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Department of Zoology



Following a degree in environmental chemistry (London University) and a D.Phil. on the biochemistry of lead poisoning from ammunition sources in birds (Oxford University) I spent four years carrying out postdoctoral research at the Station Biologique de la Tour du Valat in the Camargue, France. I then spent sixteen years in the Research Department of the RSPB where I initiated and headed up the International Research Section working on bird conservation problems in collaboration with BirdLife Partner organisations in Africa, Asia and Europe. I subsequently spent ten years as Director of Conservation at the Wildfowl and Wetlands Trust (WWT) overseeing their wetland science and delivery in the UK and overseas. This included a 6-month secondment to World Land Trust acting as CEO. Since leaving WWT in 2018 I have worked independently and am actively involved in conservation science. In 2018 I became an Honorary Research Fellow in the Department of Zoology at Cambridge University and in 2019 became an Honorary Professor in the School of Biological Sciences at the University of East Anglia.

I have previously held several Trustee roles, including for the RSPB, the Wetlands International European Association and WWT Consulting, and I have sat various technical expert committees and the Cambridge Conservation Initiative Collaborative Fund Selection Panel. My current roles include the following public appointments: member of the Darwin Expert Committee; the Darwin Plus Advisory Group; the UK REACH Independent Scientific Expert Pool (RISEP). My other roles include as a member of the Lead Ammunition Group and as vice-president of the Science and Conservation Council of Station Biologique de la Tour du Valat (research institute for the conservation of Mediterranean wetlands). I am a Fellow of WWF-UK.



I have a wide range of conservation research interests, including diagnosing the causes of declines in threatened bird species and developing and testing practical and policy solutions to reverse them. I have worked on environmental contaminants, farming systems, climate change, the wild bird trade, and protected area networks. I have a particular interest in ecotoxicology, especially lead poisoning from ammunition sources in birds.


Other publications: 

Pain, D., Green, R., Bates, N. & Allen, C. 2022. Ingestion of lead ammunition by companion animals. Letter. Vet Record 22/29 October, pages 345-346.

Green, R., Taggart, M., Pain, D., Smithson, K. 2022. Implications for food safety of the size and location of fragments of lead shotgun pellets embedded in hunted carcasses of small game animals intended for human consumption. PLoS ONE 17(8):

Pain, D.J., Green, R.E., Taggart, M. & Kanstrup, N. 2022. How contaminated with ammunition-derived lead are European small game animals? Assessing and reducing risks to human health. Ambio 51: 1772–1785.

Camacho, C., Negro, J.J., Elmberg, J., Fox, A.D., Nagy, S., Pain, D.J. & Green, A.J. 2022. Groundwater extraction poses extreme threat to Doñana World Heritage Site. Correspondence. Nature Ecology & Evolution. Published online 25 April 2022.

Thomas, V.G., Pain, D.J., Kanstrup, N. & Cromie, R. 2022. Increasing the Awareness of Health Risks from Lead Contaminated Game Meat Among International and National Human Health Organizations. European Journal of Environment and Public Health 6(2): em0110

Hampton, J.O., Dunstan, H., Toop, S.D., Flesch, J.S., Andreotti, A. & Pain, D.J. 2022. Lead ammunition residues in a hunted Australian grassland bird, the stubble quail (Coturnix pectoralis): implications for wildlife and human health. PLoS ONE 17(4): e0267401.  

Green, R.E., Taggart, M.A., Pain, D.J., Clark, N.A., Clewley, L., Cromie, R., Dodd, S.G., Elliot, B., Green, R.M.W., Huntley, B., Huntley, J., Pap, S., Porter, R., Robinson, J.A., Sheldon, R., Smith, K.W., Smith, L., Spencer, J. & Stroud, D. 2022. Effectiveness of actions intended to achieve a voluntary transition from the use of lead to non-lead shotgun ammunition for hunting in Britain. Conservation Evidence 19: 8-14.

Green, R.E., Pain, D.J. & Krone, O. 2022. The impact of lead poisoning from ammunition sources on raptor populations in Europe. Science of the Total Environment.

Thomas, V.G., Kanstrup, N & Pain, D.J. 2021. Promoting the Transition to Non-Lead Hunting Ammunition in the European Union through Regulation and Policy Options. Environmental Policy and Law l. 51 (4): 239-254.

Stroud, D., Pain, D.J. & Green, R.E. 2021. Evidence of widespread illegal hunting of waterfowl in England despite partial regulation of the use of lead shotgun ammunition. Conservation Evidence 18: 18-24

Green, R.E., Taggart, M.A., Pain, D.J., Clark, N.A., Clewley, L., Cromie, R., Elliot, B., Green, R.M.W., Huntley, B., Huntley, J., Leslie, R., Porter, R., Robinson, J., Smith, KW., Smith, L., Spencer, J. & Stroud, D. 2021. Effect of a joint policy statement by nine UK shooting and rural organisations on the use of lead shotgun ammunition for hunting pheasants in Britain. Conservation Evidence. Conservation Evidence 18:1-9.

Pain, D.J., Bardin, Ph., Hutchinson, N., Pénzesné Kónya, E. & Krause, M. 2020. A review of European progress towards the Global Strategy for Plant Conservation 2011-2020. PLANTA EUROPA and Plantlife International, 170 pp.

Taggart, M.A., Shore, R.F., Pain, D.J., Peniche, G., Martinez-Haro, M., Mateo, R., Homann, J., Raab, A., Feldmann, J., Lawlor, A.J., Potter, E.D., Walker, L.E., Braidwood, D.W., French, A.S., Parry-Jones, J., Swift, J.A. & Green, R.E. 2020. Concentration and origin of lead (Pb) in liver and bone of Eurasian buzzards (Buteo buteo) in the United Kingdom. Environmental Pollution 267 (2020) 115629 -

Green, R.E. & Pain, D.J. 2020. Additional mortality rate of wildfowl caused by lead poisoning from gunshot ingestion: a re-analysis of data from a 70-year-old field experiment on wild Mallards Anas platyrhynchos Wildfowl 70: 242–256.  

