Hila Levy, Univ Oxford, Dept Zool, Oxford, England
Stephen R. Fiddaman, Univ Oxford, Dept Zool, Oxford, England
Julianna A. Vianna, Pontificia Univ Catolica Chile, Dept Ecosistemas & Medio Ambiente, Santiago, Chile
Daly Noll, Pontificia Univ Catolica Chile, Dept Ecosistemas & Medio Ambiente, Santiago, Chile; Univ Chile, Inst Ecol & Biodiversidad, Dept Ciencias Ecol, Santiago, Chile
Gemma Clucas, Cornell Univ, Cornell Atkinson Ctr Sustainable Future, Ithaca, NY; Cornell Univ, Cornell Lab Ornithol, Ithaca, NY
Jasmine K. H. Sidhu, Univ Oxford, Dept Zool, Oxford, England
Michael J. Polito, Louisiana State Univ, Dept Oceanog & Coastal Sci, Baton Rouge, LA
Charles A. Bost, Univ La Rochelle, UMR 7372 CNRS, Ctr Etud Biol Chize CEBC, Villiers En Bois, France
Richard A. Phillips, British Antarctic Survey, Cambridge, England
Sarah Crofts, Falklands Conservat, Stanley, Falkland Island
Gary D. Miller, Univ Western Australia, Microbiol & Immunol, PALM, Crawley, WA, Australia
Pierre Pistorius, Nelson Mandela Univ, Percy FitzPatrick Inst African Ornithol, Dept Zool, DST NRF Ctr Excellence, Port Elizabeth, South Africa
Francesco Bonnadonna, Univ Montpellier, Univ Paul Valery Montpellier, EPHE, CNRS,CEFE UMR 5175, Montpellier, France
Celine Le Bohec, Univ Strasbourg, CNRS, IPHC UMR 7178, Strasbourg, France; Ctr Sci Monaco, Dept Biol Polaire, Monaco, Monaco
Andres Barbosa, CSIC, Dept Ecol Evolut, Museo Nacl Ciencias Nat, Madrid, Spain
Phil Trathan, British Antarctic Survey, Cambridge, England
Andrea Raya Rey, Consejo Nacl Invest Cient & Tecn CADIC CONICET, Ctr Austral Invest Cient, Ushuaia, Tierra Del Fueg, Argentina; Univ Nacl Tierra del Fuego, Inst Ciencias Polares Ambiente & Recursos Nat, Ushuaia, Tierra Del Fueg, Argentina; Wildlife Conservat Soc, Buenos Aires, DF, Argentina
Laurent A. F. Frantz, Queen Mary Univ London, Sch Biol & Chem Sci, London, England
Tom Hart, Univ Oxford, Dept Zool, Oxford, England
Adrian L. Smith, Univ Oxford, Dept Zool, Oxford, EnglandFollow

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Over evolutionary time, pathogen challenge shapes the immunepheno type of the host to better respond to an incipient threat. The extent and direction of this selection pressure depend on the local pathogen composition, which is in turn determined by biotic and abiotic features of the environment. However, little is known about adaptation to local pathogen threats in wild animals. The Gentoo penguin (Pygoscelis papua) is a species complex that lends itself to the study of immune adaptation because of its circumpolar distribution over a large latitudinal range, with little or no admixture between ''different clades. Inthis study, we examine the diversity in a key family of innate immune genes-theToll-like receptors (TLRs)-across the range of the Gentoo penguin. The three TLRs that we investigated present varying levels of diversity, with TLR4 and TLR5 greatly exceeding the diversity of TLR7. We present evidence of positive selection in TLR4 and TLR5, which points to pathogen-driven adaptation to the local pathogen milieu. Finally, we demonstrate that two positively selected cosegregating sites in TLR5 are sufficient to alter the responsiveness of the receptor to its bacterial ligand, flagellin. Taken together, these results suggest that Gentoo penguins have experienced distinct pathogen-driven selection pressures in different environments, which may be important given the role of the Gentoo penguin as a sentinel species in some of the world's most rapidly changing environments.

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