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Sequencing of diverse mandarin, pummelo and orange genomes reveals complex history of admixture during citrus domestication

Identifieur interne : 000709 ( Pmc/Checkpoint ); précédent : 000708; suivant : 000710

Sequencing of diverse mandarin, pummelo and orange genomes reveals complex history of admixture during citrus domestication

Auteurs : G. Albert Wu [États-Unis] ; Simon Prochnik [États-Unis] ; Jerry Jenkins [États-Unis] ; Jerome Salse [France] ; Uffe Hellsten [États-Unis] ; Florent Murat [France] ; Xavier Perrier [France] ; Manuel Ruiz [France] ; Simone Scalabrin [Italie] ; Javier Terol [Espagne] ; Marco Aurélio Takita [Brésil] ; Karine Labadie [France] ; Julie Poulain [France] ; Arnaud Couloux [France] ; Kamel Jabbari [France] ; Federica Cattonaro [Italie] ; Cristian Del Fabbro [Italie] ; Sara Pinosio [Italie] ; Andrea Zuccolo [Italie] ; Jarrod Chapman [États-Unis] ; Jane Grimwood [États-Unis] ; Francisco R. Tadeo [Espagne] ; Leandro H. Estornell [Espagne] ; Juan V. Mu Oz-Sanz [Espagne] ; Victoria Ibanez [Espagne] ; Amparo Herrero-Ortega [Espagne] ; Pablo Aleza [Espagne] ; Julián Pérez-Pérez [Espagne] ; Daniel Ram N [Espagne] ; Dominique Brunel [France] ; François Luro [France] ; Chunxian Chen [États-Unis] ; William G. Farmerie [États-Unis] ; Brian Desany [États-Unis] ; Chinnappa Kodira [États-Unis] ; Mohammed Mohiuddin [États-Unis] ; Tim Harkins [États-Unis] ; Karin Fredrikson [États-Unis] ; Paul Burns [États-Unis] ; Alexandre Lomsadze [États-Unis] ; Mark Borodovsky [États-Unis, Russie] ; Giuseppe Reforgiato [Italie] ; Juliana Freitas-Astúa [Brésil] ; Francis Quetier [France] ; Luis Navarro [Espagne] ; Mikeal Roose [États-Unis] ; Patrick Wincker [France] ; Jeremy Schmutz [États-Unis] ; Michele Morgante [Italie] ; Marcos Antonio Machado [Brésil] ; Manuel Talon [Espagne] ; Olivier Jaillon [France] ; Patrick Ollitrault [France] ; Frederick Gmitter [États-Unis] ; Daniel Rokhsar [États-Unis]

Source :

RBID : PMC:4113729

Abstract

The domestication of citrus, is poorly understood. Cultivated types are selections from, or hybrids of, wild progenitor species, whose identities and contributions remain controversial. By comparative analysis of a collection of citrus genomes, including a high quality haploid reference, we show that cultivated types were derived from two progenitor species. Though cultivated pummelos represent selections from a single progenitor species, C. maxima, cultivated mandarins are introgressions of C. maxima into the ancestral mandarin species, C. reticulata. The most widely cultivated citrus, sweet orange, is the offspring of previously admixed individuals, but sour orange is an F1 hybrid of pure C. maxima and C. reticulata parents, implying that wild mandarins were part of the early breeding germplasm. A wild “mandarin” from China exhibited substantial divergence from C. reticulata, suggesting the possibility of other unrecognized wild citrus species. Understanding citrus phylogeny through genome analysis clarifies taxonomic relationships and enables sequence-directed genetic improvement.


Url:
DOI: 10.1038/nbt.2906
PubMed: 24908277
PubMed Central: 4113729


Affiliations:


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PMC:4113729

Le document en format XML

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<nlm:aff id="A15">454 Life Sciences, A Roche Company, 15 Commercial Street, Branford CT, USA</nlm:aff>
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<nlm:aff id="A15">454 Life Sciences, A Roche Company, 15 Commercial Street, Branford CT, USA</nlm:aff>
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<nlm:aff id="A16">Wallace H. Coulter Department of Biomedical Engineering, Georgia Institute of Technology, Atlanta, GA, USA</nlm:aff>
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<nlm:aff id="A16">Wallace H. Coulter Department of Biomedical Engineering, Georgia Institute of Technology, Atlanta, GA, USA</nlm:aff>
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<affiliation wicri:level="1">
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<nlm:aff id="A19">Consiglio per la Ricerca e la Sperimentazione in Agricoltura (CRA-ACM), Acireale, Italy</nlm:aff>
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<wicri:regionArea>Embrapa Cassava and Fruits, Cruz das Almas, BA</wicri:regionArea>
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<nlm:aff id="A7">Commissariat à l'Energie Atomique (CEA), Institut de Génomique (IG), Genoscope, Evry, France</nlm:aff>
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<nlm:aff id="A22">Department of Botany and Plant Sciences, University of California, Riverside, CA, USA</nlm:aff>
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<wicri:regionArea>Department of Botany and Plant Sciences, University of California, Riverside, CA</wicri:regionArea>
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<nlm:aff id="A7">Commissariat à l'Energie Atomique (CEA), Institut de Génomique (IG), Genoscope, Evry, France</nlm:aff>
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<country xml:lang="fr">France</country>
<wicri:regionArea>Centre National de Recherche Scientifique (CNRS), Evry</wicri:regionArea>
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<name sortKey="Schmutz, Jeremy" sort="Schmutz, Jeremy" uniqKey="Schmutz J" first="Jeremy" last="Schmutz">Jeremy Schmutz</name>
<affiliation wicri:level="1">
<nlm:aff id="A2">HudsonAlpha Biotechnology Institute, Huntsville, AL, USA</nlm:aff>
<country xml:lang="fr">États-Unis</country>
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<name sortKey="Morgante, Michele" sort="Morgante, Michele" uniqKey="Morgante M" first="Michele" last="Morgante">Michele Morgante</name>
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<nlm:aff id="A5">Istituto di Genomica Applicata, Udine, Italy</nlm:aff>
<country xml:lang="fr">Italie</country>
<wicri:regionArea>Istituto di Genomica Applicata, Udine</wicri:regionArea>
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<affiliation wicri:level="1">
<nlm:aff id="A24">Department of Agriculture and Environmental Sciences, University of Udine, Udine, Italy</nlm:aff>
<country xml:lang="fr">Italie</country>
<wicri:regionArea>Department of Agriculture and Environmental Sciences, University of Udine, Udine</wicri:regionArea>
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<name sortKey="Machado, Marcos Antonio" sort="Machado, Marcos Antonio" uniqKey="Machado M" first="Marcos Antonio" last="Machado">Marcos Antonio Machado</name>
<affiliation wicri:level="2">
<nlm:aff id="A6">Centro de Citricultura Sylvio Moreira, IAC, Cordeirópolis, SP, Brazil</nlm:aff>
<country xml:lang="fr">Brésil</country>
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<nlm:aff id="A8">Centro de Genomica, Instituto Valenciano de Investigaciones Agrarias (IVIA), Valencia, Spain</nlm:aff>
<country xml:lang="fr">Espagne</country>
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<name sortKey="Jaillon, Olivier" sort="Jaillon, Olivier" uniqKey="Jaillon O" first="Olivier" last="Jaillon">Olivier Jaillon</name>
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<nlm:aff id="A7">Commissariat à l'Energie Atomique (CEA), Institut de Génomique (IG), Genoscope, Evry, France</nlm:aff>
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<wicri:regionArea>Commissariat à l'Energie Atomique (CEA), Institut de Génomique (IG), Genoscope, Evry</wicri:regionArea>
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<nlm:aff id="A21">Département de Biologie, Université d’Evry, Evry, France</nlm:aff>
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<wicri:regionArea>Département de Biologie, Université d’Evry, Evry</wicri:regionArea>
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<nlm:aff id="A23">Centre National de Recherche Scientifique (CNRS), Evry, France</nlm:aff>
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<nlm:aff id="A4">CIRAD, UMR AGAP, Montpellier, France</nlm:aff>
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<nlm:aff id="A13">Citrus Research and Education Center (CREC), Institute of Food and Agricultural Sciences (IFAS), University of Florida, Lake Alfred, FL, USA</nlm:aff>
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<nlm:aff id="A25">Division of Genetics, Genomics, and Development, University of California, Berkeley, CA, USA</nlm:aff>
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<title xml:lang="en" level="a" type="main">Sequencing of diverse mandarin, pummelo and orange genomes reveals complex history of admixture during citrus domestication</title>
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<name sortKey="Wu, G Albert" sort="Wu, G Albert" uniqKey="Wu G" first="G. Albert" last="Wu">G. Albert Wu</name>
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<name sortKey="Couloux, Arnaud" sort="Couloux, Arnaud" uniqKey="Couloux A" first="Arnaud" last="Couloux">Arnaud Couloux</name>
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<nlm:aff id="A7">Commissariat à l'Energie Atomique (CEA), Institut de Génomique (IG), Genoscope, Evry, France</nlm:aff>
<country xml:lang="fr">France</country>
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<name sortKey="Jabbari, Kamel" sort="Jabbari, Kamel" uniqKey="Jabbari K" first="Kamel" last="Jabbari">Kamel Jabbari</name>
<affiliation wicri:level="1">
<nlm:aff id="A7">Commissariat à l'Energie Atomique (CEA), Institut de Génomique (IG), Genoscope, Evry, France</nlm:aff>
<country xml:lang="fr">France</country>
<wicri:regionArea>Commissariat à l'Energie Atomique (CEA), Institut de Génomique (IG), Genoscope, Evry</wicri:regionArea>
