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Downloaded from http://rspb.royalsocietypublishing.org/ on August 19, 2016 rspb.royalsocietypublishing.org Research Cite this article: Tennant JP, Mannion PD,
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Downloaded from http://rspb.royalsocietypublishing.org/ on August 19, 2016 rspb.royalsocietypublishing.org Research Cite this article: Tennant JP, Mannion PD,