Cercosporoid Fungi (<I>Mycosphaerellaceae</I>)
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IMA FUNGUS · 7(1): 161–216 (2016) doi:10.5598/imafungus.2016.07.01.10 Cercosporoid fungi (Mycosphaerellaceae) 5. Species on dicots ARTICLE (Anacardiaceae to Annonaceae) Uwe Braun1, Pedro W. Crous2, and Chiharu Nakashima3 1Martin-Luther-Universität, Institut für Biologie, Bereich Geobotanik und Botanischer Garten, Herbarium, Neuwerk 21, 06099 Halle (Saale), Germany; corresponding author e-mail: [email protected] 2CBS-KNAW, Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands 3Graduate School of Bioresources, Mie University, 1577 Kurima-machiya, Tsu, Mie 514-8507, Japan Abstract: The present paper is a continuation of a series of comprehensive taxonomic treatments of cercosporoid Key words: fungi (formerly Cercospora s. lat.), belonging to Mycosphaerellaceae (Ascomycota). This fifth contribution of this Ascomycota series proceeds with treatments of cercosporoid fungi on dicots and comprises species occurring on hosts belonging Cercospora s. lat. to the families Anacardiaceae and Annonaceae, which are described and illustrated in alphabetical order under the hyphomycetes particular cercosporoid genera, supplemented by keys to the species concerned. A detailed introduction, a survey of taxonomy currently recognised cercosporoid genera, a key to the genera concerned, and a discussion of taxonomically relevant characters were published in the first part of this series. The following taxonomic novelties are introduced:Passalora cotini sp. nov., P. guoana nom. nov., P. rhois-aromaticae sp. nov., and Pseudocercospora rhoicola. Article info: Submitted: 1 February 2016; Accepted: 9 June 2016; Published: 16 June 2016. INTRODUCTION in this fungal group whenever possible. On the other hand, symptoms and morphological characters exhibited in vivo are One of the largest groups of fungi belonging to still important in diagnosing plant pathogenic fungi, especially Mycosphaerellaceae (Capnodiales, Ascomycota) includes for routine monitoring and identification purposes, but also genera and species akin to the genus Cercospora in light of the ecology and taxonomy of the fungi concerned, which are commonly referred to as cercosporoid fungi which cause diseases on living plants. Therefore, optimal (Crous & Braun 2003, Braun et al. 2013). The taxa descriptions of new and already known species should cover concerned comprise dematiaceous, holoblastic asexual the full range of traits ranging from characters in vivo, in morphs and at least partly mycosphaerella-like sexual vitro to phylogenetic data. Currently only a limited number morphs. Corresponding mucedinaceous fungi within of cercosporoid fungi has been cultivated and sequenced, Mycosphaerellaceae are often classified as ramularioid and hundreds of taxa are only known from characters in fungi. Ramularia, the eponymous genus, is associated vivo. However, all names have to be taken into consideration with sexual morphs belonging to Mycosphaeraella s. str. for taxonomic purposes and are subject to the rules of the The type species of the latter genus has a phylogenetically International Code of Nomenclature for algae, fungi, and proven asexual morph pertaining to Ramularia (Verkley et plants (ICN; McNeill et al. 2012), including Art. 11 dealing al. 2004) making Mycosphaerella a younger heterotypic with the priority of names. Thus, all old names are applicable synonym of Ramularia, which has been proposed as the when dealing with taxonomic-nomenclatural questions, i.e. to preferred name (Crous et al. 2009c, Wijayawardene et al. find correct names for certain taxa in accordance with the 2014, Videira et al. 2015) and recommended for protection Code. To facilitate this process, careful examinations of all old (Rossman et al. 2015). type collections and detailed descriptions are necessary and Cercosporoid fungi represent a very large group of helpful for sound taxonomic conclusions. This is precisely the plant pathogenic, leaf-spotting, economically relevant sense of this series, based on Chupp’s (1954) old, seriously species causing diseases on a wide range of hosts, outdated, Cercospora monograph, and meant to represent including numerous cultivated plants. In the last few an updated, supplementary treatment intended as a platform decades, enormous taxonomic progress has been made by for the routine identification of cercosporoid fungi, including increasing application of molecular methods in capnodialean modern molecular examinations. We have spent decades fungi, including Mycosphaerellaceae (Crous et al. 2013, examining thousands of old type collections and other Groenewald et al. 2013, Bakhshi et al. 2014, Nguanhom et al. samples of cercosporoid fungi, inter alia in preparation for the 2015, Guatimosim et al. 