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<settlement type="city">Évry (Essonne)</settlement>
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<author>
<name sortKey="Cattonaro, Federica" sort="Cattonaro, Federica" uniqKey="Cattonaro F" first="Federica" last="Cattonaro">Federica Cattonaro</name>
<affiliation wicri:level="1">
<nlm:aff id="A5">Istituto di Genomica Applicata, Udine, Italy</nlm:aff>
<country xml:lang="fr">Italie</country>
<wicri:regionArea>Istituto di Genomica Applicata, Udine</wicri:regionArea>
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<author>
<name sortKey="Del Fabbro, Cristian" sort="Del Fabbro, Cristian" uniqKey="Del Fabbro C" first="Cristian" last="Del Fabbro">Cristian Del Fabbro</name>
<affiliation wicri:level="1">
<nlm:aff id="A5">Istituto di Genomica Applicata, Udine, Italy</nlm:aff>
<country xml:lang="fr">Italie</country>
<wicri:regionArea>Istituto di Genomica Applicata, Udine</wicri:regionArea>
<wicri:noRegion>Udine</wicri:noRegion>
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</author>
<author>
<name sortKey="Pinosio, Sara" sort="Pinosio, Sara" uniqKey="Pinosio S" first="Sara" last="Pinosio">Sara Pinosio</name>
<affiliation wicri:level="1">
<nlm:aff id="A5">Istituto di Genomica Applicata, Udine, Italy</nlm:aff>
<country xml:lang="fr">Italie</country>
<wicri:regionArea>Istituto di Genomica Applicata, Udine</wicri:regionArea>
<wicri:noRegion>Udine</wicri:noRegion>
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</author>
<author>
<name sortKey="Zuccolo, Andrea" sort="Zuccolo, Andrea" uniqKey="Zuccolo A" first="Andrea" last="Zuccolo">Andrea Zuccolo</name>
<affiliation wicri:level="1">
<nlm:aff id="A5">Istituto di Genomica Applicata, Udine, Italy</nlm:aff>
<country xml:lang="fr">Italie</country>
<wicri:regionArea>Istituto di Genomica Applicata, Udine</wicri:regionArea>
<wicri:noRegion>Udine</wicri:noRegion>
</affiliation>
<affiliation wicri:level="3">
<nlm:aff id="A26">Institute of Life Sciences, Scuola Superiore Sant'Anna, Pisa, Italy</nlm:aff>
<country xml:lang="fr">Italie</country>
<wicri:regionArea>Institute of Life Sciences, Scuola Superiore Sant'Anna, Pisa</wicri:regionArea>
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<settlement type="city">Pise</settlement>
<region nuts="2">Toscane</region>
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</affiliation>
</author>
<author>
<name sortKey="Chapman, Jarrod" sort="Chapman, Jarrod" uniqKey="Chapman J" first="Jarrod" last="Chapman">Jarrod Chapman</name>
<affiliation wicri:level="1">
<nlm:aff id="A1">US-Department of Energy Joint Genome Institute, Walnut Creek, CA, USA</nlm:aff>
<country xml:lang="fr">États-Unis</country>
<wicri:regionArea>US-Department of Energy Joint Genome Institute, Walnut Creek, CA</wicri:regionArea>
<wicri:noRegion>CA</wicri:noRegion>
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</author>
<author>
<name sortKey="Grimwood, Jane" sort="Grimwood, Jane" uniqKey="Grimwood J" first="Jane" last="Grimwood">Jane Grimwood</name>
<affiliation wicri:level="1">
<nlm:aff id="A2">HudsonAlpha Biotechnology Institute, Huntsville, AL, USA</nlm:aff>
<country xml:lang="fr">États-Unis</country>
<wicri:regionArea>HudsonAlpha Biotechnology Institute, Huntsville, AL</wicri:regionArea>
<wicri:noRegion>AL</wicri:noRegion>
</affiliation>
</author>
<author>
<name sortKey="Tadeo, Francisco R" sort="Tadeo, Francisco R" uniqKey="Tadeo F" first="Francisco R." last="Tadeo">Francisco R. Tadeo</name>
<affiliation wicri:level="1">
<nlm:aff id="A8">Centro de Genomica, Instituto Valenciano de Investigaciones Agrarias (IVIA), Valencia, Spain</nlm:aff>
<country xml:lang="fr">Espagne</country>
<wicri:regionArea>Centro de Genomica, Instituto Valenciano de Investigaciones Agrarias (IVIA), Valencia</wicri:regionArea>
<wicri:noRegion>Valencia</wicri:noRegion>
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</author>
<author>
<name sortKey="Estornell, Leandro H" sort="Estornell, Leandro H" uniqKey="Estornell L" first="Leandro H." last="Estornell">Leandro H. Estornell</name>
<affiliation wicri:level="1">
<nlm:aff id="A8">Centro de Genomica, Instituto Valenciano de Investigaciones Agrarias (IVIA), Valencia, Spain</nlm:aff>
<country xml:lang="fr">Espagne</country>
<wicri:regionArea>Centro de Genomica, Instituto Valenciano de Investigaciones Agrarias (IVIA), Valencia</wicri:regionArea>
<wicri:noRegion>Valencia</wicri:noRegion>
</affiliation>
</author>
<author>
<name sortKey="Mu Oz Sanz, Juan V" sort="Mu Oz Sanz, Juan V" uniqKey="Mu Oz Sanz J" first="Juan V." last="Mu Oz-Sanz">Juan V. Mu Oz-Sanz</name>
<affiliation wicri:level="1">
<nlm:aff id="A8">Centro de Genomica, Instituto Valenciano de Investigaciones Agrarias (IVIA), Valencia, Spain</nlm:aff>
<country xml:lang="fr">Espagne</country>
<wicri:regionArea>Centro de Genomica, Instituto Valenciano de Investigaciones Agrarias (IVIA), Valencia</wicri:regionArea>
<wicri:noRegion>Valencia</wicri:noRegion>
</affiliation>
</author>
<author>
<name sortKey="Ibanez, Victoria" sort="Ibanez, Victoria" uniqKey="Ibanez V" first="Victoria" last="Ibanez">Victoria Ibanez</name>
<affiliation wicri:level="1">
<nlm:aff id="A8">Centro de Genomica, Instituto Valenciano de Investigaciones Agrarias (IVIA), Valencia, Spain</nlm:aff>
<country xml:lang="fr">Espagne</country>
<wicri:regionArea>Centro de Genomica, Instituto Valenciano de Investigaciones Agrarias (IVIA), Valencia</wicri:regionArea>
<wicri:noRegion>Valencia</wicri:noRegion>
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</author>
<author>
<name sortKey="Herrero Ortega, Amparo" sort="Herrero Ortega, Amparo" uniqKey="Herrero Ortega A" first="Amparo" last="Herrero-Ortega">Amparo Herrero-Ortega</name>
<affiliation wicri:level="1">
<nlm:aff id="A8">Centro de Genomica, Instituto Valenciano de Investigaciones Agrarias (IVIA), Valencia, Spain</nlm:aff>
<country xml:lang="fr">Espagne</country>
<wicri:regionArea>Centro de Genomica, Instituto Valenciano de Investigaciones Agrarias (IVIA), Valencia</wicri:regionArea>
<wicri:noRegion>Valencia</wicri:noRegion>
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</author>
<author>
<name sortKey="Aleza, Pablo" sort="Aleza, Pablo" uniqKey="Aleza P" first="Pablo" last="Aleza">Pablo Aleza</name>
<affiliation wicri:level="1">
<nlm:aff id="A9">Centro de Protección Vegetal y Biotecnología-IVIA, Moncada, Valencia, Spain</nlm:aff>
<country xml:lang="fr">Espagne</country>
<wicri:regionArea>Centro de Protección Vegetal y Biotecnología-IVIA, Moncada, Valencia</wicri:regionArea>
<wicri:noRegion>Valencia</wicri:noRegion>
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</author>
<author>
<name sortKey="Perez Perez, Julian" sort="Perez Perez, Julian" uniqKey="Perez Perez J" first="Julián" last="Pérez-Pérez">Julián Pérez-Pérez</name>
<affiliation wicri:level="1">
<nlm:aff id="A10">Lifesequencing SL, Valencia, Spain</nlm:aff>
<country xml:lang="fr">Espagne</country>
<wicri:regionArea>Lifesequencing SL, Valencia</wicri:regionArea>
<wicri:noRegion>Valencia</wicri:noRegion>
</affiliation>
<affiliation wicri:level="3">
<nlm:aff id="A27">Secugen SL, Madrid, Spain</nlm:aff>
<country xml:lang="fr">Espagne</country>
<wicri:regionArea>Secugen SL, Madrid</wicri:regionArea>
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<settlement type="city">Madrid</settlement>
<region nuts="2" type="region">Communauté de Madrid</region>
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</affiliation>
</author>
<author>
<name sortKey="Ram N, Daniel" sort="Ram N, Daniel" uniqKey="Ram N D" first="Daniel" last="Ram N">Daniel Ram N</name>
<affiliation wicri:level="1">
<nlm:aff id="A10">Lifesequencing SL, Valencia, Spain</nlm:aff>
<country xml:lang="fr">Espagne</country>
<wicri:regionArea>Lifesequencing SL, Valencia</wicri:regionArea>
<wicri:noRegion>Valencia</wicri:noRegion>
</affiliation>
</author>
<author>
<name sortKey="Brunel, Dominique" sort="Brunel, Dominique" uniqKey="Brunel D" first="Dominique" last="Brunel">Dominique Brunel</name>
<affiliation wicri:level="1">
<nlm:aff id="A7">Commissariat à l'Energie Atomique (CEA), Institut de Génomique (IG), Genoscope, Evry, France</nlm:aff>
<country xml:lang="fr">France</country>
<wicri:regionArea>Commissariat à l'Energie Atomique (CEA), Institut de Génomique (IG), Genoscope, Evry</wicri:regionArea>
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<settlement type="city">Évry (Essonne)</settlement>
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</affiliation>
<affiliation wicri:level="1">
<nlm:aff id="A11">INRA, US EPGV_1279, Evry, France</nlm:aff>
<country xml:lang="fr">France</country>
<wicri:regionArea>INRA, US EPGV_1279, Evry</wicri:regionArea>
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<settlement type="city">Évry (Essonne)</settlement>
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</affiliation>
</author>
<author>
<name sortKey="Luro, Francois" sort="Luro, Francois" uniqKey="Luro F" first="François" last="Luro">François Luro</name>
<affiliation wicri:level="1">
<nlm:aff id="A12">INRA GEQA, San Giuliano, France</nlm:aff>
<country xml:lang="fr">France</country>
<wicri:regionArea>INRA GEQA, San Giuliano</wicri:regionArea>
<wicri:noRegion>San Giuliano</wicri:noRegion>
<wicri:noRegion>San Giuliano</wicri:noRegion>
</affiliation>
</author>
<author>
<name sortKey="Chen, Chunxian" sort="Chen, Chunxian" uniqKey="Chen C" first="Chunxian" last="Chen">Chunxian Chen</name>
<affiliation wicri:level="1">
<nlm:aff id="A13">Citrus Research and Education Center (CREC), Institute of Food and Agricultural Sciences (IFAS), University of Florida, Lake Alfred, FL, USA</nlm:aff>
<country xml:lang="fr">États-Unis</country>
