Our Publications


Pichegru, L., Nyengera, R., Mcinnes, A. M., & Pistorius, P. (2017) Avoidance of seismic survey activities by penguins. Scientific Reports 7 (16305).

Spelt, A. & Pichegru, L. (2017). Sex allocation and sex-specific parental investment in an endangered seabird. IBIS: International journal of avian science 159 (2) 272-284.


Pichegru, L., Edwards, T. B., Dilley, B. J. & Flower, T. P. (2016) African penguin tolerance to humans depends on historical exposure at colony level. Bird Conservation International 26 (3) 307-322.


Grémillet, D., Péron, C., Kato, A,, Amélineau, F,. Ropert-Coudert, Y., Ryan, P.G. & Pichegru, L. (in press) Starving seabirds: Unprofitable foraging and its fitness consequences in Cape gannets competing with fisheries in the Benguela upwelling ecosystem. Marine Biology

van Eeden, R.B., Reid, T., Ryan, P.G. & Pichegru, L. (in press) Fine-scale foraging cues for African Penguins in a highly variable marine environment. Marine Ecology Progress Series

Pichegru, L., Edwards, T., Dilley, B., Flower, T.J. & Ryan, P.G. (in press) African penguin tolerance to humans depends on historical exposure at colony level.  Bird Conservation International

McInnes, A.M., Khoosal, A., Murrell, B., Merkle, D., Lacerda, M., Nyengera, R., Coetzee, J.C., Edwards, L.C., Ryan, P.G., Rademan, J., van der Westhuizen, J.J. & Pichegru, L. (in press) Recreational fish-finders – an inexpensive alternative to scientific echo-sounders for unravelling the links between marine top predators and their prey. PLoS ONE.

Pettex, E., Barrett, R.T., Lorentsen, S-H., Bonadonna, F., Pichegru, L., Pons, J-B. & Grémillet, D. (2015). Contrasting population trends at Norwegian Northern gannet colonies: is food limitation a factor? Journal of Ornithology 156: 397-406.


Cohen, L.A., Pichegru, L., Grémillet, D., Coetzee, J.C., Upfold, L. & Ryan, P.G. (2014) Behavioural flexibility of a seabird facing decadal fluctuations in prey availability. Marine Ecology Progress Series 505, 281-293.

Lei, B., Green, J.D. & Pichegru, L. (2014) Extreme microclimate conditions in artificial nests for endangered African penguins. Bird Conservation International 24, 201-213.

McInnes, A.M., Suarez, N., Rishworth, G.M., Green, D.B., Pistorius, P.A. & Pichegru, L. (2014). Lumbering the gauntlet – Cape gannet fledglings killed by African penguins. Ostrich 85:193-196.

Pichegru, L. & Parsons, N. (2014) Female biased mortality in Endangered penguins. African Journal of Marine Science 36: 279-282.

Sabarros, P.S., Grémillet, D., Demarcq, H., Moseley, C., Mullers, R.H.E., Pichegru, L., Stenseth, N.C. & Machu, E. (2014) Fine-scale recognition and use of submesoscale fronts by Cape gannets in the southern Benguela region. Deep Sea Research II 107:77-84.


Tew-Kai, E., van der Lingen, C.D., Coetzee, J., Benhamou, S., Pichegru, L., Ryan, P.G. & Grémillet, D. (2013) Are fishery wastes an ecological trap for Cape gannets in the Benguela upwelling ecosystem? A multi-disciplinary approach. Journal of Applied Ecology 50. 659-670.

Pichegru, L. (2013) Increasing breeding success of an Endangered penguin: artificial nests or culling predatory gulls? Bird Conservation International 23, 296-308.

Pichegru, L., Ryan, P.G., van Eeden, R., Reid, T., Grémillet, D. and Wanless, R. (2012) Industrial fishing, no-take zones and endangered penguins. Biological Conservation 156, 117-125.

Pichegru, L., Cook, T., Handley, J., Voogt, N., Nupen, L., Watermeyer, J. & McQuaid, C.D. (2013) Sex-specific foraging behaviour and a field sexing technique for Endangered African penguin. Endangangered Species Research 19, 255-264.


Pichegru, L., Ryan, P.G., van Eeden, R., Reid, T., Grémillet, D. and Wanless, R. (2012) Industrial fishing, no-take zones and endangered penguins. Biological Conservation 156:117-125.

Pichegru, L. (2012) Increasing breeding success of an Endangered penguin: artificial nests or culling predatory gulls ? Bird Conservation International doi:10.1017/S0959270912000135.

Hamann, M., Grémillet, D., Ryan, P.G., Bonadonna, F., van der Lingen, C.D. and Pichegru, L. (2012). A hard-knock life: the foraging ecology of Cape cormorants amidst shifting prey resources and industrial fishing pressure. African Journal of Marine Science 34, 233-240.

