Taxonomy, biogeography and evolution of thirty-four taxa in Phytophthora Clade 7a including eleven new species from natural forests and updated descriptions for P. fragariae, P. rubi and the 'P. ×alni hybrid complex'.
Z Á Nagy, M Horta Jung, I Milenković, T Kudláček, T Bourret, K Kageyama, A Hieno, H Masuya, S Uematsu, N M Chi, P Q Thu, T-T Chang, C-H Fu, T Corcobado, J Janoušek, T Májek, M Mullett, M Tomšovský, M Ferreira, R Singh, C Maia, Z Stanivuković, Z Tomić, J Bakonyi, J F Webber, C M Brasier, T Jung
{"title":"Taxonomy, biogeography and evolution of thirty-four taxa in <i>Phytophthora</i> Clade 7a including eleven new species from natural forests and updated descriptions for <i>P. fragariae</i>, <i>P. rubi</i> and the '<i>P. ×alni</i> hybrid complex'.","authors":"Z Á Nagy, M Horta Jung, I Milenković, T Kudláček, T Bourret, K Kageyama, A Hieno, H Masuya, S Uematsu, N M Chi, P Q Thu, T-T Chang, C-H Fu, T Corcobado, J Janoušek, T Májek, M Mullett, M Tomšovský, M Ferreira, R Singh, C Maia, Z Stanivuković, Z Tomić, J Bakonyi, J F Webber, C M Brasier, T Jung","doi":"10.3114/sim.2026.114.05","DOIUrl":null,"url":null,"abstract":"<p><p>Over the past three decades, surveys in natural and managed ecosystems, driven mainly by forest biosecurity issues, have greatly expanded knowledge of the global diversity and biogeography of the plant pathogen genus <i>Phytophthora</i>. In <i>Phytophthora</i> subclade 7a alone, 16 new species were described raising the number of validly described species in this subclade to 18. Recently, in further surveys conducted in natural forests of Japan, Vietnam, Europe and North America, 11 more new Clade 7a species and one other 7a taxon have now been identified. All are considered to be endemic to the surveyed sites. They are formally described here, including their morphological properties, colony and temperature-growth characteristics and their phylogenetic relationships. Informal, updated descriptions without nomenclatural act are also presented for the known species <i>P. fragariae</i>, <i>P. rubi</i>, <i>P. uniformis</i>, <i>P.</i> ×<i>alni</i> and <i>P.</i> ×<i>multiformis</i>. With now 34 taxa including hybrids, Clade 7a is currently the largest <i>Phytophthora</i> subclade. It is essentially a Northern Hemisphere subclade, with a disjunct distribution in North America, Europe and Asia which probably reflects an original Laurasian ancestral distribution ~90-80 Mya and subsequent post-Laurasian migrations and radiations. It is comprised of predominantly soilborne and aquatic species with non-papillate non-caducous sporangia, though some may have facultative adaptations to a degree of aerial dispersal. The largest diversity hotspots are in Southeast and East Asia, and both adaptive sympatric and non-adaptive radiations are indicated in the data. All 22 'pure' or non-hybrid species are self-fertile and exhibit relatively low nuclear heterozygosity levels. In contrast, high heterozygosity levels, relatively high numbers of tri- and partly tetra-allelic sites and maternal lines partly shared with other Clade 7a taxa are consistent with ten of the taxa (30 %) originating via interspecific hybridisation. Of these five are self-fertile, but unusually, given their likely self-fertile parents, the other five, including the '<i>P.