Pain, D.J. 2020. A review of the Hairy-nosed Otter (Lutra sumatrana) in Borneo and a recent sighting at Danum Valley Conservation Area, Sabah. IUCN Otter Specialist Group Bulletin 37(3): 167-174.

Pain, D., Swift, S., Green, R. & Cromie, R. 2020. Letters: Further news on lead ammunition. British Birds 113: 232.

Thomas, V.G., Pain, D.J., Kanstrup, N. & Green, R. E. 2020. Setting maximum levels for lead in game meat in EC regulations: an adjunct to replacement of lead ammunition. Ambio DOI:

Pain, D.J., Swift, J., Green, R. & Cromie, R. 2020. The tide is turning for lead ammunition. British Birds 113: 110-118.

Pain, D.J. & Green, R.E. 2019. Risks from Lead Ammunition: Correspondence. Nature Sustainability. Published online 14 October 2019.

Arnemo, J. M., Cromie, R., Fox, A.D., Kanstrup, N., Mateo, R., Pain, D.J. & Thomas, V.G. 2019.

Transition to lead-free ammunition benefits all. Ambio 48(9): 1097-1098. Special Issue: LEAD USE IN HUNTING. Online 18 July 2019.

Green, R.E. & Pain, D.J. 2019. Risks to human health from ammunition-derived lead in Europe. Ambio 48(9): 954-968. Special Issue: LEAD USE IN HUNTING. Online 16 May 2019. DOI: 10.1007%2Fs13280-019-01194-x.pdf

Pain, D. J., Mateo, R. & Green, R.E. 2019. Effects of lead from ammunition on birds and other wildlife: a review and update. Ambio 48(9):935-953. Special Issue: LEAD USE IN HUNTING. Online 16 March 2019.  DOI: 10.1007/s13280-019-01159-0

Pain, D.J., Dickie, I., Green, R. E., Kanstrup, N. & Cromie, R. 2019.  Wildlife, human and environmental costs of using lead ammunition: an economic review and analysis. Ambio 48(9): 969-988.  Special Issue: LEAD USE IN HUNTING. Online 16 March 2019. DOI: 10.1007/s13280-019-01157-2        

Wood, K.A., Brown, M.J., Cromie, R.L., Hilton, G.M., Mackenzie, C., Newth, J.L., Pain, D.J., Perrins C.M., & Rees E.C. 2019. Regulation of lead fishing weights results in mute swan population recovery. Biological Conservation 230:67-74

Movalli, P., Krone, I., Osborn, D. & Pain, D. 2018: Monitoring contaminants, emerging infectious diseases and environmental change with raptors, and links to human health. Bird Study

Pain, D., Hughes, B., Syroechkovskiy, E., Zöckler, C., Chowdhury, S., Anderson, G. & Clark, N. 2018. Saving the Spoon-billed Sandpiper: a conservation update. British Birds 111:323-333.

Pain, D., Green, R & Cromie, R. 2018. Ornithological Masterclass 10: Ecotoxicology: Lead and Birds. BTO News, Winter 2018: 16-19.

Arnemo, J.M., Andersen, O., Stokke, S., Thomas, V.G., Krone, O., Pain, D.J. & Mateo, R. 2016. Health and environmental risks from lead-based ammunition: science versus socio-politics, Ecohealth 13 (4): 618–622.

Green, R.E. & Pain, D.J. 2016. Possible effects of ingested lead gunshot on populations of ducks wintering in the UK. Ibis 158 (4): 699-710.

Newth, J. L., Rees, E. C., Cromie, R. L., McDonald, R. A., Bearhop, S., Pain, D. J., Norton, G. J., Deacon, C. and Hilton, G. M.. 2016.  Widespread exposure to lead affects the body condition of free-living whooper swans Cygnus cygnus wintering in Britain. Environmental Pollution Feb; 209:60-7.

Pain, D.J., Cromie, R. & Green, R.E. 2015. Poisoning of birds and other wildlife from ammunition-derived lead in the UK. Pages 58-84 In: Delahay, R.J. & Spray, C.J. (Eds.). Proceedings of the Oxford Lead Symposium. Lead Ammunition: understanding and minimising the risks to human and environmental health. Edward Grey Institute, University of Oxford, UK. 152pp.

Green, R.E., & Pain, D.J. 2015. Risks of health effects to humans in the UK from ammunition-derived lead. Pages 27-43 In: Delahay, R.J. & Spray, C.J. (Eds.) 2015. Proceedings of the Oxford Lead Symposium. Lead Ammunition: understanding and minimising the risks to human and environmental health. Edward Grey Institute, University of Oxford, UK. 152pp.

Vickery, J., Ewing, S.R., Smith, K.W., Pain, D.J., Bairlein, F., Skorpilov,J.A. & Gregory, R.D. 2014. The decline of Afro-Palaearctic migrants and an assessment of potential causes. Ibis 156:1-22. 

Clark, N, Green, R & Pain, D. 2014. Saving the Spoon-billed Sandpiper: an update on the Conservation Programme. British Birds 107: 467– 475.

Clark, N, Green, R & Pain, D. 2014. Back from the brink? An update on the Spoon-billed Sandpiper Calidris pygmeus conservation programme. BirdingASIA 22: 22-19.

Newth, J. L., Cromie, R. L., Brown, M. J., Delahay, R. J., Meharg, A. A., Deacon, C., Norton, G. J.,  O’Brien, M. F. & Pain, D. J..  2012. Poisoning from lead gunshot: still a threat to wild waterbirds in Britain. European Journal of Wildlife Research.

Green, R.E. & Pain, D.J. 2012. Potential health risks to adults and children in the UK from exposure to dietary lead in gamebirds shot with lead ammunition. Food and Chemical Toxicology, 50: 4180-4190.

