Jonsson, N.N., MacLeod, M., Hayward, A.D., McNeilly, T.N., Ferguson, K.D. & Skuce, P.J. (2022) Liver fluke in beef cattle – impact on production efficiency and associated greenhouse gas emissions estimated using causal inference methods. Preventative Veterinary Medicine 2022: 105579. Download from here.


Machín, C., Corripio-Miyar, Y., Hernández, J.N., Pérez-Hernández, T., Hayward, A.D., Wright, H.W., Price, D.R.G., Matthews, J.B., McNeilly, T.N., Nisbet, A.J. & González, J.F. (2021) Cellular and humoral immune responses associated with protection in sheep vaccinated against Teladorsagia circumcincta. Veterinary Research 52: 89. Download from here.

Hayward, A.D., Skuce, P.J. & McNeilly, T.N. (2021) The influence of liver fluke infection on production in sheep and cattle: a meta-analysis. International Journal for Parasitology 51: 913-924. Download from here.

Hayward, A.D., Skuce, P.J. & McNeilly, T.N. (2021) Tolerance of liver fluke varies between breeds and producers in beef cattle. animal 15: 100126. Download from here.

Melville, L.A., Hayward, A.D., Morgan, E.R., Shaw, D.J, McBean, D., Andrews, L., Morrison, A. & Kenyon, F. (2021) Precision worm control in grazing lambs by targeting group treatment based on performance of sentinels. animal 15: 100176. Download from here.


Sparks, A.M., Hayward, A.D., Watt, K., Pilkington, J.G., Pemberton, J.M., Johnston, S.E., McNeilly, T.N. & Nussey, D.H. (2020) Maternally-derived anti-helminth antibodies predict offspring survival in a wild mammal. Proc. R. Soc. Lond. B. 287 20201931. Download from here.


Hayward, A.D., Garnier, R., Childs, D.Z., Grenfell, B.T., Watt, K.A., Pilkington, J.G., Pemberton, J.M. & Graham, A.L. (2019) From population to individual host scale and back again: testing theories of infection and defence in the Soay sheep of St. Kilda. In Wildlife Disease Ecology: Linking Theory to Data and Application eds. Wilson, K., Fenton, A. & Tompkins, D.M. Cambridge University Press. Take a look here.

Hayward, A.D., Pilkington, J.G., Wilson, K., McNeilly, T.N. & Watt, K.A. (2019) Reproductive effort influences intra‐seasonal variation in parasite‐specific antibody responses in wild Soay sheep. Functional Ecology 33: 1307-1320. Download from here.

Leivesley, J.A., Bussière, L.F., Pemberton, J.M., Pilkington, J.G., Wilson, K. & Hayward, A.D. (2019) Survival costs of reproduction are mediated by parasite infection in wild Soay sheep. Ecology Letters 22: 1203-1213.Download from here.


Griffin, R.M., Hayward, A.D., Bolund, E., Maklakov, A.A. & Lummaa, V. (2018) Sex differences in adult mortality rate mediated by early-life environmental conditions. Ecology Letters 21: 235-242. Download from here.

Hayward, A.D., Pemberton, J.M., Bérénos, C., Wilson, A.J., Pilkington, J.G. & Kruuk, L.E.B. (2018) Evidence for selection-by-environment but not genotype-by-environment interactions for fitness-related traits in a wild mammal population. Genetics 208: 349-364. Download from here.

Nenko, I., Hayward, A.D., Simons, M.J.P. & Lummaa, V. (2018) Early-life environment and differences in costs of reproduction in a preindustrial human population. PLoS One 13: e0207236. Download Open Access PDF here.


Lynsdale, C.L., Mumby, H.S., Hayward, A.D., Mar, K.U. & Lummaa, V. (2017) Parasite-associated mortality in a long-lived mammal: Variation with host age, sex, and reproduction. Ecology and Evolution 7: 10904-10915. (download Open Access pdf from here).


Hayward, A.D., Rigby, F.L. & Lummaa, V. (2016) Early-life disease exposure and associations with adult survival, cause of death, and reproductive success in preindustrial humans. Proceedings of the National Academy of Sciences of the USA 113: 8951-8956 (download PDF from here).

Bolund, E., Hayward, A.D. & Lummaa, V. (2016) Using Quantitative Genetics to Understand Human Life-History Variation. In Encyclopedia of Evolutionary Biology (Ed: R. Kliman), pp. 328-334. Elsevier, Amsterdam. (download pdf from here)


Hayward, A.D., Moorad, J., Regan, C.E., Berenos, C., Pilkington, J.G., Pemberton, J.M. & Nussey, D.H. (2015) Asynchrony of senescence among phenotypic traits in a wild mammal population. Experimental Gerontology 7156-68. Part of a Special Issue ‘Ageing In The Wild’(download PDF from here)

Mumby, H.S., Mar, K.U., Hayward, A.D.,  Htut, W., Htut-Aung, Y. & Lummaa, V. (2015) Elephants born in the high stress season have faster reproductive ageing. Scientific Reports 513946. (download PDF from here)