2016, etc.). Cultures and molecular annotated list of Cercospora and Passalora names published sequence analyses should be used for taxonomic purposes by Crous & Braun (2003). Numerous previously unpublished © 2016 International Mycological Association You are free to share - to copy, distribute and transmit the work, under the following conditions: Attribution: You must attribute the work in the manner specified by the author or licensor (but not in any way that suggests that they endorse you or your use of the work). Non-commercial: You may not use this work for commercial purposes. No derivative works: You may not alter, transform, or build upon this work. For any reuse or distribution, you must make clear to others the license terms of this work, which can be found at http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode. Any of the above conditions can be waived if you get permission from the copyright holder. Nothing in this license impairs or restricts the author’s moral rights. VOLUME 7 · NO. 1 161 Braun et al. results have been incorporated in already published issues morphological characters of the asexual morphs traditionally of the present series. Cultures, sequence analyses and used for the separation of genera do not reflect phylogenetic epitypifications on the basis of cultured and sequenced entities (i.e. genera). A corresponding detailed phylogenetic collections represent the next phase in order to transfer the treatment of the Passalora complex is under preparation. All old morphology-based names into the era of consolidated results of modern studies dealing with cercosporoid fungi ARTICLE species concepts. in Mycosphaerellaceae show that proper taxonomic results The growing number of DNA sequences of cercosporoid and conclusions have to be based on an integrated approach fungi allows better insights in the phylogenetic structure including molecular sequence analyses. Conclusions just of this fungal group on the familial as well as the generic based on morphological analyses are unreliable and do level. On the other hand, it has gradually been pointed not allow proper generic allocations. Therefore, we have out that traditionally applied phenotypic traits of the decided to revise the focus of the present series. Efforts to sexual as well as asexual morphs of cercosporoid fungi reassess and reallocate cercosporoid genera and species (Mycosphaerellaceae) are not in full agreement with will thus in future be based on re-examinations of types and genotypic generic circumscriptions. Species only forming other collections in combination with cultures and sequence mycosphaerella-like sexual morphs, i.e. without any asexual analyses whenever possible. The present series will not be fructifications, are known in many currently recognised continued in its present form. In future, it will rather focus on cercosporoid genera (e.g. Guatimosim et al. 2016). As the revision of type collections of old cercosporoid species, a consequence, the generic affinity of mycosphaerelloid including comprehensive descriptions and illustrations, ascomycetes cannot be properly resolved without cultures, which are necessary as basis for modern examinations in sequences, and phylogenetic analyses. On the other hand, vitro including subsequent sequence analyses. the classical morphological characters of asexual morphs So far four contributions have been published: part one of cercosporoid fungi are no longer reliable for allocating dealing with cercosporoid fungi on other fungi (mycophylic certain species to particular genera. The analysis of fern- taxa), on ferns as well as gymnosperms (Braun et al. 2013); inhabiting cercospoid fungi recently examined in Brazil part two dedicated to species on monocots, excluding true (Guatimosim et al. 2016) is a striking example. The genus grasses (Braun et al. 2014); part three with a treatment of Zasmidium is known to be polyphyletic (Crous et al. 2009a, cercosporoids on hosts of Poaceae (Braun et al. 2015a); b, Braun et al. 2013). Besides genuine species of Zasmidium and part four as the first contribution to cercosporoids on s. str., zasmidium-like asexual morphs are also known for dicots covering species occurring on hosts belonging to the cercosporoid genera Neoceratosperma (Crous et al. the families Acanthaceae to Amaranthaceae (Braun et al. 2014, Guatimosim et al. 2016) and Paramycosphaerella 2015b). Part five is a continuation covering cercosporoid (widened to include species with zasmidium-like asexual fungi on hosts of the dicot families Anacardiaceae and morphs by reallocations of Zasmidium aerohyalosporium, Z. Annonaceae. Generic descriptions and keys to accepted dicranopteridis, Z. nabiacense, and Mycosphaerella parkii genera are included in the first part. The structure of the (i.e. Z. parkii; Guatimosim et