Oppel, S., Pain, D. J., Lindsell, J., Lachmann, L., Diop, I., Tegetmeyer, C., Donald, P. F., Anderson, G., Bowden,C. G. R., Tanneberger, F. & Flade, M. 2011. High variation reduces the value of feather stable isotope ratios in identifying new wintering areas for aquatic warblers in West Africa. J. Avian Biol. 42: 342-354.

Hole, D. G., Huntley, B., Arinaitwe, J., Butchart, S. H. M., Collingham, Y. C., Fishpool, L. D. C., Pain, D. J. and Willis, S. G. , 2011. Toward a Management Framework for Networks of Protected Areas in the Face of Climate Change. Conservation     Biology 25(2):    305-315.

Franson, J.C. and Pain, D.J.  2011. Lead in Birds.  Chapter 16 (pp 563-594) in Environmental Contaminants in Biota: interpreting tissue concentrations. Second edition. Ed WN Beyer & J.P Meador. CRC Press, Boca Raton. Fl. USA.

Pain, D., Green, R & Clark, N. 2011. On the Edge: can the Spoon-billed Sandpiper be saved? British Birds 104: 347-412.

Pain, D., Green, R & Clark, N. 2011. Bird on the Edge: can the Spoon-billed Sandpiper Eurynorhynchus pygmeus be saved? BirdingASIA 15:26-35.

Feber, R., Hilton, G.M., Hutchins, E., Griffin, L., Ewald, N., Pain, D., Biggs, J., Hughes, I.  & Macdonald, D. 2011. Ecology and Conservation of the Tadpole Shrimp Triops cancriformis in Britain. British Wildlife June 2011. 334-341.

Pain, D.J., Cromie, R.L., Newth, J., Brown, M.J., Crutcher, E., Hardman, L.H., Hurst, L., Mateo, R., Meharg, A.A., Moran, A.C., Raab, A., Taggart, M.A. & Green, R.E. 2010. Potential Hazard to Human Health from Exposure to Fragments of Lead Bullets and Shot in the Tissues of Game Animals. PLoS ONE 5(4):e10315.

Hole, D.G., Willis, S. G., Pain, D.J., Fishpool, L., Butchart, S.H.M., Collingham, Y.C., Rahback, C., and Huntley, B. 2009. Projected impacts of climate change on a continent-wide protected area network. Ecology Letters 12(5) 420-431.

Doswald, N., Willis, S.G., Collingham, Y.C., Pain, D.J., Green, R.E., and Huntley, B. 2009. Potential impacts of climate change on the breeding and non-breeding ranges and migration distance of European Sylvia warblers. Journal of Biogeography. 36(6): 1194-1208.

Taggart, M.,Senacha, K., Green, R., Cuthbert, R., Jhala, Y., Meharg, A., Mateo, R.& Pain, D. 2009. Analysis of nine NSAIDs in ungulate tissues available to Critically Endangered vultures in India. Environmental Science and Technology 43: 4561-4566.

Pain, D. J., I. J. Fisher, and V. G. Thomas. 2009. A global update of lead poisoning in terrestrial birds from ammunition sources. In:  R. T. Watson, M. Fuller, M. Pokras, and W. G. Hunt (Eds.). Ingestion of Lead from Spent Ammunition: Implications for Wildlife and Humans. The Peregrine Fund, Boise, Idaho, USA.

Pain, D. 2009. Commentary on research needs and remediation of lead exposure from ammunition in wildlife and humans. In R. T. Watson, M. Fuller, M. Pokras, and W. G. Hunt (Eds.). Ingestion of Lead from Spent Ammunition: Implications for Wildlife and Humans. pp. 369-371. The Peregrine Fund, Boise, Idaho, USA.

Gabremichael, M.N., Spottiswoode, C., Fishpool, L., Forsythe, E., Lewis, A., Pain, D, Thomas, R & Toye, N. 2009. Occurrence of the White-tailed Swallow Hirundo megaensis near Negele, Ethiopia. Bull. African Bird Club.16(1): 83-86.

Rattner, B.A., Whitehead, M.A., Gasper, G., Metayer, C.U., Limk, W.A., Taggart, M.A., Meharg, A.A., Pattee, O.H., & Pain, D.J. 2008. Apparent tolerance of turkey vultures (Cathares aura) to the non-steroidal anti-inflammatory drug diclofenac. Environmental Toxicology and Chemistry 27: 2341-2345.

Pain, D.J., White, R., Stevenson, J., Bell, M., Williams, K.K., Fisher, P. & Wright, G. 2008.Toxicity and persistence of sodium fluoroacetate (1080) in the land crab (Gecarcinus lagostoma) on Ascension Island. Wildlife Research. 35(1): 1-7.

Senacha, K.R., Taggart, M.A., Rahmani, A.R., Jhala, Y.V., Cuthbert, R., Pain, D.J., & Green, R.E.  2008. Diclofenac levels in livestock carcasses in India before the ban. JBNHS 105(2): 148-161.

Cuthbert, R., Prakash, V., Das, D., Green, R., Jhala, Y.V., Pain, D., Senacha, K-R., Shah, N. & Taggart, M. The Role of veterinary diclofenac in the decline of vulture populations in South Asia. Indian Journal of Veterinary Medicine 2: 80-85.

Pain, D.J., Bowden, C.G.R., Cunningham, A.A., Cuthbert, R., Das, D., Ghalsasi, D.R., Gilbert, M., Jakati, R., Jhala, Y., Khan, A.A., Naidoo, V., Oaks, J.L., Parry-Jones, J., Prakash, V., Rahmani, A., Baral, H.S., Senacha, K.R., Shah, N., Swan, G., Swarup, D., Taggart, M.A., Watson, R.T., Virani, M.Z., Wolter, K. & Green, R.E. 2008. The race to prevent the extinction of South Asian vultures. Bird Conservation International 18:S30-S48.

