Extensive morphological and behavioural diversity among fourteen new and seven described species in <i> Phytophthora</i> Clade 10 and its evolutionary implications
Thomas Jung, Ivan Milenković, Tamara Corcobado, Tomáš Májek, Josef Janoušek, Tomáš Kudláček, Michal Tomšovský, Zoltán Á. Nagy, Álvaro Durán, Marthin Tarigan, E. Sanfuentes von Stowasser, Ritika Singh, Manuela Pinheiro Ferreira, J. F. Webber, Bruno Scanu, N.M. Chi, Phạm Quang Thu, Muhammad Junaid, Ade Rosmana, B. Baharuddin, T. Kuswinanti, Nasri Nasri, Koji Kageyama, Ayaka Hieno, Hayato Masuya, Seiji Uematsu, Jonàs Oliva, Miguel Ángel Redondo, C. M. B. de F. Maia, Iryna Matsiakh, Volodymyr Kramarets, Richard O’Hanlon, Ž. Tomić, C. M. Brasier, Marília Horta Jung
Abstract
During extensive surveys of global Phytophthora diversity 14 new species detected in natural ecosystems in Chile, Indonesia, USA (Louisiana), Sweden, Ukraine and Vietnam were assigned to Phytophthora major Clade 10 based on a multigene phylogeny of nine nuclear and three mitochondrial gene regions. Clade 10 now comprises three subclades. Subclades 10a and 10b contain species with nonpapillate sporangia, a range of breeding systems and a mainly soil-and waterborne lifestyle. These include the previously described P. afrocarpa , P. gallica and P. intercalaris and eight of the new species: P. ludoviciana , P. procera , P. pseudogallica , P. scandinavica , P. subarctica , P. tenuimura , P. tonkinensis and P. ukrainensis . In contrast, all species in Subclade 10c have papillate sporangia and are self-fertile (or homothallic) with an aerial lifestyle including the known P. boehmeriae , P. gondwanensis , P. kernoviae and P. morindae and the new species P. celebensis , P. chilensis , P. javanensis , P. multiglobulosa , P. pseudochilensis and P. pseudokernoviae . All new Phytophthora species differed from each other and from related species by their unique combinations of morphological characters, breeding systems, cardinal temperatures and growth rates. The biogeography and evolutionary history of Clade 10 are discussed. We propose that the three subclades originated via the early divergence of pre-Gondwanan ancestors > 175 Mya into water-and soilborne and aerially dispersed lineages and subsequently underwent multiple allopatric and sympatric radiations during their global spread.