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Marine Apex Predator Research Unit


Recent (2014 onwards) publications by MAPRU members.


In press

Parsons NJ, Gous TA, Schaeffer AM, Vanstreels RET. In press. Health evaluation of African penguins (Speniscus demersus) in southern Africa. Onderstepoort Journal of Veterinary Research.

Grilo ML, Vanstreels RET, Wallace R, Garcia-Parraga D, Braga EM, Chitty J, Catao-Dias JL, Carvalho LM. In press. Malaria in penguins: current perceptions. Avian Pathology.

Koper RP, Karczmarski L, du Preez D, Plön S. In press. Sixteen years later: Occurrence, group size, and habitat use of humpback dolphins (Sousa plumbea) in Algoa Bay, South Africa. Marine Mammal Science.

Lingham-Soliar T. In press. Experiments on ostrich decomposition and opisthotonus with implications for theropod dinosaurs.  Journal of Zoology.

Pichegru L, Edwards T, Dilley B, Flower TJ, Ryan PG. In press. African penguin tolerance to humans depends on historical exposure at colony level. Bird Conservation International.

Reisinger RR, Gröcke DR, Lübcker N, McClymont EL, Hoelzel AR, de Bruyn PJN. In press. Variation in the diet of killer whales (Orcinus orca) at Marion Island, Southern Ocean. Marine Ecology Progress Series.

Towner AV, Leos-Barajas V, Langrock R, Schick RS, Smale MJ, Kasche T, Jewell OJD, Papamastrio YP. In press. Sex-specific and individual preferences for hunting strategies in white sharks. Functional Ecology.

Turnbull LA, Ozgul A, Accouche W, Baxter R, ChongSeng L, Currie J, Doak N, Hansen D, Pistorius P, Postma E, Richards H, Schaepman-Strub G, van de Crommenacker J, von Brandis R, Fleischer-Dogley F, Bunbury N. In press.  Distinctive subpopulations of giant tortoises on Aldabra coral atoll are stable through time and space. Ecology and Evolution.

Vanstreels RET, Braga EM, Catao-Dias JL. In press. Blood parasites of penguins: a critical review. Parasitology.

Vanstreels RET, Capellino F, Silveira P, Braga EM, Heredia SAR, Loureiro J, Catao-Dias JL. In press. Avian malaria (Plasmodium spp.) in captive Magellanic penguins (Spheniscus magellanicus) from northern Argentina. Journal of Wildlife Diseases.

Wege M, Etienne M-P, Oosthuizen WC, Reisinger RR, Bester MN, de Bruyn PJN. In press. Trend changes in sympatric Subantarctic and Antarctic fur seal pup populations at Marion Island, Southern Ocean. Marine Mammal Science.



Danckwerts DK, McQuaid CD, Connan M, Smale MJ, Le Corre M, Humeau L, Kaehler S, Juhasz CC, Orlowski S, Tourmetz J, Jaquemet S. 2016. Intra-annual variation in the foraging ecology of the endangered endemic Barau’s Petrel (Pterodroma baraui) from Réunion Island, south-western Indian Ocean: insights from a multifaceted approach. Marine Biology 163(1): Article 18.

Grémillet D, Péron C, Kato A, Amélineau F, Ropert-Coudert Y, Ryan PG, Pichegru L. 2016. Starving seabirds: Unprofitable foraging and its fitness consequences in Cape gannets competing with fisheries in the Benguela upwelling ecosystem. Marine Biology 163: 35-46.

Handley J, Baylis AMM, Brickel P, Pistorius PA. 2016. Temporal variation in the diet of Gentoo penguins at the Falkland Islands. Polar Biology 39(2): 283-296.

Handley J, Pistorius PA. 2016. Kleptoparasitism in foraging Gentoo penguins. Polar Biology 39(2): 391-395.