Brooks, T. M., Collar, N. J., Green, R. E., Marsden S. J. and D. J. Pain. 2008. The Science of Bird Conservation. Bird Conservation International, Volume 18, Supplement S1, September 2008, pp S2-S12

Prakash, V., Green, R.E., Pain, D.J., Ranade, S.P., Saravanan, S., Prakash, N., Venkitachalam, R., Cuthbert, R., Rahmani, A. & Cunningham, A. 2007 Recent changes in populations of resident Gyps vultures in India. JBNHS 104(2): 129-135.

Swarup, D., Patra, R. C., Prakash, V., Cuthbert,, R., Das,, D.,  Avari, P, Pain,, D.J., Green, R.E., Sharma, A.K.,  Saini, M.,  Das, D., Taggart, M. 2007. Safety of meloxicam to critically endangered Gyps vultures and other scavenging birds in India. Animal Conservation 10: 192-198.

Senapathi, D,.Vogiatzakis, I.N., Jeganathan, P,  Gill, J.A., Green, R.E., Bowden, C.G.R. , Rahmani, A.R. ,  Pain, D. J., & Norris, K. 2007. Use of remote sensing to measure change in the extent of habitat for the critically endangered Jerdon’s Courser Rhinoptilus bitorquatus in India. Ibis 149(2): 328-337.

Taggart, M.A., Senacha, K., Green, R.G., Jhala, Y.V, Ragathan,B, Rahmani, A.R., Cuthbert, R., , Pain, D.J. & Meharg, A.A. 2007. Diclofenac residues in carcasses of domestic ungulates available to vultures in India. Environment International. 33: 759-765.

Green, R.E., Taggart. M.A., Senacha, K.R., Pain, D.J., Jhala, Y., & Cuthbert, R. 2007 Rate of decline of the oriental white-backed vulture Gyps bengalensis population in India estimated from measurements of diclofenac in carcasses of domesticated ungulates. PloS One August 2007 (8): e686

Pain, D.J., Carter, I., Sainsbury, A. W., Shore, R.F., Eden, P, Taggart, M.A., Konstantinos, S., Walker, L.A, Meharg, A.A. & Raab, A. 2007. Lead contamination and associated disease in captive and reintroduced red kites Milvus milvus in England. Sci. Total Environ. 376:116-127.

Taggart, M.A., Cuthbert, R., Das, D., Pain, D.J., Green, R.E., Shultz, S., Cunningham, A.A., Meharg, A.A.. 2007. Diclofenac disposition in Indian cow and goat with reference to Gyps vulture population declines. Environ. Pollut. 147: 60-65. doi:10.1016/j.envpol.2006.08.017 .  

Knopp, D., Deng, A., Letzel, M., Taggart, M., Himmelsbach, M., Zhu, Q-Z., Peröbner, I., Kudlak, B., Frey, S., Sengl, M., Buchberger, W., Hutchinson, C., Cunningham, A., Pain, D., Cuthbert, R., Raab, A., Meharg, A., Swan, G., Jhala, Y., Prakash, V., Rahmani, A., Quevedo M., and Niessner, R. 2007. Immunological determination of the pharmaceutical diclofenac in environmental and biological samples. In: Kennedy, I.R., Solomon, K., Gee, S., Crossan, A., Wang, S., & Sanchez-Bayo, F (eds.) Rational Environmental Management of Agrochemicals: Risk Assessment, Monitoring and Remedial Action. Chapter 13. American Chemical Society Symposium Series. Oxford University Press. pp 203-226.

Bowden, C.G.R., Prakash, V., Lindsay, N., Jakati, R.D., Cuthbert, R., Rahmani, A., Pain, D.J., Green, R.E., Cunningham, A.A., Parry-Jones, J. 2007. Progress with a conservation breeding programme to save three Critically Endangered Gyps vultures from extinction. Proceedings of the IV ISBBC September Toronto, Canada. 

Swan, G.E., Cuthbert, R., Quevedo, M., Green, R.E., Pain, D.J., Bartels, P., Cunningham, A.A., Duncan, N., Meharg, A.A., Oaks, J.L., Parry-Jones, J., Shultz, S., Taggart, M.A., Verdoorn, G., Wolter, K. 2006. Toxicity of diclofenac to Gyps vultures. Biology Letters 

Swan, G., Naidoo, V., Cuthbert, R., Green, R.E., Pain, D.J., Swarup, D., Prakash, V., Taggart, M., Bekker L, Das D, Diekmann J, Diekmann M, Killian E, Meharg A, Patra RC, Saini M, Wolter K. 2006. Removing the Threat of Diclofenac to Critically Endangered Asian Vultures. 2006. Public Library of Science Biology. March 2006 4(3): e66.

Prakash, V., Cuthbert, R.J., Green, R.E., Pain, D.J., Saravanan, Ranade, S. and Cunningham, A.A. 2006  Rapid population declines of Egyptian Vulture and Red-Headed Vulture in India. Animal Conservation 9:349354.

Green, R.E., Taggart, M.A., Das, D., Pain, D.J., Sashikumar, C., Cunningham, A.A., Cuthbert, R. 2006. Collapse of Asian vulture populations: risk of mortality from residues of the veterinary drug diclofenac in carcasses of treated cattle. J. Appl. Ecol. 43:949-956

Fisher, I.J., Pain, D.J., Thomas, V. 2006. A review of lead poisoning from ammunition sources in terrestrial birds. Biol. Cons. 131: 421-432. 0.1016/j.biocon.2006.02.018

Pain, D.J., Martins, T.L.F., Boussekey, M., Diaz, S.H., Downs, C.T., Ekstrom, J.M.M., Garnett, S., Gilardi, J.D., McNiven, D., Primot, P., Rouys, S., Saoumoé, M., Symes, C.T., Tamungang, S.A., Theuerkauf, J.,.Villafuerte, D., Verfailles, L., Widmann, P., & Widmann, I.D.. 2006. The impact of protection on nest take and nesting success of parrots in Africa, Asia and Australasia. Animal Conserv. 9: 322-330.

