Families and Genera of Diaporthalean Fungi Associated with Canker and Dieback of Tree Hosts
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Persoonia 40, 2018: 119–134 ISSN (Online) 1878-9080 www.ingentaconnect.com/content/nhn/pimj RESEARCH ARTICLE https://doi.org/10.3767/persoonia.2018.40.05 Families and genera of diaporthalean fungi associated with canker and dieback of tree hosts X.L. Fan1, J.D.P. Bezerra2, C.M. Tian1, P.W. Crous3,4,5 Key words Abstract In this study we accept 25 families in Diaporthales based on phylogenetic analyses using partial ITS, LSU, rpb2 and tef1-α gene sequences. Four different families associated with canker and dieback of tree hosts Ascomycota are morphologically treated and phylogenetically compared. These include three new families (Diaporthostomata phylogeny ceae, Pseudomelanconidaceae, Synnemasporellaceae), and one new genus, Dendrostoma (Erythrogloeaceae). Sordariomycetes Dendrostoma is newly described from Malus spectabilis, Osmanthus fragrans and Quercus acutissima having taxonomy fusoid to cylindrical, bicellular ascospores, with three new species namely D. mali, D. osmanthi and D. quercinum. Diaporthostomataceae is characterised by conical and discrete perithecia with bicellular, fusoid ascospores on branches of Machilus leptophylla. Pseudomelanconidaceae is defined by conidiogenous cells with apical collarets and discreet annellations, and the inconspicuous hyaline conidial sheath when mature on Carya cathayensis, compared to morphologically similar families Melanconidaceae and Juglanconidaceae. Synnemasporellaceae is proposed to accommodate fungi with synnematous conidiomata, with descriptions of S. toxicodendri on Toxico dendron sylvestre and S. aculeans on Rhus copallina. Article info Received: 3 November 2017; Accepted: 9 January 2018; Published: 6 February 2018. INTRODUCTION Juglanconis juglandina and J. oblonga (in Juglanconidaceae) (Voglmayr et al. 2017), foliar diseases of Eucalyptus by Hark Diaporthales represents an important order in Sordariomy nessia spp. (Harknessiaceae) (Crous et al. 2012), and foliar, cetes containing taxa that are mainly isolated as endophytes, fruit and stem diseases by Coniella spp. (Schizoparmaceae) saprobes or plant pathogens on various hosts. The order is (Alvarez et al. 2016, Marin-Felix et al. 2017). characterised by perithecia with elongate beaks, often forming The classification of Diaporthales has changed drastically within stromatic tissues, deliquescent paraphyses and asci that over the past decades because of the plasticity and variability generally deliquesce, become detached from the perithecial in morphology. The order Diaporthales and Valsales were wall when mature, and have a characteristic refractive apical first introduced by Nannfeldt (1932), based on subfamilies annulus (Rossman et al. 2007). Members of diaporthalean fungi EuDiaportheen and Valseen in Diaportheaceae proposed are responsible for several diseases causing severe damage in by Von Höhnel (1917). Later, Diaporthaceae and ‘Valsaceae’ plants with economic importance. The most notorious is chest- (now Cytosporaceae, and referred to as such below) were nut blight caused by Cryphonectria parasitica (Cryphonectria recognised in the Diaporthales by Von Arx & Müller (1954). ceae) that devastated American chestnut (Castanea dentata) Kobayashi (1970) proposed Diaporthaceae (including Valsa = populations in North America (Anagnostakis 1987, Gryzenhout Cytospora) in a wide concept including all taxa accepted in Dia et al. 2006). Other common diseases include ash anthracnose porthales by Barr (1978). Wehmeyer & Hanlin (1975) accepted due to Gnomoniella fraxinii and birch canker caused by Crypto three families within this order, including non-allantoid spored sporella platyphylla (Gnomoniaceae) (Redlin & Stack 1988, Fan Gnomoniaceae and Diaporthaceae separated on the presence et al. 2016a), stem-end rot of citrus fruits infected by Diaporthe or absence of a stroma, and Cytosporaceae with allantoid as- citri and walnut canker by Diaporthe rostrata (Diaporthaceae) cospores. Barr (1978) arranged four families (Gnomoniaceae, (Huang et al. 2013, 2015, Fan et al. 2015b, Guarnaccia & Crous Melanconidaceae, Pseudovalsaceae and Cytosporaceae) in 2017), willow and walnut canker disease caused by Cytospora Diaporthales based on beak position of ascomata and thin or chrysosperma (Cytosporaceae) (Fan et al. 2014, 2015a), birch firm ascospore walls without special emphasis on allantoid or dieback disease