Sanchez-Sarmiento AM, Rossi S, Vilca FZ, Vanstreels RET, Monteiro SA, Vale LAS, Santos RG, Marigo J, Bertozzi CP, Grisi Filho JHH, Tornisielo VL, Matishuma ER. 2016. Organochlorine pesticides in green sea turtles (Chelonia mydas) with and without fibropapillomatosis caught at three feeding areas off Brazil. Journal of the Marine Biological Association of the United Kingdom 97: 1-9.

Thiebault A, Semeria M, Lett C, Tremblay Y. 2016. How to capture fish in a school? Effect of successive predator attacks on seabird feeding success. Journal of Animal Ecology 85(1): 157–167.

Van Eeden RB, Reid T, Ryan PG, Pichegru L. 2016. Foraging cues for African Penguins on a fine spatial-temporal scale in a thermally dynamic marine environment. Marine Ecology Progress Series 543: 257-271.



Baylis A, Orben R, Pistorius P, Brickle P, Staniland I, Ratcliffe N. 2015. Winter foraging site fidelity in king penguins at the Falkland Islands. Marine Biology 162(1): 99-110.

Cabana AL, Vanstreels RET, Xavier MO, Osorio LG, Adornes AC, Leite ATM, Soares MP, Silva-Filho RP, Catao-Dias JL, Meireles MCA. 2015. Lethal concurrent avian malaria and aspergillosis in Magellanic Penguin (Spheniscus magellanicus). Boletín Chileno de Ornitología 20: 28-32.

Connan M, Teske PR, Tree AJ, Whittington PA, McQuaid CD. 2015. The subspecies of Antarctic terns (Sterna vittata) wintering on the South African coast: evidence from morphology, genetics and stable isotopes. Emu: Austral Ornithology 115(3): 223-236.

Gonzales-Viera O, Ruoppolo V, Marigo J, Carvalho VM, Bertozzi CP, Takakura C, Namiyama G, Vanstreels RET, Catao-Dias JL. 2015. Renal lesions in cetaceans from Brazil. Journal of Comparative Pathology 152: 1-10.

Goschen WS, Bornman TG, Deyzel SHP, Schumann EH. 2015. Coastal upwelling on the far eastern Agulhas Bank associated with large meanders in the Agulhas Current. Continental Shelf Research 101: 34-46.

Green D, Coetzee J, Rishworth GM, Pistorius PA. 2015.  Foraging distribution of Cape gannets in relation to oceanographic features, prey availability and marine protected areas in the Eastern Cape, South Africa. Marine Ecology Progress Series 537: 277-88.

Green D, Klages NTW, Crawford RJM, Coetzee JC, Dyer BM, Rishworth GM, Pistorius PA. 2015. Dietary change in Cape gannets reflects distributional and demographic shifts in two South African commercial fish stocks. ICES Journal of Marine Science 72(3): 771-781.

Hurtado R, Fabrizio T, Vanstreels RET, Krauss S, Webby RJ, Webster RG, Durigon EL. 2015. Molecular Characterization of Subtype H11N9 Avian Influenza Virus Isolated from Shorebirds in Brazil. Plos One 10: e0145627.

Hurtado R, Vanstreels RET, Serafini PP, Azevedo Junior SM, Durigon EL. 2015. Aquatic wild birds kept captive at remote coastal communities in northern Brazil. Boletín Chileno de Ornitología 20: 17-22.

Kessel ST, Hussey NE, Webber DM, Gruber SH, Young JM, Smale MJ, Fisk A.  2015. Close proximity detection interference with acoustic telemetry: the importance of considering tag power output in low ambient noise environments. Animal Biotelemetry 3:5.

Lingham-Soliar T. 2015. The Vertebrate Integument Volume 2, Structure, Design and Function. Springer, Heidelberg.

Lingham-Soliar T. 2015. Swimming and Flying in Vertebrates (Fluid Mechanics). In: The Vertebrate Integument Volume 2. Springer, Heidelberg, pp 1-60.

Lingham-Soliar T. 2015. Vertebrate Swimming. In: The Vertebrate Integument Volume 2. Springer, Heidelberg, pp 61-103.