</i> ×<i>cambivora</i> hybrid complex', have an A1/A2 or outcrossing breeding system. Some of the hybrids, such as the '<i>P</i>. ×<i>alni</i> hybrid complex', are known to have emerged as a result of anthropogenic disturbance events. Numerous <i>Phytophthora</i> species, including several in Clade 7a, have become highly damaging pathogens of trees and agricultural crops as a result of historical introductions via global plant trade. However, for the 11 new species described here, no symptoms were observed at the natural sample sites and their natural and potential hosts are unknown. The same applies to well over 150 <i>Phytophthora</i> species in other Clades and subclades, representing a significant plant health evidence gap. It is recommended that the biosecurity risk to forests, natural ecosystems and agriculture posed by these previously scientifically unknown taxa be actively investigated. <b>Taxonomic novelties: New species:</b> <i>Phytophthora algida</i> T. Jung, N.M. Chi, I. Milenković & M. Horta Jung, <i>Phytophthora arta</i> T. Jung, K. Kageyama, C.M. Brasier, Z.Á. Nagy & M. Horta Jung, <i>Phytophthora facultativa</i> T. Jung, T. Corcobado, Z.Á. Nagy & S. Raghuwinder, <i>Phytophthora grandoogonia</i> T. Jung, N.M. Chi, Z.Á. Nagy & M. Horta Jung, <i>Phytophthora infundibuliformis</i> T. Jung, T. Májek, I. Milenković & M. Ferreira, <i>Phytophthora irregularis</i> T. Jung, K. Kageyama, S. Uematsu, J.F. Webber & M. Horta Jung, <i>Phytophthora laevigata</i> I. Milenković, T. Jung, Z. Tomić & Z. Stanivuković, <i>Phytophthora sessilis</i> T. Jung, N.M. Chi, I. Milenković & M. Horta Jung, <i>Phytophthora ×grandisporangia</i> T. Jung, N.M. Chi, I. Milenković & M. Horta Jung, <i>Phytophthora ×leptopleura</i> T. Jung, H. Masuya, K. Kageyama, A. Hieno & M. Horta Jung, <i>Phytophthora ×nodosa</i> T. Jung, A. Hieno, H. Masuya, K. Kageyama, & S. Uematsu <b>Citation:</b> Nagy ZÁ, Horta Jung M, Milenković I, Kudláček T, Bourret T, Kageyama K, Hieno A, Masuya H, Uematsu S, Chi NM, Thu PQ, Chang T-T, Fu C-H, Corcobado T, Janoušek J, Májek T, Mullett M, Tomšovský M, Ferreira M, Singh R, Maia C, Stanivuković Z, Tomić Z, Bakonyi J, Webber JF, Brasier CM, Jung T (2026). Taxonomy, biogeography and evolution of thirty-four taxa in <i>Phytophthora</i> Clade 7a including eleven new species from natural forests and updated descriptions for <i>P. fragariae</i>, <i>P. rubi</i> and the '<i>P. ×alni</i> hybrid complex'. <i>Studies in Mycology</i> <b>114</b>: 554-661. doi: 10.3114/sim.2026.114.05.</p>","PeriodicalId":22036,"journal":{"name":"Studies in Mycology","volume":"114 ","pages":"554-661"},"PeriodicalIF":18.3000,"publicationDate":"2026-07-28","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"https://www.ncbi.nlm.nih.gov/pmc/articles/PMC13478740/pdf/","citationCount":"0","resultStr":null,"platform":"Semanticscholar","paperid":null,"PeriodicalName":"Studies in Mycology","FirstCategoryId":"99","ListUrlMain":"https://doi.org/10.3114/sim.2026.114.05","RegionNum":1,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":null,"EPubDate":"2026/6/1 0:00:00","PubModel":"eCollection","JCR":"Q1","JCRName":"MYCOLOGY","Score":null,"Total":0}
引用次数: 0
Abstract
Over the past three decades, surveys in natural and managed ecosystems, driven mainly by forest biosecurity issues, have greatly expanded knowledge of the global diversity and biogeography of the plant pathogen genus Phytophthora. In Phytophthora subclade 7a alone, 16 new species were described raising the number of validly described species in this subclade to 18. Recently, in further surveys conducted in natural forests of Japan, Vietnam, Europe and North America, 11 more new Clade 7a species and one other 7a taxon have now been identified. All are considered to be endemic to the surveyed sites. They are formally described here, including their morphological properties, colony and temperature-growth characteristics and their phylogenetic relationships. Informal, updated