<wicri:regionArea>Citrus Research and Education Center (CREC), Institute of Food and Agricultural Sciences (IFAS), University of Florida, Lake Alfred, FL</wicri:regionArea>
<wicri:noRegion>FL</wicri:noRegion>
</affiliation>
</author>
<author>
<name sortKey="Farmerie, William G" sort="Farmerie, William G" uniqKey="Farmerie W" first="William G." last="Farmerie">William G. Farmerie</name>
<affiliation wicri:level="1">
<nlm:aff id="A14">Interdisciplinary Center for Biotechnology Research, University of Florida, Gainesville, FL, USA</nlm:aff>
<country xml:lang="fr">États-Unis</country>
<wicri:regionArea>Interdisciplinary Center for Biotechnology Research, University of Florida, Gainesville, FL</wicri:regionArea>
<wicri:noRegion>FL</wicri:noRegion>
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</author>
<author>
<name sortKey="Desany, Brian" sort="Desany, Brian" uniqKey="Desany B" first="Brian" last="Desany">Brian Desany</name>
<affiliation wicri:level="1">
<nlm:aff id="A15">454 Life Sciences, A Roche Company, 15 Commercial Street, Branford CT, USA</nlm:aff>
<country xml:lang="fr">États-Unis</country>
<wicri:regionArea>454 Life Sciences, A Roche Company, 15 Commercial Street, Branford CT</wicri:regionArea>
<wicri:noRegion>Branford CT</wicri:noRegion>
</affiliation>
</author>
<author>
<name sortKey="Kodira, Chinnappa" sort="Kodira, Chinnappa" uniqKey="Kodira C" first="Chinnappa" last="Kodira">Chinnappa Kodira</name>
<affiliation wicri:level="1">
<nlm:aff id="A15">454 Life Sciences, A Roche Company, 15 Commercial Street, Branford CT, USA</nlm:aff>
<country xml:lang="fr">États-Unis</country>
<wicri:regionArea>454 Life Sciences, A Roche Company, 15 Commercial Street, Branford CT</wicri:regionArea>
<wicri:noRegion>Branford CT</wicri:noRegion>
</affiliation>
</author>
<author>
<name sortKey="Mohiuddin, Mohammed" sort="Mohiuddin, Mohammed" uniqKey="Mohiuddin M" first="Mohammed" last="Mohiuddin">Mohammed Mohiuddin</name>
<affiliation wicri:level="1">
<nlm:aff id="A15">454 Life Sciences, A Roche Company, 15 Commercial Street, Branford CT, USA</nlm:aff>
<country xml:lang="fr">États-Unis</country>
<wicri:regionArea>454 Life Sciences, A Roche Company, 15 Commercial Street, Branford CT</wicri:regionArea>
<wicri:noRegion>Branford CT</wicri:noRegion>
</affiliation>
</author>
<author>
<name sortKey="Harkins, Tim" sort="Harkins, Tim" uniqKey="Harkins T" first="Tim" last="Harkins">Tim Harkins</name>
<affiliation wicri:level="1">
<nlm:aff id="A15">454 Life Sciences, A Roche Company, 15 Commercial Street, Branford CT, USA</nlm:aff>
<country xml:lang="fr">États-Unis</country>
<wicri:regionArea>454 Life Sciences, A Roche Company, 15 Commercial Street, Branford CT</wicri:regionArea>
<wicri:noRegion>Branford CT</wicri:noRegion>
</affiliation>
</author>
<author>
<name sortKey="Fredrikson, Karin" sort="Fredrikson, Karin" uniqKey="Fredrikson K" first="Karin" last="Fredrikson">Karin Fredrikson</name>
<affiliation wicri:level="1">
<nlm:aff id="A15">454 Life Sciences, A Roche Company, 15 Commercial Street, Branford CT, USA</nlm:aff>
<country xml:lang="fr">États-Unis</country>
<wicri:regionArea>454 Life Sciences, A Roche Company, 15 Commercial Street, Branford CT</wicri:regionArea>
<wicri:noRegion>Branford CT</wicri:noRegion>
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</author>
<author>
<name sortKey="Burns, Paul" sort="Burns, Paul" uniqKey="Burns P" first="Paul" last="Burns">Paul Burns</name>
<affiliation wicri:level="1">
<nlm:aff id="A16">Wallace H. Coulter Department of Biomedical Engineering, Georgia Institute of Technology, Atlanta, GA, USA</nlm:aff>
<country xml:lang="fr">États-Unis</country>
<wicri:regionArea>Wallace H. Coulter Department of Biomedical Engineering, Georgia Institute of Technology, Atlanta, GA</wicri:regionArea>
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</affiliation>
<affiliation wicri:level="1">
<nlm:aff id="A17">School of Computational Science & Engineering, Georgia Institute of Technology, Atlanta, GA, USA</nlm:aff>
<country xml:lang="fr">États-Unis</country>
<wicri:regionArea>School of Computational Science & Engineering, Georgia Institute of Technology, Atlanta, GA</wicri:regionArea>
<wicri:noRegion>GA</wicri:noRegion>
</affiliation>
</author>
<author>
<name sortKey="Lomsadze, Alexandre" sort="Lomsadze, Alexandre" uniqKey="Lomsadze A" first="Alexandre" last="Lomsadze">Alexandre Lomsadze</name>
<affiliation wicri:level="1">
<nlm:aff id="A16">Wallace H. Coulter Department of Biomedical Engineering, Georgia Institute of Technology, Atlanta, GA, USA</nlm:aff>
<country xml:lang="fr">États-Unis</country>
<wicri:regionArea>Wallace H. Coulter Department of Biomedical Engineering, Georgia Institute of Technology, Atlanta, GA</wicri:regionArea>
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</affiliation>
<affiliation wicri:level="1">
<nlm:aff id="A17">School of Computational Science & Engineering, Georgia Institute of Technology, Atlanta, GA, USA</nlm:aff>
<country xml:lang="fr">États-Unis</country>
<wicri:regionArea>School of Computational Science & Engineering, Georgia Institute of Technology, Atlanta, GA</wicri:regionArea>
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</affiliation>
</author>
<author>
<name sortKey="Borodovsky, Mark" sort="Borodovsky, Mark" uniqKey="Borodovsky M" first="Mark" last="Borodovsky">Mark Borodovsky</name>
<affiliation wicri:level="1">
<nlm:aff id="A16">Wallace H. Coulter Department of Biomedical Engineering, Georgia Institute of Technology, Atlanta, GA, USA</nlm:aff>
<country xml:lang="fr">États-Unis</country>
<wicri:regionArea>Wallace H. Coulter Department of Biomedical Engineering, Georgia Institute of Technology, Atlanta, GA</wicri:regionArea>
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</affiliation>
<affiliation wicri:level="1">
<nlm:aff id="A17">School of Computational Science & Engineering, Georgia Institute of Technology, Atlanta, GA, USA</nlm:aff>
<country xml:lang="fr">États-Unis</country>
<wicri:regionArea>School of Computational Science & Engineering, Georgia Institute of Technology, Atlanta, GA</wicri:regionArea>
<wicri:noRegion>GA</wicri:noRegion>
</affiliation>
<affiliation wicri:level="1">
<nlm:aff id="A18">Department of Biological and Medical Physics, Moscow Institute of Physics and Technology, Dolgoprudny, Moscow Region, Russia</nlm:aff>
<country xml:lang="fr">Russie</country>
<wicri:regionArea>Department of Biological and Medical Physics, Moscow Institute of Physics and Technology, Dolgoprudny, Moscow Region</wicri:regionArea>
<wicri:noRegion>Moscow Region</wicri:noRegion>
</affiliation>
</author>
<author>
<name sortKey="Reforgiato, Giuseppe" sort="Reforgiato, Giuseppe" uniqKey="Reforgiato G" first="Giuseppe" last="Reforgiato">Giuseppe Reforgiato</name>
<affiliation wicri:level="1">
<nlm:aff id="A19">Consiglio per la Ricerca e la Sperimentazione in Agricoltura (CRA-ACM), Acireale, Italy</nlm:aff>
<country xml:lang="fr">Italie</country>
<wicri:regionArea>Consiglio per la Ricerca e la Sperimentazione in Agricoltura (CRA-ACM), Acireale</wicri:regionArea>
<wicri:noRegion>Acireale</wicri:noRegion>
</affiliation>
</author>
<author>
<name sortKey="Freitas Astua, Juliana" sort="Freitas Astua, Juliana" uniqKey="Freitas Astua J" first="Juliana" last="Freitas-Astúa">Juliana Freitas-Astúa</name>
<affiliation wicri:level="2">
<nlm:aff id="A6">Centro de Citricultura Sylvio Moreira, IAC, Cordeirópolis, SP, Brazil</nlm:aff>
<country xml:lang="fr">Brésil</country>
<wicri:regionArea>Centro de Citricultura Sylvio Moreira, IAC, Cordeirópolis, SP</wicri:regionArea>
<placeName>
<region type="state">État de São Paulo</region>
</placeName>
</affiliation>
<affiliation wicri:level="2">
<nlm:aff id="A20">Embrapa Cassava and Fruits, Cruz das Almas, BA, Brazil</nlm:aff>
<country xml:lang="fr">Brésil</country>
<wicri:regionArea>Embrapa Cassava and Fruits, Cruz das Almas, BA</wicri:regionArea>
<placeName>
<region type="state">Bahia</region>
</placeName>
</affiliation>
</author>
<author>
<name sortKey="Quetier, Francis" sort="Quetier, Francis" uniqKey="Quetier F" first="Francis" last="Quetier">Francis Quetier</name>
<affiliation wicri:level="1">
<nlm:aff id="A7">Commissariat à l'Energie Atomique (CEA), Institut de Génomique (IG), Genoscope, Evry, France</nlm:aff>
<country xml:lang="fr">France</country>
<wicri:regionArea>Commissariat à l'Energie Atomique (CEA), Institut de Génomique (IG), Genoscope, Evry</wicri:regionArea>
<placeName>
<settlement type="city">Évry (Essonne)</settlement>
</placeName>
</affiliation>
<affiliation wicri:level="1">
<nlm:aff id="A21">Département de Biologie, Université d’Evry, Evry, France</nlm:aff>
<country xml:lang="fr">France</country>
<wicri:regionArea>Département de Biologie, Université d’Evry, Evry</wicri:regionArea>
<placeName>
<settlement type="city">Évry (Essonne)</settlement>
</placeName>
</affiliation>
</author>
<author>
<name sortKey="Navarro, Luis" sort="Navarro, Luis" uniqKey="Navarro L" first="Luis" last="Navarro">Luis Navarro</name>
<affiliation wicri:level="1">
<nlm:aff id="A9">Centro de Protección Vegetal y Biotecnología-IVIA, Moncada, Valencia, Spain</nlm:aff>
<country xml:lang="fr">Espagne</country>
<wicri:regionArea>Centro de Protección Vegetal y Biotecnología-IVIA, Moncada, Valencia</wicri:regionArea>
<wicri:noRegion>Valencia</wicri:noRegion>
</affiliation>
</author>
<author>
<name sortKey="Roose, Mikeal" sort="Roose, Mikeal" uniqKey="Roose M" first="Mikeal" last="Roose">Mikeal Roose</name>
<affiliation wicri:level="1">
<nlm:aff id="A22">Department of Botany and Plant Sciences, University of California, Riverside, CA, USA</nlm:aff>
<country xml:lang="fr">États-Unis</country>
<wicri:regionArea>Department of Botany and Plant Sciences, University of California, Riverside, CA</wicri:regionArea>
