title: A supergene determines highly divergent male reproductive morphs in the ruff.
abstract: Three strikingly different alternative male mating morphs (aggressive 'independents', semicooperative 'satellites' and female-mimic 'faeders') coexist as a balanced polymorphism in the ruff, Philomachus pugnax, a lek-breeding wading bird. Major differences in body size, ornamentation, and aggressive and mating behaviors are inherited as an autosomal polymorphism. We show that development into satellites and faeders is determined by a supergene consisting of divergent alternative, dominant and non-recombining haplotypes of an inversion on chromosome 11, which contains 125 predicted genes. Independents are homozygous for the ancestral sequence. One breakpoint of the inversion disrupts the essential CENP-N gene (encoding centromere protein N), and pedigree analysis confirms the lethality of homozygosity for the inversion. We describe new differences in behavior, testis size and steroid metabolism among morphs and identify polymorphic genes within the inversion that are likely to contribute to the differences among morphs in reproductive traits.
authors.affiliation: Department of Animal & Plant Sciences, University of Sheffield, Sheffield, UK.|Institute of Zoology, University of Graz, Graz, Austria.
authors.forename: Clemens
authors.initials: C
authors.lastname: Küpper
authors.affiliation: Department of Animal & Plant Sciences, University of Sheffield, Sheffield, UK.
authors.forename: Michael
authors.initials: M
authors.lastname: Stocks
authors.affiliation: Edinburgh Genomics, Institute of Evolutionary Biology, University of Edinburgh, Edinburgh, UK.
authors.forename: Judith E
authors.initials: JE
authors.lastname: Risse
authors.affiliation: Department of Animal & Plant Sciences, University of Sheffield, Sheffield, UK.
authors.forename: Natalie
authors.initials: N
authors.lastname: Dos Remedios
authors.affiliation: Department of Animal & Plant Sciences, University of Sheffield, Sheffield, UK.|Department of Biological Sciences, Simon Fraser University, Burnaby, British Columbia, Canada.
authors.forename: Lindsay L
authors.initials: LL
authors.lastname: Farrell
authors.affiliation: Department of Biology, East Carolina University, Greenville, North Carolina, USA.
authors.forename: Susan B
authors.initials: SB
authors.lastname: McRae
authors.affiliation: Department of Biological Sciences, Simon Fraser University, Burnaby, British Columbia, Canada.|Department of Biology and Wildlife, Institute of Arctic Biology, Fairbanks, Alaska, USA.
authors.forename: Tawna C
authors.initials: TC
authors.lastname: Morgan
authors.affiliation: Scientific and Practical Center for Bioresources, Minsk, Belarus.
authors.forename: Natalia
authors.initials: N
authors.lastname: Karlionova
authors.affiliation: Scientific and Practical Center for Bioresources, Minsk, Belarus.
authors.forename: Pavel
authors.initials: P
authors.lastname: Pinchuk
authors.affiliation: Conservation Ecology Group, Groningen Institute for Evolutionary Life Sciences, Groningen, The Netherlands.
authors.forename: Yvonne I
authors.initials: YI
authors.lastname: Verkuil
authors.affiliation: Department of Biology and Wildlife, Institute of Arctic Biology, Fairbanks, Alaska, USA.
authors.forename: Alexander S
authors.initials: AS
authors.lastname: Kitaysky
authors.affiliation: Department of Neurobiology, Physiology and Behavior, University of California Davis, Davis, California, USA.
authors.forename: John C
authors.initials: JC
authors.lastname: Wingfield
authors.affiliation: Conservation Ecology Group, Groningen Institute for Evolutionary Life Sciences, Groningen, The Netherlands.|Department of Marine Ecology, NIOZ Royal Netherlands Institute for Sea Research, Den Burg, Texel, The Netherlands.
authors.forename: Theunis
authors.initials: T
authors.lastname: Piersma
authors.affiliation: Department of Animal & Plant Sciences, University of Sheffield, Sheffield, UK.
authors.forename: Kai
authors.initials: K
authors.lastname: Zeng
authors.affiliation: Department of Animal & Plant Sciences, University of Sheffield, Sheffield, UK.
authors.forename: Jon
authors.initials: J
authors.lastname: Slate
authors.affiliation: Edinburgh Genomics, Institute of Evolutionary Biology, University of Edinburgh, Edinburgh, UK.
authors.forename: Mark
authors.initials: M
authors.lastname: Blaxter
authors.affiliation: Department of Biological Sciences, Simon Fraser University, Burnaby, British Columbia, Canada.
authors.forename: David B
authors.initials: DB
authors.lastname: Lank
authors.affiliation: Department of Animal & Plant Sciences, University of Sheffield, Sheffield, UK.
authors.forename: Terry
authors.initials: T
authors.lastname: Burke
mesh_terms: D000818:Animals; D001717:Birds / Q000235:genetics* / Q000502:physiology; D002868:Chromosomal Proteins, Non-Histone / Q000235:genetics; D005260:Female; D008040:Genetic Linkage; D055106:Genome-Wide Association Study; D006239:Haplotypes; D008297:Male; D020641:Polymorphism, Single Nucleotide; D012098:Reproduction / Q000235:genetics*; D012726:Sexual Behavior, Animal; D013256:Steroids / Q000097:blood / Q000378:metabolism; D013737:Testis / Q000502:physiology
chemical_list: D002868:Chromosomal Proteins, Non-Histone; D013256:Steroids