Lingham-Soliar T. 2015. Birds. In: The Vertebrate Integument Volume 2. Springer, Heidelberg, pp 163-262.

Lingham-Soliar T. 2015. Three stages of post mortem opisthotonus uniquely captured in the dinosaur Sinosauropteryx. Journal of Ornithology 156: 257-262.

Lowther AD, Ahonen H, Hofmeyr GJG, Oosthuizen C, de Bruyn PJN, Lydersen C, Kovacs KM. 2015. The reliability of VHF telemetry data for measuring onshore attendance patterns of marine predators: a comparison with Time Depth Recorder data. Marine Ecology Progress Series 538: 249-256.

McInnes AM, Khoosal A, Murrell B, Merkle D, Lacerda M, Nyengera R, Coetzee JC, Edwards LC, Ryan PG, Rademan J, van der Westhuizen JJ, Pichegru L. 2015. Recreational fish-finders - an inexpensive alternative to scientific echo-sounders for unravelling the links between marine top predators and their prey. PLoS ONE 10: e0140936

Moura AE, Kenny JG, Chaudhuri RR, Hughes MA, Reisinger RR, de Bruyn PJN, Dahlheim ME, Hall N, Hoelzel AR. 2015. Phylogenomics of the killer whale indicates ecotype divergence in sympatry. Heredity 114: 48-55.

Moura AE, Kenny JG, Chaudhuri RR, Hughes MA, Welch AJ, Reisinger RR, de Bruyn PJN, Dahlheim ME, Hall N, Hoelzel AR. 2015. Population genomics of the killer whale indicates ecotype evolution in sympatry involving both selection and drift. Molecular Ecology 23: 5179-5192.

Niemandt C, Kovacs KM, Lydersen C, Dyer BM, Isaksen K, Hofmeyr GJG, de Bruyn PJN. 2016. Chinstrap and macaroni penguin diet and demography at Nyrøysa, Bouvetøya, Southern Ocean. Antarctic Science 28: 91-100.

Parsons NJ, Gous TA, Van Wilpe E, Strauss V, Vanstreels RET. 2015. Herpesvirus-like respiratory infection in African penguins Spheniscus demersus admitted to a rehabilitation centre. Diseases of Aquatic Organisms 116: 149-155.

Pettex E, Barrett RT, 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.

Pistorius PA, Hindell MA, Tremblay Y, Rishworth GM. 2015. Weathering a dynamic seascape: influences of wind and rain on a seabird’s year-round activity budget. PLoS ONE 10(11): e0142623.

Plön S, Cockcroft VG, Froneman PW. 2015. The natural history and conservation of Indian Ocean humpback dolphins (Sousa plumbea) in South African waters. In: Thomas Jefferson, Barbara E. Curry (eds): Humpback Dolphins (Sousa spp.): Current Status and Conservation, Advances in Marine Biology, Vol 72, Part 1, Academic Press, UK, pp. 143-162

Plön S, De Wet, Lane E, Wohlsein P, Siebert U, Thompson P. 2015. The importance of a standardized necropsy protocol for health investigations of small cetaceans in South Africa. African Journal of Wildlife Research 45 (3):332-341.

Postma M, Tordiffe ASW, Hofmeyr MS, Reisinger RR, Bester LC, Buss PE, de Bruyn PJN. 2015. Terrestrial mammal three-dimensional photogrammetry: multispecies mass estimation. Ecosphere 6(12): 293.

Reisinger RR, Keith M, Andrews RD, de Bruyn PJN. 2015. Movement and diving of killer whales (Orcinus orca) at a Southern Ocean archipelago. Journal of Experimental Marine Biology and Ecology 473: 90-102.

Rishworth GM, Pistorius PA. 2015. Protection and provisioning: the role of parental behaviour in terms of chick growth and survival in a pelagic seabird. Marine Ecology Progress Series 530: 153-162.