<wicri:noRegion>CA</wicri:noRegion>
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</author>
<author>
<name sortKey="Wincker, Patrick" sort="Wincker, Patrick" uniqKey="Wincker P" first="Patrick" last="Wincker">Patrick Wincker</name>
<affiliation wicri:level="1">
<nlm:aff id="A7">Commissariat à l'Energie Atomique (CEA), Institut de Génomique (IG), Genoscope, Evry, France</nlm:aff>
<country xml:lang="fr">France</country>
<wicri:regionArea>Commissariat à l'Energie Atomique (CEA), Institut de Génomique (IG), Genoscope, Evry</wicri:regionArea>
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<settlement type="city">Évry (Essonne)</settlement>
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</affiliation>
<affiliation wicri:level="1">
<nlm:aff id="A21">Département de Biologie, Université d’Evry, Evry, France</nlm:aff>
<country xml:lang="fr">France</country>
<wicri:regionArea>Département de Biologie, Université d’Evry, Evry</wicri:regionArea>
<placeName>
<settlement type="city">Évry (Essonne)</settlement>
</placeName>
</affiliation>
<affiliation wicri:level="1">
<nlm:aff id="A23">Centre National de Recherche Scientifique (CNRS), Evry, France</nlm:aff>
<country xml:lang="fr">France</country>
<wicri:regionArea>Centre National de Recherche Scientifique (CNRS), Evry</wicri:regionArea>
<placeName>
<settlement type="city">Évry (Essonne)</settlement>
</placeName>
</affiliation>
</author>
<author>
<name sortKey="Schmutz, Jeremy" sort="Schmutz, Jeremy" uniqKey="Schmutz J" first="Jeremy" last="Schmutz">Jeremy Schmutz</name>
<affiliation wicri:level="1">
<nlm:aff id="A2">HudsonAlpha Biotechnology Institute, Huntsville, AL, USA</nlm:aff>
<country xml:lang="fr">États-Unis</country>
<wicri:regionArea>HudsonAlpha Biotechnology Institute, Huntsville, AL</wicri:regionArea>
<wicri:noRegion>AL</wicri:noRegion>
</affiliation>
</author>
<author>
<name sortKey="Morgante, Michele" sort="Morgante, Michele" uniqKey="Morgante M" first="Michele" last="Morgante">Michele Morgante</name>
<affiliation wicri:level="1">
<nlm:aff id="A5">Istituto di Genomica Applicata, Udine, Italy</nlm:aff>
<country xml:lang="fr">Italie</country>
<wicri:regionArea>Istituto di Genomica Applicata, Udine</wicri:regionArea>
<wicri:noRegion>Udine</wicri:noRegion>
</affiliation>
<affiliation wicri:level="1">
<nlm:aff id="A24">Department of Agriculture and Environmental Sciences, University of Udine, Udine, Italy</nlm:aff>
<country xml:lang="fr">Italie</country>
<wicri:regionArea>Department of Agriculture and Environmental Sciences, University of Udine, Udine</wicri:regionArea>
<wicri:noRegion>Udine</wicri:noRegion>
</affiliation>
</author>
<author>
<name sortKey="Machado, Marcos Antonio" sort="Machado, Marcos Antonio" uniqKey="Machado M" first="Marcos Antonio" last="Machado">Marcos Antonio Machado</name>
<affiliation wicri:level="2">
<nlm:aff id="A6">Centro de Citricultura Sylvio Moreira, IAC, Cordeirópolis, SP, Brazil</nlm:aff>
<country xml:lang="fr">Brésil</country>
<wicri:regionArea>Centro de Citricultura Sylvio Moreira, IAC, Cordeirópolis, SP</wicri:regionArea>
<placeName>
<region type="state">État de São Paulo</region>
</placeName>
</affiliation>
</author>
<author>
<name sortKey="Talon, Manuel" sort="Talon, Manuel" uniqKey="Talon M" first="Manuel" last="Talon">Manuel Talon</name>
<affiliation wicri:level="1">
<nlm:aff id="A8">Centro de Genomica, Instituto Valenciano de Investigaciones Agrarias (IVIA), Valencia, Spain</nlm:aff>
<country xml:lang="fr">Espagne</country>
<wicri:regionArea>Centro de Genomica, Instituto Valenciano de Investigaciones Agrarias (IVIA), Valencia</wicri:regionArea>
<wicri:noRegion>Valencia</wicri:noRegion>
</affiliation>
</author>
<author>
<name sortKey="Jaillon, Olivier" sort="Jaillon, Olivier" uniqKey="Jaillon O" first="Olivier" last="Jaillon">Olivier Jaillon</name>
<affiliation wicri:level="1">
<nlm:aff id="A7">Commissariat à l'Energie Atomique (CEA), Institut de Génomique (IG), Genoscope, Evry, France</nlm:aff>
<country xml:lang="fr">France</country>
<wicri:regionArea>Commissariat à l'Energie Atomique (CEA), Institut de Génomique (IG), Genoscope, Evry</wicri:regionArea>
<placeName>
<settlement type="city">Évry (Essonne)</settlement>
</placeName>
</affiliation>
<affiliation wicri:level="1">
<nlm:aff id="A21">Département de Biologie, Université d’Evry, Evry, France</nlm:aff>
<country xml:lang="fr">France</country>
<wicri:regionArea>Département de Biologie, Université d’Evry, Evry</wicri:regionArea>
<placeName>
<settlement type="city">Évry (Essonne)</settlement>
</placeName>
</affiliation>
<affiliation wicri:level="1">
<nlm:aff id="A23">Centre National de Recherche Scientifique (CNRS), Evry, France</nlm:aff>
<country xml:lang="fr">France</country>
<wicri:regionArea>Centre National de Recherche Scientifique (CNRS), Evry</wicri:regionArea>
<placeName>
<settlement type="city">Évry (Essonne)</settlement>
</placeName>
</affiliation>
</author>
<author>
<name sortKey="Ollitrault, Patrick" sort="Ollitrault, Patrick" uniqKey="Ollitrault P" first="Patrick" last="Ollitrault">Patrick Ollitrault</name>
<affiliation wicri:level="1">
<nlm:aff id="A4">CIRAD, UMR AGAP, Montpellier, France</nlm:aff>
<country xml:lang="fr">France</country>
<wicri:regionArea>CIRAD, UMR AGAP, Montpellier</wicri:regionArea>
<placeName>
<settlement type="city">Montpellier</settlement>
</placeName>
</affiliation>
</author>
<author>
<name sortKey="Gmitter, Frederick" sort="Gmitter, Frederick" uniqKey="Gmitter F" first="Frederick" last="Gmitter">Frederick Gmitter</name>
<affiliation wicri:level="1">
<nlm:aff id="A13">Citrus Research and Education Center (CREC), Institute of Food and Agricultural Sciences (IFAS), University of Florida, Lake Alfred, FL, USA</nlm:aff>
<country xml:lang="fr">États-Unis</country>
<wicri:regionArea>Citrus Research and Education Center (CREC), Institute of Food and Agricultural Sciences (IFAS), University of Florida, Lake Alfred, FL</wicri:regionArea>
<wicri:noRegion>FL</wicri:noRegion>
</affiliation>
</author>
<author>
<name sortKey="Rokhsar, Daniel" sort="Rokhsar, Daniel" uniqKey="Rokhsar D" first="Daniel" last="Rokhsar">Daniel Rokhsar</name>
<affiliation wicri:level="1">
<nlm:aff id="A1">US-Department of Energy Joint Genome Institute, Walnut Creek, CA, USA</nlm:aff>
<country xml:lang="fr">États-Unis</country>
<wicri:regionArea>US-Department of Energy Joint Genome Institute, Walnut Creek, CA</wicri:regionArea>
<wicri:noRegion>CA</wicri:noRegion>
</affiliation>
<affiliation wicri:level="1">
<nlm:aff id="A25">Division of Genetics, Genomics, and Development, University of California, Berkeley, CA, USA</nlm:aff>
<country xml:lang="fr">États-Unis</country>
<wicri:regionArea>Division of Genetics, Genomics, and Development, University of California, Berkeley, CA</wicri:regionArea>
<wicri:noRegion>CA</wicri:noRegion>
</affiliation>
</author>
</analytic>
<series>
<title level="j">Nature biotechnology</title>
<idno type="ISSN">1087-0156</idno>
<idno type="eISSN">1546-1696</idno>
<imprint>
<date when="2014">2014</date>
</imprint>
</series>
</biblStruct>
</sourceDesc>
</fileDesc>
<profileDesc>
<textClass></textClass>
</profileDesc>
</teiHeader>
<front>
<div type="abstract" xml:lang="en">
<p id="P4">The domestication of citrus, is poorly understood. Cultivated types are selections from, or hybrids of, wild progenitor species, whose identities and contributions remain controversial. By comparative analysis of a collection of citrus genomes, including a high quality haploid reference, we show that cultivated types were derived from two progenitor species. Though cultivated pummelos represent selections from a single progenitor species,
<italic>C. maxima</italic>
, cultivated mandarins are introgressions of
<italic>C. maxima</italic>
into the ancestral mandarin species,
<italic>C. reticulata</italic>
. The most widely cultivated citrus, sweet orange, is the offspring of previously admixed individuals, but sour orange is an F1 hybrid of pure
<italic>C. maxima</italic>
and
<italic>C. reticulata</italic>
parents, implying that wild mandarins were part of the early breeding germplasm. A wild “mandarin” from China exhibited substantial divergence from
<italic>C. reticulata</italic>
, suggesting the possibility of other unrecognized wild citrus species. Understanding citrus phylogeny through genome analysis clarifies taxonomic relationships and enables sequence-directed genetic improvement.</p>
</div>
</front>
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</TEI>
<pmc article-type="research-article">
<pmc-dir>properties open_access</pmc-dir>
<pmc-dir>properties manuscript</pmc-dir>
<front>
<journal-meta>
<journal-id journal-id-type="nlm-journal-id">9604648</journal-id>
<journal-id journal-id-type="pubmed-jr-id">20305</journal-id>
<journal-id journal-id-type="nlm-ta">Nat Biotechnol</journal-id>
<journal-id journal-id-type="iso-abbrev">Nat. Biotechnol.</journal-id>
<journal-title-group>
<journal-title>Nature biotechnology</journal-title>
</journal-title-group>
<issn pub-type="ppub">1087-0156</issn>
<issn pub-type="epub">1546-1696</issn>
</journal-meta>
<article-meta>
<article-id pub-id-type="pmid">24908277</article-id>
<article-id pub-id-type="pmc">4113729</article-id>
<article-id pub-id-type="doi">10.1038/nbt.2906</article-id>
<article-id pub-id-type="manuscript">NIHMS586218</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Article</subject>
</subj-group>
</article-categories>
<title-group>
<article-title>Sequencing of diverse mandarin, pummelo and orange genomes reveals complex history of admixture during citrus domestication</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name>
<surname>Wu</surname>