Lynsdale, C.L., Franco dos Santos, D.J., Hayward, A.D., Mar, K.U., Htut, W., Aung, H.H., Soe, A.T. & Lummaa, V. (2015) A standardised faecal collection protocol for intestinal helminth egg counts in Asian elephants, Elephas maximusInternational Journal for Parasitology: Parasites and Wildlife 4: 307-315. (download PDF from here)

Hayward, A.D., Lummaa, V. & Bazykin, G.A. (2015) Fitness consequences of advanced ancestral age over three generations in humans. PLoS One10: e0128197. (download the Open Access PDF here) 

Hayward, A.D., Nenko, I. & Lummaa, V. (2015) Early-life reproduction is associated with increased mortality risk but enhanced lifetime fitness in pre-industrial humans. Proceedings of the Royal Society of London B 282:  20143053. (download PDF from here)

Bolund, E., Hayward, A.D., Pettay, J.E. & Lummaa, V. (2015) Effects of the demographic transition on the genetic variances and covariances of human life history traits. Evolution 69: 747-755. (download PDF from here)


Hayward, A.D., Nussey, D.H., Wilson, A.J., Berenos, C., Pilkington, J.G., Watt, K.A., Pemberton, J.M. & Graham, A.L. (2014) Natural selection on individual variation in tolerance of gastrointestinal nematode infection. PLoS Biology 12:e1001917. (download the Open Access PDF here)

Hayward, A. D., Garnier, R., Watt, K.A., Pilkington, J.G., Grenfell, B.T., Matthews, J.B., Pemberton, J.M., Nussey D.H. & Graham, A.L. (2014) Heritable, heterogeneous and costly resistance of sheep against nematodes and potential feedbacks to epidemiological dynamics. American Naturalist.184: S58-S76. (download PDF from here)

Hayward, A.D., Mar, K.U., Lahdenpera, M. & Lummaa, V. (2014) Early reproductive investment, senescence and lifetime reproductive success in female Asian elephants. Journal of Evolutionary Biology27: 772-783. (download the Open Access PDF here)

Nenko, I., Hayward, A.D. & Lummaa, V. (2014) The effect of socio-economic status and food availability on first birth interval in a preindustrial human population. Proceedings of the Royal Society of London B 281. (download PDF from here)


Hayward, A.D. (2013) Causes and consequences of intra- and inter-host heterogeneity in defence against nematodes. Parasite Immunology 35: 362-373. (download PDF from here)

Hayward, A.D., Rickard, I.J. & Lummaa, V. (2013) The influence of early-life nutrition on mortality and reproductive success during a subsequent famine in a pre-industrial population. Proceedings of the National Academy of Sciences of the USA 110: 13886-13891. (download the Open Access PDF here)

Hayward, A.D. & Lummaa, V. (2013) Testing the evolutionary basis of the Predictive Adaptive Response hypothesis in a preindustrial human population. Evolution, Medicine and Public Health 1: 106-117. (download the Open Access PDF here)

Hayward, A.D., Wilson, A.J., Pilkington, J.G., Pemberton, J.M. & Kruuk, L.E.B. (2013) Reproductive senescence in female Soay sheep: variation across traits and contributions of individual ageing and selective disappearance. Functional Ecology 27: 184-195. (download PDF here)

Brown, E.A., Pilkington, J.G., Nussey, D.H., Watt, K.A., Hayward, A.D., Tucker, R., Graham, A.L., Paterson, S., Beraldi, D., Pemberton, J.M. & Slate, J. (2013) Detecting genes for parasite burden and immunological traits in a wild population: testing the candidate gene approach. Molecular Ecology 22: 757-773. (download PDF here)


Hayward, A.D., Holopainen, J., Pettay, J. & Lummaa, V. (2012) Food and fitness: associations between crop yields and life-history traits in a longitudinally-monitored preindustrial human population. Proceedings of the Royal Society of London B 279: 4165-4173. (download the Open Access PDF here 

Hayward, A.D., Wilson, A.J., Pilkington, J.G., Clutton-Brock, T.H., Pemberton, J.M. & Kruuk, L.E.B. (2011) Natural selection on parasite burden varies across ages and environmental conditions in a wild mammal. Journal of Evolutionary Biology 24: 1664-1676. (download PDF here)

Graham, A.L., Hayward, A.D., Watt, K.A., Pilkington, J.G., Pemberton, J.M. & Nussey, D.H. (2010) Fitness correlates of heritable variation in antibody responsiveness in a wild mammal. Science 330: 662-665. (download PDF here)

Hayward, A.D., Pilkington, J.G., Pemberton, J.M. & Kruuk, L.E.B. (2010) Maternal effects and early-life performance are associated with parasite resistance across life in free-living Soay sheep. Parasitology 137: 1261-1273. (download PDF here 

Hayward, A.D., Wilson, A.J., Pilkington, J.G., Pemberton, J.M. & Kruuk, L.E.B. (2009) Ageing in a variable habitat: environmental stress affects senescence in parasite resistance in St Kilda Soay sheep. Proceedings of the Royal Society of London B 276: 3477-3485. (download the Open Access PDF here)