Smale MJ, Dicken ML, Booth AJ. 2015. Seasonality, behaviour and philopatry of raggedtooth sharks, Carcharias taurus in Eastern Cape nursery areas, South Africa. African Journal of Marine Science 37: 219-231.

Van der Vyver JSF, De Bruyn PJN, Lipinski MR, Leslie RW. 2015. First record of a porbeagle shark Lamna nasus (Bonnaterre, 1788) stranding at sub-Antarctic Marion Island. Marine Biodiversity Records 8: 1-5.

Van der Vyver JSF, Sauer WHH, McKeown NJ, Yemane D, Shaw PW, Lipinski MR. 2015. Phenotypic divergence despite high gene flow in chokka squid Loligo reynaudi (Cephalopoda: Loligonidae): implications for fishery management. Marine Biological Association of the United Kingdom 1-9.

Vanstreels RET, Silva-Filho RP, Kolesnikovas CKM, Bhering RCC, Ruoppolo V, Epiphanio S, Amaku M, Ferreira Junior FC, Braga EM, Catao-Dias JL. 2015. Epidemiology and pathology of avian malaria in penguins undergoing rehabilitation in Brazil. Veterinary Research 46: 30.

Vanstreels RET, Woehler EJ, Ruoppolo V, Vertigan P,Carlile N, Priddel D, Finger A, Dann P, Herrin KV, Thompson P, Ferreira Junior FC, Braga EM, Hurtado R, Epiphanio S, Catao-Dias JL. 2015. Epidemiology and molecular phylogeny of Babesia sp. in Little Penguins Eudyptula minor in Australia. International Journal of Parasitology: Parasites and Wildlife 4: 198-205.

Whitehead O, Rollinson DP, Reisinger RR. 2015. Pseudostalked barnacles Xenobalanus globicipitis attached to killer whales Orcinus orca in South African waters. Marine Biodiversity 45: 873-876.

Whittington PA, Tree AJ, Connan M, Watkins EG. 2015. The status of the Damara tern in the Eastern Cape. Ostrich: Journal of African Ornithology 86(1-2): 65-73.

Windberger UB, Auer R, Smale MJ. 2015. Blood rheology in Cape fur seals and bottlenose dolphins: Implications for muscle perfusion. Biodiversity & Endangered Species 3:1.



Best PB, Smale MJ, Glass J, Herian K, von der Heyden S. 2014. Identification of stomach contents from a Shepherd’s beaked whale Tasmacetus shepherdi. Journal of the Marine Biological Association of the United Kingdom. 94(6): 1093-1097.

Best PB, Meÿer MA, Thornton M, Kotze PGH, Seakamela SM, Hofmeyr GJG, Wintner S, Weland CD, Steinke D. 2014. Confirmation of the occurrence of a second killer whale morphotype in South African waters. African Journal of Marine Science 36: 215-224.

Bizelli CC, Silva SR, Costa JD, Vanstreels RET, Atzingen M, Santoro ML, Fernandes I, Catao-Dias JL, Faquim-Mauro EL. 2014. Isolation and characterization of IgM and IgG antibodies from plasma of Magellanic penguins (Spheniscus magellanicus). Avian Diseases 59: 1-10.

Blanchet M-A, Lydersen C, Biuw M, de Bruyn PJN, Hofmeyr GJG, Krafft BA, Kovacs KM. 2014. Instrumentation and handling effects on Antarctic fur seals (Arctocephalus gazella). Polar Research 33: 21630.

Bouveroux Th., Kiszka J, Heithaus MR, Jauniaux T, Pezeril S. 2014. Direct evidence for gray seal (Halichoerus grypus) predation and scavenging on harbor porpoise (Phocoena phocoena). Marine Mammals Science: 30(4): 1542-1548.

Bouveroux Th., Tyson RB, Nowacek DP. 2014. Abundance and site fidelity of bottlenose dolphins in coastal waters and bays in Panama City, Florida. Journal of Cetacean Research and Management 14: 37-42.