<given-names>G. Albert</given-names>
</name>
<xref ref-type="aff" rid="A1">1</xref>
<xref ref-type="author-notes" rid="FN3"></xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Prochnik</surname>
<given-names>Simon</given-names>
</name>
<xref ref-type="aff" rid="A1">1</xref>
<xref ref-type="author-notes" rid="FN3"></xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Jenkins</surname>
<given-names>Jerry</given-names>
</name>
<xref ref-type="aff" rid="A2">2</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Salse</surname>
<given-names>Jerome</given-names>
</name>
<xref ref-type="aff" rid="A3">3</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Hellsten</surname>
<given-names>Uffe</given-names>
</name>
<xref ref-type="aff" rid="A1">1</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Murat</surname>
<given-names>Florent</given-names>
</name>
<xref ref-type="aff" rid="A3">3</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Perrier</surname>
<given-names>Xavier</given-names>
</name>
<xref ref-type="aff" rid="A4">4</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Ruiz</surname>
<given-names>Manuel</given-names>
</name>
<xref ref-type="aff" rid="A4">4</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Scalabrin</surname>
<given-names>Simone</given-names>
</name>
<xref ref-type="aff" rid="A5">5</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Terol</surname>
<given-names>Javier</given-names>
</name>
<xref ref-type="aff" rid="A8">8</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Takita</surname>
<given-names>Marco Aurélio</given-names>
</name>
<xref ref-type="aff" rid="A6">6</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Labadie</surname>
<given-names>Karine</given-names>
</name>
<xref ref-type="aff" rid="A7">7</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Poulain</surname>
<given-names>Julie</given-names>
</name>
<xref ref-type="aff" rid="A7">7</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Couloux</surname>
<given-names>Arnaud</given-names>
</name>
<xref ref-type="aff" rid="A7">7</xref>
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<contrib contrib-type="author">
<name>
<surname>Jabbari</surname>
<given-names>Kamel</given-names>
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<xref ref-type="aff" rid="A7">7</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Cattonaro</surname>
<given-names>Federica</given-names>
</name>
<xref ref-type="aff" rid="A5">5</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Del Fabbro</surname>
<given-names>Cristian</given-names>
</name>
<xref ref-type="aff" rid="A5">5</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Pinosio</surname>
<given-names>Sara</given-names>
</name>
<xref ref-type="aff" rid="A5">5</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Zuccolo</surname>
<given-names>Andrea</given-names>
</name>
<xref ref-type="aff" rid="A5">5</xref>
<xref ref-type="aff" rid="A26">26</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Chapman</surname>
<given-names>Jarrod</given-names>
</name>
<xref ref-type="aff" rid="A1">1</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Grimwood</surname>
<given-names>Jane</given-names>
</name>
<xref ref-type="aff" rid="A2">2</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Tadeo</surname>
<given-names>Francisco R.</given-names>
</name>
<xref ref-type="aff" rid="A8">8</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Estornell</surname>
<given-names>Leandro H.</given-names>
</name>
<xref ref-type="aff" rid="A8">8</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Muñoz-Sanz</surname>
<given-names>Juan V.</given-names>
</name>
<xref ref-type="aff" rid="A8">8</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Ibanez</surname>
<given-names>Victoria</given-names>
</name>
<xref ref-type="aff" rid="A8">8</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Herrero-Ortega</surname>
<given-names>Amparo</given-names>
</name>
<xref ref-type="aff" rid="A8">8</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Aleza</surname>
<given-names>Pablo</given-names>
</name>
<xref ref-type="aff" rid="A9">9</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Pérez-Pérez</surname>
<given-names>Julián</given-names>
</name>
<xref ref-type="aff" rid="A10">10</xref>
<xref ref-type="aff" rid="A27">27</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Ramón</surname>
<given-names>Daniel</given-names>
</name>
<xref ref-type="aff" rid="A10">10</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Brunel</surname>
<given-names>Dominique</given-names>
</name>
<xref ref-type="aff" rid="A7">7</xref>
<xref ref-type="aff" rid="A11">11</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Luro</surname>
<given-names>François</given-names>
</name>
<xref ref-type="aff" rid="A12">12</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Chen</surname>
<given-names>Chunxian</given-names>
</name>
<xref ref-type="aff" rid="A13">13</xref>
<xref ref-type="author-notes" rid="FN2"></xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Farmerie</surname>
<given-names>William G.</given-names>
</name>
<xref ref-type="aff" rid="A14">14</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Desany</surname>
<given-names>Brian</given-names>
</name>
<xref ref-type="aff" rid="A15">15</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Kodira</surname>
<given-names>Chinnappa</given-names>
</name>
<xref ref-type="aff" rid="A15">15</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Mohiuddin</surname>
<given-names>Mohammed</given-names>
</name>
<xref ref-type="aff" rid="A15">15</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Harkins</surname>
<given-names>Tim</given-names>
</name>
<xref ref-type="aff" rid="A15">15</xref>
<xref ref-type="author-notes" rid="FN1"></xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Fredrikson</surname>
<given-names>Karin</given-names>
</name>
<xref ref-type="aff" rid="A15">15</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Burns</surname>
<given-names>Paul</given-names>
</name>
<xref ref-type="aff" rid="A16">16</xref>
<xref ref-type="aff" rid="A17">17</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Lomsadze</surname>
<given-names>Alexandre</given-names>
</name>
<xref ref-type="aff" rid="A16">16</xref>
<xref ref-type="aff" rid="A17">17</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Borodovsky</surname>
<given-names>Mark</given-names>
</name>
<xref ref-type="aff" rid="A16">16</xref>
<xref ref-type="aff" rid="A17">17</xref>
<xref ref-type="aff" rid="A18">18</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Reforgiato</surname>
<given-names>Giuseppe</given-names>
</name>
<xref ref-type="aff" rid="A19">19</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Freitas-Astúa</surname>
<given-names>Juliana</given-names>
</name>
<xref ref-type="aff" rid="A6">6</xref>
<xref ref-type="aff" rid="A20">20</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Quetier</surname>
<given-names>Francis</given-names>
</name>
<xref ref-type="aff" rid="A7">7</xref>
<xref ref-type="aff" rid="A21">21</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Navarro</surname>
<given-names>Luis</given-names>
</name>
<xref ref-type="aff" rid="A9">9</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Roose</surname>
<given-names>Mikeal</given-names>
</name>
<xref ref-type="aff" rid="A22">22</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Wincker</surname>
<given-names>Patrick</given-names>
</name>
<xref ref-type="aff" rid="A7">7</xref>
<xref ref-type="aff" rid="A21">21</xref>
<xref ref-type="aff" rid="A23">23</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Schmutz</surname>
<given-names>Jeremy</given-names>
</name>
<xref ref-type="aff" rid="A2">2</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Morgante</surname>
<given-names>Michele</given-names>
</name>
<xref ref-type="aff" rid="A5">5</xref>
<xref ref-type="aff" rid="A24">24</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Machado</surname>
<given-names>Marcos Antonio</given-names>
</name>
<xref ref-type="aff" rid="A6">6</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Talon</surname>
<given-names>Manuel</given-names>
</name>
<xref ref-type="aff" rid="A8">8</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Jaillon</surname>
<given-names>Olivier</given-names>
</name>
<xref ref-type="aff" rid="A7">7</xref>
<xref ref-type="aff" rid="A21">21</xref>
<xref ref-type="aff" rid="A23">23</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Ollitrault</surname>
<given-names>Patrick</given-names>
</name>
<xref ref-type="aff" rid="A4">4</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Gmitter</surname>
<given-names>Frederick</given-names>
</name>
<xref ref-type="aff" rid="A13">13</xref>
<xref ref-type="corresp" rid="cor1">*</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Rokhsar</surname>
<given-names>Daniel</given-names>
</name>
<xref ref-type="aff" rid="A1">1</xref>
<xref ref-type="aff" rid="A25">25</xref>
<xref ref-type="corresp" rid="cor1">*</xref>
</contrib>
</contrib-group>
<aff id="A1">
<label>1</label>
US-Department of Energy Joint Genome Institute, Walnut Creek, CA, USA</aff>
<aff id="A2">
<label>2</label>
HudsonAlpha Biotechnology Institute, Huntsville, AL, USA</aff>
<aff id="A3">
<label>3</label>
INRA/UBP UMR 1095 GDEC, Clermont Ferrand, France</aff>
<aff id="A4">
<label>4</label>
CIRAD, UMR AGAP, Montpellier, France</aff>
<aff id="A5">
<label>5</label>
Istituto di Genomica Applicata, Udine, Italy</aff>
<aff id="A6">
<label>6</label>
Centro de Citricultura Sylvio Moreira, IAC, Cordeirópolis, SP, Brazil</aff>
<aff id="A7">
<label>7</label>
