Families of Diaporthales based on morphological and phylogenetic evidence
Abstract
Diaporthales is an important ascomycetous order comprising phytopathogenic, saprobic, and endophytic fungi, but interfamilial taxonomic relationships are still ambiguous. Despite its cosmopolitan distribution and high diversity with distinctive morphologies, this order has received relativelyiaceae, Macrohilaceae, Melanconidaceae, Pseudoplagiostomaceae, Schizoparmaceae, Stilbosporaceae and Sydowiellaceae. Taxonomic uncertainties among genera are also clarified and recurrent discrepancies in the taxonomic position of families within the Diaporthales are discussed. An updated outline and key to families and genera of the order is presented.
Introduction
The Diaporthales is a distinct order in the subclass Diaporthomycetidae (Sordariomycetes) and it includes pathogens, saprobes and endophytes, with no known coprophilous, hypersaprobes or mycophylic species (Barr, 1978, Rossman et al., 2007, Vasilyeva et al., 2007, Maharachchikumbura et al., 2015, Maharachchikumbura et al., 2016). Taxa of this order inhabit a wide diversity of hosts and substrates, including most economically and ecologically important trees and crops, soil and living animal and human tissues (Barr, 1978, Gryzenhout et al., 2006c). Species in Diaporthales form solitary or aggregated, immersed to erumpent, rarely superficial, orange, brown to black perithecial ascomata, with short or long necks, that are located in stromatic tissues or substrates, with a centrum (or hamathecium) lacking or with few paraphyses (Alexopoulus and Mims, 1978, Barr, 1978, Castlebury et al., 2002). Asci are unitunicate with a conspicuous refractive ring (Hawksworth et al., 1995, Rossman et al., 2007). Ascospore morphology is diverse, ranging from short to elongate and aseptate or septate with hyaline or pigmented walls. The asexual morphs of Diaporthales are generally coelomycetous (Rossman et al. 2007), producing acervuli or pycnidial conidiomata, with or without a well-developed stroma. Conidiogenesis is phialidic or rarely annellidic and conidia are usually unicellular or 1-septate (Rossman et al. 2007).
Fungal taxa placed in “Diaporthaceae sensu lato” were divided into two groups (von Höhnel 1917) as “Eu-Diaportheen”, to accommodate genera without allantoid ascospores and “Valseen” to accommodate genera with allantoid ascospores. Nannfeldt (1932) introduced the order Diaporthales to accommodate von Höhnel's Eu-Diaportheen group. Luttrell (1951) described Diaporthales as an order comprising species that have a “Diaporthe-type centrum” and “Endothia-type ascus”. Chadefaud (1960) analysed characters of stromatic tissues in diaporthoid taxa and recognised families as Diaporthaceae or Cytosporaceae (= Valsaceae), Melanconidaceae and Gnomoniaceae. Wehmeyer (1975) classified the Diaporthales to include three families: Diaporthaceae, Gnomoniaceae and Cytosporaceae. Barr (1978) revised the order Diaporthales accepting Gnomoniaceae and Cytosporaceae in the suborder Gnomoniineae. Melanconidaceae and Pseudovalsaceae were accommodated in the suborder Melanconidineae. To differentiate genera, Barr (1978) used characters such as presence or absence of stromata, stromatic development and tissue types, the position of the perithecia and perithecial necks relative to the substrate, as well as ascospore shape; and Monod (1983) distinguished genera within Gnomoniaceae based on characters of the stromatic tissues, asexual morphs and ascospores. Three families were recognised in Diaporthales by Eriksson (2001), including Cytosporaceae, Melanconidaceae and Vialaeaceae. Based on analysis of LSU nrDNA sequence data, Castlebury et al. (2002) accepted Diaporthaceae, Gnomoniaceae, Melanconidaceae and Cytosporaceae in Diaporthales. Gnomoniaceae was revised by several recent studies and new taxa were introduced (Sogonov et al., 2008, Walker et al., 2010, Walker et al., 2012, Mejía et al., 2011). Castlebury et al. (2002) did not confirm Vialaeaceae as a family in Diaporthales and therefore excluded it from Diaporthales. Réblová et al. (2004) introduced Togniniaceae to this order based on small subunit (SSU) nrDNA; however, Mostert et al. (2006) concluded that its placement was ambiguous based on large subunit (LSU) nrDNA. Maharachchikumbura et al. (2015) excluded Togniniaceae from Diaporthales and accommodated it in Togniniales. Gryzenhout et al. (2006c) introduced the Cryphonectria–Endothia complex as the family Cryphonectriaceae. Sydowiellaceae and the Schizoparme–Pilidiella complex with the genus Coniella were introduced as Schizoparmaceae in Diaporthales (Rossman et al., 2007, Alvarez et al., 2016). Harknessiaceae was introduced into Diaporthales accommodating Harknessia with wuestneia-like sexual morphs (Crous et al. 2012b). Pseudoplagiostomaceae was introduced by Cheewangkoon et al. (2010) to accommodate Pseudoplagiostoma. Voglmayr & Jaklitsch (2014) resurrected Stilbosporaceae in Diaporthales based on phylogenetic analysis of LSU nrDNA sequence data and transferred the genera Stegonsporium and Stilbospora to this family. Macrohilaceae was introduced by Crous et al. (2015), based on an analysis of LSU nrDNA to accommodate Macrohilum. Suetrong et al. (2015) introduced Tirisporellaceae into Diaporthales; however, Jones et al. (2015) excluded this family from Diaporthales. Norphanphoun et al. (2016) introduced Lamproconiaceae to accommodate Lamproconium and Hercospora. Juglanconidaceae was introduced in the Diaporthales by Voglmayr et al. (2017). However, molecular data suggest that additional families still remain to be elucidated (Gryzenhout et al., 2006c, Crous et al., 2012a, Crous et al., 2015, Voglmayr et al., 2017). Currently there are 14 families accepted in the Diaporthales.
Given the taxonomic discrepancies within Diaporthales, the present study uses a combined taxonomic approach based on morphology and DNA sequence analyses of the partial 28S nrDNA (LSU), the internal transcribed spacer regions and intervening 5.8S nrDNA (ITS), DNA-directed RNA polymerase II second largest subunit (rpb2), and translation elongation factor 1-alpha (tef1) gene regions to investigate phylogenetic relationships of all genera in Diaporthales to update their classification. All taxonomic novelties and present taxonomic families are re-described and illustrated where necessary. We also present new data on each family to provide a better taxonomic understanding.
Materials and methods
Isolates and specimens
Specimens were collected from Germany, Italy, Russia, Thailand and the UK. They were placed in paper bags and collection details noted. Specimens were brought to the laboratory in Zip-lock plastic bags and examined with a Motic SMZ 168 stereomicroscope. Rehydrated fruiting bodies were used to observe morphological characteristics of ascomata, asci, ascospores and other tissues and characters were photographed with a Canon 550D digital camera fitted to the Nikon ECLIPSE 80i compound microscope. Photomicrographs were arranged with Adobe Photoshop v. CS6 and all measurements were made with Tarosoft v. 0.9.0.7. Specimens were preserved and are deposited at the BBH and MFLU fungaria. Taxonomic novelties and descriptions were deposited in MycoBank (Crous et al. 2004), and new species were established using modern criteria and standards (Taylor et al., 2000, Seifert and Rossman, 2010, Jeewon and Hyde, 2016).
Sporocarps were removed from the substrate using a sterilised needle and placed in a few drops of sterilised distilled water on a sterilised cavity slide and a spore suspension was prepared as described in Chomnunti et al. (2014). Germinating ascospores were aseptically transferred to Petri dishes containing Potato Dextrose Agar (PDA) or Malt Extract Agar (MEA) (Crous et al. 2009). Colonies were photographed and characters were noted. Colony colour on PDA and MEA was determined with the colour charts of Rayner (1970). Living cultures are deposited at Mae Fah Luang University (MFLU) and the Westerdijk Fungal Biodiversity Institute (CBS) culture collections. Autoclaved pine needles were placed on water agar (WA) to observe conidiomatal development and sporulating (Crous et al. 2009).
Types and other relevant authentic specimens were loaned from accessible fungaria [New York State Museum (NY), Naturhistorisches Museum Wien (W), Swedish Museum of Natural History (S), Royal Botanic Gardens, Kew (K), Universität Wien (WU)]. A small part of the fungarium specimen was cut and rehydrated in water or 5 % KOH. Micro-morphological characters were observed from rehydrated ascomata and photography was done as previously described.
DNA extraction, PCR amplification and phylogeny
Fresh fungal mycelia grown on MEA for 4 wk at 18 °C was scraped from the colony margin and sometimes perithecial content of fresh specimens were used for genomic DNA extraction following the protocol outlined by Jeewon et al. (2002). PCR amplification and sequencing of the LSU nrDNA region using the primer pair LROR/LR5 (Vilgalys & Hester 1990, Rehner & Samuels 1994), ITS nrDNA region using primer pair ITS5/ITS4 (White et al. 1990), rpb2 region using the primer pair fRPB2-5F/fRPB2-7cR (Liu et al. 1999), and tef1 region using primer pair EF1-728F/EF1-986R (Carbone & Kohn 1999) were performed.
Each amplification reaction contained 0.125 μL of 5 units/μL Ex-Taq DNA polymerase (TaKaRa), 2.5 μL of 10 × PCR buffer, 2 μL of 2 mM MgCl2, 2.5 μL of 2 mM dNTPs, 1 μL of 0.2–1.0 μM primer, <500 ng DNA template and was adjusted with double-distilled water to a total volume of 25 mL. Amplification reactions were performed in a thermal-cycler (BIORAD 1000™ Thermal Cycler, Bio-Rad Laboratories, Hercules, California). The temperature profile for both ITS nrDNA and LSU nrDNA was an initial denaturing step for 2 min at 94 °C, followed by 35 amplification cycles of denaturation at 94 °C for 60 s, annealing at 58 °C for 60 s and extension at 72 °C for 90 s and a final extension step of 72 °C for 10 min (Phillips et al. 2008). The temperature profile for the rpb2 was: initial denaturation at 94 °C for 120 s, followed by 35 amplification cycles of denaturation at 95 °C for 45 s, annealing at 57 °C for 50 s and extension at 72 °C for 90 s (Liu et al. 1999). The temperature profile for tef1 was: initial denaturation at 94 °C for 120 s, followed by 35 amplification cycles of denaturation at 95 °C for 30 s, 58 °C for 50 s, 72 °C 60 s (Glass & Donaldson 1995). All PCR products with a DNA ladder were determined by electrophoresis at 120 V/cm for 20 min in 1 % agarose gel stained with ethidium bromide (0.5 mg/mL). The gel was visualised under a UV transilluminator to estimate the fragment size. PCR products were purified and sequenced with both primers at the Sunbiotech Company, Beijing, China. Sequences were edited and condensed with DNASTAR Lasergene v. 7.1. The sequences generated in this study were supplemented with additional sequences obtained from GenBank (Table 1) based on blast searches and published literature. Multiple sequence alignments were generated with MAFFT v. 7 (http://mafft.cbrc.jp/alignment/server/index.html) and the alignment was manually improved with BioEdit v. 7.0.5.2 (Hall 1999).
Table 1
Details of the strains included for molecular and/or morphological study.
| Fungal species | Culture accession no. | Specimen voucher no. | Host/substrate | GenBank accession numbers | |||
|---|---|---|---|---|---|---|---|
| ITS | LSU | tef1-α | rpb2 | ||||
| Alnecium auctum | CBS 124263 | WU 30206 | Alnus glutinosa | KF570154 | KF570154 | KF570200 | KF570170 |
| Ambarignomonia petiolorum | CBS 121227 | BPI 844274 | Liquidambar styraciflua | EU254748 | EU255070 | EU221898 | EU219307 |
| CBS 116866 | BPI 843530 | Liquidambar styraciflua | EU199193 | AY818963 | – | EU199151 | |
| Amphilogia gyrosa | CBS 112922 | AFTOL-ID 1985 | Elaeocarpus dentatus | – | FJ176889 | – | FJ238374 |
| YMJ 91123101 | HAST 91123101 | Elaeocarpus japonicus | EF026147 | – | KC465404 | – | |
| CMW 10470 | – | Elaeocarpus japonicus | – | AY194108 | – | – | |
| Amphiporthe hranicensis | CBS 119289 | BPI 843515 | Tilia platyphyllos | EU199178 | EU199122 | – | EU199137 |
| Anisogramma anomala | 529478 | – | Corylus avellana | EU683064 | EU683066 | – | – |
| Anisogramma virgultorum | 529479 | – | Betula pendula | EU683062 | EU683065 | – | – |
| Apiognomonia errabunda | AR 2813 | AR 2813 | Fagus sylvatica | DQ313525 | NG027592 | DQ313565 | DQ862014 |
| Apiognomonia veneta | MFLUCC 16–1193 | MFLU 17–0896 | Platanus acerifolia | MF190114 | MF190056 | – | – |
| MFLUCC 17–1656 | MFLU 17–0896B | Platanus acerifolia | MF190115 | MF190057 | – | – | |
| CBS 897.79 | Monod LAU | Platanus orientalis | – | EU255195 | EU221910 | EU219259 | |
| Apioplagiostoma populi | ID 858501 | ApLA2 | Populus tremuloides | KP637024 | – | – | – |
| Apiosporopsis carpinea | CBS 771.79 | – | Carpinus betulus | – | AF277130 | – | – |
| Apiosporopsis sp. | Masuya 11Af2–1 | – | Alnus firma | – | AB669034 | – | – |
| Apoharknessia insueta | CBS 111377 | CPC 1451 | Eucalyptus pellita | JQ706083 | AY720814 | – | – |
| CBS 114575 | CPC 10947 | Eucalyptus pellita | JQ706082 | AY720813 | – | – | |
| Ascitendus austriacus | CBS 131685 | – | Decayed driftwood of Alnus glutinosa | – | AF261067 | – | JQ429257 |
| CBS 102665 | – | Submerged stems of Fagus sylvatica | AF242263 | AF242263 | – | – | |
| Asteroma alneum | CBS 109840 | – | Alnus glutinosa | EU167609 | EU167609 | – | – |
| Asteroma sp. | Masuya 8Ah9–1 | – | Alnus hirsuta | – | AB669035 | – | – |
| Asterosporium asterospermum | – | MFLU 15–3555 | Fagus sylvatica | – | MF190062 | – | MF377615 |
| KT 2125 | HHUF 30038 | Fagus crenata | – | AB553743 | – | – | |
| CBS 112404 | – | Fagus sylvatica | – | AB553745 | – | – | |
| KT 2101 | HHUF 30037 | Fagus crenata | – | AB553742 | – | – | |
| KT 2138 | HHUF 30039 | Fagus crenata | – | AB553744 | – | – | |
| Aurantioporthe corni | MNA 1003 | – | Cornus alternifolia | KF495043 | KF495058 | – | – |
| SDS 1001 | – | Cornus alternifolia | KF495046 | KF495061 | – | – | |
| Aurantiosacculus acutatus | CPC 13704 | CBS H-20933 | Eucalyptus viminalis | JQ685514 | JQ685520 | – | – |
| Aurantiosacculus eucalyptorum | CPC 13229 | – | Eucalyptus globulus | JQ685515 | JQ685521 | – | – |
| Aurapex penicillata | CMW 11296 | – | Myrica faya | AY214315 | AY194090 | – | – |
| CMW 10032 | – | Miconia theaezans | AY214312 | AY194104 | – | – | |
| Auratiopycnidiella tristaniopsis | CBS 132180 | CBS H-20932 | Tristaniopsis laurina | JQ685516 | JQ685522 | – | – |
| Aurifilum marmelostoma | CBS 124930 | CMW 28288 | Terminalia ivorensis | FJ882856 | HQ730874 | – | – |
| CBS 124929 | PREM 60257 | Terminalia mantaly | FJ882855 | HQ730873 | – | – | |
| Brachysporium nigrum | MR 1346 | – | – | – | KT991662 | – | KT991652 |
| Cainiella johansonii | Kruys 727 (UPS) | – | Dryas sp. | JF701922 | – | – | – |
| Kruys 731 (UPS) | – | Dryas octopetala | JF701922 | JF701920 | – | – | |
| Calosphaeria pulchella | CBS 115999 | JF 3200 | Prunus sp. | EU367451 | AY761075 | – | GU180661 |
| Celoporthe dispersa | CMW 9978 | – | Syzygium cordatum | AY214316 | HQ730854 | HQ730841 | – |
| Celoporthe eucalypti | CMW 26913 | – | Eucalyptus EC48 clone | HQ730839 | HQ730865 | HQ730852 | – |
| Chaetoconis polygoni | – | MFLU 17–0965 | Rumex acetosa | – | MF190063 | – | – |
| CBS 405.95 | – | Polygonum sachalinense | – | EU754141 | – | – | |
| Chapeckia nigrospora | CBS 125532 | BPI 863766 | Betula sp. | JF681957 | EU683068 | – | – |
| Chiangraiomyces bauhiniae | MFLUCC 17–1669 | MFLU 17–0964 | Bauhinia sp. | MF190119 | MF190064 | MF377598 | MF377603 |
| MFLUCC 17–1670 | CHUNI 81 | Bauhinia sp. | MF190118 | MF190065 | MF377599 | MF377604 | |
| Chromendothia citrina | AR 3445 | – | Quercus mongolica | – | EU255074 | EU222013 | EU219342 |
| CBS 109758 | BPI 747935 | Quercus mongolica | – | AF408335 | – | – | |
| Chrysocrypta corymbiae | CBS 132528 | CPC 19279 | Corymbia sp. | JX069867 | JX069851 | – | – |
| Chrysofolia barringtoniae | TBRC 5647 | SDBR-CMUENBA048 | Barringtonia sp. | KU948046 | KU948045 | – | – |
| Chrysofolia colombiana | CPC 24986 | CBS 139909 | Eucalyptus urophylla | KR476738 | KR476771 | – | – |
| Chrysoporthe cubensis | CMW 14394 | – | Eucalyptus sp. | JN942342 | JN940856 | GQ290137 | – |
| Chrysoporthella hodgesiana | CMW 10641 | CBS 115854 | Tibouchina semidecandra | AY692322 | – | – | – |
| Coniella africana | CBS 114133 | CBS H-22706 | Eucalyptus nitens | AY339344 | AY339293 | KX833600 | KX833421 |
| Coniella australiensis | IMI 261318 | BPI 748425 | Leaf litter | AF408336 | AF408336 | KX833692 | KX833497 |
| Coniella crousii | NFCCI 2213 | AMH 9406 | Terminalia chebula | HQ264189 | – | – | – |
| Coniella fragariae | CBS 110394 | RMF 74.01 | Forest soil | KJ710463 | KJ710441 | KX833695 | KX833499 |
| CBS 172.49 | STE-U 3930 | Fragaria sp. | AY339317 | AY339282 | AY339352 | – | |
| Coniella koreana | CBS 143.97 | CBS H-22710 | – | KX833584 | AF408378 | KX833684 | KX833490 |
| Coniella pseudokoreana | MFLUCC 12–0427 | MFLU 13–0282 | – | MF190145 | – | – | – |
| MFLUCC 17–1673 | MFLU 13–0282B | – | MF190146 | – | – | – | |
| Coniella pseudostraminea | CBS 814.71 | IMI 233050 | Fragaria sp. | KX833582 | – | KX833682 | – |
| Coniella quercicola | CBS 283.76 | – | Eucalyptus nitens | AY339344 | AY339293 | AY339364 | – |
| Coniella straminea | CBS 149.22 | STE U 3932 | Fragaria sp. | AY339348 | AY339296 | AY339366 | KX833506 |
| Coniella tibouchinae | CPC 18511 | BECM1 | Tibouchina granulosa | JQ281774 | JQ281776 | JQ281778 | KX833507 |
| Coniella wangiensis | CPC 19397 | CPC 19397 | Eucalyptus sp. | JX069873 | JX069857 | KX833705 | KX833509 |
| Coryneum longipes | AR 3541 | BPI 872021 | Quercus cerris | – | EU683072 | – | – |
| Coryneum modonia | AR 3558 | BPI 749131 | Castanea sativa | – | EU683073 | – | – |
| Coryneum umbonata | AR 3897 | BPI 843585 | Quercus cerris | – | EU683074 | – | – |
| Corynym arausiaca | MFLUCC 13–0658 | MFLU 17–0875 | Quercus sp. | MF190120 | MF190066 | MF377574 | MF377609 |
