OE, Jones, F, Lee, WM, Amos, W, Harwood, J & Nichols, RA 2002, 'Patterns of colonization in a metapopulation of grey seals' Nature, vol 416, no. 6879, pp. 424-427. DOI: 10.1038/416424a
Harwood, J Mass die-offs.
McLaren, I, Brault, S, Harwood, J & Vardy, D Report of the Eminent panel on seal management.
Harwood, J 2001, 'Marine mammal and their environments in the twenty-first century' Journal of Mammalogy, vol 82, no. 3, pp. 630-640.
Harwood, J & Wilson, B 2001, 'The implications of developments on the Atlantic frontier for marine mammals' Continental Shelf Research, vol 21, no. 8-10, pp. 1073-1093. DOI: 10.1016/S0278-4343(00)00125-4
Harwood, J Coping with uncertainty when managing marine resources..
Bowen, WD, Harwood, J, Goodman, D & Swartzman, GL Review of the November 2000 Biological Opinion and Incidental Take Statement with Respect to the Western stock of the steller Sea Lion, with comments on draft August 2001 Biological Opinion..
Harwood, J Seals and sealions.
Hess, GR, Randolph, SE, Arneberg, P, Chemini, C, Furlanello, C, Harwood, J, Roberts, M & Swinton, J Spatial aspects of disease dynamics.
Grenfell, BT, Amos, W, Arneberg, P, Bjornstadt, ON, Greenman, JV, Harwood, J, Lanfranchi, P, McLean, A, Norman, RA, Read, AF & Skorping, A Visions for future research in wildlife epidemiology.
Hall, AJ & Harwood, J Available information on the distribution and abundance of marine mammals in the Skagerrak and Kattegat..
Harwood, J 2000, 'Risk assessment and decision analysis in conservation' Biological Conservation, vol 95, no. 2, pp. 219-226.
Harwood, J 1999, 'A risk assessment framework for the reduction of cetacean by-catches' Aquatic Conservation: Marine and Freshwater Ecosystems, vol 9, no. 6, pp. 593-599.
Swinton, J, Gilligan, CA, Harwood, J & Hall, AJ 1999, 'Scaling of phocine distemper virus transmission with harbour seal community size.' Ecologie, vol 30, no. 4, pp. 231-240.
Hall, AJ, Green, NJL, Jones, KC, Pomeroy, P & Harwood, J 1998, 'Thyroid hormones as biomarkers in grey seals' Marine Pollution Bulletin, vol 36, no. 6, pp. 424-428.
Harwood, J 1998, 'What killed the monk seals?' Nature, vol 393, pp. 17-18.
Harwood, J & Amos, W 1998, 'Factors affecting levels of genetic diversity in natural populations' Philosophical Transactions of the Royal Society of London Series B, vol 353, pp. 177-186.
Swinton, JJ, Harwood, J, Grenfell, BT & Gilligan, CA 1998, 'Persistence thresholds for PDV infection in harbour seal metapopulations' Journal of Animal Ecology, vol 66, pp. 54-68.
Harwood, J 1998, 'Indirect effects in marine ecosystems: implications for management' Environmental Conservation, vol 23, no. 2, pp. 173-174.
Hall, AJ, Pomeroy, P, Green, N, Jones, K & Harwood, J 1997, 'Infection, haematology and biochemistry in grey seal pups exposed to chlorinated biphenyls' Marine Environmental Research, vol 43, no. 1/2, pp. 81-98.
Beudels, RC, Devillers, P & Harwood, J 1997, 'Estimating the size of the population of sitatunga (Tragelaphus spekei) in the "Parc National de lAkagera", Rwanda' Journal of African Ecology, vol 111, pp. 1-10.
Harwood, J, Pomeroy, P, Green, N, Hall, AJ & 2, O 1996, 'Congener-specific exposure of grey seals (Halichoerus grypus) pups to chlorinated biphenyls during lactation' Canadian Journal of Fisheries and Aquatic Sciences, vol 53, no. 7, pp. 1526-1534.
HARWOOD, J, ANDERSON, SS, FEDAK, MA, HAMMOND, PS, HIBY, AR, MCCONNELL, BJ, MARTIN, AR & THOMPSON, D 1989, 'NEW APPROACHES FOR FIELD STUDIES OF MAMMALS - EXPERIENCES WITH MARINE MAMMALS' Biological Journal of the Linnean Society, vol 38, no. 1, pp. 103-111. DOI: 10.1111/j.1095-8312.1989.tb01567.x
MCCANN, TS, FEDAK, MA & HARWOOD, J 1989, 'PARENTAL INVESTMENT IN SOUTHERN ELEPHANT SEALS, MIROUNGA-LEONINA' Behavioral Ecology and Sociobiology, vol 25, no. 2, pp. 81-87. DOI: 10.1007/BF00302924

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