Commissariat à l'Energie Atomique (CEA), Institut de Génomique (IG), Genoscope, Evry, France</aff>
<aff id="A8">
<label>8</label>
Centro de Genomica, Instituto Valenciano de Investigaciones Agrarias (IVIA), Valencia, Spain</aff>
<aff id="A9">
<label>9</label>
Centro de Protección Vegetal y Biotecnología-IVIA, Moncada, Valencia, Spain</aff>
<aff id="A10">
<label>10</label>
Lifesequencing SL, Valencia, Spain</aff>
<aff id="A11">
<label>11</label>
INRA, US EPGV_1279, Evry, France</aff>
<aff id="A12">
<label>12</label>
INRA GEQA, San Giuliano, France</aff>
<aff id="A13">
<label>13</label>
Citrus Research and Education Center (CREC), Institute of Food and Agricultural Sciences (IFAS), University of Florida, Lake Alfred, FL, USA</aff>
<aff id="A14">
<label>14</label>
Interdisciplinary Center for Biotechnology Research, University of Florida, Gainesville, FL, USA</aff>
<aff id="A15">
<label>15</label>
454 Life Sciences, A Roche Company, 15 Commercial Street, Branford CT, USA</aff>
<aff id="A16">
<label>16</label>
Wallace H. Coulter Department of Biomedical Engineering, Georgia Institute of Technology, Atlanta, GA, USA</aff>
<aff id="A17">
<label>17</label>
School of Computational Science & Engineering, Georgia Institute of Technology, Atlanta, GA, USA</aff>
<aff id="A18">
<label>18</label>
Department of Biological and Medical Physics, Moscow Institute of Physics and Technology, Dolgoprudny, Moscow Region, Russia</aff>
<aff id="A19">
<label>19</label>
Consiglio per la Ricerca e la Sperimentazione in Agricoltura (CRA-ACM), Acireale, Italy</aff>
<aff id="A20">
<label>20</label>
Embrapa Cassava and Fruits, Cruz das Almas, BA, Brazil</aff>
<aff id="A21">
<label>21</label>
Département de Biologie, Université d’Evry, Evry, France</aff>
<aff id="A22">
<label>22</label>
Department of Botany and Plant Sciences, University of California, Riverside, CA, USA</aff>
<aff id="A23">
<label>23</label>
Centre National de Recherche Scientifique (CNRS), Evry, France</aff>
<aff id="A24">
<label>24</label>
Department of Agriculture and Environmental Sciences, University of Udine, Udine, Italy</aff>
<aff id="A25">
<label>25</label>
Division of Genetics, Genomics, and Development, University of California, Berkeley, CA, USA</aff>
<aff id="A26">
<label>26</label>
Institute of Life Sciences, Scuola Superiore Sant'Anna, Pisa, Italy</aff>
<aff id="A27">
<label>27</label>
Secugen SL, Madrid, Spain</aff>
<author-notes>
<corresp id="cor1">
<label>*</label>
Correspondence to: D.S.R. (
<email>dsrokhsar@gmail.com</email>
); F.G.G. (
<email>fgmitter@ufl.edu</email>
)</corresp>
<fn id="FN1" fn-type="present-address">
<label></label>
<p id="P1">Current address: Life Technologies Corp., Grand Island, NY 14072, USA.</p>
</fn>
<fn id="FN2" fn-type="present-address">
<label></label>
<p id="P2">Current address: USDA, ARS, Southeastern Fruit and Tree Nut Research Laboratory, Byron, GA, USA</p>
</fn>
<fn id="FN3" fn-type="equal">
<label></label>
<p id="P3">These authors contributed equally.</p>
</fn>
</author-notes>
<pub-date pub-type="nihms-submitted">
<day>21</day>
<month>5</month>
<year>2014</year>
</pub-date>
<pub-date pub-type="epub">
<day>08</day>
<month>6</month>
<year>2014</year>
</pub-date>
<pub-date pub-type="ppub">
<month>7</month>
<year>2014</year>
</pub-date>
<pub-date pub-type="pmc-release">
<day>01</day>
<month>1</month>
<year>2015</year>
</pub-date>
<volume>32</volume>
<issue>7</issue>
<fpage>656</fpage>
<lpage>662</lpage>
<pmc-comment>elocation-id from pubmed: 10.1038/nbt.2906</pmc-comment>
<permissions>
<license xlink:href="http://www.nature.com/authors/editorial_policies/license.html#terms">
<license-p>Users may view, print, copy, and download text and data-mine the content in such documents, for the purposes of academic research, subject always to the full Conditions of use:
<ext-link ext-link-type="uri" xlink:href="http://www.nature.com/authors/editorial_policies/license.html#terms">http://www.nature.com/authors/editorial_policies/license.html#terms</ext-link>
</license-p>
</license>
</permissions>
<abstract>
<p id="P4">The domestication of citrus, is poorly understood. Cultivated types are selections from, or hybrids of, wild progenitor species, whose identities and contributions remain controversial. By comparative analysis of a collection of citrus genomes, including a high quality haploid reference, we show that cultivated types were derived from two progenitor species. Though cultivated pummelos represent selections from a single progenitor species,
<italic>C. maxima</italic>
, cultivated mandarins are introgressions of
<italic>C. maxima</italic>
into the ancestral mandarin species,
<italic>C. reticulata</italic>
. The most widely cultivated citrus, sweet orange, is the offspring of previously admixed individuals, but sour orange is an F1 hybrid of pure
<italic>C. maxima</italic>
and
<italic>C. reticulata</italic>
parents, implying that wild mandarins were part of the early breeding germplasm. A wild “mandarin” from China exhibited substantial divergence from
<italic>C. reticulata</italic>
, suggesting the possibility of other unrecognized wild citrus species. Understanding citrus phylogeny through genome analysis clarifies taxonomic relationships and enables sequence-directed genetic improvement.</p>
</abstract>
</article-meta>
</front>
<floats-group>
<fig id="F1" orientation="portrait" position="float">
<label>Figure 1</label>
<caption>
<p id="P40">A selection of mandarin, pummelo and orange fruits, including cultivars sequenced in this study. Pummelos (numbered 1, 2 in outline, on left) are large trees that produce very large fruit, with white, pink or red flesh color (2) and yellow or pink rinds. Most cultivars have large leaves having petioles with prominent wings. Apomictic reproduction is absent and most selections are self-incompatible. Mandarins (3–7) are smaller trees bearing smaller fruit, with orange flesh (9, 11) and rind color. Mandarins have both apomictic and zygotic reproduction and some are self-compatible. Oranges (8, 10) are generally intermediate in tree and fruit size, flesh (10) and rind color is commonly orange, and apomictic reproduction is always present. (The sour orange shown (12) is immature.)</p>
</caption>
<graphic xlink:href="nihms586218f1"></graphic>
</fig>
<fig id="F2" orientation="portrait" position="float">
<label>Figure 2</label>
<caption>
<p id="P41">Nucleotide diversity distribution in citrus.(
<bold>a</bold>
) Nucleotide heterozygosity distribution computed in overlapping 100kb windows (with 5 kb step size) across the Low acid (LAP) and Chandler (CHP) pummelo genomes and between the non-shared haplotypes of this parent-child pair (LAP/CHP) is shown. The peak at ~6 heterozygous sites/kb in all three pairwise comparisons represents the characteristic nucleotide diversity of the species
<italic>C. maxima</italic>
; the peak near ~1 heterozygous site/kb reflects a bottleneck in the ancestral
<italic>C. maxima</italic>
population after divergence from
<italic>C. reticulata</italic>
(
<xref ref-type="supplementary-material" rid="SD1">Supplementary Note 10</xref>
).
<bold>(b)</bold>
Nucleotide heterozygosity for the traditional Willowleaf mandarin (WLM) plotted along chromosome 6, computed in overlapping windows of 200 kb (with 100 kb step size). This chromosome shows an example of the clear discontinuity in single nucleotide variant heterozygosity levels between ~5/kb in the M/M segment (orange bar) and ~17/kb in the M/P segment (blue bar).
<bold>(c)</bold>
Nucleotide heterozygosity distribution computed in overlapping 500kb windows (with 5 kb step size) in Ponkan (PKM, solid line) and Willowleaf (WLM, dashed line) mandarins. Genomic segments are designated M/M, M/P or P/P based on a set of 1,537,264 SNPs that differentiate
<italic>C. reticulata</italic>
(M) from C.
<italic>maxima</italic>
(P). Both mandarins contain admixed segments from
<italic>C. maxima</italic>
introgression (M/P) as well as M/M segments, and these are plotted and normalized separately for easy comparison..
<bold>(d)</bold>
Nucleotide heterozygosity distribution computed in overlapping windows of 500kb (5 kb offsets) for sweet orange (SWO) and sour orange (SSO). The three different genotypes of the SWO genome (M/M, P/P and M/P), and the SSO genotype M/P are normalized and plotted separately</p>
</caption>
<graphic xlink:href="nihms586218f2"></graphic>
</fig>
<fig id="F3" orientation="portrait" position="float">
<label>Figure 3</label>
<caption>
<p id="P42">Admixture patterns and nucleotide diversity in cultivated citrus. For each of the three groups of sequenced citrus, variation in nucleotide diversity (averaged over 500kb windows with step size 250kb) is shown across the genome for one representative cultivar above genotype maps (horizontal bars: green =
<italic>C. maxima/C. maxima</italic>
; blue =
<italic>C. maxima/C. reticulata</italic>
; orange=
<italic>C. reticulata/C. reticulata</italic>
; grey=unknown; the 9 chromosomes are numbered at the top).
<bold>(a)</bold>
Sweet orange (SWO) nucleotide diversity with genotype maps for SWO and sour orange (SSO). Note the
<italic>C. maxima/C. maxima</italic>
genotype (green segments present on chromosomes 2 and 8) in SWO.
<bold>(b)</bold>
Willowleaf mandarin (WLM) nucleotide diversity and genotype maps for three traditional mandarins (Ponkan mandarin (PKM), WLM, Huanglingmiao (HLM)) and three recent mandarin types (Clementine (CLM), W. Murcott mandarin (WMM), haploid Clementine reference (HCR)). For the haploid Clementine reference sequence (HCR), red and green segments indicate
<italic>C.reticulata</italic>
and
<italic>C. maxima</italic>
haplotypes, respectively. All five mandarin types show pummelo introgressions (blue or green segments).