| MFLUCC 15–1110 | BBH 42437 | Quercus sp. | MF190121 | MF190067 | MF377575 | MF377610 | |
| Crinitospora pulchra | CBS 138014 | CBS H-21729 | Mangifera indica | KJ710466 | KJ710443 | – | – |
| Cryphonectria parasitica | ATCC 38755 | – | Castanea dentata | AY141856 | EU199123 | EU222014 | – |
| AFTOL ID 2123 | ATCC 38755 | – | – | – | DQ862033 | DQ862017 | |
| Cryptodiaporthe aesculi | AR 3580 | BPI 748430 | Aesculus hippocastanum | EU199179 | AF408342 | – | EU199138 |
| CBS 109765 | AFTOL-ID 1238 | Aesculus hippocastanum | – | DQ836905 | – | DQ836892 | |
| Cryptometrion aestuescens | CMW 18790 | PREM 60249 | Eucalyptus grandis | GQ369458 | HQ730869 | – | – |
| CMW 18793 | – | Eucalyptus grandis | GQ369459 | HQ730870 | – | – | |
| Cryptosporella hypodermia | CBS 116866 | BPI 748432 | Ulmus minor | EU199181 | AF408346 | – | EU199140 |
| Cryptosporella suffusa | CBS 121077 | BPI 871231 | Alnus incana | EU199184 | EU199124 | – | EU199142 |
| Cytospora ambiens | ATCC 52280 | ATCC 52280 | Acer rubrum | AY347345 | AF277146 | – | – |
| Cytospora austromontana | Willow21 | – | – | KM669911 | – | KM669767 | – |
| Cytospora carbonacea | CFCC 50056 | – | Ulmus pumila | KP281263 | KP310809 | KP310852 | – |
| Cytospora cedri | CBS 196.50 | – | – | AF192311 | – | JX438575 | – |
| Cytospora centrivillosa | MFLUCC 16–1206 | MFLU 17–0887 | Sorbus domestica | MF190122 | MF190068 | – | MF377600 |
| MFLUCC 17–1660 | BBH 42449 | Sorbus domestica | MF190123 | MF190069 | – | MF377601 | |
| – | MFLU 17–0999 | Sorbus domestica | MF190124 | MF190070 | – | – | |
| Cytospora ceratosperma | AR 3426 | – | – | – | EU255209 | – | – |
| Cytospora chrysosperma | CFCC 89630 | – | Salix psammophila | KF765674 | KF765690 | – | KF765706 |
| Cytospora fraxinigena | MFLUCC 14–0868 | BBH 42442 | Fraxinus ornus | MF190133 | MF190078 | – | – |
| – | MFLU 17–0880 | Fraxinus ornus | MF190134 | MF190079 | – | – | |
| Cytospora germanica | CXY 217 | – | Populus sp. | JQ086564 | JX524618 | – | – |
| Cytospora hippophaes | CFCC 89640 | – | Hippophae rhamnoides | KF765682 | KF765698 | KP310865 | KF765714 |
| Cytospora junipericola | – | BBH 42444 | Juniperus communis | MF190126 | MF190071 | MF377579 | – |
| – | MFLU 17–0882 | Juniperus communis | MF190125 | MF190072 | MF377580 | – | |
| Cytospora mali | CFCC 50044 | – | Malus baccata | KR045637 | KR045717 | – | – |
| Cytospora malicola | SXFX–V2 | – | Malus pumila | GU174579 | – | JQ900335 | – |
| Cytospora melanodiscus | Jimslanding2 | – | Alnus tenuifolia | JX438621 | – | JX438605 | – |
| Worrall2b | – | Alnus tenuifolia | JX438620 | – | JX438606 | – | |
| Cytospora melnikii | MFLUCC 16–0635 | T 1104 | Populus nigra | KY417736 | KY417770 | – | – |
| Cytospora nivea | CFCC 89643 | – | Salix psammophila | KF765685 | KF765701 | – | KF765717 |
| Cytospora punicae | CBS 199.50 | Punica granatum | JX438622 | – | JX438568 | – | |
| Cytospora quercicola | MFLUCC 14–0867 | BBH 42443 | Quercus sp. | MF190129 | MF190073 | – | – |
| – | MFLU 17–0881 | Quercus sp. | MF190128 | MF190074 | – | – | |
| Cytospora ribis | CFCC 50027 | – | Ulmus pumila | KP281268 | KP310814 | KP310857 | – |
| Cytospora rosae | MFLUCC 14–0845 | MFLU 17–0885 | Rosa canina | MF190131 | MF190075 | – | – |
| MFLUCC 17–1664 | BBH 42447 | Rosa canina | MF190130 | MF190076 | – | – | |
| Cytospora sacculus | CFCC 89625 | – | Juglans regia | KR045646 | KR045725 | KP310861 | – |
| Cytospora salicina | MFLUCC 16–0637 | T-1017 | Salix fragilis | KY417751 | KY417785 | – | – |
| MFLUCC 16–1190 | MFLU 17–1655 | Cornus sanguinea | MF190132 | MF190077 | – | – | |
| Cytospora sordida | HMBF 159 | – | Juglans regia | KF225613 | KF225627 | – | – |
| Cytospora sp. | CMON41 | – | Phaseolus vulgaris | JQ753989 | JQ754081 | – | – |
| Cytospora translucens | CZ320 | – | – | FJ755269 | FJ755269 | – | – |
| Diaporthe azadirachtae | TN 01 | – | Azadirachta indica | KC631323 | – | – | – |
| Diaporthe cassines | CBS 136440 | CPC 21916 | Cassine peragua | KF777155 | KF777208 | KF777244 | – |
| Diaporthe cynaroidis | 150e | – | Myrtus communis | KC959207 | – | – | – |
| CBS 122676 | CMW 22190 | Protea cynaroides | KC343058 | EU552122 | – | – | |
| Diaporthe decedens | CBS 114281 | UPSC 2957 | Corylus avellana | KC343059 | AF408348 | – | – |
| Diaporthe eres | MFLUCC 17–1667 | T400 | Fraxinus pennsylvanica | MF190137 | MF190080 | MF377594 | – |
| MFLUCC 17–1668 | MFLU 17–0890 | Fraxinus pennsylvanica | MF190138 | MF190081 | MF377595 | – | |
| MFLUCC 14–0862 | T98 | Catalpa bignonioides | MF190135 | MF190082 | MF377596 | – | |
| MFLUCC 17–1661 | MFLU 17–0889 | Catalpa bignonioides | MF190136 | MF190083 | MF377597 | – | |
| AR 5193 | – | Ulmus sp. | KJ210529 | – | KJ210550 | – | |
| PS57 | – | Glycine max | JF430494 | JF704176 | – | – | |
| Diaporthe eucalyptorum | MFLUCC 12–0306 | – | Leaf litter | KT459419 | – | KT459453 | – |
| Diaporthe litoricola | MFLUCC 16–1195 | BBH 42436 | Stem of sea-shore plant | MF190139 | MF190086 | – | – |
| MFLUCC 17–1657 | MFLU 17–0894 | Stem of sea-shore plant | MF190140 | MF190087 | – | – | |
| Diaporthe maytenicola | CBS 136441 | CPC 21896 | Maytenus acuminata | KF777157 | KF777210 | – | – |
| Diaporthe nobilis | Napa911 | – | – | KM669958 | – | KM669814 | – |
| Diaporthe rudis | – | IT 1526 | Acer campestre | MF190141 | MF190088 | MF377576 | – |
| – | MFLU 17–0895 | Acer campestre | MF190142 | MF190089 | MF377577 | – | |
| MFLUCC 16–1197 | BBH 42452 | Umbelliferous stem | MF190143 | MF190085 | – | – | |
| MFLUCC 17–1658 | MFLU 15–2661 | Umbelliferous stem | MF190144 | MF190084 | – | – | |
| LC6147 | – | Dendrobenthamia japonica | KY011890 | KY011864 | KY011901 | – | |
| BPI 748231 | – | – | – | AF362560 | – | – | |
| CBS 113201 | CBS H-7950 | Vitis vinifera | KC343234 | – | KC343960 | – | |
| Diaporthella corylina | CBS 121124 | BPI 871218 | Corylus sp. | KC343004 | – | – | – |
| Diaporthella sp. | CN5 | – | Corylus avellana | KP205483 | – | KP205456 | – |
| CN13 | – | Corylus avellana | KP205484 | – | KP205457 | – | |
| Dicarpella dryina | ICMP 14042 | – | Quercus sp. | KC145909 | – | KC145954 | – |
| ICMP 14043 | – | Quercus ilex | KC145858 | – | KC145955 | – | |
| Diplodina microsperma | CBS 114545 | CPC 2336 | Protea sp. | JN712461 | JN712525 | – | – |
| Discula destructiva | CBS 109771 | BPI 1107757 | Cornus nuttallii | EU199186 | AF408359 | – | EU199144 |
| MD 254 | BPI 1107741 | Cornus florida | AF429741 | AF429721 | AF429732 | – | |
| Disculoides eucalypti | CPC 17650 | – | Eucalyptus sp. | JQ685517 | JQ685523 | – | – |
| Disculoides eucalyptorum | CBS 132184 | CPC 17648 | Eucalyptus viminalis | NR120090 | – | – | – |
| Ditopella biseptata | – | MFLU 17–0884B | Alnus glutinosa | MF190147 | MF190091 | – | MF377616 |
| – | MFLU 17–0884 | Alnus glutinosa | MF190148 | MF190090 | – | MF377617 | |
| Ditopella ditopa | CBS 109748 | BPI 748439 | Alnus glutinosa | DQ323526 | EU199126 | – | EU199145 |
| Ditopellopsis sp. | CBS 121471 | BPI 872061 | Clethra alnifolia | EU254763 | EU255088 | EU221936 | EU219254 |
| Dwiroopa lythri | AR 3383 | BPI 747560 | Lythrum salicaria | – | AF408364 | – | – |
| Endothia gyrosa | CMW 2091 | CRY 1515 | Quercus palustris | AF046905 | AY194114 | – | – |
| Endothiella gyrosa | CMW 10436 | AFTOL-ID 1223 | Quercus sp. | AF452117 | – | – | – |
| Erythrogloeum hymenaeae | CPC 18819 | – | Hymenaea courbaril | JQ685519 | JQ685525 | – | – |
| Eutypella sp. | MFLUCC 16–1215 | BBH 42446 | Alnus cordata | MF190165 | MF190112 | MF377578 | MF377618 |
| Foliocryphia eucalypti | CBS 124779 | CPC 12494 | Eucalyptus coccifera | GQ303276 | GQ303307 | – | – |
| Gnomonia gnomon | CBS 829.79 | Monod 267 LAU | Populus sp. | AY818957 | AY818964 | EU221905 | – |
| CBS 199.53 | – | Corylus avellana | DQ491518 | AF408361 | EU221885 | EU219295 | |
| Gnomoniella fraxini | AR 3999 | BPI 843391 | Fraxinus americana | AY455814 | AY455818 | – | – |
| AR 2793 | BPI 746411 | Fraxinus americana | AY455813 | AY455817 | – | – | |
| Gnomoniopsis agrimoniae | MFLUCC 14–0844 | MFLU 17–0888 | Agrimonia eupatoria | – | MF190093 | MF377585 | – |
| MFLUCC 17–1662 | BBH 42450 | Agrimonia eupatoria | – | MF190092 | MF377586 | – | |
| Gnomoniopsis alderdunensis | CBS 125680 | BPI 879186 | Rubus parviflorus | GU320825 | – | – | – |
| Gnomoniopsis chamaemori | CBS 803.79 | Monod 345 LAU | Rubus chamaemorus | EU254808 | EU255107 | – | – |
| Gnomoniopsis racemula | AR 3892 | BPI 871003 | Epilobium angustifolium | EU254841 | EU255122 | EU221889 | EU219241 |
| Greeneria saprophytica | MFLUCC 12–0298 | MFLU 13–0255 | Syzygium cumini | KJ021933 | KJ021935 | – | – |
| Greeneria uvicola | FI1 2007 | – | Vitis sp. | HQ586009 | GQ870619 | – | – |
| FI1 2008 | – | Vitis sp. | HQ586010 | GQ870620 | – | – | |
| Hapalocystis berkeleyi | AR 3851 | – | – | – | EU683069 | – | – |
| MFLUCC 13–0662 | IT 1187 | Platanus sp | – | KP744486 | – | – | |
| Harknessia eucalypti | CBS 342.97 | – | Eucalyptus regnans | AY720745 | AF408363 | – | – |
| CPC 13643 | – | Eucalyptus regnans | JQ706089 | JQ706215 | – | – | |
| Harknessia karwarrae | CPC 10928 | – | Eucalyptus botryoides | AY720748 | AY720841 | – | – |
| Harknessia molokaiensis | CBS 114877 | – | Eucalyptus robusta | AY720749 | AY720842 | – | – |
| CPC 19269 | – | Eucalyptus cypellocarpa | JQ706127 | JQ706248 | – | – | |
| Harknessia weresubiae | CBS 113075 | – | Eucalyptus sp. | AY720741 | AY720835 | – | – |
| CPC 5109 | – | Eucalyptus sp. | AY720744 | AY720838 | – | – | |
| Hercospora tiliae | AR 3526 | – | Tilia tomentosa | – | AF408365 | – | – |
| Holocryphia eucalypti | CBS 115852 | CMW 14545 | Eucalyptus sp. | JQ862840 | JQ862797 | JQ863037 | – |
| CMW 7033 | PREM 56305 | Eucalyptus saligna | JQ862838 | JQ862795 | JQ863035 | – | |
| Hyaliappendispora galii | MFLUCC 16–1208 | MFLU 17–0893 | Galium sp. | MF190150 | MF190095 | MF377587 | – |
| MFLUCC 17–1761 | MFLU 17–0966 | Galium sp. | MF190149 | MF190094 | MF377588 | – | |
| Hyalorostratum brunneisporum | A573 2b | ILL 40792 | – | – | HM191720 | – | – |
| Immersiporthe knoxdaviesiana | CMW 37314 | PREM 60740 | Rapanea melanophloeos | JQ862770 | JQ862760 | – | – |
| CMW 37319 | PREM 60739 | Rapanea melanophloeos | JQ862765 | JQ862755 | – | – | |
| Juglanconis juglandina | D96 | WU 35960 | Juglans regia | KY427145 | KY427145 | KY427214 | KY427195 |
| AR 3860 | WU 35959 | Juglanconis juglandina | KY427149 | – | KY427218 | KY427199 | |
| CBS 121083 | BPI 843622 | Juglans regia | KY427148 | KY427148 | KY427217 | KY427198 | |
| Juglanconis oblonga | MAFF 410216 | TFM FPH 2623 | Juglans ailanthifolia | KY427153 | KY427153 | KY427222 | KY427203 |
| MAFF 410217 | TFM FPH 3599 | Juglans ailanthifolia | KY427154 | KY427154 | KY427223 | KY427204 | |
| Juglanconis pterocaryae | MAFF 410079 | TFM FPH 3373 | Pterocarya rhoifolia | KY427155 | KY427155 | KY427224 | KY427240 |
| Lamproconium desmazieri | AR 3525 | BPI 748445 | Tilia sp. | – | AF408372 | – | – |
| MFLUCC 15–0870 | MFLU 15–1940 | Tilia sp. | KX430134 | KX430135 | MF377591 | MF377605 | |
| MFLUCC 14–1047 | MFLU 14–0780 | Tilia sp. | KX430132 | KX430133 | MF377592 | ||
| MFLUCC 15–0872 | MFLU 15–2111 | Tilia sp. | KX430139 | AF408372 | MF377593 | MF377606 | |
| MFLUCC 15–0873 | MFLU 15–2192 | Tilia sp. | KX430140 | KX430141 | – | – | |
| MFLUCC 15–0871 | MFLU 15–2037 | Tilia sp. | KX430136 | KX430137 | – | – | |
| Lanspora coronata | AFTOL-ID 736 | JK 5839A | – | – | U46889 | – | DQ470899 |
| Lasmenia sp. | CBS 124122 | LMS 2011b | Nephelium lappaceum | GU797405 | JF838337 | – | – |
| CBS 124123 | LMS 2011c | Nephelium lappaceum | GU797406 | JF838338 | – | – | |
| CBS 124124 | LMS 2011d | Nephelium lappaceum | JF838336 | JF838341 | – | – | |
| CBS 124125 | LMS 2011a | Nephelium lappaceum | GU797407 | JF838340 | – | – | |
| Latruncellus aurorae | CBS 125526 | PREM 60348 | Galpinia transvaalica | HQ171209 | HQ171214 | – | – |
| CBS 124904 | PREM 60349 | Galpinia transvaalica | GU726946 | HQ171213 | – | – | |
| Leucostoma kunzei | ATCC 64881 | ATCC 64881 | Picea pungens | AY347320 | – | JX438595 | – |
| Luteocirrhus shearii | CBS 130776 | PERTH 08439362 | Banksia baxteri | KC197021 | KC197019 | – | – |
| CBS 130775 | PERTH 08355312 | Banksia baxteri | KC197024 | KC197018 | – | – | |
| Macrohilum eucalypti | CPC 10945 | – | Eucalyptus sp. | DQ195781 | DQ195793 | – | – |
| CPC 19421 | CBS H-22279 | Eucalyptus piperita | KR873244 | KR873275 | – | – | |
| Mamianiella coryli | – | BPI 877578 | Corylus californica | EU254862 | – | – | – |
| Marsupiomyces epidermoidea | – | MFLU 15–2921 | Quercus robur | – | MF190058 | – | – |
| – | BBH 42451 | Quercus robur | – | MF190059 | – | – | |
| Marsupiomyces quercina | MFLUCC 13–0664 | MFLU 17–0876 | Quercus sp. | MF190116 | MF190061 | – | – |
| MFLUCC 14–0566 | BBH 42438 | Quercus sp. | MF190117 | MF190060 | – | – | |
| Mastigosporella anisophylleae | CPC 22461 | – | Anisophyllea sp. | KF779492 | KF777221 | – | – |
| CBS 136421 | CBS H-21429 | Anisophyllea sp. | NR137844 | – | – | – | |
| Mazzantia napelli | AR 3498 | BPI 748443 | Aconitum lycoctonum | – | AF408368 | EU222017 | EU219345 |
| AFTOL-ID 2126 | AR 3498 | – | – | – | – | DQ862020 | |
| Melanconiella chrysodiscosporina | MFLUCC 17–1671 | MFLU 16–1309 | Fagus sylvatica | MF190166 | – | – | – |
| Melanconiella chrysomelanconium | – | MFLU 17–0879 | Carpinus betulus | MF190167 | MF190113 | – | MF377619 |
| Melanconiella ellisii | – | BPI 878343 | Carpinus caroliniana | JQ926271 | – | JQ926406 | JQ926339 |
| Melanconiella spodiaea | SPOD1 | WU 31854 | Carpinus betulus | JQ926301 | – | – | JQ926367 |
| Melanconis alni | AR 3748 | BPI 872035 | Alnus viridis | EU199195 | EU199130 | – | EU199153 |
| AR 3500 | BPI 748444 | Alnus viridis | – | AF408371 | EU221896 | EU219300 | |
| Melanconis italica | MFLUCC 17–1659 | MFLU 15–1112 | Alnus cordata | MF190152 | MF190097 | – | MF377602 |
| MFLUCC 16–1199 | MFLU 17–0883 | Alnus cordata | MF190151 | MF190096 | – | – | |
| Melanconis marginalis | AR 3442 | BPI 748446 | Alnus rubra | EU199197 | AF408373 | EU221991 | EU219301 |
| Melanconis stilbostoma | E01051 | – | – | AY577814 | AY577813 | – | – |
| E00153 | – | – | AY577811 | AY577810 | – | – | |
| AR 3548 | – | Betula sp. | – | AF362567 | – | – | |
| AR 3501 | BPI 748234 | Betula alba | – | – | EU221886 | EU219299 | |
| Microascospora fragariae | 1.1 | – | – | HM854850 | – | – | – |
| 1.3 | – | – | HM854852 | – | – | – | |
| 12 | – | – | HM854849 | – | – | – | |
| Microascospora rubi | – | MFLU 17–0886 | Rubus ulmifolia | MF190154 | MF190098 | MF377581 | MF377611 |
| – | BBH 42448 | Rubus ulmifolia | MF190153 | MF190099 | MF377582 | MF377612 | |
| Microthia havanensis | CMW 11298 | PREM 57518 | Eucalyptus saligna | AY214320 | AY194091 | – | – |
| CMW 38367 | – | Psidium cattleianum | KJ027495 | – | – | – | |
| Myrmecridium montsegurinum | JF 13180 | PRM 934684 | Fraxinus excelsior | KT991674 | KT991664 | – | KT991654 |
| Occultocarpon ailaoshanense | LCM 524.01 | BPI 879253 | Alnus nepalensis | JF779849 | JF779853 | – | JF779856 |
| LCM 522.01 | BPI 879254 | Alnus nepalensis | JF779848 | JF779852 | JF779862 | JF779857 | |
| Ophiodiaporthe cyatheae | YMJ 1364 | HAST 1364 | Cyathea lepifera | JX570889 | JX570891 | KC465406 | JX570893 |
| Ophiognomonia melanostyla | LCM 389.01 | BPI 879257 | Tilia cordata | JF779850 | JF779854 | – | JF779858 |
| Ophiognomonia vasiljevae | AR 4298 | BPI 877671 | Juglans nigra | EU254977 | EU255162 | EU221999 | EU219331 |
| Ophiostoma gemellus | CMW 23059 | – | Tarsonemus sp. | DQ821562 | DQ821533 | – | – |
| Pachytrype princeps | Rogers s.n. | – | – | – | FJ532382 | – | – |
| Pachytrype rimosa | FF1066 | – | – | – | FJ532381 | – | – |
| Papulosa amerospora | AFTOL-ID 748 | JK 5547F | – | – | DQ470950 | – | DQ470901 |
| Paradiaporthe artemisiae | MFLUCC 14–0850 | MFLU 12–2131 | Artemisia sp. | MF190155 | MF190100 | MF377583 | – |
| MFLUCC 17–1663 | BBH 42448 | Artemisia sp. | MF190156 | MF190101 | MF377584 | – | |
| Phaeoacremonium aleophilum | CBS 631.94 | – | Vitis vinifera | AF266647 | AB278175 | KF764643 | – |
| Phaeoacremonium vibratilis | CBS 117115 | BPI 2460 | Fagus sylvatica | KF764573 | DQ649065 | KF764645 | HQ878611 |
| Phaeoappendispora thailandensis | MFLUCC 13–0161 | MFLU 17–0873 | Quercus sp. | MF190157 | MF190102 | – | MF377613 |
| MFLUCC 17–1762 | BBH 42435 | Quercus sp. | MF190158 | MF190103 | – | MF377614 | |
| Phaeocytostroma ambiguum | CPC 17077 | – | Zea mays | FR748041 | FR748100 | FR748073 | – |
| CPC 17078 | – | Zea mays | FR748044 | FR748101 | FR748074 | – | |
| Phaeodiaporthe appendiculata | CBS 123821 | WU 32449 | Acer campestre | KF570156 | KF570156 | – | – |
| CBS 123809 | WU 32448 | Acer campestre | KF570155 | KF570155 | – | – | |
| Phragmoporthe conformis | MFLUCC 14–0567 | MFLU 15–2662 | Alnus glutinosa | KU315388 | KU315389 | KU315391 | – |
| AR 3632 | BPI 748450 | Alnus rubra | – | AF408377 | – | – | |
| Plagiostoma dilatatum | LCM 402.02 | BPI 878957 | Salix irrorata | GU367070 | – | – | GU367104 |
| Plagiostoma jonesii | MFLUCC 16–1189 | MFLU 17–0878 | Umbelliferous stem | MF190159 | MF190104 | MF377589 | – |
| MFLUCC 17–1654 | BBH 42440 | Umbelliferous stem | MF190160 | MF190105 | MF377590 | – | |
| Plagiostoma salicellum | CBS 109755 | BPI 843490 | Salix sp. | EU255047 | – | EU221912 | – |
| Plagiostoma salicicola | MFLUCC 13–0656 | MFLU 17–0877 | Salix sp. | MF190161 | MF190106 | – | – |
| MFLUCC 17–1666 | BBH 42439 | Salix sp. | MF190162 | MF190107 | – | – | |
| Pleuroceras capreae | CBS 372.69 | – | Salix sp. | – | AF277143 | – | – |
| Pleuroceras oregonense | AR 4333 | BPI 877719 | Salix sitchensis | EU255060 | EU255196 | EU221931 | EU219313 |
| Pleuroceras pleurostylum | CBS 906.79 | Monod 469/LAU | Salix helvetica | EU255061 | EU255197 | EU221962 | EU219311 |
| Pleuroceras tenellum | CBS 121082 | BPI 871059 | Acer rubrum | EU199199 | EU255202 | EU221907 | EU199155 |
| Prosopidicola mexicana | CBS 113529 | CBS-H 7948 | Prosopis glandulosa | AY720709 | – | – | – |
| CBS 113530 | C 158 | Prosopis glandulosa | AY720710 | – | – | – | |
| Pseudoplagiostoma corymbiae | CPC 19287 | CBS H- 20957 | Corymbia sp. | JX069861 | JX069845 | – | – |