resulting from Melanconis stilbostoma (Melan non-allantoid ascospores. Families within the Diaporthales conidaceae) (Fan et al. 2016b), walnut dieback disease by have been segregated by several mycologists to utilise vari- 1 The Key Laboratory for Silviculture and Conservation of the Ministry of ous criteria: stromatic tissues, arrangement of ascomata in the Education, Beijing Forestry University, Beijing 100083, China; stroma or substrate, and ascospore shape, e.g., four families corresponding author e-mail: Chengming Tian, [email protected]. (Cytosporaceae, Endoxylaceae, Gnomoniaceae and Melan 2 Departamento de Micologia Prof. Chaves Batista, Universidade Federal de conidaceae) by Monod (1983), three families (Cytosporaceae, Pernambuco, Av. Prof. Moraes Rego, s/n, Centro de Biociências, Cidade Melanconidaceae and Phyllachoraceae) by Cannon (1988), Universitária, CEP: 50670-901, Recife, PE, Brazil. 3 Westerdijk Fungal Biodiversity Institute, Uppsalalaan 8, 3584 CT Utrecht, while Hawksworth et al. (1995) merged the Cytosporaceae, The Netherlands. Gnomoniaceae and Melanconidaceae proposed by Barr (1990) 4 Wageningen University and Research Centre (WUR), Laboratory of Phyto- to Cytosporaceae and Melanconidaceae. These changes and pathology, Droevendaalsesteeg 1, 6708 PB Wageningen, The Netherlands. confusions suggested that phenotypic characters alone were 5 Department of Microbiology and Plant Pathology, Forestry and Agricultural Biotechnology Institute (FABI), University of Pretoria, Pretoria 0028, South unable to provide sufficient evidence to resolve phylogenetic Africa. and evolutionary patterns among these fungi. © 2018 Naturalis Biodiversity Center & Westerdijk Fungal Biodiversity Institute 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. 120 Persoonia – Volume 40, 2018 Molecular studies on fungi began in the early 1990s, and since definition colour camera, using differential interference contrast then ribosomal DNA sequence data were accepted as the (DIC) illumination and the Nikon software NIS-Elements D standard gene loci for fungi (Berbee & Taylor 1992, Schoch et Package v. 3.00. Adobe Bridge CS v. 6 and Adobe Photoshop al. 2012). Zhang & Blackwell (2001) recognised three lineages CS v. 5 were used for the manual editing. Nomenclatural novel- in Diaporthales, while Castlebury et al. (2002) postulated six ties and descriptions were deposited in MycoBank (Crous et major lineages, namely the Cryphonectria-Endothia complex, al. 2004). Colony diameters were measured, and the colony Cytosporaceae s.str., Diaporthaceae s.str., Gnomoniaceae colours described after 3 wk according to the colour charts of s.str., Melanconidaceae s.str. and the Schizoparme complex. Rayner (1970). When Rossman et al. (2007) reviewed the Diaporthales, nine families were recognised, i.e., Cryphonectriaceae, Cyto DNA extraction, amplification and sequencing sporaceae, Diaporthaceae, Gnomoniaceae, Melanconidaceae, Genomic DNA was extracted using a modified CTAB method, Pseudovalsaceae, Schizoparmeaceae, Sydowiellaceae and with fungal mycelium harvested from PDA plates with cel- Togniniaceae. Kirk et al. (2008) added Melogrammataceae and lophane (Doyle & Doyle 1990). The ITS region was amplified listed 10 families in this order, whereas Jaklitsch & Voglmayr with the primers ITS1 and ITS4 (White et al. 1990), the LSU (2012) placed Melogrammataceae within Xylariales rather than region with the primers LR0R and LR5 (Vilgalys & Hester 1990), Diaporthales. Subsequently, the Pseudoplagiostomataceae, the rpb2 region with primers fRPB2-5F and fRPB2-7cR (Liu et Harknessiaceae, Macrohilaceae and Tirisporellaceae were also al. 1999), and the tef1-α gene with the primers EF1-728F and added to the Diaporthales (Cheewangkoon et al. 2010, Crous EF1-986R (Carbone & Kohn 1999). The PCR mixture for all et al. 2012, 2015, Suetrong et al. 2015). Voglmayr & Jaklitsch regions consisted of 1 μL genomic DNA, 3 mM MgCl2, 20 μM (2014) resurrected Stilbosporaceae, while the Togniniaceae of each dNTP, 0.2 μM of each primer and 0.25 U BIOTAQ and Tirisporellaceae were reallocated to the Togniniales and DNA polymerase (Bioline). Conditions for PCR of ITS and LSU Tirisporellales (Gramaje et al. 2015, Jones et al. 2015). Later, genes constituted an initial denaturation step of 2 min at 95 °C, Lamproconiaceae and Juglanconidaceae were proposed as followed by 35 cycles of 30 s at 95 °C, 45 s at 51 °C and 1 min new families in this order (Norphanphoun et al. 2016, Voglmayr at 72 °C, and a final