Cuthbert, R., Pain, D.J., Green. R.E, Swan, G. & Swarup, D. 2006. Comparative toxicity studies of NSAIDs in birds: A criticism of Reddy et al. Environ. Toxicol. Pharmacol. 23(2): 254-255

Tushabe, H, Kalema, J., Byaruhanga, A.,  Asasira, J.,  Ssegawa, P, Balmford, A.,  Davenport, T, Fjeldså, J., Friis,  I.,  Pain, D.J., Pomeroy, D,  Williams, P., Williams, C. 2006. A Nationwide Assessment of the Biodiversity Value of Uganda's Important Bird Areas Network Conservation Biology 20(1): 85-99

Cuthbert, R., Parry-Jones, J, Green, R.E. and Pain, D.J. 2006. NSAIDs and scavenging birds: potential impacts beyond Asia’s critically endangered vultures. Biol. Lett. 3: 90-93

Martins, T.L.F., de L. Brooke, M., Hilton, G., Famsworth, F., Gould, J, Pain, D.J. 2006. Costing eradications of alien mammals from islands. Animal Conserv. 9(4): 439-444.

Jeganathan, P., Green, R.E., Norris, K., Wotton, S.R., Bowden, C.G.R., Pain, D. & Rahmani, A.R. 2006.Conservation of the Critically Endangered Jerdon’s Courser Rhinoptilus Bitorquatus in India. JBNHS 103(2-3):227-230.

Sanderson, F.J., Donald, P.F., Pain D.J., Burfield, I.J., van Bommel, F.P.J. 2006. Long-term population declines in Afro-Palearctic migrant birds. Biol. Conserv. 131: 93-105.

Naidoo, V., Swan, G., Cuthbert, R., Green, R., Pain, D., Swarup, D., Prakash, V., Taggart, M., Bekker, L., Das, D., Diekmann, J., Diekmann, M., Killian, E., Meharg, A., Patra, R.C., Saini, M., Wolter, K., & Bartels, P. 2006. Could meloxicam be the saviour of the Asian white-backed vulture? Proceedings of the 17th Conference of the African Association of Zoos and Aquaria, Gonubie, South Africa 10-12 May 2006. pp 29-35.

Cardoso M, Hyatt A, Selleck P, Lowther S, Prakash V, Pain D, Cunningham AA, Boyle D. 2005. Phylogenetic analysis of the DNA polymerase gene of a novel alphaherpesvirus isolated from an Indian gyps vulture. Virus Genes. 2005 May. 30(3):371-81.

Prakash, V., Green, R.E., Rahmani, A.R., Pain, D.J., Virani, M.Z., Khan, A.A., Baral, H.S., Jhala, Y.V.,Naoroji, R., Shah, N., Bowden, C.G.R., Choudhury, B.C., Narayan, G., Gautam, P. 2005. Evidence to support that diclofenac caused catastrophic vulture population decline Current Sci., vol. 88(10): 1533-1534

Pain, D.J., Meharg, A.A., Ferrer, M., Taggart, M., Pentariani, V. 2005. Lead concentrations in bones and feather of the globally threatened Spanish imperial eagle. Biol. Cons. 121: 603-610.

Taggart, M.A., Carlisle, M, Pain, D.J., Williams, R., Green, D., Osborn, D., Meharg, A.A. 2005. Arsenic levels in the soils and macrophytes of the ‘Entremuros’ after the Aznalcóllar mine spill. Environ. Pollut.  133 (1): 129-138.

Pain, D.J., Fishpool, L., Byaruhanga, A., Arinaitwe, J., and Balmford, A. 2005. Biodiversity representation in Uganda’s forest IBAs. Biol.  Conserv. 125:133-138.

Pain, D.J., Cuthbert, R., Green, R.E., Bowden, C., Taggart, M., Swan, G. E., Naidoo, V., Wolter, K., Cunningham, A.A, Parry-Jones, J., Jhala, Y., Swarup, D., Rahmani. A., & Prakash, V. Gyps species vulture declines across South Asia: the role of the veterinary drug diclofenac. Extended abstract for plenary lecture at the 27th IUGB Conference, Hannover Germany August 2005.

Jeganathan, P., Green, R.E., Norris, K. Vogiatzakis, I. N., Bartsch, A., Wotton, S.R., Bowden, C.G.R, Griffiths, G.H., Pain, D. & Rahmani, A.B. 2004. Modelling habitat selection and distribution of the Critically Endangered Jerdon’s courser Rhinoptilus bitorquatus in scrub jungle: an application of a new tracking method.  J. Appl. Ecol. 41: 224-237

Shultz, S., Baral, H.S., Charman, S., Cunningham, A.A., Das, D., Ghalsasi, D.R., Goudar, M.S., Green, R.E., Jones, A., Nighot, P., Pain, D.J., Prakash, V. 2004. Diclofenac poisoning is widespread in declining vulture populations across the Indian subcontinent. Proceedings of the Royal Society of London B (Supplement), 271:S458-S460.

Green, R.E., Newton, I., Shultz, S., Cunningham, A.A., Gilbert, M., Pain, D.J., & Prakash, V.. 2004. Diclofenac poisoning as a cause of vulture population declines across the Indian subcontinent. J. Appl. Ecol.  41: 793-800.

Taggart, M.A., Carlisle, M., Pain, D.J., Williams, R., Osborn, D., Joyson, A., Meharg, A.A. 2004. The distribution of arsenic in soils affected by the Aznalcóllar mine spill, SW Spain. Sci. Total Environ. 323:137-152.

Pain, D.J., Gargi, R., Cunningham, A.A., Jones, A. & Prakash, V. 2004. Mortality of globally threatened Sarus cranes Grus antigon from monocrotophos poisoning in India. Sci. Total Environ. 326: 55-61.