| Pseudoplagiostoma eucalypti | CBS 124807 | CBS H-20303 | Eucalyptus urophylla | GU973512 | GU973606 | GU973542 | – |
| CPC 14161 | – | Eucalyptus camaldulensis | GU973510 | GU973604 | GU973540 | – | |
| Pseudoplagiostoma oldii | CBS 115722 | – | Eucalyptus camaldulensis | GU973535 | GU973610 | GU973565 | – |
| CBS 124808 | CBS H-20300 | Eucalyptus camaldulensis | GU973534 | GU973609 | GU973564 | – | |
| Pseudoplagiostoma variabile | CBS 113067 | CBS H-20304 | Eucalyptus globulus | GU973536 | GU973611 | GU973566 | – |
| Pustulomyces bambusicola | MFLUCC 11–0436 | MFLU 13–0369 | Bambusa sp. | KF806752 | KF806753 | KF806755 | – |
| Pyricularia bothriochloae | CPC 21650 | CBS-H 21436 | Bothriochloa bladhii | KF777186 | KF777238 | – | – |
| Pyricularia oryzae | REA 8401 | PH0063 | Ophiuros exaltatus | KM484916 | KM485022 | – | – |
| Rossmania ukurunduensis | AR 3484 | BPI 747566 | Acer ukurunduense | – | EU683075 | – | – |
| Rostraureum tropicale | CMW 9972 | PREM 57519 | Terminalia ivorensis | AY167436 | AY194092 | – | – |
| CMW 9975 | – | Terminalia ivorensis | AY167439 | – | – | – | |
| Sillia ferruginea | AR 3440 | BPI 843619 | Corylus avellana | JF681959 | EU683076 | – | – |
| CBS 126567 | BPI 843619 | Corylus avellana | JF681959 | EU683076 | – | – | |
| Sirococcus tsugae | CBS 119626 | BPI 871167 | Tsuga mertensiana | EU199203 | EU199136 | EF512534 | EU199159 |
| AR 4010 | – | Cedrus deodara | EF512478 | EU255207 | EU221928 | EU219289 | |
| Sordaria fimicola | CBS 508.50 | – | Dung | AY681188 | AY681160 | – | DQ368647 |
| Stegonsporium acerophilum | CBS 117025 | WU 28050 | Acer saccharum | EU039982 | EU039993 | EU040027 | KF570173 |
| Stenocarpella macrospora | CBS 117560 | MRC 8615 | Zea mays | FR748048 | EU754219 | – | – |
| Stenocarpella maydis | CBS 117559 | MRC 8614 | Zea mays | FR748052 | DQ377937 | – | – |
| Stilbospora ellipsosporum | WJ 1840 | – | Carpinus betulus | – | AY616229 | – | – |
| Stilbospora macrosperma | CBS 121883 | – | Carpinus betulus | JX517290 | JX517299 | – | KF570196 |
| Sydowiella depressula | CBS 813.79 | – | Rubus sp. | EU552156 | – | – | – |
| Sydowiella fenestrans | CBS 125530 | BPI 843503 | Chamerion angustifolium | JF681956 | EU683078 | – | – |
| Sydowiella urticicola | MFLUCC 13–0665 | MFLU 13–0260 | Urtica dioica | – | MF190108 | – | – |
| MFLUCC 17–1665 | BBH 42439 | Urtica dioica | – | MF190109 | – | – | |
| Thyridium vestitum | AFTOL-ID 172 | OSC 100064 | – | – | AY544671 | – | DQ470890 |
| Tubakia seoraksanensis | CBS 127490 | – | Quercus mongolica | HM991734 | KP260499 | – | – |
| BJFCCC140824–15 | – | – | KP260502 | KP260501 | – | – | |
| Tubakia thailandensis | MFLUCC 12–0303 | MFLU 13–0260 | Decaying leaf | MF190163 | MF190110 | – | – |
| MFLUCC 17–1672 | MFLU 13–0260B | Decaying leaf | MF190164 | MF190111 | – | – | |
| Ursicollum fallax | CMW 18119 | PREM 58840 | Coccoloba uvifera | DQ368755 | EF392860 | – | – |
| Valsalnicola oxystoma | AR 4833 | BPI 884137 | Alnus viridis | JX519559 | JX519563 | – | – |
| AR 5137 | BPI 884135 | Alnus tenuifolia | JX519561 | – | – | – | |
| Valsella salicis | AR 3514 | BPI 748461 | Salix fragilis | – | EU255210 | EU222018 | EU219346 |
| Valseutypella multicollis | CBS 105.89 | – | Quercus ilex | DQ243803 | – | – | – |
| Waydora typica | PDD 103894 | PDD 103894 | – | KF727412 | KF727413 | – | – |
AFTOL: Assembling the Fungal Tree of Life culture collection; AMH: Ajrekar Mycological Herbarium, India; ATCC: American Type Culture Collection, Virginia, USA; BBH: National Science and Technology Development Agency, Thailand; BECM: British Empire and Commonwealth Museum, UK; BJFCCC: Beijing Forestry University, China; BPI: U.S. National Fungus Collections, Systematic Botany and Mycology Laboratory, USA; CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, The Netherlands; CFCC: China Forestry Culture Collection Center, Beijing, China; CMW: Forestry and Agricultural Biotechnology Institute, University of Pretoria, South Africa; CPC: Culture collection of Pedro Crous, The Netherlands; FI: Museo di Storia Naturale dell'Universita, Italy; HAST: Herbarium, Biodiversity Research Center, Academia Sinica, Taiwan; HHUF: Hirosaki University, Japan; ICMP: International Collection of Micro-organisms from Plants, New Zealand; ILL: University of Illinois, USA; IMI: International Mycological Institute, Kew, UK; JF: Jonkershoek Forestry Research Centre, South Africa; LCM: Universidad de Chile, Laboratorio de Citogenetica de Mamiferos, Chile; MAFF: MAFF Genebank, Ministry of Agriculture Forestry and Fisheries, USA; MFLU: Mae Fah Luang University herbarium, Thailand; MFLUCC: Mae Fah Luang University Culture Collection, Thailand; MNA: Museum of Northern Arizona, USA; NFCCI: National Fungal Culture Collection, India; OSC: Oregon State University, Corvallis, USA; PERM: University of Perm, South Africa; PERTH: Western Australian Herbarium, Australia; PH: Academy of Natural Sciences of Drexel University, Philadelphia, PA; RMF: Rocky Mountain Herbarium, University of Wyoming, USA; SDSU: San Diego State University, USA; STE-U: University of Stellenbosch, Plant Pathology Department, South Africa; TBRC: Thailand Bioresource Research Center, Bangkok, Thailand; TFM: Forestry and Forest Products Research Institute, Matsunosato, Japan; UPSC: Fungal Culture Collection at the Botanical Museum, Uppsala University, Sweden; WU: Universitaet Wien, Austria.
Maximum likelihood analysis (ML) was performed by RAxMl GUI v. 1.3 (Stamatakis et al., 2008, Silvestro and Michalak, 2012). The search strategy was set to rapid bootstrapping and the analysis was carried out with 1 000 replicates using the GTRGAMMAI model of nucleotide substitution, which was the best model predicted for the concatenated LSU nrDNA, ITS nrDNA, rpb2 and tef1 alignment by MrModeltest v. 2.3 (Nylander 2004).
For the Bayesian analyses (BI) of the individual loci and concatenated LSU nrDNA, ITS nrDNA, rpb2 and tef1 alignment, MrModeltest v. 2.3 (Nylander 2004) was used to determine the best nucleotide substitution model settings for MrBayes. A dirichlet state frequency was predicted for all four data partitions and GTR+I+G as best model for LSU nrDNA, ITS nrDNA, and rpb2; for tef1 the best model was GTR+G. The heating parameter was set to 0.2 and trees were saved every 1 000 generations (Ronquist et al. 2012). The Markov Chain Monte Carlo (MCMC) analysis of four chains started in parallel from a random tree topology.
The maximum parsimony analysis (MP) was performed with PAUP v. 4.0b10 (Swofford 2003). Ambiguously aligned regions were excluded and all characters were unordered and given equal weight. Alignment gaps were treated as a fifth character state. Trees were inferred using the heuristic search option with TBR branch swapping and 100 random sequence additions. MaxTrees were set to 1 000, branches of zero length were collapsed and all multiple parsimonious trees were saved. Tree length (TL), consistency index (CI), retention index (RI), rescaled consistency index (RC), homoplasy index (HI), and log likelihood (-ln L) were calculated for trees generated under different optimality criteria. The robustness of the most parsimonious trees was evaluated by 1 000 bootstrap replications resulting from the maximum parsimony analysis, each with 10 replicates of random step-wise addition of taxa (Felsenstein 1985). The Kishino–Hasegawa tests (Kishino & Hasegawa 1989) were performed to determine whether the trees inferred under different optimality criteria were significantly different.
Trees were viewed in FigTree v. 1.4.3 (Rambaut 2012). The final alignments and the trees obtained were deposited in TreeBASE (http://purl.org/phylo/treebase/phylows/study/TB2:S21148) and are available under study accession no. S21148.
Results
To reveal the phylogenetic position of genera, families and genera incertae sedis within the order Diaporthales, a phylogenetic analysis was performed with LSU nrDNA, ITS nrDNA, rpb2 and tef1 sequence data. Sequences of representative species were selected from Maharachchikumbura et al., 2016, Norphanphoun et al., 2016, Voglmayr et al., 2017 and supplemented with sequences from GenBank. The LSU nrDNA, ITS nrDNA, rpb2, tef1 and combined data matrices contained 1 423, 735, 1 064, 427 and 3 652 characters with gaps, respectively. The alignment comprised 310 strains and Eutypella sp. (MFLUCC 16–1215) was selected as the outgroup.
The same concatenated alignment was subjected to phylogenetic analyses, including a Bayesian analysis, a maximum parsimony analysis and a maximum likelihood analysis. The concatenated sequence alignment contained 2 027 parsimony-informative characters, 385 were variable and parsimony uninformative and 1 241 were constant. The parsimony analysis yielded the maximum of 1 000 equally most parsimonious trees (TL = 16 973 steps; CI = 0.278; RI = 0.728; RC = 0.202; HI = 0.722). The ML analysis yielded a tree with a likelihood value of ln: −75295.054554 and the following model parameters: alpha: 0.368178; Π(A): 0.246723, Π(C): 0.249231, Π(G): 0.277805, and Π(T): 0.226241. The Bayesian analysis lasted 72 151 000 generations (average standard deviation of split frequencies value = 0.016671) and the consensus trees and posterior possibilities were calculated from the 103 301 trees in each of the two run files, of which a total of 72 152 trees in each of the two run files, of which a total of 108 228 were sampled after discarding the first 25 % of generations for burn-in. The different data partitions contained 787, 529, 761 and 390 unique site patterns (LSU nrDNA, ITS nrDNA, rpb2 and tef1, respectively).
The phylogeny resulting from the analysis of combined gene sequence data is shown in Fig. 1. Overall, the topologies obtained from the different phylogenetic analyses were mostly similar and the best scoring RAxML tree is illustrated here. The bootstrap support values of maximum likelihood analysis (MLB), maximum parsimony analysis (MPB) and Bayesian posterior probability scores (PP) are noted at the nodes. The separation of Diaporthales from other Sordariomycetes taxa is well-supported (MLB/MPB/PP = 100/96/1). The order separates into 21 familial clades with good support values and two genera incertae sedis clades labelled as 5 and 18.
Consensus tree resulting from a maximum likelihood analysis of a combined LSU nrDNA, ITS nrDNA, rpb2 and tef1 sequence alignment for taxa of Diaporthales and other species in Sordariomycetes. Families are indicated in coloured blocks. RAxML bootstrap support values (MLB above 50 %), maximum parsimony bootstrap support values (MPB above 50 %) and Bayesian posterior probabilities (BPP above 0.90) are given at the nodes (MLB/MPB/BPP). The scale bar represents the expected number of changes per site. The tree is rooted to Eutypella sp (MFLUCC 16–1215). All the new sequences used in this study are in blue bold and type sequences are in black bold. The nodes that received maximum support (MLB/MPB/BPP = 100/100/1) are indicate by ∗
Clade 1 is represented by Gnomoniaceae with moderate support values (MLB/MPB/PP = 60/–/0.9) and comprised Alnecium, Ambarignomonia, Amphiporthe, Anisogramma, Apiognomonia, Apioplagiostoma, Asteroma, Cryptodiaporthe, Cryptosporella, Discula, Ditopella, Ditopellopsis, Gnomonia, Gnomoniella, Gnomoniopsis, Mamianiella, Marsupiomyces, Occultocarpon, Ophiognomonia, Phragmoporthe, Plagiostoma, Pleuroceras, Sirococcus and Valsalnicola. Anisogramma and Mamianiella formed a distinct clade with high support value. Morphologically they are similar and these genera appear to be congeneric. Therefore we synonymise Anisogramma under Mamianiella giving priority to the older name and its taxonomic stability. In addition, Mamianiella is nested in between Anisogramma species and this supports that both genera should be synonymised. Here we introduce one new genus and six new species to Gnomoniaceae. We introduce Plagiostoma salicicola based on morphology and phylogeny. Plagiostoma jonesii, the second species, was a fully-supported clade sister to Plagiostoma salicellum, P. dilatatum and P. salicicola. It is morphologically distinct from other species in Plagiostoma. Sequences of the asexual morph of Apiognomonia veneta (= Discula nervisequa) is included here and clustered with other Apiognomonia veneta strains. Ditopella biseptata is introduced based on phylogeny as well as morphology. Ditopella biseptata formed a fully-supported clade sister to D. ditopa. We introduce a new genus Marsupiomyces with two phylogenetically well-supported species, M. quercina and M. epidermoidea. Gnomoniopsis agrimoniae represents a new species distinct from other species of Gnomoniopsis.
Clade 2 is represented by Melanconidaceae sensu stricto with good support values (MLB/MPB/PP = 93/91/0.9) and it is considered as Melanconidaceae. Most genera listed under Melanconidaceae in recent publications are excluded from this family, based on morphology and available sequence data. We introduce a new Melanconis species as M. italica.
Clade 3 (MLB/MPB/PP = 100/92/1) represents the new family Apiosporopsidaceae which is introduced to accommodate a single genus, Apiosporopsis.
Clade 4 is represented by Juglanconidaceae (MLB/MPB/PP = 99/94/1), which was recently introduced by Voglmayr et al. (2017) based on a fungal species isolated from Juglans nigra.
Clade 5 is represented by Diaporthella species with low bootstrap support and is considered here as Diaporthales genera incertae sedis, pending the availability of sequence data for the type species Diaporthella aristata.
The family Cryphonectriaceae is represented by Clade 6 which is phylogenetically poorly-supported (MLB/MPB/PP = 63/–/0.9) but morphologically distinct from other families in Diaporthales. Almost all taxa in this family have molecular data.
Clade 7 is represented by the Harknessiaceae, which is phylogenetically poorly-supported (MLB/MPB/PP = 60/68/–). Species of Harknessia and wuestneia-like sexual morphs have been linked by morphological characteristics (Crous and Rogers, 2001, Crous et al., 2012b), but the types of both genera have not been linked by molecular data. Dwiroopa lythri clusters basal to Harknessia species with low support values (MLB/MPB/PP = 60/68/–; Fig. 1).
Schizoparmaceae (Clade 8) is phylogenetically well-supported (MLB/MPB/PP = 94/88/–). We introduce a new Coniella species as Coniella pseudokoreana and it clusters sister to Coniella straminea.
Clade 9 (MLB/MPB/PP = 91/71/1) represents the new family Erythrogloeaceae which is introduced in this study to accommodate Chrysocrypta, Disculoides and Erythrogloeum.
Clade 10 (MLB/MPB/PP = 93/–/1) currently encompassies 6 genera within the Melanconiellaceae. In particular, Melanconiella sisters to Microascospora where as Greeneria is nested in between Dicarpella and Tubakia. Microascospora is introduced here based on Microascospora rubi collected from Italy and Microascospora fragariae (= Sphaeronaemella fragariae) which was already placed in Microascales. Other genera were previously placed in Melanconidaceae, but phylogenetically they do not cluster with M. stilbostoma, which is the family type of the Melanconidaceae. Dicarpella and Tubakia form a distinct clade within this family to represent both genera as holomorphs. Greeneria saprophytica is distant from Greeneria uvicola, which is the type of this genus. We introduce a new Tubakia species as T. thailandensis. A new genus Microascospora based on M. rubi is introduced here and M. rubi strains have high support as a distinct species. Microascospora rubi forms a sister clade to Microascospora fragariae (= Sphaeronaemella fragariae) and high bootstrap support values confirmed it as a species. In addition, we include sequences of Melanconiella chrysodiscosporina and M. chrysomelanconium from recently collected specimens.
Clade 11 is represented by the monotypic family Auratiopycnidiellaceae, which is newly introduced in this study based on Auratiopycnidiella tristaniopsis, and it is morphologically and phylogenetically well-supported (MLB/MPB/PP = 95/–/1).
Clade 12 comprises the monotypic family Pseudoplagiostomaceae with full-support (MLB/MPB/PP = 100/100/1).
Clade 13 (MLB/MPB/PP = 100/68/1) represents Apoharknessiaceae to accommodate Apoharknessia and Lasmenia.
Clade 14 is represented by the family Diaporthaceae with good support (MLB/MPB/PP = 86/91/0.9). We also introduce several new genera such as Chiangraiomyces which is typified by C. bauhiniae; Paradiaporthe, typified by P. artemisiae and Hyaliappendispora typified by H. galii. Paradiaporthe is nested with reliable support between Chiangraiomyces and Phaeocytostroma while Chiangraiomyces, clusters sister to Ophiodiaporthe. Hyaliappendispora grouped sister to Phaeodiaporthe. Here we included several new isolates of Diaporthe species and Diaporthe litoricola forms a fully-supported clade sister to Diaporthe maytenicola. In addition, we include a new collection of Diaporthe eres and a new collection of Diaporthe rudis.
Clade 15 represents the family Macrohilaceae with full-support (MLB/MPB/PP = 100/100/0.9).
Clade 16 is represented by Cytosporaceae with good support (MLB/MPB/PP = 88/51/1). In addition to Cytospora, we include Waydora and Pachytrype in Cytosporaceae based on molecular data. Here we introduce five Cytospora species viz., Cytospora centrivillosa, Cytospora fraxini, Cytospora junipericola, Cytospora quercicola, and Cytospora rosae. Cytospora centrivillosa forms a distinct clade that is sister to Cytospora melanodiscus and Cytospora mali. Cytospora melanodiscus is morphologically quite different having 1-septate ascospores. Here we added sequences of C. salicina from freshly collected specimens.
Prosopidicolaceae (Clade 17) is introduced to accommodate Prosopidicola mexicana.
Phaeoappendicospora thailandensis (Clade 18) forms separate fully-supported (MLB/MPB/PP = 100/100/0.9) clade. It currently does not have a high affinity with any known family in Diaporthales, therefore we consider this species as Diaporthales genera incertae sedis.