<bold>(c)</bold>
Low acid pummelo (LAP) nucleotide diversity and genotype maps for two pummelos (LAP, Chandler pummelo (CHP)).</p>
</caption>
<graphic xlink:href="nihms586218f3"></graphic>
</fig>
<fig id="F4" orientation="portrait" position="float">
<label>Figure 4</label>
<caption>
<title>Mangshan mandarin is a species distinct from
<italic>C. maxima</italic>
and
<italic>C. reticulata</italic>
</title>
<p id="P43">(
<bold>a</bold>
) Midpoint-rooted neighbor-joining phylogenetic tree of citrus chloroplast genomes. (
<bold>b</bold>
) The frequency distributions of the pairwise sequence divergences (across 100 kb windows) between Mangshan mandarin (CMS) and
<italic>C. maxima</italic>
(green), CMS and
<italic>C. reticulata</italic>
(orange),
<italic>C. reticulata</italic>
and
<italic>C. maxima</italic>
(light blue), as well as the distinctly lower CMS intrinsic nucleotide diversity (dashed blue). (c) The first two coordinates of principal coordinate analysis of the citrus nuclear genomes, based on pairwise distances and the metric multidimensional scaling. The
<italic>C. maxima</italic>
-
<italic>C. reticulata</italic>
axis (Principle coordinate 1, 47.5% variance) separates pummelos (green) from mandarins (orange), with oranges (blue) lying in between; Principle coordinate 2 (19.6% of variance) separates CMS (purple) from the others.</p>
</caption>
<graphic xlink:href="nihms586218f4"></graphic>
</fig>
<table-wrap id="T1" position="float" orientation="landscape">
<label>Table 1</label>
<caption>
<title>Sequenced cultivars and proportions derived from the ancestral species
<italic>C. reticulata</italic>
and
<italic>C. maxima</italic>
</title>
<p id="P44">Three letter abbreviations as used throughout this work and common systematic designation are shown. Sequence depth reported as count of aligned reads to reference, after removal of duplicate reads. Chloroplast genome type inferred from shotgun reads aligning to the sweet orange chloroplast genome
<sup>
<xref rid="R38" ref-type="bibr">38</xref>
</sup>
, with M indicating mandarin type and P indicating pummelo type. Diploid nuclear genotype proportions refer to fraction of genome in megabases using the HCR physical map (proportions of unknown genotype are not shown but can be inferred by subtracting the three genotype proportions from 100%). The last two columns show proportions of
<italic>C. maxima</italic>
and
<italic>C. reticulata</italic>
haplotypes, and are derived from the three genotype proportions. max. =
<italic>C. maxima</italic>
; ret. =
<italic>C.reticulata</italic>
.</p>
</caption>
<table frame="box" rules="all">
<thead>
<tr>
<th align="left" valign="top" rowspan="1" colspan="1">Cultivar</th>
<th align="left" valign="top" rowspan="1" colspan="1">Abbr.</th>
<th align="left" valign="top" rowspan="1" colspan="1">Common
<break></break>
designation</th>
<th align="left" valign="top" rowspan="1" colspan="1">Sequence
<break></break>
generated</th>
<th align="center" valign="top" rowspan="1" colspan="1">Cp
<break></break>
type</th>
<th align="center" valign="top" rowspan="1" colspan="1">ret./
<break></break>
ret.</th>
<th align="center" valign="top" rowspan="1" colspan="1">ret./
<break></break>
max.</th>
<th align="center" valign="top" rowspan="1" colspan="1">max./
<break></break>
max.</th>
<th align="center" valign="top" rowspan="1" colspan="1">ret.</th>
<th align="center" valign="top" rowspan="1" colspan="1">max.</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left" rowspan="1" colspan="1">Haploid Clementine</td>
<td align="left" rowspan="1" colspan="1">HCR</td>
<td align="left" rowspan="1" colspan="1">
<italic>C</italic>
. ×
<italic>clementina</italic>
</td>
<td align="left" rowspan="1" colspan="1">7× Sanger</td>
<td align="center" rowspan="1" colspan="1">M</td>
<td align="center" rowspan="1" colspan="1">n/a</td>
<td align="center" rowspan="1" colspan="1">n/a</td>
<td align="center" rowspan="1" colspan="1">n/a</td>
<td align="center" rowspan="1" colspan="1">89%</td>
<td align="center" rowspan="1" colspan="1">11%</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">Clementine mandarin</td>
<td align="left" rowspan="1" colspan="1">CLM</td>
<td align="left" rowspan="1" colspan="1">
<italic>C</italic>
. ×
<italic>clementina</italic>
</td>
<td align="left" rowspan="1" colspan="1">110× Illumina</td>
<td align="center" rowspan="1" colspan="1">M</td>
<td align="center" rowspan="1" colspan="1">58%</td>
<td align="center" rowspan="1" colspan="1">42%</td>
<td align="center" rowspan="1" colspan="1">0%</td>
<td align="center" rowspan="1" colspan="1">79%</td>
<td align="center" rowspan="1" colspan="1">21%</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">Ponkan mandarin</td>
<td align="left" rowspan="1" colspan="1">PKM</td>
<td align="left" rowspan="1" colspan="1">
<italic>C</italic>
.
<italic>reticulata</italic>
<xref ref-type="table-fn" rid="TFN1">*</xref>
</td>
<td align="left" rowspan="1" colspan="1">55× Illumina</td>
<td align="center" rowspan="1" colspan="1">M</td>
<td align="center" rowspan="1" colspan="1">85%</td>
<td align="center" rowspan="1" colspan="1">14%</td>
<td align="center" rowspan="1" colspan="1">0.7%</td>
<td align="center" rowspan="1" colspan="1">92%</td>
<td align="center" rowspan="1" colspan="1">8%</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">Willowleaf mandarin</td>
<td align="left" rowspan="1" colspan="1">WLM</td>
<td align="left" rowspan="1" colspan="1">
<italic>C</italic>
. ×
<italic>deliciosa</italic>
</td>
<td align="left" rowspan="1" colspan="1">110× Illumina</td>
<td align="center" rowspan="1" colspan="1">M</td>
<td align="center" rowspan="1" colspan="1">91%</td>
<td align="center" rowspan="1" colspan="1">8.8%</td>
<td align="center" rowspan="1" colspan="1">0%</td>
<td align="center" rowspan="1" colspan="1">95%</td>
<td align="center" rowspan="1" colspan="1">4.4%</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">W. Murcott mandarin</td>
<td align="left" rowspan="1" colspan="1">WMM</td>
<td align="left" rowspan="1" colspan="1">
<italic>C. reticulata</italic>
</td>
<td align="left" rowspan="1" colspan="1">25× Illumina</td>
<td align="center" rowspan="1" colspan="1">M</td>
<td align="center" rowspan="1" colspan="1">69%</td>
<td align="center" rowspan="1" colspan="1">30%</td>
<td align="center" rowspan="1" colspan="1">0.4%</td>
<td align="center" rowspan="1" colspan="1">85%</td>
<td align="center" rowspan="1" colspan="1">15%</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">Chandler pummelo</td>
<td align="left" rowspan="1" colspan="1">CHP</td>
<td align="left" rowspan="1" colspan="1">
<italic>C. maxima</italic>
</td>
<td align="left" rowspan="1" colspan="1">22× Illumina</td>
<td align="center" rowspan="1" colspan="1">P</td>
<td align="center" rowspan="1" colspan="1">0%</td>
<td align="center" rowspan="1" colspan="1">0.4%</td>
<td align="center" rowspan="1" colspan="1">99.6%</td>
<td align="center" rowspan="1" colspan="1">0.2%</td>
<td align="center" rowspan="1" colspan="1">99.8%</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">Low acid pummelo</td>
<td align="left" rowspan="1" colspan="1">LAP</td>
<td align="left" rowspan="1" colspan="1">
<italic>C. maxima</italic>
</td>
<td align="left" rowspan="1" colspan="1">17× Illumina</td>
<td align="center" rowspan="1" colspan="1">P</td>
<td align="center" rowspan="1" colspan="1">0%</td>
<td align="center" rowspan="1" colspan="1">0%</td>
<td align="center" rowspan="1" colspan="1">100%</td>
<td align="center" rowspan="1" colspan="1">0%</td>
<td align="center" rowspan="1" colspan="1">100%</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">Sweet orange</td>
<td align="left" rowspan="1" colspan="1">SWO</td>
<td align="left" rowspan="1" colspan="1">
<italic>C</italic>
. ×
<italic>sinensis</italic>
</td>
<td align="left" rowspan="1" colspan="1">80× Illumina</td>
<td align="center" rowspan="1" colspan="1">P</td>
<td align="center" rowspan="1" colspan="1">14%</td>
<td align="center" rowspan="1" colspan="1">82%</td>
<td align="center" rowspan="1" colspan="1">3%</td>
<td align="center" rowspan="1" colspan="1">55%</td>
<td align="center" rowspan="1" colspan="1">44%</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">Seville sour orange</td>
<td align="left" rowspan="1" colspan="1">SSO</td>
<td align="left" rowspan="1" colspan="1">
<italic>C</italic>
. ×
<italic>aurantium</italic>
</td>
<td align="left" rowspan="1" colspan="1">36× Illumina</td>
<td align="center" rowspan="1" colspan="1">P</td>
<td align="center" rowspan="1" colspan="1">0%</td>
<td align="center" rowspan="1" colspan="1">98%</td>
<td align="center" rowspan="1" colspan="1">0%</td>
<td align="center" rowspan="1" colspan="1">49%</td>
<td align="center" rowspan="1" colspan="1">49%</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn id="TFN1">
<label>*</label>
<p id="P45">Ponkan mandarin is widely assumed to represent
<italic>C. reticulata</italic>
, but as shown here it has substantial admixture from
<italic>C. maxima</italic>
.</p>
</fn>
</table-wrap-foot>
</table-wrap>
</floats-group>
</pmc>
<affiliations>
<list>
<country>
<li>Brésil</li>
<li>Espagne</li>
<li>France</li>
<li>Italie</li>
<li>Russie</li>
<li>États-Unis</li>
</country>
<region>
<li>Bahia</li>
<li>Communauté de Madrid</li>
<li>Toscane</li>
<li>État de São Paulo</li>
</region>
<settlement>
<li>Clermont-Ferrand</li>
<li>Madrid</li>
<li>Montpellier</li>
<li>Pise</li>
<li>Évry (Essonne)</li>
</settlement>
</list>
<tree>
<country name="États-Unis">
<noRegion>
<name sortKey="Wu, G Albert" sort="Wu, G Albert" uniqKey="Wu G" first="G. Albert" last="Wu">G. Albert Wu</name>