Shultz, S., Pain, D.J., Cunningham, A., Green, R.E., Parry-Jones, J & Prakash, V.  2004. Current conservation priorities for the Asian Gyps vulture crisis. pp. 208-213 In: Monadjem, A., Anderson, M.D., Piper, S.E. & Boshoff, A.F. (Eds). Vultures in The Vultures of Southern Africa – Quo Vadis? Proceedings of a workshop on vulture research and conservation in southern Africa. Birds of Prey Working Group, Johannesburg.

Prakash, V., Pain, D.J., Shultz, S., and Cunningham A.A. 2004. Saving Asia's Gyps Vultures: The "Vulture Rescue" Team's Conservation Programme. Pages 245-255 in ‘Raptors Worldwide’ edited by R.D. Chancellor and B-U. Meyburg. WWGBP/MME.  

Jeganathan, P.; Rahmani, A. R.; Green, R.E.; Norris, K.; Bowder, C.; Wotton, S. R.; Pain, D. 2004. Conservation of the critically endangered Jerdon's Courser Rhinoptilus bitorquatus in India: final report. Bombay Natural History Society, Mumbai.

Pain, D.J., Meharg, A., Sinclair, G., Powell, N., Finnie, J., Williams, R. & Hilton, G. 2003. Levels of Cadmium and Zinc in soil and plants following the toxic spill of a pyrite mine (Aznalcollar, Spain). Ambio 32(1): 52-57.

Prakash, V., Pain, D.J., Cunningham, A.A., Donald, P.F., Prakash, N., Verma, A., Gargi, R., Sivakumar, S., Rahmani, A. R. 2003. Catastrophic collapse of Indian white-backed Gyps bengalensis and long-billed Gyps indicus vulture populations. Biological Conservation 109(3): 381-390.

Pain, D.J., Cunningham, A.A., Donald, P. F., Duckworth, J. W., Houston, D. C., Katzner, T., Parry-Jones, J., Poole, C., Prakash, V., Round, P., Timmins, R. 2003. Gyps vulture declines in Asia; temporospatial trends, causes and impacts.   Conserv. Biol. 17 (3): 661-671.

Pattee, O.H. and Pain, D.J. 2003 Lead in the Environment. P373-408 Chapter 15 in Hoffman, D.J., Rattner, B.A., Allen Burton, G. Jr. & Cairns, J. Jr. Handbook of Ecotoxicology. Lewis Publishers USA. 

Jeganathan, P., Green, R.E., Bowden, C.G.R., Norris, K., Pain, D & Rahmani, A. 2002. Use of tracking strips and automatic cameras for detecting Critically Endangered Jerdon’s coursers Rhinoptilus bitorquatus in scrub jungle in Andhra Pradesh, India. Oryx 36(2):182-188. 

Meharg, A.A., Pain, D.J., Ellam, R.M., Baos, R., Olive, V., Joyson, A., Powell, N., Green, A.J., & Hiraldo, F. 2002. Isotopic identification of the sources of lead contamination for white storks (Ciconia ciconia) in a marshland ecosystem (Doñana, S.W. Spain). Sci. Total Environ. 300: 81-86.

Donald, P.D., Pisano, G., Rayment, M.D. & Pain, D.J. 2002. The Common Agricultural Policy, EU enlargement and the conservation of Europe’s farmland birds. Agricul. Ecosyst. Environ. , 89: 167-179.

Pain, D.J. & Donald, P.F. 2002. Outside the Reserve: pandemic threats to bird biodiversity. Chapter 8 in: Norris, K & Pain, D.J. (Eds). 2002. Conserving Bird Biodiversity: General Principles and their Application. Cambridge University Press, Conservation Biology Series.

Pain, D.J., Cunningham., A.A., Prakash, V. & Ghalsasi, G.R.  2002. Vulture declines in India: patterns, causes and spread. In Proceedings of a workshop on Asian vulture declines at the NAOC, New Orleans, September 2002.

Sim, I.M.W, Cross, A.V., Lamacraft D.L & Pain, D.J.  2001. Correlates of Common Buzzard Buteo buteo density and breeding success in the West Midlands. Bird Study (48): 317-329

Cunningham, A. A., Prakash, V., Ghalsasi, G.R. & Pain, D. 2001. Investigating the Cause of Catastrophic Declines in Asian Griffon Vultures (Gyps indicus and G. bengalensis).  Reports from the workshop on Indian Gyps vultures. Katzner, T. & Parry-Jones, J. (eds.) 4th Eurasian Congress on Raptors, p 133, Seville, Spain. Estación Biológica Donaña, Raptor Research Foundation. 

Pain, D.J. 2001. Conservation of critically endangered Gyps vultures in India. In: Bennun & Virani (eds). Responding to the Asian vulture crisis: Planning for vulture monitoring and conservation in Kenya. Proceedings and recommendations of a seminar and workshop held at the National Museums of Kenya, 10 May 2001. NMK Ornithology Research Report 41: June 2001.

Pain, D.J., Rose, L. & Castro, F. 2001. The Doñana Disaster. RSPB Cons. Rev. No 13.

Sim, I.M.W., Campbell, L., Pain, D.J. & Wilson, J.D. 2000. Correlates of the population increase of common buzzard Buteo buteo in the West Midlands between 1983 and 1996. Bird Study 47:154-164.

Pain, D.J., Brooke, M, Finnie, J. & Jackson, A., 2000. A. Effects of brodifacoum on the land crab on Ascension Island. J. Wildl. Manage. 64(2):380-387.

Pain, D.J., Bavoux, C. & Burneleau, G. & Wyatt, C. 1999.  Levels of Polychlorinated Biphenyls, Organochlorine Pesticides, Mercury and Lead in relation to shell thickness in Marsh harriers (Circus aeruginosus) eggs from Charente-Maritime, France. Environ. Pollut.104: 61-68.