Clade 19 represents the family Stilbosporaceae that is fully-supported (MLB/MPB/PP = 100/100/1). Although Crinitospora is morphologically different from Stegonsporium and Stilbospora, inclusion of this genus in the family is phylogenetically well-supported.
Clade 20 comprises the family Coryneaceae and includes molecular data for Coryneum arausiaca collected from Italy. Coryneum arausiaca has high support (MLB/PP = 100/96/1) as a separate species.
Clade 21 represents the family Sydowiellaceae with very good support (MLB/MPB/PP = 98/79/1) and here we introduce a new species Sydowiella urticicola. Sydowiella urticicola clade received high support values.
Lamproconiaceae is represented by clade 22 and comprises Lamproconium and Hercospora.
The new family Asterosporiaceae (23) is introduced to accommodate Asterosporium asterospermum. This monogeneric family received high support (MLB/MPB/PP = 100/96/1) and is sister to the Sydowiellaceae and Lamproconiaceae.
Taxonomy
Diaporthales Nannf., Nova Acta R. Soc. Scient. Upsal. 8: 53. 1932.
Saprobic or pathogenic in plants, and animals, including humans or inhabiting soil. Sexual morph: Pseudostromata or ascostromata well-developed, poorly developed or absent, scattered, immersed or erumpent, solitary to aggregated, valsoid to diatrypoid, broadly elliptical, oval to circular from above, yellowish orange, pale brown, dark brown to black, some species turning purple or umber in KOH. Entostroma normally limited to the region near the perithecial walls, prosenchymatous, pale-coloured, and slightly differentiated from the surrounding bark tissue. Ectostromatic disc well- or poorly developed, subhyaline, yellowish white, pale brown, rarely dark brown to black, pulvinate, flat or slightly convex, orbicular, circular or somewhat irregular, with or without black zone or a crust consisting of fungus tissue, sclerotioid, coriaceous. Central column present or absent, if present beneath the disc more or less conical, comprising hyaline or pigmented hyphae mixed with a pigmented, cream, yellow, olive, brownish or grey, powdery amorphous substance. Ascomata perithecial, scattered, solitary or aggregated, immersed to erumpent, rarely superficial, globose to subglobose, sometimes circinate, arranged in a valsoid to diatrypoid configuration or single, coriaceous, sometimes with plate-like ornamentation around ostiole, black to brown, ostiolate, papillate. Papilla lacking or upright, long or short, one or more, central or eccentric, slanted to horizontal on host tissue, sometimes converging, with neck swollen at the tips, fuscous black to umber, ostiole with hyaline periphyses. Peridium thin or thick, comprising outer, dark, thick-walled, cells of textura angularis and inner, mostly small, hyaline, thin-walled, flattened cells of textura angularis. Hamathecium aparaphysate or comprising few broad cellular, filiform to cylindrical, septate to aseptate, branched to unbranched, hyaline paraphyses and sometimes parenchymatous cells attached at the base and asci dissolving at maturity. Asci generally 2–32-spored, unitunicate, ellipsoid, cylindrical, fusiform, clavate, oblong-clavate, broadly fusoid to cylindrical-fusoid, short pedicellate, apex blunt, usually with distinct, J- refractive ring. Ascospores overlapping uniseriate, biseriate, partially biseriate to fasciculately arrange, ovoid, ellipsoid, oblong, fusoid, cylindrical, filamentous or allantoid, aseptate to multi-septate, rarely distoseptate, constricted or not at the septa, hyaline, olivaceous to brown, smooth- to sometimes ornamented walled, ends mostly rounded, rarely pointed, multi-guttulate, straight or curved, smooth- to sometimes ornamented walled to rarely ornamented, hyaline to dark brown. Appendages absent or present; if present, apical or basal, subulate, navicular or whip-shaped, smooth, hyaline. Asexual morph: Coelomycetous. Stroma present or absent, immersed to superficial, opening by irregular rupture, globose, subglobose to irregular, solitary to gregarious, orange, brown to dark brown, sometimes loculate. Conidiomata amphigenous, eustromatic, punctiform, pycnidial or acervular, sometimes pyriform in section and divided into compartments by bending of peridium, subcuticular, peridermal or subepidermal, brown to black or orange with dark brown border, sometimes with a central, well-developed, pale brown, pseudoparenchymatous layer, becoming thinner or absent at the margin of the conidiomata, sometimes with pale coloured, ectostromatic disc and central column or with radiate scutella. Scutella convex, membranous, brown, somewhat translucent, with a central hyaline or pale disc, giving rise to radiating hyphae, thick-walled cells radiating from a central point, rounded to pointed at the tips. Peridium comprising pale to dark brown cells of textura angularis to textura globulosa. Paraphyses present or absent. If present, hyaline, cellular, subcylindrical, branched or not, with obtuse apex, septate, constricted at septa. Conidiophores reduced to conidiogenous cells or arising from the upper most cells of basal and parietal tissue or under the developing scutellum, densely aggregated or few, filiform, fusiform, cylindrical to globose, simple or branched, septate or aseptate, sometimes septate only at the base, smooth, hyaline or hyaline at the top, pale brown at the base, sometimes dimorphic. Alpha conidiophores tightly aggregated, subcylindrical, branched in mid region, consisting of few supporting cells, giving rise to septate, ampulliform, cylindrical to irregular conidiogenous cells or paraphyses, straight to sinuous, septate, cylindrical, hyaline to pale brown, branched only at the base, smooth, formed from the innermost layer cells of the conidiomatal wall, sometimes with terminal and lateral apex, with minute periclinal thickening and collarette. Beta conidiophores interspersed among alpha conidiophores, hyaline, subcylindrical, branched, septate. Conidiogenous cells lining the inner cavity of conidioma, enteroblastic to holoblastic, annellidic or phialidic, discrete or integrated, hyaline to olivaceous, smooth, lageniform, subcylindrical to ampulliform, with terminal truncate locus, simple or branched, proliferating several times percurrently near apex, with flaring collarettes or apex truncate, with minute periclinal thickening or terminal truncate locus. Conidia broadly ellipsoid, oval, obovoid, allantoid, fusoid to sigmoid, sinuate to slightly angular, hyaline to brown, hyaline when immature, becoming medium brown to dark brown at maturity, smooth-walled, guttulate, aseptate to septate or distoseptate, apex obtuse, base truncate with a visible scar or a flat protruding scar at the base, sometimes the apical and basal cell darker than other cells or with hyaline tip in apical cell, sometimes with or without a longitudinal germ slit, sometimes with marginal frill or becoming golden brown at germination, with solitary, brown, wavy germ tubes.
Notes: The order Diaporthales was introduced to accommodate “true” diaportheen taxa and Eriksson & Winka (1997) accommodated Diaporthales in Sordariomycetidae. Barr, 1978, Monod, 1983, Castlebury et al., 2002, Rossman et al., 2007, Maharachchikumbura et al., 2015, Maharachchikumbura et al., 2016, Rossman et al., 2015 and Voglmayr et al. (2017) clarified the taxonomic and phylogenetic concepts. Maharachchikumbura et al. (2015) introduced the subclass Diaporthomycetidae to accommodate the order Diaporthales. Morphologically and phylogenetically this is a well-supported order comprising Apiosporopsidaceae, Apoharknessiaceae, Asterosporiaceae, Auratiopycnidiellaceae, Coryneaceae, Cryphonectriaceae, Cytosporaceae, Diaporthaceae, Erythrogloeaceae, Gnomoniaceae, Harknessiaceae, Juglanconidaceae, Lamproconiaceae, Macrohilaceae, Melanconidaceae, Melanconiellaceae, Prosopidicolaceae, Pseudoplagiostomaceae, Schizoparmaceae, Stilbosporaceae, and Sydowiellaceae.
Apiosporopsidaceae Senan., Maharachch. & K.D. Hyde, fam. nov. MycoBank MB821538. Facesoffungi number FoF03455. Clade 3.
Parasitic on living leaves and twigs. Sexual morph: Ascomata scattered, black, oval to almost spherical, immersed in the leaf tissue beneath a thin, well-developed clypeus, neck lacking or only slightly papillate, periphysate. Peridium comprises 5–6 outer layers of dark, thick-walled cells of textura angularis and inner, thin-walled, strongly flattened cells of textura angularis. Hamathecium aparaphysate. Asci 8-spored, unitunicate, short-pedicellate, apex blunt with J- apical ring. Ascospores 1–2-seriate, elliptical to fusoid, often slightly flattened on one side, unicellular, hyaline. Asexual morph: Coelomycetous. Stroma loculate, globose to irregular, sometimes with beaks. Conidiogenous cells phialidic, short to elongate, simple or branched. Conidia oblong or cylindrical to allantoid, 1-celled, hyaline.
Type genus: Apiosporopsis (Traverso) Mariani.
Type species: Apiosporopsis saccardoana Mariani.
Apiosporopsis carpinea (Fr.) Mariani, Atti Soc. ital. Sci. nat. (Modena) 50: 165. 1911. Facesoffungi number FoF03456. Fig. 2.
Apiosporopsis carpinea (IMI 11662). A. Packet of the herbarium specimen. B. Herbarium specimen. C. Ascomata on substrate. D, E. Vertical section of ascomata. F–I. Asci. J–M. Ascospores. Scale bars: C = 500 µm, D, E = 50 µm, F–M = 10 µm.
Basionym: Xyloma carpini Fr., Observ. mycol. (Havniae) 2: 363. 1818.
Illustration: For asexual morph see Potebnia (1910).
Saprobic on over-wintered plants. Sexual morph: Clypeus 70–140 μm wide, 50–70 μm high, slight, prosenchymatous. Ascomata 112–250 μm diam, 140–170 μm high, globose or depressed, immersed, usually hypophyllous, apapillate, apex rounded with plane pore or short papillate or conic. Peridium 10–20 μm wide, comprising thick-walled, brown cells of textura angularis. Asci 40–75 × 8–14 μm, 8-spored, unitunicate, cylindrical, sessile, apical ring bilobed, distinct, shallow. Ascospores 10–15 × 3.5–6.5 μm, overlapping uniseriate, ellipsoid, ovoid or fusoid, straight or often inequilateral, guttulate, hyaline, aseptate. Asexual morph: Conidiomata acervular, superficial, black, coriaceous. Conidiophores reduced to conidiogenous cells. Conidiogenous cells 5–10 μm long, conical, wide, aseptate, hyaline. Conidia 12–15 × 8–9 μm, oblong to ellipsoid, hyaline, aseptate, with two small guttules (description of asexual morph from Potebnia 1910).
Material examined: Austria, Sonntagberg, New Rosenau, July, on leaves of Carpinus betulus (Betulaceae), P.P. Strasser, IMI 11662.
Notes: Traverso (1907) erected Apiosporopsis as a subgenus of Guignardia to accommodate Guignardia carpinea and G. veneta based on their distinct morphological characters. Mariani (1911) raised Apiosporopsis to generic rank describing A. saccardiana as a third species. Von Höhnel (1917) proposed Sphaerognomonia to accommodate Apiosporopsis carpinea. Reid & Dowser (1990) evaluated this genus and proposed Apiosporopsis as the correct name for Sphaerognomonia, retaining the type species as Apiosporopsis carpinea. Index Fungorum (2017) and MycoBank (2017) list another two species of Apiosporopsis as A. saccardoana and A. coronillae.
Apiosporopsis carpinea was recorded only on over-wintered living leaves. Gloeosporium robergei was reported as the asexual morph of A. carpinea (Potebnia, 1910, Treigien and Markovskaja, 2007). However, there are no molecular data to prove this. Gloeosporium robergei was reported as the causal agent of bud mortality and twig cankers on Ostrya virginiana (Sinclair & Hudler 1980). Sequences of this species (CBS 617.72 and CBS 738.68) placed the genus in the Diaporthales, but not in the Gnomoniaceae or Melanconidaceae (Sogonov et al. 2008). The molecular analysis of this study revealed that Apiosporopsis species formed a separate, well-supported clade (Fig. 1, Clade 3). Morphologically this clade is distinct from other families of Diaporthales having ascospores with pseudo-septate, sharply pointed ends, sessile unitunicate asci with a bilobed apical ring, and apapillate, immersed ascomata. Hence, we introduce the family Apiosporopsidaceae to accommodate these species.
Apoharknessiaceae Senan., Maharachch. & K.D. Hyde, fam. nov. MycoBank MB821881. Facesoffungi number FoF03457. Clade 13.
Endophytic, saprobic or pathogenic. Sexual morph: Undetermined. Asexual morph: Conidiomata stromatic or eustromatic, subepidermal to immersed, solitary to gregarious, subglobose to irregular, unilocular, pale brown. Conidiomata wall outer layer composed of thin-walled, pale brown cells of textura angularis, inner layer pale yellow to hyaline. Conidiophores reduced to conidiogenous cells or hyaline, septate, cylindrical, and sparingly branched. Conidiogenous cells holoblastic, cylindrical, lageniform to ampulliform, hyaline, smooth, invested in mucus. Conidia obclavate, conical, aseptate, pale brown, with a longitudinal band on the flat surface, thick and smooth-walled, guttulate, with short hyaline apiculus, with small globule of mucus on base or obtuse apex with a scar at the base.
Type genus: Apoharknessia Crous & S.J. Lee.
Type species: Apoharknessia insueta (B. Sutton) Crous & S.J. Lee.
Notes: Apoharknessia displays similar morphological characters to Harknessia but differs in having a hyaline, apical apiculus. Nag Raj (1993) listed Mastigonetron, as a synonym for Harknessia. Mastigonetron is typified by M. fuscum (= H. insueta). However, this species has a Wuestneia sexual morph, W. fusca, and it does not cluster with other Harknessia species. Therefore, Apoharknessia was introduced to accommodate H. insueta (Lee et al. 2004). The genus Apoharknessia presently accommodates two species (Crous et al. 2017).
Lasmenia species cause rachis necrosis, flower abortion and necrotic spots on leaves of Nephelium lappaceum. Several Lasmenia species associated with tropical fruits as pathogens have been isolated. DNA-based studies report a close affinity of Lasmenia to Cryphonectriaceae (Serrato-Diaz et al. 2011). Lasmenia was introduced in 1886 without designating any type species and L. balansae was selected as the lectotype species by von Höhnel (1910). There are 12 species recorded under Lasmenia in Index Fungorum (2017). Lasmenia species are reported as the causative agents of rachis necrosis, flower abortion, fruit rot, and leaf spots on Nephelium lappaceum (Serrato-Diaz et al. 2011). A few species have been transferred to Lasmeniella, but some species remain doubtful.
Phylogenetic analysis in the present study indicates that Apoharknessia and Lasmenia clearly belong to the Diaporthales in a well-supported clade (Fig. 1, Clade 13). However, the sequences of Lasmenia which are included in this study are not of a known species and given the sparse taxa in this family, any affinity between the two genera can not be ascertained.
Hence, we introduce a new family Apoharknessiaceae to accommodate these two genera. Morphologically species of this clade are distinct from other families of Diaporthales in having eustromatic to stromatic pycnidial conidiomata, blastic or phialidic conidiogenesis and ellipsoid to conical conidia with a longitudinal band on the flat surface or small globule of mucus at the base.
Apoharknessia insueta (B. Sutton) Crous & S.J. Lee, Stud. Mycol. 50: 240. 2004. Facesoffungi number FoF03458.
Illustration: See Lee et al. (2004).
Foliicolous forming bleached spots or saprobic on various substrates. Sexual morph: Undetermined. Asexual morph: Conidiomata stromatic, subepidermal to immersed, solitary to gregarious, subglobose to irregular, unilocular, pale brown. Conidiomata wall outer layer composed of thin-walled, pale brown cells of textura angularis, inner layer pale yellow to hyaline. Conidiophores reduced to conidiogenous cells. Conidiogenous cells 5–15 × 4–6 μm ( = 9 × 4.8 μm), lageniform to ampulliform, hyaline, smooth, invested in mucus. Conidia 10–12 × 7.5–9 μm ( = 10.5 × 8 μm), conical, aseptate, brown, with a longitudinal band on the flat surface, thick and smooth-walled, guttulate, with short hyaline apiculus, with small globule of mucus on base. Basal appendage 2 × 1–1.5 μm, often gelatinising and resulting in a minute marginal frill on the truncate base of the conidia (description based on Nag Raj 1993).
Notes: Apoharknessia was introduced and typified by Apoharknessia insueta and it clustered distant from Harknessia sensu stricto (Clade 7) (Lee et al. 2004). Apoharknessia is morphologically similar to Harknessia but distinct in having a hyaline apical apiculus in conidia and cultures on oatmeal or malt extract agar not forming fluffy aerial mycelium. In addition, it grows within the medium and sporulates directly on hyphae without forming conidiomata. Crous et al. (2017) introduced a new species as Apoharknessia eucalyptorum.
Asterosporiaceae Senan. Maharachch. & K.D. Hyde, fam. nov. MycoBank MB821539. Facesoffungi number FoF03459. Clade 23.
Endophytic or saprobic on Betulaceae, Fagaceae, Juglandaceae and Sapindaceae. Sexual morph: Undetermined. Asexual morph: Conidiomata acervular, subepidermal, erumpent at maturity, solitary, or occasionally confluent, unilocular, dark brown to black. Conidiomata wall composed of thin-walled, brown cells of textura angularis. Conidiophores cylindrical, branched at the base, septate, hyaline to pale brown. Conidiogenous cells holoblastic, cylindrical, unbranched, integrated, determinate, hyaline to pale brown, smooth. Conidia terminal, transversely distoseptate, consisting of four arms, with reduced lumina, brown, smooth-walled.
Type genus: Asterosporium Kunze.
Type species: Asterosporium hoffmannii Kunze.
Notes: A molecular phylogenetic analysis based on SSU nrDNA, LSU nrDNA, ITS nrDNA and beta-tubulin positions Asterosporium species within Sordariomycetes (Tanaka et al. 2010). Wijayawardene et al. (2016) showed that Asterosporium species are related to Diaporthales forming a sister clade to species in Sydowiellaceae based on combined ITS nrDNA and LSU nrDNA sequence analyses. In this study, Asterosporium species are positioned in Diaporthales (Fig. 1, Clade 23) and constitute a well-supported sister clade to Sydowiellaceae and Lamproconiaceae. Morphologically, Asterosporium species are distinct from other members of Diaporthales in having star-like, brown conidia. Hence, we introduce a novel family Asterosporiaceae to accommodate Asterosporium species. We illustrate Asterosporium asterospermum collected from Italy.
Asterosporium asterospermum (Pers.) Hughes, Canad. J. Bot. 36: 738. 1958. Fig. 3.
Asterosporium asterospermum (MFLU 15−3555). A. Conidiomata on host substrate. B, C. Vertical section of conidiomata. D–H. Different stages of conidiogenesis. I–M. Conidia. Scale bars: A = 1 mm, B = 400 μm, C = 50 μm, D–H = 20 μm, I–M = 30 μm.
Basionym: Stilbospora asterosperma Pers. [as ‘asterospora’], Syn. meth. fung. (Göttingen) 1: 96. 1801.
Saprobic on twigs and branches of Fagus sylvatica. Sexual morph: Undetermined. Asexual morph: Conidiomata 2–2.5 mm high, 0.8–1 mm diam ( = 2.1 × 0.86 mm, n = 15), acervular, subepidermal, erumpent at maturity, solitary, or occasionally confluent, unilocular, dark brown to black. Conidiomata wall 25–30 μm ( = 29, n = 20), composed of thin-walled, brown cells of textura angularis. Conidiophores 30–35 μm high, 5–8 μm wide ( = 29 × 7 μm, n = 20), cylindrical, branched at the base, septate, hyaline to pale brown. Conidiogenous cells 70–100 μm high, 4–7 μm wide ( = 80 × 5 μm, n = 20), holoblastic, cylindrical, unbranched, integrated, determinate, hyaline to pale brown, smooth-walled. Conidia 65–75 × 90–115 μm ( = 68 × 100 μm, n = 20), terminal, transversely distoseptate, consisting of four arms, with reduced lumina, brown, smooth-walled.
Specimen examined: Italy, Forlì-Cesena Province, Santa Sofia, near Passo la Calla, on dead branch of Fagus sylvatica (Fagaceae), 29 Sep. 2012, E. Camporesi, IT 805, MFLU 15-3555, HKAS 92536.
Notes: Asterosporium was introduced and typified by Asterosporium asterospermum (= Stilbospora asterosperma and Asterosporium hoffmannii) and there are five species listed in Index Fungorum (2017), namely A. acerinum, A. asterospermum, A. attenuatum, A. hoffmannii and A. strobilorum. However, only A. asterospermum has DNA sequence data in GenBank. There are no records for the sexual morph of Asterosporium (Tanaka et al. 2010). Species of this genus are associated with twigs and stems of overwintered plants as endophytes.
Auratiopycnidiellaceae Senan., Maharachch. & K.D. Hyde, fam. nov. MycoBank MB821540. Facesoffungi number FoF03460. Clade 11.
Foliicolous. Sexual morph: Undetermined. Asexual morph: Conidiomata amphigenous, pycnidia, globose, orange on leaves with dark brown border. Peridium comprises pale brown cells of textura angularis. Paraphyses hyaline, cellular, subcylindrical, branched or not, with obtuse apex, septate, constricted at septa. Conidiophores reduced to conidiogenous cells. Conidiogenous cells hyaline, smooth, lageniform to ampulliform, with terminal truncate locus, thick-walled, sometimes appearing to proliferate percurrently. Conidia ellipsoid, smooth, solitary, median 1-septate, constricted at septum, apex obtuse, base truncate, thickened, at times with marginal frill, becoming golden brown at germination with solitary, brown, wavy germ tubes.