Cherel Y, Connan M, Jaeger A, Richard P. 2014. Seabird year-round and historical feeding ecology: blood and feather δ13C and δ15N values document foraging plasticity of small sympatric petrels. Marine Ecology Progress Series 505: 267-280.

Cohen LA, Pichegru L, Grémillet D, Coetzee JC, Upfold L, Ryan PG. 2014. Behavioural flexibility of a seabird facing decadal fluctuations in prey availability. Marine Ecology Progress Series 505: 281-293.

Connan M, McQuaid CD, Bonnevie BT, Smale MJ, Cherel Y. 2014.  Combined stomach content, lipid and stable isotope analyses reveal spatial and trophic partitioning among three sympatric albatrosses from the Southern Ocean. Marine Ecology Progress Series 497: 259-272.

Connan M, Hofmeyr GJG, Smale MJ, Pistorius PA. 2014. Trophic investigations of Cape fur seals at the eastern-most extreme of their distribution. African Journal of Marine Science 36: 331-344.

Daly R, Smale MJ, Cowley PD, Froneman PW. 2014. Residency patterns and migration dynamics of adult bull sharks (Carcharhinus leucas) on the east coast of southern Africa. PLoS ONE 9(10):e109357.

Jauniaux T, Garigliany M-M, Loos P, Bourgain J-L, Bouveroux Th., Coignoul F, Haelters J, Karpouzopoulos J, Pezeril S, Desmecht D. 2014. Bite injuries of grey seals (Halichoerus grypus) on harbour porpoises (Phocoena phocoena): the DNA proof. PLoS ONE 9(12): e108993.

Kohler SA, Connan M, Kolasinski J, Cherel Y, McQuaid CD, Jaquemet S. 2014. Trophic overlap between sexes in the dimorphic African Black Oystercatcher foraging on an alien mussel. Austral Ecology 39: 567-578.

Lane EP, de Wet M, Thompson P, Siebert U, Wohlsein P, Plön S. 2014. A systematic health assessment of Indian Ocean bottlenose (Tursiops aduncus) and Indo-Pacific humpback (Sousa plumbea) dolphins, incidentally caught in shark nets off the KwaZulu-Natal coast, South Africa. PloS One 9(9): e0107038.

Lei B, Green JD, Pichegru L. 2014. Extreme microclimate conditions in artificial nests for endangered African penguins. Bird Conservation International 24: 201-213.

Lett C, Semeria M, Thiebault A, Tremblay Y. 2014. Effects of successive predator attacks on prey aggregations. Theoretical Ecology 7(3): 239–252.

Lingham-Soliar T. 2014. Comment on ‘‘A Jurassic ornithischian dinosaur from Siberia with both feathers and scales”. Science 346: 434-435.

Lingham-Soliar T. 2014. Feather structure, biomechanics and biomimetics: the incredible lightness of being. Journal of Ornithology 155: 323–336.

Lingham-Soliar  T. 2014. Response to comments by C. Palmer on my paper, feather structure, biomechanics and biomimetics: the incredible lightness of being. Journal of Ornithology.

Lingham-Soliar T. 2014. The Vertebrate Integument Volume 1. Origin and Evolution. Springer, Heidelberg.

Lingham-Soliar T. 2014. The First Vertebrates, Jawless Fishes, the Agnathans. In: The Vertebrate Integument Volume 1. Springer, Heidelberg, pp 11-32.

Lingham-Soliar T. 2014. The Earliest Jawed Vertebrates, the Gnathostomes. In: The Vertebrate Integument Volume 1. Springer, Heidelberg, pp 33-58.

Lingham-Soliar T. 2014. Evolution of Modern Fishes: Critical Biological Innovations. In: The Vertebrate Integument Volume 1. Springer, Heidelberg, pp 59-98.

Lowther AD, Lydersen C, Biuw M, de Bruyn PJN, Hofmeyr GJG, Kovacs KM. 2014. Post-breeding at-sea movement of three central place foragers in relation to submesoscale fronts in the Southern Ocean around Bouvetøya. Antarctic Science 26: 533-544.