Moseley, C., Grémillet, D., Connan, M., Ryan, P.G., Mullers, R.H.E., van der Lingen, C.D., Miller, T.W., Coetzee, J.C., Crawford, R.J.M., Sabarros, P., McQuaid, C.D. and Pichegru, L. (2012) Body condition and foraging ecology of two populations of seabirds historically feeding on different diet. Journal of Experimental Marine Biology and Ecology 422-423, 29-38.

Ryan, P.G., Edwards, L.C. and Pichegru, L. (2012) African penguins Spheniscus demersus, bait balls and the Allee effect. Ardea 100, 89-94.

Cook, T.C., Hamann, M., Pichegru, L., Bonadonna, F., Grémillet, D. and Ryan, P.G. (2012) When seabirds hit the bottom: investigating fine-scale four-dimensional foraging behaviour in small flighted divers. Marine Biology. 159, 373–387.


Write, K.L.B.,Pichegru, L. and Ryan, P.G. (2011) Penguins are attracted to dimethyl sulphide at sea. Journal of Experimental Biology. 214, 2509-2511.

Crawford, R.J.M,, Altwegg, R., Barham, B.J., Barham, P.J., Durant, J.M., Dyer, B.M., Makhado, A.B., Pichegru, L., Ryan, P.G., Underhill, L.G., Upfold, L., Visagie. J,, Waller, L.J. and Whittington, P.A. (2011) Collapse of South Africa’s penguins in the early 21st century: a consideration of food availability. African Journal of Marine Science 33, 139-156.

Pichegru, L., Ropert-Coudert, Y., Kato, A., Takahashi, A., Dyer, B.M. and Ryan, P.G. (2011) Diving patterns of female Macaroni Penguins breeding on Marion Island, South Africa. Polar Biology 34, 945-954.


Grémillet, D., Mullers, R.H.E., Moseley, C., Pichegru, L., Coetzee, J.C., Sabarros, S., van der Lingen, C.D., Ryan, P.G., Kato, A. and Ropert-Coudert, Y. (2010) Seabirds, fisheries and cameras. Frontiers for Ecology 8, 401-402.

Pichegru, L., Grémillet, D., Crawford, R.J.M. and Ryan P.G. (2010) Marine no-take zone rapidly benefit threatened penguin. Biology Letters 6, 498-501.

Ryan, P.G.,Pichegru, L., Ropert-Coudert, Y., Grémillet, D, and Kato, A. (2010) On a wing and a prayer: the foraging ecology of breeding Cape Cormorants. Journal of Zoology 280, 25-32.

Pichegru, L., Ryan, P.G., Crawford, R.J.M. and Grémillet, D. (2010) Behavioural inertia places a top marine predator at risk from environmental change in the Benguela upwelling system. Marine Biology 157, 537-544.


Pichegru, L., Ryan, P.G., Le Bohec, C., van der Lingen, C.D., Navarro, R., Petersen, S., Lewis, S., van der Westhuizen, J. and Grémillet, D. (2009) Overlap between vulnerable top predators and fisheries in the Benguela upwelling system: implications for marine protected areas. Marine and Ecological Progress Series 391, 199-208.

Okes, N.C., Hockey, P.A.R., Pichegru, L., van der Lingen, C.D., Crawford, R.J.M. and Grémillet, D. (2009) Competition for shifting resources in the southern Benguela upwelling: seabirds versus purse-seine fisheries. Biological Conservation 142, 2361-2368.

Grémillet, D., Pichegru, L., Kuntz, G., Woakes, A.G., Wilkinson, S., Crawford, R.J.M. and Ryan, P.G. (2008). A Junk-food hypothesis for Cape gannets feeding on fishery waste. Proceedings of the Royal Society of London, 18, 1-9.

Pichegru, L., Ryan, P.G., van den Lingen, C.D., Ropert-Coudert, Y., Coetzee, J. and Grémillet, D. (2007) Foraging strategies and energetics of Cape gannets Morus capensis feeding on live prey and fishery discards in the Benguela upwelling system. Marine and Ecological Progress Series 350, 127-136.

Grémillet, D., Pichegru, L., Siorat, F. and Georges, J-Y. (2006). Conservation implications of the mismatch between population dynamics and foraging effort in Northern gannets from the English Channel. Marine and Ecological Progress Series319, 15-25.

Massemin-Challet, S., Gendner, J-P., Samtmann, S., Pichegru, L., Wulgué, A. and Le Maho, Y. (2006) The effect of migration strategy and food availability on White Stork Ciconia ciconia breeding success. Ibis 148, 503-508.