Type genus: Auratiopycnidiella Crous & Summerell.
Type species: Auratiopycnidiella tristaniopsidis Crous & Summerell.
Notes: Crous et al. (2012a) described Auratiopycnidiella as a genus with subepidermal, orange, pycnidial conidiomata, forming hyaline, holoblastic conidiogenous cells, with or without a thickened scar and hyaline, ellipsoid, 1-septate conidia having a thickened hilum or minute marginal frill. Crous et al. (2012a) reported that the genus is phylogenetically distant to Melanconidaceae based on LSU nrDNA sequence data and treated this genus as Diaporthales genera incertae sedis pending the availability of more molecular data. A megablast search of NCBI's GenBank nucleotide database using the calmodulin, ITS nrDNA, and beta-tubulin sequences retrieved sequence similarities with Harknessiaceae and Cryphonectriaceae (Crous et al. 2012a). Our phylogenies generated herein indicate that Auratiopycnidiella forms a single branch which is phylogenetically distinct from all other included families (Fig. 1, Clade 11) and hence we introduce Auratiopycnidiellaceae to accommodate Auratiopycnidiella. Auratiopycnidiella currently comprises a single species with a single isolate.
Auratiopycnidiella tristaniopsidis Crous & Summerell [as ‘tristaniopsis’], Persoonia 28: 69. 2012. Facesoffungi number FoF03461.
Illustration: See Crous et al. (2012a).
Foliicolous. Sexual morph: Undetermined. Asexual morph: Conidiomata up to 200 μm diam, amphigenous, pycnidia, globose, orange on leaves with dark brown border, with irregular central opening. Peridium up to 25 μm thick, comprising 4–7 layers of pale brown cells of textura angularis. Paraphyses hyaline, cellular, subcylindrical, branched or not, with obtuse apex, 2–6-septate, constricted at septa Conidiophores 10–25 × 3–6 μm, reduced to conidiogenous cells. Conidiogenous cells hyaline, smooth, lageniform to ampulliform, with terminal truncate locus, thick-walled, sometimes appearing to proliferate percurrently. Conidia 13–15 × 5–5.5 μm, ellipsoid, smooth, solitary, medially 1-septate, constricted at septum, obtuse at apex, truncate at base, thickened at times with marginal frill, hyaline becoming golden brown during germination with solitary, brown, wavy germ tubes 90° to the long axis of the spore (description based on Crous et al. 2012a).
Notes: Auratiopycnidiella was introduced and typified by Auratiopycnidiella tristaniopsis. This is a monotypic genus comprising only the type species, A. tristaniopsis. Auratiopycnidiella tristaniopsis forms leaf spots on its host species. Morphologically this taxon shows some similarities to taxa of the Cryphonectriaceae in having orange conidiomata. However, phylogenetically it is distinct from Cryphonectriaceae.
Coryneaceae Corda, Icon. fung. (Prague) 3: 36. 1839. Clade 20.
Synonym: Pseudovalsaceae M.E. Barr, Mycol. Mem. 7: 151. 1978.
Saprobic on dead wood or pathogenic. Sexual morph: Stromata solitary, erumpent, comprising pseudoparenchymatous cells. Ectostromatic disc well or poorly developed, brown to black, comprising small cells of textura prismatica cells. Ascomata perithecial, arranged in valsoid configuration, immersed, aggregated, globose to subglobose, coriaceous, brown to black, papillate, ostiolate. Papilla upright, central, broad, sometimes converging, comprising brown cells of textura porrecta. Peridium comprising outer, thick-walled, brown cells of textura angularis and inner, thick-walled, hyaline, compressed cells of textura angularis. Hamathecium comprising broad, cellular, septate paraphyses, attached to base, longer than asci. Asci 8-spored, unitunicate, ellipsoid to cylindrical, thin-walled, pedicellate, apex rounded with a J- apical ring. Ascospores overlapping uni- to biseriate, hyaline or initially hyaline, brown at maturity, irregularly fasciculate, ellipsoid, fusoid or elongate, 1–3-septate, often distoseptate, end cells pale brown or hyaline, sometimes end cells pointed, straight or curved not constricted at the septa, guttulate, smooth-walled. Asexual morph: Coelomycetous. Conidiomata acervular, solitary, erumpent through the outer periderm layers of host or immersed, scattered, surface tissues above slightly dome-shaped. Conidiomatal wall composed of thin-walled, vertically arranged dark brown cells of textura angularis. Conidiophores branched at the base or not, cylindrical to globose, septate or aseptate, hyaline or hyaline at the apex, pale brown at the base. Conidiogenous cells terminal, hyaline, annellidic, cylindrical, sometimes with setulose apical appendages. Conidia hyaline to dark brown, curved, broadly fusiform to cylindrical or clavate, smooth-walled, 4–6-distoseptate, sometimes the apical and basal cell darker than other cells with hyaline tip in apical cell.
Type genus: Coryneum Nees.
Type species: Coryneum umbonatum Nees.
Notes: The family Coryneaceae (Fig. 1, Clade 20) was introduced by Corda (1839) based on Coryneum. However, Barr (1978) introduced the family Pseudovalsaceae based on Pseudovalsa lanciformis, which is the sexual morph of Coryneum umbonatum. Hence Pseudovalsaceae must be synonymised under Coryneaceae giving priority to the older name. Rossman et al. (2015) protected the earliest name Coryneum (1816) over Pseudovalsa (1863) and conserved Coryneum umbonatum as the type species. This family comprises fungal taxa with upright, erumpent perithecia and central beaks. However, many genera previously included in Coryneaceae have been placed in various other families (Castlebury et al. 2002) and the only genus remaining in the family is Coryneum.
Coryneum arausiaca (Fabre) Senan., Maharachch. & K.D. Hyde, comb. nov. MycoBank MB821543. Facesoffungi number FoF03462. Fig. 4.
Coryneum arausiaca (MFLU 14–0796). A, B. Ascostromata on substrate. C, D. Vertical section of ascoma. E. Peridium. F. Periphyses. G. Paraphyses. –J. Asci. K–S. Ascospores. T. Conidiomata on substrate. U. Vertical section of conidiomata. V. Conidiophores, conidiogeneous cells with conidia. W–ZB. Conidia. Scale bars: A, B = 500 μm, C, D, S = 100 μm, E–G, K, Q, U–Z = 10 μm, H–J = 20 μm, R = 1 mm, T = 50 μm.
Basionym: Pseudovalsa arausiaca Fabre, Sphér. Vaucl.: 56. 1883.
Saprobic on branches of Quercus sp. Sexual morph: Stromata comprising loosely packed, black, hyphae mostly around the neck. Ascomata 600–700 μm high, 315–365 μm diam ( = 640 × 340 μm, n = 20), immersed, 5–10 aggregated in one group, visible only as ostiolar opening through cracks in bark, valsoid, globose, brown to black, papillate, ostiolate, ostiole periphysate, periphyses hyaline, long. Peridium 25–50 μm ( = 45 μm, n = 10), 10–15 layers of thick-walled, brown-walled cells of textura angularis and papilla comprising brown cells of textura porrecta. Paraphyses 5–10 μm ( = 7.6 μm, n = 20), few, hyaline, septate, attached at base, longer than asci. Asci 145–155 × 25–30 μm ( = 146 × 25.6 μm, n = 10), 8-spored, unitunicate, clavate, short pedicellate, apically rounded, narrow, J-, without an obvious apical ring. Ascospores 70–90 × 6.5–8.5 μm ( = 77 × 7.5 μm, n = 10), 2–3-seriate, broadly ellipsoidal, ends pointed, 1–3-septate, not constricted at the septa, hyaline, guttulate, smooth-walled. Asexual morph: Conidiomata acervular, 1–1.3 mm wide, 0.5–0.55 mm high ( = 1.1 × 0.51 mm, n = 20), solitary, erumpent through the outer periderm layers of host, scattered, surface tissues above slightly domed. Conidiomatal wall 100–150 μm ( = 135 μm, n = 20), composed of thin-walled, vertically arranged, dark brown cells of textura epidermis. Conidiophores 20–35 μm long, 4–7 μm wide ( = 30 × 6 μm, n = 20), branched at the base, cylindrical, septate, hyaline at the top, pale brown at the base. Conidiogenous cells 4–7 μm long, 4.5–6 μm wide ( = 6 × 5 μm, n = 20), formed from the apical cell of the conidiophore, holoblastic, cylindrical, hyaline. Conidia 42–56 × 13–16 μm ( = 48 × 14 μm, n = 20), curved, broadly fusiform to fusiform-cylindrical or clavate (rather variable in form), dark brown, smooth-walled, 4–6-disto-septate, with apical and basal cells darker than other cells, apical cell with a hyaline tip, truncate and black at base.
Culture characteristics: Ascospores germinating on MEA within 12 h and germ tubes produced from both ends, fast growing on MEA at 25 ºC, after 1 wk reaching 3 cm diam, white, cottony, margin wavy, superficial, slightly effuse, radially striated, edges with more aerial mycelium than centre.
Specimens examined: Italy, Province of Forlì-Cesena, Civitella di Romagna, Pian di Spino, on branch of Quercus sp. (Fagaceae), 25 Mar. 2013, E. Camporesi, IT 1144, (neotype designated here MFLU 14–0796, cultures ex-neotype, MFLUCC 13–0658); Province of Forlì-Cesena, Civitella di Romagna, Pian di Spino, on branch of Quercus sp. (Fagaceae), 16 Feb. 2015, E. Camporesi, IT 1144A, paraneotype HKAS83943, cultures ex-paraneotype, MFLUCC 15–1110.
Notes: We have re-collected and neotypified Pseudovalsa arausiaca. Pseudovalsa arausiaca has immersed, globose perithecia in a valsoid configuration with broadly ellipsoidal, 1–3-septate, hyaline ascospores. The neotype is morphologically identical to Pseudovalsa arausiaca described by Fabre (1883). However, we could not locate the type specimens and assume that they are lost. Fortunately, we obtained fresh material from the same host genus and location. Therefore, a neotype is designated here with sequence data. Rossman et al. (2015) protected Coryneum over Pseudovalsa. Hence, we propose a new combination for Pseudovalsa arausiaca as Coryneum arausiaca. Both sexual and asexual morphs of Coryneum arausiaca were obtained from the same specimen as well as cultures which indicate a holomorph connection. We illustrate both sexual and asexual morphs of Coryneum arausiaca and the combined gene analysis of LSU nrDNA, ITS nrDNA, rpb2 and tef1 shows the distinct placement of C. arausiaca within Coryneaceae (Fig. 1, Clade 20).
Cryphonectriaceae Gryzenh. & M.J. Wingf., Mycologia 98: 246. 2006. Clade 6.
Saprobic or pathogenic in forest trees and economic crops. Sexual morph: Ascostromata scattered, immersed or erumpent, aggregated, oval to circular from above, comprising two layers, upper layer of yellowish orange to pale brown cells, purpling in KOH and inner layer of hyaline cells, mixed with plant cells. Ascomata immersed, aggregated, several in one stroma, globose to subglobose, fuscous black to umber, with long neck, or ostiolar canal sometimes immersed in stromatic tissues, or superficial, necks covered in umber stromatic tissue of textura porrecta, inner wall of the necks or ostiolar canal covered with hyaline, filamentous periphyses. Peridium comprising inner layer of small, hyaline cells of textura angularis and outer layer of small, brown cells of textura angularis. Hamathecium comprising a few cellular paraphyses and parenchymatous cells, attached at the base and asci dissolving at maturity. Asci 8-spored, unitunicate, cylindrical-fusoid to clavate, pedicellate, with distinct, J- refractive ring. Ascospores overlapping uniseriate to biseriate, ellipsoid, fusoid to cylindrical, aseptate to multi-septate, not constricted at the septa, hyaline, sometimes brown, smooth-walled. Asexual morph: Coelomycetous. Conidiomata occurring as a part of ascomata as conidial locules or solitary structures, uni- to multi-loculate, pyriform, subglobose to pulvinate, necks absent or present, if present, with one to several attenuated necks, superficial or semi-immersed, orange to fuscous-black. Conidiophores cylindrical, aseptate, hyaline, sometimes reduced to conidiogenous cells. Conidiogenous cells lining the inner cavity of the conidiomata, phialidic, sometimes within flattened bases, ampulliform, inconspicuous, with attenuated or truncate apices, hyaline, smooth. Conidia minute, sometimes both micro- and macro-conidia present, sigmoid, broadly ellipsoid to fusoid, obovoid-cylindrical to allantoid, aseptate, hyaline.
Type genus: Cryphonectria (Sacc.) Sacc. & D. Sacc.
Type species: Cryphonectria parasitica D. Sacc.
Notes: Cryphonectriaceae (Fig. 1, Clade 6) is mostly a pathogenic family comprising some of the world's most important tree pathogens (Vermeulen et al. 2011). Cryphonectriaceous species are saprobes, endophytes and phytopathogens. They cause cankers, blights and dieback of economically important plants and forest trees. Castlebury et al. (2002) recognised the Cryphonectria-Endothia complex (a precursor to the Cryphonectriaceae) as a separate clade in Diaporthales based on analysis of LSU nrDNA sequence data. Cryphonectriaceae was formally established by Gryzenhout et al. (2006c) to accommodate the Cryphonectria-Endothia complex and other allied genera when analysing LSU nrDNA sequence data of fungal taxa in Diaporthales. Species of this family can be distinguished from other families of Diaporthales by orange stromatic tissues, which turn purple in KOH and yellow in lactic acid. Initially Amphilogia, Chrysoporthe, Cryphonectria, Endothia and Rostraureum were placed in the family (Gryzenhout et al. 2006c). Subsequently, several additional genera were added to the family, some associated with serious canker or foliar diseases, namely: Aurantiosacculus, Aurapex, Aurifilum, Celoporthe, Chromendothia, Chrysocrypta, Chrysofolia, Cryptometrion, Diversimorbus, Foliocryphia, Holocryphia, Immersiporthe, Lasmenia, Latruncellus, Luteocirrhus, Mastigosporella, Microthia, Prosopidicola and Ursicollum (Vasilyeva, 1993, Gryzenhout et al., 2006a, Gryzenhout et al., 2006b, Nakabonge et al., 2006, Begoude et al., 2010, Gryzenhout et al., 2010, Vermeulen et al., 2011, Crous et al., 2012a, Chen et al., 2013, Crane and Burgess, 2013, Crous et al., 2013).
Endothia (1849) is typified by E. gyrosa and the asexual morph of this genus was reported as an Endothiella species (Barr 1978). However, Endothiella is congeneric with Cryphonectria and Endothiella eucalypti is the asexual morph of type species of Cryphonectria, C. eucalypti (Jackson 2003). Endothiella (1906) is based on the type species, Endothiella gyrosa, now placed in Cryphonectria as C. decipiens (Gryzenhout et al. 2009). Barr (1978) observed several specimens of Cryphonectria and Endothia and she used stromatic configuration and ascospore characters to differentiate these two genera. According to Barr (1978), Cryphonectria has a valsoid configuration of perithecia in prosenchymatous stromata and ellipsoid or ovoid, 1-septate ascospores, while Endothia has a diatrypoid configuration of perithecia in pseudoparenchymatous stromata and allantoid, unicellular ascospores. Based on these characters, most Endothia species have been moved to Cryphonectria and the generic name Endothia was restricted to the species with a diatrypoid configuration of the perithecia and allantoid, unicellular ascospores. Combined analysis of LSU nrDNA, ITS nrDNA, rpb2 and tef1 sequence data in the present study shows Cryphonectriaceae is not well-supported (Fig. 1, Clade 6). Phylogenetic analyses of this study also place Cryphonectria and Endothia as two separate genera, as well as Chrysocrypta (Fig. 1, Clade 9), Lasmenia (Fig. 1, Clade 13) and Prosopidicola (Fig. 1, Clade 17) outside of Cryphonectriaceae. Hence, currently this family comprises Amphilogia, Aurantioporthe, Aurantiosacculus, Aurapex, Aurifilum, Celoporthe, Chromendothia, Chrysofolia, Chrysoporthe, Chrysoporthella, Cryphonectria, Cryptometrion, Diversimorbus, Endothia, Foliocryphia, Holocryphia, Immersiporthe, Latruncellus, Luteocirrhus, Mastigosporella, Microthia, Rostraureum and Ursicollum.
Cryphonectria parasitica (Murrill) M.E. Barr, Mycol. Mem. 7: 143. 1978. Facesoffungi number FoF03463. Fig. 5.
Cryphonectria parasitica (NY 01293321). A. Herbarium packet. B. Herbarium specimen. C. Ascostromata on substrate. D. Horizontal section of ascostroma. E. Vertical cross section of ascoma. F–I. Asci. J–M. Ascospores. N. Horizontal cross section of conidiomata. O. Vertical cross section of conidioma. P, Q. Conidia attached to the conidiogenous cells and conidiophore. R. Conidia. Scale bars: C, D = 1 mm, F–I, P–R = 10 µm, E, Q, O = 100 µm, N = 200 µm, J–M = 5 µm.
Basionym: Diaporthe parasitica Murrill, Torreya 6: 189. (1906).
Pathogenic on branches of Castanea dentata. Sexual morph: Ascostromata 6.5–1 mm diam ( = 0.8 mm, n = 20), comprising erumpent to superficial, orange epistromatic portion and immersed, hyaline, parenchymatous portion. Ascomata 650–715 μm high, 210–220 μm diam ( = 685 × 216 μm, n = 20), perithecial, immersed, globose to subglobose, with black to brown ostiole, ostiolar canal slender, covered with orange to fuscous-black stromatic tissue. Hamathecium aparaphysate, comprising parenchymatous tissues. Asci 20–35 × 5–8 μm ( = 28 × 6.4 μm, n = 20), 8-spored, unitunicate, fusiform to cylindrical base with small pedicel, apex oblong. Ascospores 5–6 × 2–2.5 μm ( = 5.5 × 2.2 μm, n = 20), overlapping uni- or biseriate, hyaline, ellipsoid to fusiform, 1-septate. Asexual morph: Conidiomata 250–300 μm high, 180–200 μm diam ( = 280 × 185 μm, n = 20), eustromatic, erumpent, pyriform to pulvinate, orange to fuscous black, occurring in the same stroma as perithecia. Conidiophores 3–4 × 1–1.5 μm ( = 3.4 × 1.1 μm, n = 20), cylindrical, unbranched, hyaline. Conidiogenous cells 2.5–5 × 0.5–1 μm ( = 3.3 × 0.8 μm, n = 20), phialidic, simple or branched. Conidia 1.8–2.5 × 0.5–1 μm ( = 2.1 × 0.9 μm, n = 20), hyaline, minute, allantoid to cylindrical, aseptate.
Materials examined: USA, New York. Bronx Co. Bronx. North of Botanical Museum, Bronx Park, on Castanea dentata (Fagaceae), 26 Nov. 1905, W.A. Murrill (holotype 01293321, as Diaporthe parasitica, NY).
Notes: American chestnut blight, caused by Cryphonectria parasitica, destroyed American chestnut trees in the USA and Canada at the end of the 19th century. Scientists believed Cryphonectria parasitica arrived from north-east Asia in the late 19th century and they discovered that Japanese and Chinese chestnut varieties showed resistance to C. parasitica. Spores of this fungus are highly resistant to unfavourable environmental conditions and they can be produced at any time of year when conditions are suitable. The fungus can exist as a saprobe and a parasite. Mycelium can survive more than 10 mo in dried bark and soil (Hepting 1974). Conidia and ascospores of C. parasitica are sometimes forcibly ejected and spread in wind and rain. Spores of Cryphonectria parasitica are also dispersed by beetles and birds. In addition to chestnut species, some oak species and Chinquapin also are infected by Cryphonectria parasitica.
Cytosporaceae Fr. [as ‘Cytisporei’], Syst. orb. veg. (Lundae) 1: 118. 1825. Clade 16.
Synonym: Valsaceae Tul. & C. Tul. [as ‘Valsarum’], Select. fung. carpol. (Paris) 1: 180. 1861.
Pathogenic or saprobic on plant tissues. Sexual morph: Stromata well or poorly developed. Ectostroma circular or irregular, usually well developed in the upper regions. Entostroma normally limited to the region near the perithecial walls. Ascomata perithecia, immersed to erumpent, solitary or 6–10 ascomata aggregated in valsoid configuration, globose to oblong, coriaceous, black to brown, with long neck swollen at the tips, ostiolate. Ostiole periphysate, open through the neck. Peridium thin, comprising outer, 4–6 layers of, dark brown, thick-walled, cells of textura angularis and 5–7 layers of, inner, small, hyaline, thin-walled, cells of textura angularis. Hamathecium comprising few, hyaline paraphyses limited only at young stage. Asci unitunicate, 8-spored, clavate, short-pedicellate, apex round, with apical ring. Ascospores uni- to biseriate, unicellular or rarely bicellular, allantoid or ellipsoid, hyaline, smooth-walled. Asexual morph: Stromata uniloculate, black, circular in shape. Locule composed of numerous inter connecting chambers arranged radially or irregularly within a continuous mass of ectostromatic tissue, one conidioma per locule. Conidiomata pyriform in section, brown, divided into compartments by bending of peridium. Peridium consists of brown, 5–7 layers of textura angularis cells. Conidiophores reduced to conidiogenous cells. Conidiogenous cells arising from conidiomatal wall, phialidic, simple or branched, hyaline, cylindrical. Conidia unicellular, allantoid, hyaline, smooth-walled.
Type genus: Cytospora Ehrenb.