Mahfouz C, Henry F, Courcot L, Pezeril S, Bouveroux Th., Dabin W, Jauniaux T, Khalaf G, Amara R. 2014. Could the chemical contamination be the cause of the increase in the number of stranded harbour porpoises (Phocoena phocoena) along the southern North Sea. Environmental Research 133: 266-273.

McInnes AM, Suarez N, Rishworth GM, Green DB, Pistorius PA, Pichegru L. 2014. Lumbering the gauntlet - Cape Gannet fledglings killed by African Penguins. Ostrich 85(2): 193–196.

Moura AE, Janse van Rensburg C, Pilot M, Tehrani A, Best PB, Thornton M, Plön S, de Bruyn PJN, Dahlheim ME, Hoelzel AR. 2014. Killer whale genome reveals historical population decline correlated with climate change. Molecular Biology and Evolution.

Perissinotto R, Bornman TG, Steyn P, Miranda NAF, Dorrington RA, Matcher GF, Strydom N, Peer N. 2014. Tufa stromatolite ecosystems on the South African south coast. South African Journal of Science 110 (9/10): 1-8.

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

Reisinger RR, Oosthuizen WC, Péron G, Cory Toussaint D, Andrews RD, de Bruyn PJN. 2014. Satellite tagging and biopsy sampling of killer whales at Subantarctic Marion Island: effectiveness, immediate reactions and long-term responses. PLoS One 9:e111835.

Rishworth GM, Tremblay Y, Green DB, Connan M, Pistorius PA. 2014. Drivers of time-activity budget variability during breeding in a pelagic bird. PLoS One 9(12): e116544.

Rishworth G, Tremblay Y, Green D, Pistorius PA. 2014. An automated approach towards measuring time-activity budgets in colonial seabirds. Methods in Ecology and Evolution 5: 854-863.

Rishworth G, Connan M, Green D, Pistorius PA. 2014. Sex differentiation based on the gular stripe in the apparently monomorphic Cape gannet. African Zoology 49: 107-112.

Ryklief R, Pistorius PA, Johnson R. 2014. Spatial and seasonal patterns in sighting rate and life-history composition of the white shark Carcharodon carcharias at Mossel Bay, South Africa. African Journal of Marine Science 36: 449-453.

Sabarros PS, Grémillet D, Demarcq H, Moseley C, Mullers RHE, Pichegru L, Stenseth NC, Machu E. 2014. Fine-scale recognition and use of submesoscale fronts by Cape gannets in the southern Benguela region. Deep Sea Research Part II Topical Studies in Oceanography 107: 77-84.

Thiebault A, Mullers R, Pistorius PA, Meza Torres MA, Dubroca L, Tremblay Y. 2014. From colony to first patch: processes of prey searching and social information in Cape gannets. The AUK: Ornithological Advances 131(4): 595-609.

Thiebault A, Mullers R, Pistorius PA, Tremblay Y. 2014. Local enhancement in a seabird: reaction distances and foraging consequence of predator aggregations. Behavioural Ecology 25: 1302-1310.

Toms J, Compton J, Smale MJ, von der Heyden S. 2014. Variation in paleo-shorelines explains contemporary population genetic patterns of rocky shore species. Biology Letters 10: 20140330.

Tremblay Y, Thiebault A, Mullers R, Pistorius PA. 2014. Bird-borne cameras show that seabird movement patterns relate to previously unrevealed proximate environment, not prey. PLoS ONE 9(2): e88424.

Vanstreels RET, Kolesnikovas CKM, Sandri S, Silveira P, Belo NO, Ferreira Junior FC, Epiphanio S, Steindel M, Braga EM, Catao-Dias JL. 2014. Outbreak of avian malaria associated to multiple species of Plasmodium in Magellanic penguins undergoing rehabilitation in Southern Brazil. Plos One 9: e94994.