descriptions without nomenclatural act are also presented for the known species P. fragariae, P. rubi, P. uniformis, P. ×alni and P. ×multiformis. With now 34 taxa including hybrids, Clade 7a is currently the largest Phytophthora subclade. It is essentially a Northern Hemisphere subclade, with a disjunct distribution in North America, Europe and Asia which probably reflects an original Laurasian ancestral distribution ~90-80 Mya and subsequent post-Laurasian migrations and radiations. It is comprised of predominantly soilborne and aquatic species with non-papillate non-caducous sporangia, though some may have facultative adaptations to a degree of aerial dispersal. The largest diversity hotspots are in Southeast and East Asia, and both adaptive sympatric and non-adaptive radiations are indicated in the data. All 22 'pure' or non-hybrid species are self-fertile and exhibit relatively low nuclear heterozygosity levels. In contrast, high heterozygosity levels, relatively high numbers of tri- and partly tetra-allelic sites and maternal lines partly shared with other Clade 7a taxa are consistent with ten of the taxa (30 %) originating via interspecific hybridisation. Of these five are self-fertile, but unusually, given their likely self-fertile parents, the other five, including the 'P. ×cambivora hybrid complex', have an A1/A2 or outcrossing breeding system. Some of the hybrids, such as the 'P. ×alni hybrid complex', are known to have emerged as a result of anthropogenic disturbance events. Numerous Phytophthora species, including several in Clade 7a, have become highly damaging pathogens of trees and agricultural crops as a result of historical introductions via global plant trade. However, for the 11 new species described here, no symptoms were observed at the natural sample sites and their natural and potential hosts are unknown. The same applies to well over 150 Phytophthora species in other Clades and subclades, representing a significant plant health evidence gap. It is recommended that the biosecurity risk to forests, natural ecosystems and agriculture posed by these previously scientifically unknown taxa be actively investigated. Taxonomic novelties: New species:Phytophthora algida T. Jung, N.M. Chi, I. Milenković & M. Horta Jung, Phytophthora arta T. Jung, K. Kageyama, C.M. Brasier, Z.Á. Nagy & M. Horta Jung, Phytophthora facultativa T. Jung, T. Corcobado, Z.Á. Nagy & S. Raghuwinder, Phytophthora grandoogonia T. Jung, N.M. Chi, Z.Á. Nagy & M. Horta Jung, Phytophthora infundibuliformis T. Jung, T. Májek, I. Milenković & M. Ferreira, Phytophthora irregularis T. Jung, K. Kageyama, S. Uematsu, J.F. Webber & M. Horta Jung, Phytophthora laevigata I. Milenković, T. Jung, Z. Tomić & Z. Stanivuković, Phytophthora sessilis T. Jung, N.M. Chi, I. Milenković & M. Horta Jung, Phytophthora ×grandisporangia T. Jung, N.M. Chi, I. Milenković & M. Horta Jung, Phytophthora ×leptopleura T. Jung, H. Masuya, K. Kageyama, A. Hieno & M. Horta Jung, Phytophthora ×nodosa T. Jung, A. Hieno, H. Masuya, K. Kageyama, & S. Uematsu Citation: Nagy ZÁ, Horta Jung M, Milenković I, Kudláček T, Bourret T, Kageyama K, Hieno A, Masuya H, Uematsu S, Chi NM, Thu PQ, Chang T-T, Fu C-H, Corcobado T, Janoušek J, Májek T, Mullett M, Tomšovský M, Ferreira M, Singh R, Maia C, Stanivuković Z, Tomić Z, Bakonyi J, Webber JF, Brasier CM, Jung T (2026). Taxonomy, biogeography and evolution of thirty-four taxa in Phytophthora Clade 7a including eleven new species from natural forests and updated descriptions for P. fragariae, P. rubi and the 'P. ×alni hybrid complex'. Studies in Mycology114: 554-661. doi: 10.3114/sim.2026.114.05.