Meharg, A.A., Osborn, D, Pain, D.J., Sanchez, A. & Naveso, M.1999. A challenge to ecotoxicology and environmental chemistry: The Doñana mining catastrophe. Letter to the Editor:  Environ. Toxicol. Chem.  18: 811-812.

Meharg, A.A., Osborn, D., Pain, D.J, Sánchez, A. & Naveso, M.A. 1999.  Contamination of Doñana food-chains after the Aznalcóllar mine disaster. Environ. Pollut. 105(3): 387-390.

D.W. Gibbons, K.W. Smith, P. Atkinson, D.J. Pain, W.J. Arendt, G. Grey, J. Hartley, A. Owen, C., Clubbe 1999. Montserrat oriole: island endemic threatened by volcanic eruption. RSPB Conserv. Rev. No 12 p97-101.

Pain, D., A. Sánchez, A.A. Meharg 1998. The Doñana Ecological Disaster: contamination of World Heritage estuarine marsh ecosystem with acidified pyrite mine waste. Sci. Total Environ. 222: 45-54.

Pain, D.J. & Pienkowski, M.W. 1998. Some main themes and questions arising from “Farming and Birds in Europe”. In Proceedings of the Fifth meeting on the European Forum on Nature Conservation and Pastoralism, 18-21st September 1996, Cogne, Italy. pages 152-160.

Pain, D.J., Bavoux, C. & Burneleau, G. 1997. Seasonal blood lead concentrations in marsh harriers Circus aeruginosus from Charente-Maritime, France: relationship with the hunting season. Biological Conservation 81:1-7.

Pain, D.J., Hill, D. & McCracken, D.I. 1997 Impact of agricultural intensification of pastoral systems on bird distribution in Britain 1970-1990. Agric. Ecosyst. Environ. 64:19-32.

Pain, D.J. & Dixon, J. 1997. Why farming and Birds? Chapter 1, pp 1-24, In: Pain, D.J. & Pienkowski, M.W. (eds) Farming and Bird in Europe: the Common Agricultural Policy and its Implications for Bird Conservation. Academic Press, London. 

Pain, D.J. & Pienkowski, M.W. 1997. A future for farming and Birds? Chapter 13, pages 358-388, In Pain, D.J. & Pienkowski, M.W. (eds) Farming and Bird in Europe: the Common Agricultural Policy and its Implications for Bird Conservation. Academic Press, London. 

Batty, J., Pain, D.J., & Caurant, F. 1996. Metal concentrations in eels (Anguilla anguilla) from the Camargue region of France. Biol. Conserv. 76: 17-23.

Pain, D.J. 1996. Lead in waterfowl. Chapter 10 pages 251-264 In: W.N. Beyer, G.H. Heinz and A.W. Redmon (Eds.) Environmental Contaminants in Wildlife: interpreting tissue concentrations. SETAC CRC Lewis Publishers, Boca Raton.

Siva-Jothy, M.T., Gibbons, D. W., & Pain, D. 1995. Female oviposition-site preference and egg hatching success in the damselfly Calopteryx splendens xanthostoma. Behav. Ecol. Sociobiol.37: 39-44.

Pain, D.J., Sears, J., & Newton, I. 1995. Lead concentrations in birds of prey in Britain. Environ. Pollut. 87(2):173180.

Pain, D.J. 1995. Lead in the environment. Chapter 16, pages 356-391 In: Hoffman, D.J., Rattner, B.A., Allen Burton, G. Jr. & Cairns, J. Jr. Handbook of Ecotoxicology. CRC Press, Lewis Publishers USA. 755p

Pain, D.J. & Dunn, E. 1995. The effects of agricultural intensification upon pastoral birds: lowland wet grasslands (the Netherlands) and transhumance farming (Spain). pp. 90-98 In: McCracken, D.I. & Bignal, E. Farming on the edge: the nature of traditional farmland in Europe. JNCC, Peterborough 1995. 216p.

Pain, D.J. & Amiard-Triquet, C. 1993. Lead poisoning of raptors in France and elsewhere.  Ecotox. Environ. Safety 25(2):183-192.

Pain, D.J., Amiard-Triquet, C., Bavoux, C., Burneleau, G., Eon, L., & Nicolau-Guillaumet, P. 1993. Lead poisoning in wild populations of marsh harriers Circus aeruginosus in the Camargue and Charente-Maritime, France. Ibis 135:379-386.

Pain, D.J., Metayer, C. & Amiard, J-C. 1993. Lead determination in avian blood: application to a study of lead contamination in raptors from France. Int. J. Environ. Analyt. Chem. 53:29-35.

Pain, D.J. & Eon, L. 1993. Methods of investigating the presence of ingested lead shot in waterfowl: an improved visual technique. Wildfowl 44:184-187.

Pain, D.J., Amiard-Triquet, C. & Sylvestre, C. 1992. Tissue lead concentrations and shot ingestion in nine species of waterbirds from the Camargue, France. Ecotox. Environ. Safety. 24:217-233.

Gibbons, D.W. & Pain, D.J. 1992. The influence of river flowrate on the breeding behaviour of Calopteryx damselflies. J. Anim. Ecol. 61: 238-289.

Amiard-Triquet, C., Pain, D.J., Mauvais, G. & Pinault, L. 1992. Lead poisoning in waterfowl: field and experimental data. Book Chapter 25pp. in: Vernet, J.P. (ed.) Impact of heavy metals on the environment. Elsevier Science Publishers, North Holland.

Pain, D.J. 1992 (Ed.). Lead poisoning in waterfowl. Proceedings of an IWRB Workshop, Brussels, Belgium. June 1991. IWRB Special Publication No 16. Slimbridge, UK. 105p.