Type species: Cytospora chrysosperma (Pers.) Fr.
Notes: The Cytosporaceae (Fig. 1, Clade 16) comprises phytopathogenic species and saprobes. Most Cytospora species are plant pathogens and cause cankers and dieback of many hardwoods and coniferous trees, as well as rarely on herbaceous plants. Generally, Cytospora cankers are known as valsa-canker, Leucostoma-canker or perennial canker (Farr et al. 1989). Cytospora species have been reported as highly virulent and destructive pathogens on Prunus and Populus trees (Biggs, 1989, Kepley and Jacobi, 2000). A few Cytospora species are considered as facultative wound parasites that attack damaged or weakened plants.
Maharachchikumbura et al., 2015, Maharachchikumbura et al., 2016 listed 13 genera under this family as Amphicytostroma, Chadefaudiomyces, Cryptascoma, Cytospora, Ditopellina, Durispora, Harpostroma, Hypospilina, Kapooria, Leptosillia, Maculatipalma, Pachytrype, and Paravalsa. However, the type species of Amphicytostroma, A. tiliae is the asexual morph of the type species of Amphiporthe, A. hranicensis, and these generic names are synonyms (Sutton 1980). Amphiporthe is more widely used than Amphicytospora and it seems best to protect the former (Rossman et al. 2015). However, Amphiporthe belongs in Gnomoniaceae (Sogonov et al. 2008; Fig. 1, Clade 1) and we exclude this genus from Cytosporaceae. Rossman et al. (2015) proposed to use Cytospora (1818) rather than Valsa (1825), Valsella (1870), Leucostoma (1917), Valseutypella (1919), or Leucocytospora (1927). Xenotypa is a genus in Gnomoniaceae and typified by Xenotypa aterrima. This genus is characterised by having solitary or aggregated, erumpent, globose, papillate ascomata with allantoid to cylindrical, unicellular, hyaline ascospores. Morphologically this is similar to Paravalsa and Valsella. Ananthapadmanaban (1990) described the relationship between Xenotypa and Paravalsa, accommodating Paravalsa in Valsaceae. However, many of the fungal taxa listed in Maharachchikumbura et al. (2015) do not share similar morphological characters and it is necessary to restrict this family to Cytospora sensu-lato. Cytospora, Valsella, Leucostoma, Valsa and Pachytrype have sequence data in accessible data bases. Hence, we suggest to accommodate Cytospora, Paravalsa, Pachytrype, Waydora and Xenotypa in Cytosporaceae. However, the Cytospora sensu-lato complex still needs to be resolved using high resolution genes as it seems to comprise several genera.
Cytospora centrivillosa Senan., Camporesi & K.D. Hyde, sp. nov. MycoBank MB821567. Facesoffungi number FoF03464. Fig. 6.
Cytosporacentrivillosa (MFLU 17–0887). A, B. Stromata on substrate. C, D. Vertical cross section of ascomata. E. Peridium. F–H. Asci. I. Paraphyses. J. Ascospores. K. Conidiomata on MEA. L–N. Conidiogenous cells, conidiophores, conidia. O. Conidia. Scale bars: A = 500 µm, B = 200 µm, C, D = 50 µm, E = 5 µm, I = 20 µm, F–H, J, L–O = 10 µm, K = 500 µm.
Etymology: Name based on two Latin words “centrum” and “villos” meaning hamathecium comprising filiform paraphyses.
Saprobic on dead branch of Sorbus domestica. Sexual morph: Stromata poorly developed, comprising loosely packed parenchymatous cells, black. Ascomata 550–725 μm high, 160–215 μm diam ( = 611 × 190 μm, n = 20), aggregated, immersed, globose to subglobose, dark brown, coriaceous, ostiolate, papillate. Papilla 285–430 μm high, 90–130 μm diam ( = 340 × 101 μm, n = 20), long, central or asymmetrically located, wall thick, internally covered by hyaline periphyses. Peridium comprises brown, thick-walled cells of textura angularis. Asci 75–85 × 15–19 μm ( = 79 × 18 μm, n = 20), 8-spored, unitunicate, clavate to fusiform, without apical ring and pedicel. Ascospores 16–20 × 4–6 μm ( = 17 × 5 μm, n = 20), biseriate, allantoid, hyaline, smooth. Asexual morph: Coelomycetous. Conidiomata on MEA appears as pale yellow, slimy heads of conidial mass, immersed, black. Conidiophores 6.5–8 × 3–3.5 μm ( = 7.4 × 3.1 μm, n = 20), cylindrical, unbranched, hyaline. Conidiogenous cells 10–13.5 × 1–2 μm ( = 11.7 × 1.6 μm, n = 20), cylindrical, tapering towards the apices, bearing single conidia at each tip, hyaline. Conidia 4–6 × 1–1.5 μm ( = 5.1 × 1.1 μm, n = 20), eguttulate, allantoid, aseptate, hyaline.
Culture characteristics: Colonies growing on MEA attenuated 1 cm incubated at 18 °C within 4 d, fast growing, circular, flat, entire, white, thin, tightly attached to the media, mycelia clots arrange radially from centre to margin.
Specimens examined: Italy, Province of Forlì-Cesena, Predappio, Monte Mirabello, on dead and aerial branch of Sorbus domestica (Rosaceae), 1 Oct. 2014, E. Camporesi, IT 2132 (holotype MFLU 17–0887, isotype BBH 42449, culture ex-type MFLUCC 16–1206); Province of Forlì-Cesena, Predappio, Monte Mirabello, on dead and aerial branch of Sorbus domestica (Rosaceae), 13 Oct. 2014, E. Camporesi, IT 2132B, MFLU 17–0999, culture MFLUCC 17–1660.
Note: Cytospora centrivillosa is morphologically and phylogenetically distinct from other species in Cytospora and our analysis results in a distinct clade with full support (Fig. 1, Clade 16).
Cytospora fraxinigena Senan., Camporesi & K.D. Hyde, sp. nov. MycoBank MB821568. Facesoffungi number FoF03465. Fig. 7.
Cytosporafraxinigena (MFLU 17–0880).A. Ascomata on substrate. B. Vertical cross section of ascoma. C. Peridium. D–F. Asci. G–J. Ascospores. Scale bars: B = 100 µm, C = 20 µm, D–F = 10 µm, G–J = 5 µm.
Etymology: Named after the host genus Fraxinus.
Saprobic on dead branch of Fraxinus ornus. Sexual morph: Stromata poorly developed, comprising loosely packed parenchymatous cells, black. Ascomata 350–500 × 150–230 μm ( = 429 × 189 μm, n = 20), immersed in stromatic tissues, globose to subglobose, dark brown, coriaceous, ostiolate, papillate. Papilla 185–200 × 60–95 μm ( = 193 × 79 μm, n = 20), long, central, wide, thick-walled, internally covered by hyaline periphyses. Peridium comprises brown, thick-walled cells of textura angularis. Asci 26–33 × 6.2–7.5 μm ( = 30 × 6.7 μm, n = 20), 8-spored, unitunicate, clavate to fusiform, without apical ring and pedicel. Ascospores 5.5–7.5 × 1.5–2 μm ( = 6.4 × 1.7 μm, n = 20), biseriate, allantoid, hyaline, smooth. Asexual morph: Not observed.
Culture characteristics: Colonies growing on MEA attenuated 1 cm incubated at 18 °C within 7 d, moderate fast growing, irregular, flat, undulate, white, woolly, loosely attached to the media.
Specimen examined: Italy, Province of Forlì-Cesena, Santa Sofia, near Corniolo, dead branch of Fraxinus ornus (Oleaceae), 6 Dec. 2013, E. Camporesi, IT 1562 (holotype MFLU 17–0880, isotype BBH 42442, culture ex-type MFLUCC 14–0868).
Notes: Cytospora fraxinigena forms a distinct clade which is sister to Cytospora cedri and Cytospora rosae (Fig. 1, Clade 16). Morphologically, Cytospora fraxinigena differs from those species in having slightly horizontal necks closely arranged at apex and hamathecium without paraphyses.
Cytospora junipericola Senan., Camporesi & K.D. Hyde, sp. nov. MycoBank MB821569. Facesoffungi number FoF03466. Fig. 8.
Cytospora junipericola (MFLU 17–0882). A. Ascomata on substrate. B. Vertical cross section of ascoma. C. Peridium. D–G. Asci. H. Ascospores. Scale bars: A = 500 µm, B = 100 µm, C = 20 µm, D–H = 10 µm.
Etymology: Named after the host genus Juniperus.
Saprobic on dead branch of Juniperus sp. Sexual morph: Stromata poorly developed, comprising loosely packed parenchymatous cells, black. Ascomata 630–700 μm high, 150–250 μm diam ( = 670 × 170 μm, n = 20), immersed in stromatic tissues, globose to subglobose, dark brown, coriaceous, ostiolate, papillate. Papilla 300–500 μm high, 45–65 μm diam ( = 440 × 58 μm, n = 20), long, central, wide, thick-walled, internally covered by hyaline periphyses. Peridium comprises brown, thick-walled cells of textura angularis. Asci 30–35 × 5.5–7 μm ( = 32 × 6 μm, n = 20), 8-spored, unitunicate, clavate to fusiform, without apical ring and pedicel. Ascospores 5–10 × 1–2 μm ( = 7 × 1.5 μm, n = 20), biseriate, allantoid, hyaline, smooth. Asexual morph: Not observed.
Culture characteristics: Colonies growing on MEA attenuated 1 cm incubated at 18 °C within 7 d, moderate fast growing, irregular, flat, undulate, greenish ash, woolly, curled, loosely attached to the media.
Specimen examined: Italy, Province of Forlì-Cesena, Santa Sofia, near Cabelli, dead branch of Juniperus communis (Cupressaceae), 13 Jan. 2014, E. Camporesi, IT 1643 (holotype MFLU 17–0882, isotype BBH42444).
Notes: Cytospora junipericola forms a distinct clade that is sister to Cytospora quercicola with high bootstrap support (Fig. 1, Clade 16). Morphologically Cytospora junipericola produces tightly packed aggregated ascomata in poorly developed stromatic tissues. Papilla are asymmetrically located and only the ostiolar openings are close together.
Cytospora quercicola Senan., Camporesi, & K.D. Hyde, sp. nov. MycoBank MB821570. Facesoffungi number FoF03467. Fig. 9.
Cytospora quercicola (MFLU 17–0881). A. Ascomata on substrate. B. Cross section of ascoma. C. Peridium. D–G. Asci. H. Ascospores. Scale bars: A = 200 µm, B = 100 µm, C = 20 µm, D–H = 10 µm.
Etymology: Named after the host genus Quercus.
Saprobic on dead branch of Quercus sp. Stromata poorly developed, spread around the papilla, black. Ascomata 550–725 μm high, 160–215 μm diam ( = 611 × 190 μm, n = 20), scattered, aggregated, immersed, globose to subglobose, dark brown, coriaceous, ostiolate, papillate. Papilla 285–430 μm high, 90–130 μm diam ( = 340 × 101 μm, n = 20), long, central or asymmetrically located, papilla close to each other when open to host surface. Peridium comprises brown, thick-walled cells of textura angularis. Asci 75–85 × 15–19 μm ( = 79 × 18 μm, n = 20), 8-spored, unitunicate, clavate to fusiform, without apical ring and pedicel. Ascospores 16–20 × 4–6 μm ( = 17 × 5 μm, n = 20), biseriate, allantoid, hyaline, smooth.
Culture characteristics: Colonies growing on MEA becoming 1 cm within 7 d incubated at 18 °C, circular, flat, smooth colony with white mycelium, mycelia loosely attached to the substrate.
Specimen(s) examined: Italy, Province of Forlì-Cesena, Santa Sofia, near Camposonaldo, on dead branch of Quercus sp. (Fagaceae), 10 Dec. 2013, E. Camporesi, IT 1568 (holotype MFLU 17-0881, isotype BBH 42443, culture ex-type MFLUCC 14-0867).
Notes: The Cytospora quercicola clade is fully-supported by the multi-gene phylogenetic analyses (Fig. 1, Clade 16). This species is sister to Cytospora junipericola.
Cytospora rosae Senan., Camporesi, & K.D. Hyde, sp. nov. MycoBank MB821571. Facesoffungi number FoF03468. Fig. 10.
Cytospora rosae (MFLU 17–0885). A. Ascomata on substrate. B. Cross section of ascoma. C. Peridium. D–I. Asci. J. Paraphysis. K. Ascospores. L. Colony on MEA upper surface. M. Colony on MEA lower surface. N–P. Conidiomata on MEA. P. Horizontal cross section of conidioma. Q–R. Peridium with conidiogeneous cells, conidiophores and conidia. S. Conidia. Scale bars: A, P–N = 500 µm, B = 50 µm, C, I, Q–R = 10 µm, D–H, J, K, S = 5 µm, P =100 µm, O = 1 mm.
Etymology: Named after the host genus Rosa.
Saprobic on Rosa canina. Sexual morph: Stromata restricted to around the ostiolar neck, black. Ascomata 235–255 μm high, 130–150 μm diam ( = 240 × 140 μm, n = 20), solitary to rarely aggregated, scattered, immersed, globose, brown, coriaceous, ostiolate, papillate. Papilla 127–140 μm high, 70–90 μm diam ( = 135 × 87 μm, n = 20), straight or curved, long, brown, internally covered by hyaline periphyses, wall comprising elongated, thick-walled cells. Peridium 16–23 μm diam ( = 20 μm, n = 20), comprising outer, thick-walled, brown cells of textura angularis and inner, compressed, thick-walled, hyaline cells of textura angularis. Hamathecium comprising septate, hyphae-like, hyaline, 1.5–2.7 μm diam ( = 2.5 μm, n = 20) paraphyses. Asci 20–23 × 3.2–3.7 μm ( = 21 × 3.7 μm, n = 20), unitunicate, 8-spored, clavate, short-pedicellate, apex round, with apical ring. Ascospores 4.2–6.3 × 1–1.5 μm ( = 5.5 × 1.3 μm, n = 20), uni- to biseriate, unicellular, allantoid, or ellipsoid, hyaline, smooth-walled. Asexual morph: Conidiomata 100–200 μm diam ( = 150 μm, n = 20), solitary to aggregate, immersed, pyriform to subglobose, multi-loculate, black, coriaceous, ostiolate, papillate, peridium folded into centrum. Pycnidial walls 4–7 μm diam ( = 6 μm, n = 20), comprising small, thick-walled, brown cells of textura angularis. Conidiophores 8–12 × 1.5–2.5 μm ( = 11 × 2 μm, n = 20), cylindrical, shorter than conidiogenous cells, branched, hyaline. Conidiogenous cells 10–15 × 1–1.5 μm ( = 12 × 1.2 μm, n = 20), phialidic, cylindrical, tapering towards the apices, bearing single conidia at each tip. Conidia 3–5 × 0.5–1 μm ( = 2 × 1 μm, n = 20), aguttulate, elongated to allantoid, slightly curved, aseptate, hyaline.
Culture characteristics: Colonies growing on MEA attained 2 cm within 7 d incubated at 18 °C, filamentous, flat, filiform, middle blackish ash, margin off white, cottony, tiny mycelium clots arrange radially from centre to margin.
Specimen(s) examined: Italy, Province of Forlì-Cesena, Galeata, near Passo delle Forche, on dead branch of Rosa canina (Rosaceae), 15 Apr. 2014, E. Camporesi, IT 1814 (holotype MFLU 17-0885, isotype BBH 42447, cultures ex-type MFLUCC 14–0845; Province of Forlì-Cesena, Galeata, near Passo delle Forche, on dead branch of Rosa canina (Rosaceae), 4 Jan. 2016, E. Camporesi, IT 1814 (paratype MFLU 15–3596, cultures ex-paratype MFLUCC 17–1664).
Notes: Combined ITS nrDNA, LSU nrDNA, rpb2 and tef1 sequence data in the current study shows that Cytospora rosae forms a distinct clade with high bootstrap support, basal to Cytospora fraxinigena (Fig. 1, Clade 16). Morphologically, Cytospora rosae has unique characters of solitary ascomata and small asci with septate, wide, hyaline, hyphae-like paraphyses.
Cytospora salicina Norphanphoun et al., Mycosphere 8: 80. 2017. Fig. 11.
Cytosporasalicina (MFLU 17–0891). A. Conidiomata on substrate. B. Horizontal cross section of conidioma. C, D. Vertical cross section of conidiomata. E. Conidiophore, conidiogenous cells arrangement. F. Peridium. G–M. Conidiophores, conidiogenous cells, conidia. N. Conidia. Scale bars: A = 500 µm, B = 200 µm, C, D = 100 µm, E, F = 15 µm, G–M = 10 µm, N = 5 µm.
Saprobic on twigs and branches of Cornus sanguinea. Sexual morph: Undetermined. Asexual morph: Stromata appear as black pinhead spots surrounding by yellow to pale brown tissues on the substrate, immersed, rosette, labyrinthine, pale brown to black, 1–5 pycnidia in a stroma, comprising loosely packed, pale brown cells of textura globosa, ostiole. Papilla narrow, short, internally covered by periphyses, converged, black, furfuraceous. Pycnidial locules multi-chambered, subdivided by invaginations of common pycnidial walls. Conidiomata 530–600 μm high, 600–870 μm diam ( = 570 × 705 μm, n = 20), solitary to aggregate, immersed, pyriform to subglobose, black, coriaceous, ostiolate, papillate, peridium folded into centrum. Papilla 200–300 μm high, 60–95 μm diam ( = 210 × 80 μm, n = 20), internally covered by hyaline filiform periphyses. Pycnidial walls 7–11 μm diam ( = 9.2 μm, n = 20), comprising small, thick-walled, brown cells of textura angularis, separates from stromata at maturity. Conidiophores 9–15 × 1.5–2.5 μm ( = 11.4 × 2 μm, n = 20), cylindrical, shorter than conidiogenous cells, branched, hyaline. Conidiogenous cells 10–20 μm high, 1–1.5 μm diam ( = 16 × 1.3 μm, n = 20), phialidic, cylindrical, tapering towards the apices, bearing single conidia at each tip. Conidia 4.5–6 × 0.5–1.5 μm ( = 5 × 1.3 μm, n = 20), eguttulate, elongated to allantoid, slightly curved, aseptate, hyaline.
Culture characteristics: Colonies growing on PDA attenuated 2 cm incubated at 18 °C within 10 d, circular, flat, entire, white, thin, slightly aerial mycelia, loosely attached to the media.
Specimen examined: Russia, Rostov Region, Krasnosulinsky District, Donskoye forestry, Kabanya Balka (Boar gully), twigs and branches of Cornus sanguinea subsp. australis (Cornaceae), 27 Oct. 2015, T.S. Bulgakov, R1111, MFLU 17–0891, living culture MFLUCC 16–1190.
Notes: Cytospora salicina was introduced by Norphanphoun et al. (2017) from Russia causing canker on Salix sp. However, we collected this specimen from Russia associated with twigs and branches of Cornus sanguinea. Cytospora salicina is closely related to C. chrysosperma, C. melnikii, and C. sordida (Fig. 1, Clade 16).
Diaporthaceae Höhn. ex Wehm., Am. J. Bot. 13: 638. 1926. Clade 14.
Pathogenic, endophytic or saprobic on terrestrial and rarely submerged plants. Sexual morph: Pseudostromata well- or poorly developed, pulvinate, erumpent, flat or slightly convex, orbicular, circular or somewhat irregular, sclerotioid, coriaceous, whitish to brownish black, with or without black zone or a crust consisting of fungus tissue, solitary or containing up to 10 ascomata in a stroma. Ectostromatic disk subhyaline to brown. Ascomata perithecial, immersed to erumpent, solitary or aggregated in a valsoid configuration, globose or compressed, coriaceous, black, ostiolate, papillate. Papilla short or long, erumpent, convergent, cylindrical to conical, black, internal wall covered by hyaline periphyses, composed of vertically arranged parenchymatous tissues. Peridium comprising outer layer of flattened, thick-walled, dark-brown cells of textura angularis and inner, hyaline, thin-walled cells of textura angularis. Hamathecium comprising septate, unbranched, cylindrical paraphyses. Asci 8-spored, unitunicate, clavate, oblong-clavate to broadly fusoid, sessile, with a distinct apical ring. Ascospores biseriate to partially biseriate, ellipsoid, oblong to fusoid, unicellular or 1-septate, constricted at septum, with or without appendages at both ends, hyaline, dark brown, sometimes narrowly rounded ends and multi-guttulate, smooth-walled. Asexual morph: Conidiomata acervular or pycnidial, globose, initially immersed, erumpent at maturity, solitary, scattered, coriaceous, black, elongated ostiolar neck, sometime becoming multi-loculate with one to several clearly defined black necks extending above the stroma, often with yellowish, conidial mass extruding from ostiole. Peridium comprising 3–4 layers of pale brown cells of textura intricata to textura angularis. Conidiophores sometimes dimorphic. Alpha conidiophores tightly aggregated, subcylindrical, branched in mid region, consisting of 2–3 supporting cells, giving rise to septate, ampulliform, cylindrical to irregular conidiogenous cells or paraphyses, straight to sinuous, 1–5-septate, cylindrical, hyaline to pale brown, branched only at the base, smooth, formed from the inner most cell layers of the conidiomatal wall, sometimes terminal and lateral, apex with minute periclinal thickening and collarette. Beta conidiophores interspersed among alpha conidiophores, hyaline, subcylindrical, branched, 1–3-septate. Alpha conidiogenous cells enteroblastic, phialidic, cylindrical or subcylindrical, terminal and lateral, slightly tapering towards the apex or sometimes apex with minute periclinal thickening and collarette. Beta conidiogenous cells phialidic, integrated, terminal and lateral. Alpha conidia abundant, fusiform, ovate, subcylindrical to narrowly ellipsoid, straight or curved, occasionally irregular, smooth-walled, 0–2-septate, hyaline, base truncate to sub-truncate, apex obtuse, straight to curved, occasionally slightly sigmoid, pale to medium brown, with many guttules, sometimes short, hyaline, appendages at both ends. Beta conidia subcylindrical, fusiform to hooked, straight to slightly curved, aseptate, hyaline, smooth, base sub-truncate, sometimes widest in middle or in upper third, tapering to acutely rounded apex, truncate at base.