</noRegion>
<name sortKey="Borodovsky, Mark" sort="Borodovsky, Mark" uniqKey="Borodovsky M" first="Mark" last="Borodovsky">Mark Borodovsky</name>
<name sortKey="Borodovsky, Mark" sort="Borodovsky, Mark" uniqKey="Borodovsky M" first="Mark" last="Borodovsky">Mark Borodovsky</name>
<name sortKey="Burns, Paul" sort="Burns, Paul" uniqKey="Burns P" first="Paul" last="Burns">Paul Burns</name>
<name sortKey="Burns, Paul" sort="Burns, Paul" uniqKey="Burns P" first="Paul" last="Burns">Paul Burns</name>
<name sortKey="Chapman, Jarrod" sort="Chapman, Jarrod" uniqKey="Chapman J" first="Jarrod" last="Chapman">Jarrod Chapman</name>
<name sortKey="Chen, Chunxian" sort="Chen, Chunxian" uniqKey="Chen C" first="Chunxian" last="Chen">Chunxian Chen</name>
<name sortKey="Desany, Brian" sort="Desany, Brian" uniqKey="Desany B" first="Brian" last="Desany">Brian Desany</name>
<name sortKey="Farmerie, William G" sort="Farmerie, William G" uniqKey="Farmerie W" first="William G." last="Farmerie">William G. Farmerie</name>
<name sortKey="Fredrikson, Karin" sort="Fredrikson, Karin" uniqKey="Fredrikson K" first="Karin" last="Fredrikson">Karin Fredrikson</name>
<name sortKey="Gmitter, Frederick" sort="Gmitter, Frederick" uniqKey="Gmitter F" first="Frederick" last="Gmitter">Frederick Gmitter</name>
<name sortKey="Grimwood, Jane" sort="Grimwood, Jane" uniqKey="Grimwood J" first="Jane" last="Grimwood">Jane Grimwood</name>
<name sortKey="Harkins, Tim" sort="Harkins, Tim" uniqKey="Harkins T" first="Tim" last="Harkins">Tim Harkins</name>
<name sortKey="Hellsten, Uffe" sort="Hellsten, Uffe" uniqKey="Hellsten U" first="Uffe" last="Hellsten">Uffe Hellsten</name>
<name sortKey="Jenkins, Jerry" sort="Jenkins, Jerry" uniqKey="Jenkins J" first="Jerry" last="Jenkins">Jerry Jenkins</name>
<name sortKey="Kodira, Chinnappa" sort="Kodira, Chinnappa" uniqKey="Kodira C" first="Chinnappa" last="Kodira">Chinnappa Kodira</name>
<name sortKey="Lomsadze, Alexandre" sort="Lomsadze, Alexandre" uniqKey="Lomsadze A" first="Alexandre" last="Lomsadze">Alexandre Lomsadze</name>
<name sortKey="Lomsadze, Alexandre" sort="Lomsadze, Alexandre" uniqKey="Lomsadze A" first="Alexandre" last="Lomsadze">Alexandre Lomsadze</name>
<name sortKey="Mohiuddin, Mohammed" sort="Mohiuddin, Mohammed" uniqKey="Mohiuddin M" first="Mohammed" last="Mohiuddin">Mohammed Mohiuddin</name>
<name sortKey="Prochnik, Simon" sort="Prochnik, Simon" uniqKey="Prochnik S" first="Simon" last="Prochnik">Simon Prochnik</name>
<name sortKey="Rokhsar, Daniel" sort="Rokhsar, Daniel" uniqKey="Rokhsar D" first="Daniel" last="Rokhsar">Daniel Rokhsar</name>
<name sortKey="Rokhsar, Daniel" sort="Rokhsar, Daniel" uniqKey="Rokhsar D" first="Daniel" last="Rokhsar">Daniel Rokhsar</name>
<name sortKey="Roose, Mikeal" sort="Roose, Mikeal" uniqKey="Roose M" first="Mikeal" last="Roose">Mikeal Roose</name>
<name sortKey="Schmutz, Jeremy" sort="Schmutz, Jeremy" uniqKey="Schmutz J" first="Jeremy" last="Schmutz">Jeremy Schmutz</name>
</country>
<country name="France">
<noRegion>
<name sortKey="Salse, Jerome" sort="Salse, Jerome" uniqKey="Salse J" first="Jerome" last="Salse">Jerome Salse</name>
</noRegion>
<name sortKey="Brunel, Dominique" sort="Brunel, Dominique" uniqKey="Brunel D" first="Dominique" last="Brunel">Dominique Brunel</name>
<name sortKey="Brunel, Dominique" sort="Brunel, Dominique" uniqKey="Brunel D" first="Dominique" last="Brunel">Dominique Brunel</name>
<name sortKey="Couloux, Arnaud" sort="Couloux, Arnaud" uniqKey="Couloux A" first="Arnaud" last="Couloux">Arnaud Couloux</name>
<name sortKey="Jabbari, Kamel" sort="Jabbari, Kamel" uniqKey="Jabbari K" first="Kamel" last="Jabbari">Kamel Jabbari</name>
<name sortKey="Jaillon, Olivier" sort="Jaillon, Olivier" uniqKey="Jaillon O" first="Olivier" last="Jaillon">Olivier Jaillon</name>
<name sortKey="Jaillon, Olivier" sort="Jaillon, Olivier" uniqKey="Jaillon O" first="Olivier" last="Jaillon">Olivier Jaillon</name>
<name sortKey="Jaillon, Olivier" sort="Jaillon, Olivier" uniqKey="Jaillon O" first="Olivier" last="Jaillon">Olivier Jaillon</name>
<name sortKey="Labadie, Karine" sort="Labadie, Karine" uniqKey="Labadie K" first="Karine" last="Labadie">Karine Labadie</name>
<name sortKey="Luro, Francois" sort="Luro, Francois" uniqKey="Luro F" first="François" last="Luro">François Luro</name>
<name sortKey="Murat, Florent" sort="Murat, Florent" uniqKey="Murat F" first="Florent" last="Murat">Florent Murat</name>
<name sortKey="Ollitrault, Patrick" sort="Ollitrault, Patrick" uniqKey="Ollitrault P" first="Patrick" last="Ollitrault">Patrick Ollitrault</name>
<name sortKey="Perrier, Xavier" sort="Perrier, Xavier" uniqKey="Perrier X" first="Xavier" last="Perrier">Xavier Perrier</name>
<name sortKey="Poulain, Julie" sort="Poulain, Julie" uniqKey="Poulain J" first="Julie" last="Poulain">Julie Poulain</name>
<name sortKey="Quetier, Francis" sort="Quetier, Francis" uniqKey="Quetier F" first="Francis" last="Quetier">Francis Quetier</name>
<name sortKey="Quetier, Francis" sort="Quetier, Francis" uniqKey="Quetier F" first="Francis" last="Quetier">Francis Quetier</name>
<name sortKey="Ruiz, Manuel" sort="Ruiz, Manuel" uniqKey="Ruiz M" first="Manuel" last="Ruiz">Manuel Ruiz</name>
<name sortKey="Wincker, Patrick" sort="Wincker, Patrick" uniqKey="Wincker P" first="Patrick" last="Wincker">Patrick Wincker</name>
<name sortKey="Wincker, Patrick" sort="Wincker, Patrick" uniqKey="Wincker P" first="Patrick" last="Wincker">Patrick Wincker</name>
<name sortKey="Wincker, Patrick" sort="Wincker, Patrick" uniqKey="Wincker P" first="Patrick" last="Wincker">Patrick Wincker</name>
</country>
<country name="Italie">
<noRegion>
<name sortKey="Scalabrin, Simone" sort="Scalabrin, Simone" uniqKey="Scalabrin S" first="Simone" last="Scalabrin">Simone Scalabrin</name>
</noRegion>
<name sortKey="Cattonaro, Federica" sort="Cattonaro, Federica" uniqKey="Cattonaro F" first="Federica" last="Cattonaro">Federica Cattonaro</name>
<name sortKey="Del Fabbro, Cristian" sort="Del Fabbro, Cristian" uniqKey="Del Fabbro C" first="Cristian" last="Del Fabbro">Cristian Del Fabbro</name>
<name sortKey="Morgante, Michele" sort="Morgante, Michele" uniqKey="Morgante M" first="Michele" last="Morgante">Michele Morgante</name>
<name sortKey="Morgante, Michele" sort="Morgante, Michele" uniqKey="Morgante M" first="Michele" last="Morgante">Michele Morgante</name>
<name sortKey="Pinosio, Sara" sort="Pinosio, Sara" uniqKey="Pinosio S" first="Sara" last="Pinosio">Sara Pinosio</name>
<name sortKey="Reforgiato, Giuseppe" sort="Reforgiato, Giuseppe" uniqKey="Reforgiato G" first="Giuseppe" last="Reforgiato">Giuseppe Reforgiato</name>
<name sortKey="Zuccolo, Andrea" sort="Zuccolo, Andrea" uniqKey="Zuccolo A" first="Andrea" last="Zuccolo">Andrea Zuccolo</name>
<name sortKey="Zuccolo, Andrea" sort="Zuccolo, Andrea" uniqKey="Zuccolo A" first="Andrea" last="Zuccolo">Andrea Zuccolo</name>
</country>
<country name="Espagne">
<noRegion>
<name sortKey="Terol, Javier" sort="Terol, Javier" uniqKey="Terol J" first="Javier" last="Terol">Javier Terol</name>
</noRegion>
<name sortKey="Aleza, Pablo" sort="Aleza, Pablo" uniqKey="Aleza P" first="Pablo" last="Aleza">Pablo Aleza</name>
<name sortKey="Estornell, Leandro H" sort="Estornell, Leandro H" uniqKey="Estornell L" first="Leandro H." last="Estornell">Leandro H. Estornell</name>
<name sortKey="Herrero Ortega, Amparo" sort="Herrero Ortega, Amparo" uniqKey="Herrero Ortega A" first="Amparo" last="Herrero-Ortega">Amparo Herrero-Ortega</name>
<name sortKey="Ibanez, Victoria" sort="Ibanez, Victoria" uniqKey="Ibanez V" first="Victoria" last="Ibanez">Victoria Ibanez</name>
<name sortKey="Mu Oz Sanz, Juan V" sort="Mu Oz Sanz, Juan V" uniqKey="Mu Oz Sanz J" first="Juan V." last="Mu Oz-Sanz">Juan V. Mu Oz-Sanz</name>
<name sortKey="Navarro, Luis" sort="Navarro, Luis" uniqKey="Navarro L" first="Luis" last="Navarro">Luis Navarro</name>
<name sortKey="Perez Perez, Julian" sort="Perez Perez, Julian" uniqKey="Perez Perez J" first="Julián" last="Pérez-Pérez">Julián Pérez-Pérez</name>
<name sortKey="Perez Perez, Julian" sort="Perez Perez, Julian" uniqKey="Perez Perez J" first="Julián" last="Pérez-Pérez">Julián Pérez-Pérez</name>
<name sortKey="Ram N, Daniel" sort="Ram N, Daniel" uniqKey="Ram N D" first="Daniel" last="Ram N">Daniel Ram N</name>
<name sortKey="Tadeo, Francisco R" sort="Tadeo, Francisco R" uniqKey="Tadeo F" first="Francisco R." last="Tadeo">Francisco R. Tadeo</name>
<name sortKey="Talon, Manuel" sort="Talon, Manuel" uniqKey="Talon M" first="Manuel" last="Talon">Manuel Talon</name>
</country>
<country name="Brésil">
<region name="État de São Paulo">
<name sortKey="Takita, Marco Aurelio" sort="Takita, Marco Aurelio" uniqKey="Takita M" first="Marco Aurélio" last="Takita">Marco Aurélio Takita</name>
</region>
<name sortKey="Freitas Astua, Juliana" sort="Freitas Astua, Juliana" uniqKey="Freitas Astua J" first="Juliana" last="Freitas-Astúa">Juliana Freitas-Astúa</name>
<name sortKey="Freitas Astua, Juliana" sort="Freitas Astua, Juliana" uniqKey="Freitas Astua J" first="Juliana" last="Freitas-Astúa">Juliana Freitas-Astúa</name>
<name sortKey="Machado, Marcos Antonio" sort="Machado, Marcos Antonio" uniqKey="Machado M" first="Marcos Antonio" last="Machado">Marcos Antonio Machado</name>
</country>
<country name="Russie">
<noRegion>
<name sortKey="Borodovsky, Mark" sort="Borodovsky, Mark" uniqKey="Borodovsky M" first="Mark" last="Borodovsky">Mark Borodovsky</name>
</noRegion>
</country>
</tree>
</affiliations>
</record>

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