Pain, D.J. 1992. Lead poisoning in birds: a Southern European perspective. Pages 109-114 in: Finlayson, C.M., Hollis, G.E., and Davis, T.J. (eds.) Managing Mediterranean wetlands and their birds. Proc. Symp. Grado, Italy, 1991. IWRB Special Pub.No 20. Slimbridge, UK. 285pp.

Pain, D.J. 1992. Lead poisoning in waterfowl: a review. pages 7-14 in: Pain, D.J. 1992 (ed.). Lead poisoning in waterfowl. Proceedings of an IWRB Workshop, Brussels, Belgium. June 1991. IWRB Special Publication No 16. Slimbridge, UK. 105p.

Pain, D.J. 1992. Lead poisoning in waterfowl: National reports. pages 86-94 in: Pain, D.J. 1992 (ed.). Lead poisoning in waterfowl. Proceedings of an IWRB Workshop, Brussels, B  elgium. June 1991. IWRB Special Publication No 16. Slimbridge, UK. 105p.

Pain, D.J. 1991. Lead shot densities and settlement rates in Camargue marshes, France. Biol. Conserv. 57:273-286.

Pain, D.J. 1991. L'intoxication saturnine de l'avifaune; une synthese des travaux Francais. Gibier Faune Sauvage. 8:79-92.

Amiard-Triquet, C., Pain, D.J. & Delves, H.T. 1991. Exposure to trace elements of flamingos living in a biosphere reserve, the Camargue, France. Environ. Pollut. 69:193-201.

Pain, D.J. 1991. Why are lead poisoned waterfowl rarely seen? The disappearance of waterfowl carcasses in the Camargue, France. Wildfowl. 42:118-122.

Pain, D.J. 1991. Lead poisoning in birds: an international perspective. Acta XX Congressus Internationalis Ornithologici. IV. 2343-2352.

Amiard-Triquet, C., Pain, D.J. & Sylvestre, C. 1991. Lead contamination in nine species of waterbirds living in the Rhone river delta, France. P. 179-182. In: Proc. of the 8th International Conference of Heavy Metals in the Environment.  Sept. 16-20th 1991. Edinburgh, Scotland.  Vol. 2:179-182.

Pain, D.J. 1990. Lead shot ingestion by waterbirds in the Camargue, France: an investigation of levels and interspecific differences. Environ. Pollut. 66:273-285.

Pain, D.J. & Handrinos, G.I. 1990. The incidence of ingested lead shot in ducks of the Evros delta, Greece. Wildfowl. 41:167-170.

Pain, D.J. 1989. Haematological parameters as predictors of blood lead and indicators of lead poisoning in the black duck Anas rubripes. Environ. Pollut. 60:67-81.

Pain D.J. 1989. Lead poisoning in waterfowl: a review. In: G. Matthews (ed.). Proceedings of the IWRB Symposium on Managing Waterfowl Populations. Astrakhan, USSR, 29 Sept.-9 October 1989. 230p.

Pain, D.J. & Rattner, B.A. 1988. Hematology and mortality of black duck Anas rubripes after the ingestion of one number four lead shot. Bull. Environ. Contam. Toxicol. 40:159-164.




Donald, P.F., Collar, N. J., Marsden, S.J.  & Pain, D.J. 2013. Facing Extinction: The world’s rarest birds and the race to save them. Second Edition (updated). Christopher Helm, Bloomsbury Publishing plc., London pp 320.

Donald, P.F., Collar, N. J., Marsden, S.J.  & Pain, D.J. 2010. Facing Extinction: The world’s rarest birds and the race to save them. T & A.D. Poyser., London pp 312.

Norris, K & Pain, D.J. (Eds.). 2002. Conserving Bird Biodiversity: General Principles and their Application. Cambridge University Press, Conservation Biology 7. Cambridge University Press.  pp 337.

Pain, D.J. & Pienkowski, M.W. (Eds.) 1997. Farming and Bird in Europe: the Common Agricultural Policy and its Implications for Bird Conservation. Academic Press, London. pp 436.



Expert groups


Lead Ammunition Group 2018. Lead from Ammunition: Risks to Wildlife, Captive Animals and People and Risk Mitigation: Update Report Executive Summary. Prepared by D Pain on behalf of the LAG. 12pp. Available at:

Green, R. E. & Pain, D. J. 2015. An evaluation of the risks to human health in the UK from lead derived from ammunition. In: Lead Ammunition Group (2015). Lead Ammunition, Wildlife and Human Health. A report prepared for the Department for Environment, Food and Rural Affairs and the Food Standards Agency in the United Kingdom. Appendix 1, 93-148.  

Pain, D. J.  & Green, R. E. 2015. An evaluation of the risks to wildlife in the UK from lead derived from ammunition. In: Lead Ammunition Group (2015). Lead Ammunition, Wildlife and Human Health. A report prepared for the Department for Environment, Food and Rural Affairs and the Food Standards Agency in the United Kingdom. Appendix 1, 263-382.





Anderson, O., ….. Pain, D.J. et al. 2020. Co-author and co-signatory. EUROPEAN SCIENTISTS’ OPEN LETTER ON THE RISKS OF LEAD AMMUNITION ECHA proposal under REACH Regulations[1] Restriction proposal on the use of lead gunshot in or over wetlands. August 2020.

Andersen, O., ……. Pain, D. et al… 2018.  Co-author and co-signatory. European Scientists’ Open Letter on the Risks of Lead Ammunition. ECHA PROPOSAL UNDER REACH REGULATIONS[1] Restriction proposal on the use of lead gunshot in or over wetlands.




Pain, D., Green, R. and Taylor, C. 2020. Children are at risk from game shot with lead. Guardian Letters 10 August 2020.

Boobis, A.R., Browne, D., Cromie, R., Green, R.E., Krebs, J., Newton, I., Pain, D., Perrins, C., Stroud, D. & Swift, J. 2019. Use safe alternative to toxic lead gunshot.

Guardian Letters Mon 29 Jul 2019 14.11 BST


Honorary Research Fellow