Type genus: Diaporthe Nitschke.
Type species: Diaporthe eres Nitschke.
Notes: The family Diaporthaceae (Fig. 1, Clade 14) comprises many endophytic and phytopathogenic fungal species (Udayanga et al. 2011) and it was introduced and accommodated in Diaporthales by von Höhnel (1917). Wehmeyer (1975) confined this family to Diaporthe and Mazzantia. However, Barr (1978) synonymised Diaporthaceae under Valsaceae. Castlebury et al. (2002) analysed LSU nrDNA sequence data of diaporthoid taxa and showed the distinct placement of Diaporthaceae in Diaporthales, forming a well-supported clade. Diaporthaceae previously comprised only Diaporthe (Phomopsis) and Mazzantia based on phylogenetic analysis (Castlebury et al. 2002). However, Lumbsch & Huhndorf (2010) included Apioporthella and Leucodiaporthe in this family. A LSU nrDNA sequences analysis by Lamprecht et al. (2011) indicates placement of Stenocarpella and Phaeocytostroma within Diaporthaceae. Pustulomyces was introduced based on a combined gene analysis of LSU nrDNA, SSU nrDNA and tef1 sequence data (Dai et al. 2014). Voglmayr & Jaklitsch (2014) confirmed the phylogenetic placement of Phaeodiaporthe in Diaporthaceae based on analysis of LSU nrDNA sequence data. Maharachchikumbura et al. (2015) listed Allantoporthe, Apioporthella, Clypeoporthella, Diaporthe, Diaporthella, Diaporthopsis, Leucodiaporthe, Mazzantia, Mazzantiella, Ophiodiaporthe and Pustulomyces as genera of Diaporthaceae. Rossman et al. (2015) synonymised Mazzantiella under Mazzantia based on greater usage of Mazzantia. The genus Clypeoporthella is based on C. brencklei, and a recently collected C. brencklei (BPI 843482) specimen was grown in culture and sequenced. DNA sequence data showed that C. brencklei clustered together with Diaporthe and it has a Phomopsis asexual morph. Thus, Clypeoporthella is considered as a synonym of Diaporthe (Sogonov et al. 2008). The genus Diaporthopsis was introduced to accommodate species that are similar to Diaporthe, with unicellular ascospores and was typified by D. angelicae. Molecular analysis of LSU nrDNA sequence data showed that D. angelicae clustered within the Diaporthe. In addition, Diaporthopsis angelicae has similar morphological characters of stromata, perithecia, and centrum to species of Diaporthe. Based on morphology and molecular data, Diaporthopsis was synonymised under Diaporthe (Castlebury et al., 2003, Gomes et al., 2013). The genus Diaporthella has aggregated perithecia within well-developed stromata and median, 1-septate ascospores. Diaporthella corylina is strongly parasitic and causes dieback of Corylus stems. Morphologically Diaporthella corylina shows similar characters to Anisogramma anomala. Anisogramma based on A. virgultorum is known to belong in the Gnomoniaceae (Castlebury et al., 2002, Vasilyeva et al., 2007). However, the LSU nrDNA, ITS nrDNA, rpb2 and tef1 combined gene analyses in the current study show (Fig. 1, Clade 5) the phylogenetic placement of Diaporthella is outside of Diaporthaceae and it does not show affinities with any families in Diaporthales. Hence Diaporthaceae comprises Allantoporthe, Apioporthella, Chaetoconis, Diaporthe, Leucodiaporthe, Mazzantia, Ophiodiaporthe, Phaeocytostroma, Phaeodiaporthe, Pustulomyces and Stenocarpella. Based on an LSU nrDNA phylogeny, Gao et al. (2017) showed Diaporthe sensu lato to be polyphyletic, including genera such as Mazzantia, Ophiodiaporthe, Pustulomyces, Phaeocytostroma, and Stenocarpella. In the present study, we address this situation by proposing Chiangraiomyces, Paradiaporthe, Hyaliappendispora as new genera in Diaporthaceae. We collected and illustrate here several taxa in Diaporthaceae that are new to science or are poorly studied.
Chiangraiomyces Senan. & K.D. Hyde, gen. nov. MycoBank MB821544. Facesoffungi number FoF03469.
Etymology: Name related to the collection locality of Chiang Rai, Thailand.
Saprobic on dead wood. Sexual morph: Ascomata solitary, scattered, immersed to erumpent, globose to subglobose, coriaceous, black, papillate, ostiolate. Papilla long, internally covered by hyaline, periphyses. Peridium comprising outer, thick-walled, brown cells of textura angularis and inner, hyaline, thick-walled, compressed cells of textura angularis. Hamathecium comprising hyaline, aseptate, filamentous paraphyses. Asci unitunicate, 8-spored, fusiform, sessile to short pedicellate, with J-, funnel-shaped, apical ring. Ascospores biseriate to overlapping uniseriate, fusiform to ellipsoid, hyaline, smooth-walled, 1-septate, with two large guttules in the centre and two small guttules at the ends. Asexual morph: Conidiomata produced on PDA when incubated at 18 °C after 2 wk, pycnidial, globose, erumpent at maturity, black, coriaceous, short neck. Conidiomatal wall comprising pale brown, thick-walled cells of textura angularis. Conidiophores ampulliform, straight, branched, septate, hyaline, smooth. Conidiogenous cells phialidic, terminal, cylindrical, slightly tapering towards the apex. Hamathecium aparaphysate. Alpha conidia aseptate, hyaline, smooth, ovate to ellipsoidal, less in amount. Beta conidia fusiform to hooked, base subtruncate, aseptate, hyaline, smooth.
Type species: Chiangraiomyces bauhiniae Senan. & K.D. Hyde.
Chiangraiomyces bauhiniae Senan. & K.D. Hyde, sp. nov. MycoBank MB821545. Facesoffungi number FoF03470. Fig. 12.
Chiangraiomyces bauhiniae (MFLU 17–0964). A, B. Ascomata on substrate. C. Cross section of ascoma. D. Peridium. E. Paraphyses. F–H. Asci. I–N. Ascospores. O. Conidiomata on MEA. P, Q. Alpha and beta conidiogeneous cells attached to conidiophores. R. Beta conidia. Scale bars: A, O = 500 μm, B = 200 μm, C = 100 μm, D–H, P–R = 10 μm, I–N = 5 μm.
Etymology: Name based on the host Bauhinia, from which it was collected.
Saprobic on Bauhinia sp. Sexual morph: Ascomata 200–300 μm high, 150–180 μm diam ( = 230 × 240 μm, n = 20), solitary, scattered, immersed to erumpent, globose to subglobose, coriaceous, black, papillate, ostiolate. Papilla 115–140 μm high, 75–90 μm diam ( = 130 × 85 μm, n = 20), long, internally covered by hyaline, periphyses. Peridium 11–14 μm wide ( = 12.5 μm, n = 20), comprising outer, thick-walled, brown cells of textura angularis and inner, hyaline, thick-walled, compressed cells of textura angularis. Hamathecium 2.5–3 μm wide ( = 2.8 μm, n = 20), comprising hyaline, aseptate, filamentous paraphyses. Asci 75–90 × 12–13 μm ( = 78 × 12.5 μm, n = 20), unitunicate, 8-spored, fusiform, with J-, funnel-shaped, apical ring, sessile to short pedicellate. Ascospores 17–18 × 3–4 μm ( = 17.8 × 3.6 μm, n = 20), biseriate to overlapping uniseriate, fusiform to ellipsoid, hyaline, smooth-walled, 1-septate, with two large guttules in the centre and two small guttules at the ends. Asexual morph: Conidiomata 300–500 μm diam ( = 450 μm, n = 20), produced on PDA when incubated at 18 °C after 2 wk, pycnidial, globose, erumpent at maturity, black, coriaceous, short neck. Conidiomatal wall comprising pale brown, thick-walled cells of textura angularis. Conidiophores 4–6 × 2–4 μm ( = 5 × 3 μm, n = 20), ampulliform, straight, branched, septate, hyaline, smooth. Conidiogenous cells 7–10 × 2–3 μm ( = 8 × 2.3 μm, n = 20), phialidic, terminal, cylindrical, slightly tapering towards the apex. Hamathecium aparaphysate. Alpha conidia 3–5 × 2–4 μm ( = 4.7 × 3.3 μm, n = 20), aseptate, hyaline, smooth, ovate to ellipsoidal, less in amount. Beta conidia 18–38 × 1.5–2 μm ( = 24 × 1.7 μm, n = 20), fusiform to hooked, base sub-truncate, aseptate, hyaline, smooth.
Culture characteristics: Colonies growing on MEA attained 1 cm within 7 d when incubated 25 °C, fast growing, circular, irregular, flat, white, forming aerial mycelia with hyphae loosely attached to the medium.
Specimen examined: Thailand, Chiang Rai, Mae Fah Luang University, near University President's house, on dead twigs of Bauhinia sp. (Fabaceae), I.C. Senanayake, 25 Dec. 2014, CHUNI 81 (holotype MFLU 17-0964, cultures ex-type MFLUCC 17–1669, MFLUCC 17–1670).
Notes: Chiangraiomyces bauhiniae has immersed, solitary ascomata, fusiform asci, with a J-, funnel-shaped, apical ring, and oval to fusiform ascospores with 2 large central guttules and 2 small marginal guttules. Phylogenetically, Chiangraiomyces bauhiniae forms a fully-supported clade that is sister to Ophiodiaporthe cyatheae (Fig. 1, Clade 14). Hence, we introduce Chiangraiomyces to accommodate this taxon.
Paradiaporthe Senan., Camporesi & K.D. Hyde, gen. nov. MycoBank MB821546. Facesoffungi number FoF03471.
Etymology: The name reflects the morphological similarity to Diaporthe.
Saprobic on dead twigs of Artemisia sp. Sexual morph: Ascomata solitary, scattered, immersed, becoming erumpent when mature, globose to subglobose, black, coriaceous, ostiolate, papillate. Papilla periphysate with short, wide, prominent ostiole. Peridium thin at the base, gradually thickening towards the neck, comprising inner, hyaline, compressed, thin-walled cells of textura angularis and outer, thick-walled, brown cells of textura angularis. Hamathecium aparaphysate. Asci 8-spored, unitunicate, fusiform to clavate, sessile, apex rounded with a J-, apical ring. Ascospores biseriate, fusiform with pointed ends, medianly 1-septate, hyaline, smooth-walled. Asexual morph: Undetermined.
Type species: Paradiaporthe artemisiae Senan., Camporesi & K.D. Hyde.
Paradiaporthe artemisiae Senan., Camporesi & K.D. Hyde, sp. nov. MycoBank MB821547. Facesoffungi number FoF03472. Fig. 13.
Paradiaporthe artemisiae (MFLU 17–0886). A. Ascomata on substrate. B. Cross section of ascoma. C. Peridium. D–G. Asci. H–L. Ascospores. Scale bars: A = 200 μm, B = 100 μm, C = 20 μm, D–G, H–L = 10 μm.
Etymology: The name reflects the host genus Artemisia.
Saprobic on dead twigs of Artemisia sp. Sexual morph: Ascomata 280–300 μm high, 180–200 μm wide ( = 290 × 190 μm, n = 10), solitary, scattered, immersed, becoming erumpent when mature, globose to subglobose, black, coriaceous, ostiolate, papillate. Papilla 135–138 μm high, 110–140 μm wide, ( = 136 × 115 μm, n = 10), comprising filiform, hyaline periphyses with short, wide, prominent ostiole. Peridium 8–13 μm ( = 10 μm, n = 10), thin at the base, gradually thickened towards the neck, comprising inner, hyaline, compressed, thin-walled cells of textura angularis and outer, thick-walled, brown cells of textura angularis. Hamathecium aparaphysate. Asci 45–60 × 11–14 μm ( = 51 × 13.5 μm, n = 20) 8-spored, unitunicate, fusiform to clavate, sessile, apex rounded, with a J-, bi-lobed, apical ring. Ascospores 14–18.5 × 4–5 μm ( = 16 × 4.2 μm, n = 20) biseriate to overlapping uniseriate, fusiform with two small globules at the ends and two large globules at the middle of spore, medianly 1-septate, hyaline, smooth-walled. Asexual morph: Undetermined.
Culture characteristics: Colonies growing on MEA attained 1 cm within 7 d when incubated at 18 °C, irregular, circular, flat, woolly, white, mycelia loosely attached to the substrate.
Specimen examined: Italy, Province of Forlì-Cesena, Bagno di Romagna, Valbonella, on dead stem of Artemisia sp. (Asteraceae), E. Camporesi, 9 Jul. 2014, IT 1982 (holotype MFLU 17–0886, isotypes BBH 42448, cultures ex-type MFLUCC 14–0850, MFLUCC 17–1663).
Notes: Paradiaporthe artemisiae has erumpent, solitary ascomata with prominent, wide papilla. Morphologically, Paradiaporthe is similar to Diaporthe. However, Paradiaporthe artemisiae forms a distinct clade which is sister to Phaeocytostroma artemisiae (Fig. 1, Clade 14). Hence, we introduce Paradiaporthe as a new genus based on morphology and phylogeny.
Hyaliappendispora Senan., Camporesi & K.D. Hyde, gen. nov. MycoBank MB821548. Facesoffungi number FoF03473.
Etymology: Name reflects hyaline ascospores with long appendages.
Saprobic on dead stems. Sexual morph: Ascomata solitary to aggregate, immersed, globose to subglobose, black to brown, coriaceous, ostiolate, papillate. Papilla short, wide, internally covered by hyaline periphyses. Peridium comprising outer, dark brown, thick-walled cells of textura angularis and inner, thin-walled, hyaline, compressed cells of textura angularis. Hamathecium comprising filiform, septate, hyaline paraphyses which are longer than asci. Asci 8-spored, unitunicate, cylindrical to fusiform, short pedicellate, apex rounded with a J- apical ring. Ascospores biseriate to overlapping biseriate, oval to ellipsoid, hyaline, medianly 1-septate, multiguttulate, with appendages. Appendages at both apical and basal ends, long, thread-like, covered by loose capsule. Asexual morph: Ceolomycetous. Sporulate on PDA at 20 °C after 1 mo, crowded at colony margin, appears at pale yellow bubbles when release the conidial mass. Conidiomata globose, erumpent, black. Peridium comprising thick-walled, pale brown cells of textura angularis. Conidiophores ampulliform, septate, branched, hyaline. Conidiogeneous cells phialidic, terminal, cylindrical, elongate, hyaline. Conidia fusiform, unicellular, hyaline, smooth.
Type species: Hyaliappendispora galii Senan., Camporesi & K.D. Hyde.
Hyaliappendispora galii Senan., Camporesi & K.D. Hyde, sp. nov. MycoBank MB821549. Facesoffungi number FoF03474. Fig. 14.
Hyaliappendispora galii (MFLU 15–2269). A–C. Ascomata on substrate. D. Cross section of ascoma. E. Peridium. F. Paraphyses. G–J. Asci. K–N. Ascospores. O. Culture in upper surface view. P. Culture in lower surface view. Q. Conidiomata on PDA. R, S. Conidiogeneous cells, conidiophores and conidia. T. Conidia. Scale bars: A–C, Q = 200 μm, D = 100 μm, E = 20 μm, F–N, R–T = 10 μm.
Etymology: The name reflects the host genus Galium.
Saprobic on dead stem of Galium sp. Sexual morph: Ascomata 395–450 μm high, 180–200 μm wide ( = 419 × 190 μm, n = 10), solitary to aggregated, immersed, globose to subglobose, black to brown, coriaceous, ostiolate, papillate. Papilla 160–210 μm high, 100–185 μm wide ( = 172 × 158 μm, n = 10), short, wide, internally covered by hyaline periphyses. Peridium 15–25 μm wide ( = 20 μm, n = 10), comprising outer, dark brown, thick-walled cells of textura angularis and inner, thin-walled, hyaline, compressed cells of textura angularis. Hamathecium comprising filiform, septate paraphyses 1.5–3.5 μm wide ( = 2 μm, n = 10), which are longer than asci. Asci 110–125 × 20–25 μm ( = 116 × 21 μm, n = 20), 8-spored, unitunicate, cylindrical to fusiform, short pedicellate, apex rounded with a J- apical ring. Ascospores 20–25 × 7–10 μm ( = 22 × 9 μm, n = 20), biseriate to overlapping biseriate, oval to ellipsoid, hyaline, medianly 1-septate, multiguttulate, with appendages. Appendages 6–11 × 2–3 μm ( = 8 × 2.3 μm, n = 10), at both ends, long, thread-like, covered by loose capsule. Asexual morph: Ceolomycetous. Sporulate on PDA at 18 °C after 1 mo, crowded at colony margin, appears at pale yellow bubbles when release the conidial mass. Conidiomata globose, erumpent, black. Peridium comprising thick-walled, pale brown cells of textura angularis. Conidiophores 10–15 × 1.5–2.5 μm ( = 13 × 2.1 μm, n = 10), ampulliform, septate, branched, hyaline. Conidiogeneous cells 8–16 × 1.5–3 μm ( = 11 × 2.5 μm, n = 20), phialidic, terminal, cylindrical, elongate, hyaline. Conidia 7.5–9.5 × 1.5–2.5 μm ( = 8.3 × 2.2 μm, n = 20), fusiform, unicellular, hyaline, smooth.
Culture characteristics: Colonies growing on PDA incubated at 18 °C attaining 1 cm diam within 14 d, irregular, undulate, umbonate, whitish ash clots with tightly arranged, short, aerial mycelium, erumpent, globose, pale brownish, with viscous droplets produced after 7 d, when colonies incubate further, conidiomata arised on culture media, concentrated at colony margin, appears as black, coriaceous bubbles at the beginning and later become yellow, slimy bubbles with conidial mass.
Specimen examined: Italy, Province of Arezzo, Quota, near Casuccia di Micheli, on dead stem of Galium sp. (Rubiaceae), E. Camporesi, 8 Jun. 2015, IT 2925 (holotype MFLU 15–2269, isotype BBH 42450, culture ex-type MFLUCC 16–1208).
Notes: Hyaliappendispora is morphologically distinct from other genera in Diaporthaceae in having biguttulate, uniseptate, hyaline ascospores with long filamentous apical and basal appendages and wall of the appendages makes a ring-like ornamentation at the proximal end. Phylogenetically Hyaliappendispora galii forms a fully-supported distinct clade that is sister to Phaeodiaporthe (Fig. 1, Clade 14).
Chaetoconis polygoni (Ellis & Everh.) Clem., Gen. fung. (Minneapolis): 176. 1909. Facesoffungi number FoF03475. Fig. 15.
Chaetoconis polygoni (MFLU 17–0965). A. Conidiomata on substrate. B, D. Cross section of conidioma. C. Peridium. E–H. Conidiogenous cells attached to conidia. I–K. Conidia. Scale bars: A = 1 mm, B, D = 100 µm, C, E–H = 50 µm, I–K = 10µm.
Synonym: Amphorula polygoni (Ellis & Everh.) Petr., Sydowia 13: 181. 1959.
Saprobic on stem of Rumex acetosa. Sexual morph: Undetermined. Asexual morph: Conidiomata 175–250 μm high, 200–275 μm diam ( = 200 × 250 μm, n = 20), pycnidial, scattered, immersed to erumpent, globose to sub-globose, dark brown, unilocular or multilocular, ostiolate, papillate. Peridium 20–30 μm thick, comprising several layers of inner thin-walled, hyaline, compressed cells of textura angularis and outer, thick-walled, dark brown cells of textura angularis. Ostiole one or more, circular. Conidiophores 12–25 × 2–3.5 μm ( = 20 × 3 μm, n = 20), hyaline, branched, septate, smooth, with acropleurogenous conidia, formed from the inner pycnidial wall cells. Conidiogenous cells 30–45 × 9–11 μm ( = 32 × 9.5 μm, n = 20), enteroblastic, phialidic, determinate, integrated, cylindrical, hyaline, smooth, with minute channel and collarette. Conidia 35–50 × 4–5 μm ( = 37 × 4.5 μm, n = 20), hyaline, 2-euseptate, continuous, base obtuse, apex extended into a filiform, cellular, unbranched appendage, thin-walled, smooth, guttulate, obclavate.
Specimen examined: Germany, on the edge of a mixed forest, 39 m asl, sandy, acid, fresh, mesotroph, on stem of Rumex acetosa (Polygonaceae), 9 May 2013, RK. Schumacher, CHUNI 73, MFLU 17–0965.
Notes: Chaetoconis polygoni has quite different morphological characteristics compared to other taxa in Diaporthaceae. Molecular analyses in this study showed that our collection clustered together with C. polygoni (CBS 405.95; Fig 1, Clade 14). However, we could not obtain a culture and therefore extracted DNA directly from the sporocarps. The sexual morph of Chaetoconis polygoni was reported as Ceriospora polygonacearum (Barney et al. 2006) which was assigned to Sordariales (Campbell et al. 2003) and later Senanayake et al. (2015) reassigned it to Xylariales. However, morphologically Ceriospora polygonacearum does not show any affinity to Diaporthales.
