1/24/2017 The origin of citrus Guohong Albert Wu, Javier Terol, Victoria Ibanez, Antonio López-García, Estela GENOMICS ILLUMINATES THE ORIGIN AND Pérez-Román, Carles Borredá, Concha Domingo, Francisco R. Tadeo, José Carbonell-Caballero, Roberto Alonso, Franck Curk, Dongliang Du, Patrick Ollitrault, DISPERSAL OF CITRUS Mikeal L. Roose , Joaquin Dopazo, Frederick G. Gmitter Jr., Daniel S. Rokhsar and Manuel Talon Instituto Valenciano de Investigaciones Agrarias (IVIA) Pure species, admixtures and hybrids in the Genus Citrus Pure species, admixtures and hybrids in the Genus Citrus Common name Species name Common name Species name Corsican citron C. medica L. Guan-xi-mi-you pummelo C. maxima [(Burm.) Merr]. Mac Veu citron C. medica L C. grandis (Swingle, Tanaka) Citrus lumia Risso & Poit . Sha-tian-you pummelo C. maxima [(Burm.) Merr]. PURE SPECIES HYBRIDS ADMIXTURES Buddha’s hand citron C. medica L. (Noot.) Swingle C. grandis (Swingle, Tanaka) var. Sarcodactylus Grapefruit, C. x paradisi (Macfadyen) Citrons Sour oranges Mandarins Humpang citron C. medica L. cv. Marsh Sun Chu Sha Kat mandarin C. reticulata (Blanco) C. reticulata var. Sweet orange, C. x sinensis L. (Osbeck) Pummelos Rangpur limes Sweet oranges austera (Swingle) C. erythrosa (Tanaka) cv. Washington Navel Tachibana mandarins Rough limes Sour orange, C. x aurantium L. Grapefruits Tachibana mandarin C. tachibana (Mak.) Tan cv.Sevillano C. reticulata (Blanco) Sun Chu Sha Kat mandarins Mexican limes Lemons Lemon, cv.Eureka C. x limon L. (Burm. f.) Sunki mandarin C. sunki (Hayata, Hort. ex Tanaka Eremorages Australian desert lime Eremocitrus glauca (Lindl.) Swingle Australian desert limes (sour mandarin, suanju) C. reticulata (Blanco) C. * glauca (Lindl.) Burkill Calamondins Cleopatra mandarin C. reshni (Hort. ex Tanaka) Eremocitrus glauca (Lindl.) Swingle Australian finger limes C. reticulata (Blanco) Eremorange. Eremocitrus glauca x Citrus sinnensis. Australian round limes Changsha mandarin Citrus reticulata (Blanco) Australian desert lime hybrid Huanglingmiao mandarin C.* reticulata (Hort. ex Tanaka) Australian finger lime Microcitrus australasica (F.Muell.) Swingle Kumquats Satsuma (unshiu) mandarin, C. * unshiu [(Mak.) Marc] C. australasica F. Muell. cv. Owari C. *reticulata (Swingle) Micranthas Ponkan C. *reticulata (Blanco, Swingle) Australian finger lime Microcitrus australasica (F.Muell.) Swingle Mangshan mandarins (Chinese honey orange) C. australasica F. Muell Dancy mandarin, C. * tangerina (Tanaka) Papedas Dancy tangerine C. *reticulata (Swingle) Australian round lime Microcitrus australis Swingle Willowleaf mandarin C. deliciosa (Ten. Hort. ex Tanaka) C. australis (A. Cunn. ex Mudie) Clementine mandarin, C. x clementina (Hort. ex Tanaka) Australian round lime Microcitrus australis Swingle Poncirus cv. Clementina de Nules C.x reticulata (Swingle) C. australis (A. Cunn. ex Mudie) King mandarin C. * nobilis (Lour.) Kumquat, Nagami Fortunella margarita (Lour.) Swingle C. *reticulata (Swingle) Calamondin C. madurensis (Lour.) W. Murcott mandarin C. reticulata (Blanco) Mexican/key lime C. * aurantifolia (Christm.) Swingle Rangpur lime C. x limonia (Osbeck) Micrantha, Biasong C. micrantha (Wester) Mangshan mandarin C. mangshanensis Red rough lemon C x jambhiri (Lush) Ichang papeda C. ichangensis (Swingle) Low acid pummelo C. maxima [(Burm.) Merr]. (Siamese Sweet) C. grandis (Swingle, Tanaka) Trifoliate orange Poncirus trifoliata (L.) Raf. Chinese box orange Severinia buxifolia (Poir.)Tenore Chandler pummelo C. maxima [(Burm.) Merr]. Genealogy of domesticated citrus Chloroplast phylogenetic tree of the genus Citrus 1 1/24/2017 Nuclear phylogenetic tree of the genus Citrus Nuclear phylogenetic tree of the genus Citrus The oldest citrus fossil specimen Chronogram of citrus speciation Entire margin Late Miocene Pliocene Pleistocene Citrus maxima ASIAN RADIATION Intramarginal vein 8.0-6.0 Ma Late Miocene Citrus reticulata Citrus aurantifolia AUSTRALIAN RADIATION 4.6-4-1 Ma Early Pliocene Articulated and winged petiole TACHIBANA SPECIATION Citrus maxima Citrus ichangensis 2,4-2.3 Ma Citrus micrantha Early Pleistocene Ma Citrus linczangensis (~ 8 Ma, midpoint of Late Miocene) Yunnan, China. Sanping Xie et al.(2013) The center of origin of citrus and its dispersal Biogeography of the genus Citrus Biogeography of the genus Citrus 2 1/24/2017 Biogeography of the genus Citrus Biogeography of the pure Citrus species 22 22 7 25. 7 25. 24 24 26 26 20 20 42 14 5 3130 21 145 6 39 9 9 41 372740 356 2 3629 2 18 1 18 1 23 1213 15 8 28 32 23 1213 158 10 38 44 10. 19 34 33 19 4 4 16 45 16 11 11 17 17 17 43 Blue = pure species Black = hybrids Red = admixtures 3 3 The center of origin of citrus Biogeography of the pure Citrus species C. mangshanensis Tachibana Sun Chu Cha Kat C. linczanensis C. ichangensis Citron Pummelo Kumquat Micrantha Australian limes The Asian radiation and the monsoon Radiations during Late Miocene in the foothills of the Himalaya Late Miocene 8,0-6,4 Ma Ma 3 1/24/2017 The Australian radiation. Land bridges between The Tachibana speciation. Land bridges between main Wallacea and Australia land and Taiwan, the Ryukyu archipelago and Japan WEST EAST Aglaia Alocasia Begonia Pseuduvaria Neonauclea Uvaria Palm lineages Bridelia Begonia … etc The center of origin of citrus and ancient dispersal routes Summary 2,4 Ma Genomic analysis has revealed that the genus Citrus is formed by pure species, admixtures and hybrids. Based on the analyses of the pure species, we propose that: 8-6 Ma • The center of origin of citrus was located in Southeast Asia between the provinces of Assam and Yunnan and northern Myanmar. • The dispersal of citrus followed a predominant west to east direction with three main radiations: • The Asian radiation occurred in the Late Miocene and was driven by a dramatic climate change of the intensity of the Indian summer monsoon. • The Australian radiation was enabled by the Early Pliocene 4,6 Ma orogeny that provoked the emergence of submerged land and corridors between the Wallacea and Australia • Radiations in islands of Eastern Asian during the Early Pleistocene were due to expansion of glaciers that reduced sea levels rising land bridges between continental Asia and Japan. 4.