在过去三十年中,主要由森林生物安全问题推动的自然和管理生态系统调查极大地扩展了对植物病原体疫霉属的全球多样性和生物地理学的认识。仅在疫霉亚枝7a中就发现了16个新种,使该亚枝的有效描述种数增加到18个。最近,通过对日本、越南、欧洲和北美的天然林的进一步调查,又发现了11个新的7a枝新种和1个7a分类单元。所有这些都被认为是调查地点特有的。本文正式介绍了它们的形态特征、菌落和温度生长特征及其系统发育关系。非正式的,没有命名行为的更新描述也提出了已知物种P. fragariae, P. rubi, P. uniformis, P. ×alni和P. ×multiformis。7a枝是目前最大的疫霉亚枝,包括杂种在内共有34个类群。它本质上是一个北半球亚分支,在北美、欧洲和亚洲的分布不连贯,可能反映了90- 80mya的原始劳氏祖先分布和后来的劳氏迁移和辐射。它主要由土生和水生物种组成,具有非乳头状非早落孢子囊,尽管有些可能对一定程度的空中传播具有兼性适应。最大的多样性热点在东南亚和东亚,数据中显示了自适应同域辐射和非自适应辐射。所有22个“纯”或非杂交物种都是自交的,并且表现出相对较低的核杂合性水平。相比之下,高杂合性水平、相对较多的三等位基因位点和部分四等位基因位点以及与其他Clade 7a类群部分共享的母系,与通过种间杂交起源的10个类群(30%)一致。其中五个是自育的,但不寻常的是,考虑到它们可能具有自育能力的父母,其他五个,包括“P. ×cambivora杂交复合体”,具有A1/A2或异交繁殖系统。一些杂交,如“P. ×alni杂交复合体”,已知是由于人为干扰事件而出现的。由于全球植物贸易的历史引入,许多疫霉菌物种,包括进化枝7a中的一些物种,已经成为树木和农作物的高度破坏性病原体。然而,对于这里描述的11个新种,在自然采样点未观察到任何症状,它们的自然宿主和潜在宿主尚不清楚。这同样适用于其他分支和亚分支中超过150种疫霉物种,这代表了一个重要的植物健康证据差距。建议积极调查这些以前科学上未知的分类群对森林、自然生态系统和农业构成的生物安全风险。分类新发现:新种:藻疫霉T. Jung, N.M. Chi, I. milenkoviki & M. Horta Jung, arta疫霉T. Jung, K. Kageyama, C.M. Brasier, Z.Á。Nagy & M. Horta Jung,兼性疫霉T. Jung, T. Corcobado, Z.Á。Nagy & S. Raghuwinder,大疫霉T. Jung, N.M. Chi, Z.Á。Nagy & M. Horta Jung,疫病疫霉T. Jung, T. Májek, I. milenkoviki & M. Ferreira,非常规疫霉T. Jung, K. Kageyama, S. Uematsu, J.F. Webber & M. Horta Jung, laevigata I. milenkoviki, T. Jung, Z. tomiki & Z. stanivukoviki,候疫霉T. Jung, N.M. Chi, I. milenkoviki & M. Horta Jung,疫霉×grandisporangia T. Jung, N.M. Chi, I. milenkoviki & M. Horta Jung,疫霉×leptopleura T. Jung, H. Masuya, K. Kageyama,引用本文:Nagy ZÁ, Horta Jung M, milenkovovic I, Kudláček T, Bourret T, Kageyama K, Hieno A, Masuya H, Uematsu S, Chi NM, Thu PQ, Chang T-T, Fu C-H, Corcobado T, Janoušek J, Májek T, Mullett M, Tomšovský M, Ferreira M, Singh R, Maia C, stanivukoviki Z, tomiki Z, Bakonyi J, Webber JF, Brasier CM, Jung T(2026)。Phytophthora Clade 7a 34个分类群的分类、生物地理和进化,包括11个天然林新种,以及P. fragariae、P. rubi和P. ×alni杂交复合体的更新描述。真菌学研究114:554-661。doi: 10.3114 / sim.2026.114.05。
期刊介绍:
The international journal Studies in Mycology focuses on advancing the understanding of filamentous fungi, yeasts, and various aspects of mycology. It publishes comprehensive systematic monographs as well as topical issues covering a wide range of subjects including biotechnology, ecology, molecular biology, pathology, and systematics. This Open-Access journal offers unrestricted access to its content.
Each issue of Studies in Mycology consists of around 5 to 6 papers, either in the form of monographs or special focused topics. Unlike traditional length restrictions, the journal encourages submissions of manuscripts with a minimum of 50 A4 pages in print. This ensures a thorough exploration and presentation of the research findings, maximizing the depth of the published work.