Diaporthe litoricola Senan., E.B.G. Jones & K.D. Hyde, sp. nov. MycoBank MB821550. Facesoffungi number FoF03476. Fig. 16.
Diaporthelitoricola (MFLU 17–0874). A–C. Ascomata on substrate. D. Cross section of ascomata. E. Peridium. F–I. Asci. J–L. Ascospores. M, N. Conidioma. O, P. Conidiophores, conidiogenous cells and conidia arrangement. Q. Alpha conidia. R, S. Beta conidia. Scale bars: A = 500 µm, B–D, M, N = 200 µm, E = 50 µm, F–I, O, P = 20 µm, J–L, Q, S = 10 µm.
Etymology: The name is based on the Latin words “litore” and “cola” meaning “beach-loving” since this fungus was collected from dead branches of beach plants.
Saprobic on dead stem of sea-shore plants. Sexual morph: Ascomata 800–900 μm high, 450–600 μm diam ( = 880 × 475 μm, n = 20), solitary, scattered, immersed, globose to subglobose, dark brown, coriaceous, ostiolate, papillate. Papilla 380–430 μm high, 110–140 μm diam ( = 420 × 130 μm, n = 20), conspicuous, long, black, with pale yellow apex, brown, unbranched seta in apex, internally covered by hyaline, filamentous periphyses. Peridium 7–12 μm wide, ( = 9.3 μm, n = 20), comprising several layers of compressed, thick-walled, olivaceous to brown cells of textura angularis. Hamathecium aparaphysate or sometimes with a few cellular paraphyses. Asci 80–90 × 11–12 μm ( = 87.5 × 11.1 μm, n = 20), 8-spored, unitunicate, cylindrical, pedicellate, apex rounded, with bilobed, distinct apical ring. Ascospores 16–19 × 4.5–5 μm ( = 18 × 4.8 μm, n = 20), biseriate, fusiform to ellipsoid, 1-septate, hyaline, guttulate. Asexual morph: Conidiomata 500–900 μm high, 800–1 000 μm diam ( = 880 × 900 μm, n = 20), produced on PDA when incubated at 18 °C after 4 wk, pycnidial, globose, initially immersed, erumpent at maturity, black, coriaceous, elongated neck, often yellowish white, with conidial cirrus extruding from ostiole. Conidiomatal wall comprising pale brown, thick-walled cells of textura angularis. Conidiophores 5–7 × 4–7 μm ( = 6.3 × 5.3 μm, n = 20), ampulliform, straight to sinuous, unbranched, hyaline to olivaceous, smooth. Conidiogenous cells 14.5–21 × 1.8–2.8 μm ( = 17.3 × 2.3 μm, n = 20), phialidic, terminal, cylindrical, slightly tapering towards the apex. Hamathecium aparaphysate. Alpha conidia 13–16 × 2.8–3.8 μm ( = 14.7 × 3.3 μm, n = 20), aseptate, hyaline, smooth, ovate to ellipsoidal, base subtruncate, often biguttulate. Beta conidia 1.5–2 × 18–38 μm ( = 1.7 × 24 μm, n = 20), fusiform to hooked, base sub-truncate, aseptate, hyaline, smooth.
Culture characteristics: Colonies growing on PDA attained 1 cm diam within 7 d when incubated at 18 °C, flat, circular, smooth, white, slightly woolly, tightly attached to media, mycelial ends unbranched.
Specimen examined: UK, Hampshire, Eastney shore, on stem of undetermined sea-shore plant, 20 Mar. 2016, E.B.G. Jones, GJ 242 (holotype MFLU 17–0874, isotype BBH 42436, cultures ex-type MFLUCC 16–1195, MFLUCC 17–1657).
Notes: Diaporthe litoricola differs morphologically from D. maytenicola in having large, multi-guttulate ascospores, cylindrical asci, deeply immersed, long papillate, solitary ascomata and elongate, fusiform to cylindrical alpha conidia. Phylogenetically this fungus is closely related to Diaporthe maytenicola, D. decedens and D. nobilis. Diaporthe litoricola forms a moderately-supported clade in this study (Fig 1, Clade 14).
Diaporthe rudis (Fr.) Nitschke, Pyrenomycetes Germanici 2: 282. 1870. Facesoffungi number FoF03477. Fig. 17.
Diaporthe rudis (MFLU 17–0895). A–C. Ascomata on substrate. D–E. Cross sections of ascoma. F. Peridium. G–J. Asci. K–P. Ascospores. Scale bars: A = 500 μm, B, C = 200 μm, D, E = 100 μm, F = 20 μm, G–P = 10 μm.
Saprobic on dead umbelliferous stems. Sexual morph: Clypeus appears as black, wide patches, forming a black mat on substrate connecting all the ascomata and spread around the individual ascomata. Ascomata 540–620 μm high, 250–275 μm wide ( = 590 × 260 μm, n = 10), solitary or rarely aggregated, erumpent, globose to subglobose, black, coriaceous, ostiolate, papillate. Papilla 290–375 μm high, 75–95 μm wide ( = 330 × 85 μm, n = 10), long, asymmetrically located, straight or curved, internally covered by hyaline periphyses, with apex of papilla pale brown, swollen, blunt, sometimes slightly covered by black, mycelial mat. Peridium 11–16 μm wide ( = 14 μm, n = 10), comprising thick-walled, brown, compressed cells of textura angularis. Hamathecium aparaphysate. Asci 43–46 × 11–12 μm ( = 43 × 11.6 μm, n = 10), 8-spored, unitunicate, clavate to fusiform, sessile, apex rounded, with a characteristic, bilobed, J- apical ring. Ascospores 11–13 × 3–4.5 μm ( = 12 × 3.9 μm, n = 10), biseriate, fusiform to elongate ellipsoid, 1-median septate, with each cell containing two guttules, hyaline, smooth-walled. Asexual morph: Undetermined.
Culture characteristics: Colonies growing on PDA attaining 2.5 cm diam within 10 d when incubated at 18 °C, circular, entire, flat, white, tightly attached to the media, aerial mycelia less or sparse, forming few, erumpent, globose, black, viscous droplets after 7 d.
Specimen examined: UK, Hampshire, Winchester, Whiteley, Botley Wood, on umbelliferous stem, 25 May 2016, E.B.G. Jones, GJ 301 (MFLU 17–0895, BBH 42452, living cultures MFLUCC 16–1197, MFLUCC 17–1658).
Notes: Diaporthe rudis was epitypified by Udayanga et al. (2014) based on morphology and phylogeny. Diaporthe rudis has a broad host range. This collection was obtained from umbelliferous woody stems and it forms very long, curved, narrow, papilla deeply immersed in substrate. They appear as pale yellow spots with black margins. Ostioles are blunt and covered by pale yellow cells. However, the base of the ascomata is immersed in deep layers of substrate. All D. rudis cluster together and phylogenetically related to D. cynaroidis and D. cassines. The phylogenetic affinities of these species are still unclear, but morphologically they are differing in terms of asci and ascospore morphology and size.
Diaporthe eres Nitschke, Pyrenomycetes Germanici 2: 245. 1870. Fig. 18.
Diaporthe eres (MFLU 15–2104). A, B. Conidiomata on substrate. C. Cross section of conidioma. D. Wall of conidioma. E–H. Conidiophore, conidiogenous cell attached to conidia. I, J. Conidia. Scale bars: A = 1 mm, B = 200 μm, C = 100 μm, D = 20 μm, E–H = 10 μm, I, J = 5 μm.
Saprobic on stem of Fraxinus pennsylvanica. Sexual morph: Not observed. Asexual morph: Conidiomata 125–140 μm high, 265–300 μm diam at base ( = 135 × 280 μm, n = 10), pycnidial, pyriform, initially immersed, erumpent at maturity, globose to pyriform, black, coriaceous, elongated neck, often with yellowish white, conidial cirrus extruding from ostiole. Conidiomatal wall 34–36 μm diam ( = 35 μm, n = 10), parenchymatous, consisting of 4–7 layers of pale brown, thick-walled cells of textura angularis. Conidiophores 4–6 × 4.5–8 μm ( = 4.6 × 6.6 μm, n = 20), ampulliform, straight to sinuous, unbranched, hyaline, smooth. Conidiogenous cells 8–14 × 1.5–3 μm ( = 11.2 × 2.2 μm, n = 20), phialidic, terminal, cylindrical, slightly tapering towards the apex. Hamathecium aparaphysate. Alpha conidia 5.8–7.5 × 2.5–3.5 μm ( = 6.4 × 2.8 μm, n = 10), aseptate, hyaline, smooth, ovate to ellipsoidal, base subtruncate, often biguttulate. Beta conidia not seen.
Culture characteristics: Colonies growing on MEA attenuated 2 cm within 10 d incubated at 18 °C, fast growing, entire, flat circular, white, with radially arranged minute mycelium clots later becoming creamy or pale yellow.
Specimen examined: Russia, Cotton Fabric urban micro district, on stem of Fraxinus pennsylvanica (Oleaceae), 14 May 2015, T.S. Bulgakov, T-400 (MFLU 15–2104, MFLU 17–0890, living cultures MFLUCC 17–1667, MFLUCC 17–1668).
Notes: Phylogeny depicts a close association between the two D. eres strains collected from Fraxinus pennsylvanica. In this study, D. eres has been treated as a “complex”. It is noted herein that combined gene phylogeny also support such a complex as strains from different hosts/ regions are phylogenetically apart. We did not see beta conidia for this strain on the host or in culture.
Erythrogloeaceae Senan., Maharachch. & K.D. Hyde, fam. nov. MycoBank MB821551. Facesoffungi number FoF03478. Clade 9.
Foliicolous associated with leaf spots. Sexual morph: Undetermined. Asexual morph: Conidiomata epiphyllous, subepidermal, sometime eustromatic, acervular or subglobose, brown to black or yellow-orange, amphigenous, opening by irregular rupture, wall of 2–6 layers of orange-brown textura angularis, exuding slimy orange masses of conidia. Conidiophores reduced to conidiogenous cells. Conidiogenous cells lining the inner cavity of conidioma, hyaline to olivaceous, smooth, subcylindrical to ampulliform, tapering to a long, thin neck, at times apical part elongated into a long neck, proliferating several times percurrently near apex, with flaring collarettes, or apex truncate, with minute periclinal thickening. Conidia hyaline to olivaceous, smooth, guttulate or not, thin-walled, ellipsoid, fusoid, ovoid to somewhat obclavate, straight to curved, apex subobtuse, obtusely rounded, base truncate, with prominent marginal frill, or dimorphic, intermixed in same conidiomata. Macroconidia broadly ellipsoid to obovoid, hyaline, smooth, granular to guttulate, thick-walled, apex obtuse, base flattened. Microconidia hyaline, smooth, guttulate, fusoid-ellipsoid, acutely rounded at apex, truncate at base.
Type genus: Erythrogloeum Petr.
Type species: Erythrogloeum hymenaeae Gonz. Frag. & Cif. ex Petr.
Notes: Phylogenetic analyses from the current study based on combined LSU nrDNA, ITS nrDNA, rpb2, and tef1 sequences showed that Chrysocrypta is basal to Disculoides and Erythrogloeum (Fig. 1, Clade 9). Chrysocrypta has previously been accommodated in the Cryphonectriaceae but the latter is distantly related (Fig. 1, Clade 6). Morphologically members of clade 9 depicts distinct characters in having epiphyllous acervuli, and subcylindrical to ampulliform conidiogenous cells. The sexual morphs of those taxa have not been reported. Disculoides was introduced and typified by Disculoides eucalyptorum (Crous et al. 2012a). Disculoides eucalyptorum was shown to be distinct from Erythrogloeum hymenaeae, which was sister to the Greeneria-Melanconiella complex based on rDNA sequence gene analyses (Crous et al. 2012a).
Chrysocrypta was introduced based on Chrysocrypta corymbiae, which was isolated from leaves of Corymbia species. Chrysocrypta is similar to Foliocryphia (Cryphonectriaceae), but is distinct in forming dimorphic conidia. Crous et al. (2012c) accommodated this taxon in Cryphonectriaceae based on morphology and rDNA sequence phylogeny. However, stromatic tissues of Chrysocrypta corymbiae do not turn purple with KOH, which is a basic characteristic of Cryphonectriaceae. In addition, DNA sequence data herein indicate that Chrysocrypta corymbiae does not belong in Cryphonectriaceae. Hence given the morphological distinctiveness and strongly supported clade (9), a new family Erythrogloeaceae is introduced to accommodate Chrysocrypta, Disculoides and Erythrogloeum.
Erythrogloeum hymenaeae Gonz. Frag. & Cif. ex Petr., Sydowia 7: 379. 1953. Facesoffungi number FoF03479. Fig. 19.
Erythrogloeum hymenaeae (F45467). A. Packet of specimen. B. Herbarium specimen. C–D. Conidiomata on substrate. E. Peridium in KOH. F–H. Conidiophores, Conidiogeneous cells and conidia. I. Conidia. Scale bars: C = 100 μm, D = 200 μm, E–G = 10 μm, H–I = 5 μm.
Foliicolous, associated with leaf spots. Sexual morph: Undetermined. Asexual morph: Conidiomata up to 250 μm diam, acervular, epiphyllous, eustromatic, subepidermal, solitary, rupturing surface by irregular splits. Peridium comprises thin-walled cells of textura angularis. Conidiophores reduced to conidiogenous cells. Conidiogenous cells 5–10 × 2.5–4 μm, hyaline, smooth, phialidic with periclinal thickening, discrete, lageniform to cylindrical, lining the inner walls of cavity. Conidia 7–9 × 2.5–3 μm, hyaline, smooth, guttulate or not, thin-walled, ellipsoid to ovoid, apex obtusely rounded, tapering to a truncate base (description based on Crous et al. 2012a).
Specimen examined: Costa Rica, San José, on leaves of Hymenaea courbaril, Nov. 1929, H. Schmidt (F45468 syntype).
Notes: The monotypic genus Erythrogloeum comprises the type species Erythrogloeum hymenaeae, which is validly described based on Phyllosticta hymenaeae by Petrak (1953). Erythrogloeum hymenaeae is associated with a severe anthracnose of apical twigs and seedlings of Hymenaeae species (Ferreira et al. 1992). This fungus has been reported from in Brazil and Costa Rica.
Gnomoniaceae G. Winter [as ‘Gnomonieae’], Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1.2: 570. 1886. Clade 1.
Saprobic on bark and leaves of overwintered plants. Sexual morph: Stromata lacking, or poorly to well-developed, scattered, erumpent, pustuliform with one or rarely two ascomata or valsoid, broadly elliptic to rounded, large. Ectostromata well-developed, brown to black, thick ectostromatic disc at perithecial necks. Ascomata immersed to erumpent, solitary or aggregated, globose to subglobose, black, coriaceous, thin-walled, with one or more long, central or eccentric necks with hyaline periphyses. Peridium comprising few layers of brown, thick-walled cells of textura angularis. Hamathecium comprising few hyaline, septate, cellular paraphyses. Asci 8–32-spored, unitunicate, oval, fusiform to almost filiform, short pedicellate, with a distinct, J- apical ring. Ascospores biseriate, overlapping uniseriate to fasciculate, oval, fusiform, ovoid to subulate, small, unicellular to 1-septate, rarely multi-septate, ends mostly rounded, rarely pointed, appendages absent or subulate, navicular or whip-shaped, smooth. Asexual morph: Conidiomata acervulal or pycnidial, subcuticular, papillate or not, oblate to globose, black, thick-walled, with one chamber containing whitish conidial mass. Conidiophores simple, filiform to fusiform, annellation visible or invisible, densely branched. Conidiogenous cells usually phialidic, rarely with a few annellidic scars, irregular in shape, lageniform to cylindrical, gradually tapering to ends for one quarter to three-quarters of their length, or abruptly narrowing to long neck at about half of the phialide length, or abruptly narrowing at apex, straight or curved, sometimes asymmetric swollen nodes, proliferating into other conidiogenous cells at basal or middle part. Conidia broadly ellipsoid to oval, sometimes obovoid, allantoid, occasionally curved or sinuate to slightly angular, hyaline, often unicellular.
Type genus: Gnomonia Ces. & De Not.
Type species: Gnomonia gnomon (Tode) J. Schröt.
Notes: Gnomoniaceae (Fig. 1, Clade 1) was introduced by Winter (1886). This family is characterised by immersed, rarely erumpent or superficial ascomata, without a stroma or aggregated with a rudimentary stroma. Species in Gnomoniaceae inhabit various hosts and substrates, including herbaceous plants, shrubs and trees as endophytes, pathogens and saprobes (Rossman et al., 2007, Walker et al., 2012). Pathogenicity of gnomoniaceous taxa is quite diversified, causing various diseases on plants. However, most gnomoniaceous species are restricted to overwintered plants in temperate and subtropical biomes.
Maharachchikumbura et al. (2015) accommodated 33 genera in Gnomoniaceae. Additionally, we introduce a new genus Marsupiomyces based on M. quercina and the second species M. epidermoidea. However, we exclude five genera from the family based on morphology and phylogeny and also included the additional genera Mamianiella and Marsupiomyces within this family. Hence, we accept 30 genera in this family: Alnecium, Ambarignomonia, Amphiporthe, Anisomyces, Apiognomonia, Apioplagiostoma, Asteroma, Bagcheea, Cryptosporella, Cylindrosporella, Diplacella, Ditopella, Ditopellopsis, Gloeosporidina, Gnomonia, Gnomoniella, Gnomoniopsis, Mamianiella, Marsupiomyces, Millerburtonia, Occultocarpon, Ophiognomonia, Phragmoporthe, Phylloporthe, Plagiostoma, Pleuroceras, Sirococcus, Spataporthe, Uniseta and Valsalnicola. Here we introduce, describe and illustrate new fungal taxa which belong to Gnomoniaceae.
Doubtful genera or genera excluded from Gnomoniaceae
Anisogramma was introduced and typified by Anisogramma virgultorum, and almost all characters of this genus are similar to Mamianiella. De Silva et al. (2009) analysed the phylogenetic relationship of Anisogramma species based on LSU nrDNA sequence data and reported its placement outside of Gnomoniaceae. Both Mamianiella and Anisogramma commonly occur on Corylus species. Combined LSU nrDNA, ITS nrDNA, rpb2 and tef1 gene analyses of the present study show that Mamianiella is nested in between Anisogramma species. Morphological comparison also reveals that both genera are characterised by Mamianiella and hence Mamianiella does not warrant generic status with high bootstrap support value (Fig. 1, Clade 1). Mamianiella is an older name than Anisogramma. Therefore, we synonymise Anisogramma under Mamianiella giving priority to the older name. Mamianiella Höhn. was introduced and typified by M. coryli, (based on Sphaeria coryli) and Mamiania was introduced and typified by M. fimbriata (based on Sphaeria fimbriata). Von Arx & Muller (1954) suggested to retain both genera as one genus. However, Barr (1978) separated these two genera based on ascospore morphology as Mamianiella has unicellular ascospores, while Mamiania produces apiosporous ascospores. This is, however, not a strong character to differentiate these two genera, while almost all other characters are similar to each other. Hence, we synonymise Mamiania under Mamianiella giving priority to the older name Mamianiella.
Clypeoporthe, was reduced to synonymy in Gnomonia by Monod (1983). However, some species in this genus have eutypelloid configuration of ascomata in parenchymatous stromatic tissues. However, it is necessary to obtain DNA sequence data to resolve this genus. Depazea was typified by D. frondicola and it was assigned to Mycosphaerellaceae as Sphaerulina frondicola (Verkley et al. 2013). Hence, we exclude Depazea form Gnomoniaceae.
Phylloporthe, a plant parasitic, monotypic genus was introduced and is typified by P. vernoniae. There is no molecular data for P. vernoniae and it is not clear whether this genus belongs to Gnomoniaceae or not. Hence, we maintain this genus in Gnomoniaceae until molecular data for the type species are available.
Skottsbergiella was introduced and typified by Skottsbergiella diaporthoides which has large perithecia immersed in massive, externally crustose, pseudoparenchymatous stromata. Petrak (1971) assigned this genus to eutypoid fungi based on its stromatic consistency. This genus is morphologically similar to Diaporthella, which is placed in Diaporthales incertae sedis (Barr 1978). Skottsbergiella diaporthoides was renamed as Diaporthe diaporthoides and accommodated in Diaporthaceae (Barr 1978). Hence Skottsbergiella is not a valid genus.
Xenotypa is typified by X. aterrima. We observed a specimen of X. aterrima (as Hydnum aterrima, from S under accession no: F130640) on account of the elongated allantoid ascospores and solitary to aggregated ascomata, this taxon has closer affinity to Cytosporaceae than Gnomoniaceae. Hence, we exclude Xenotypa from Gnomoniaceae and include it in Cytosporaceae, until molecular data is available to confirm the placement.
Zythia is typified by Z. resinae which is synonymised under Sarea resinae. Molecular data demonstrate a placement of Sarea resinae within Trapeliaceae (Baeomycetales, Ostropomycetidae). Therefore, here we exclude Zythia from Gnomoniaceae. However, Z. fragariae shows an affinity to Gnomoniaceae. It is a common parasite on strawberry and Shipton (1967) reported Zythia fragariae as the asexual morph of Gnomonia fragariae. Walker et al. (2010) synonymised Gnomonia fragariae in Gnomoniopsis as G. comari. Hence Zythia is not considered to be a genus in Gnomoniaceae.
Plagiostoma salicicola Senan., Camporesi & K.D. Hyde, sp. nov. MycoBank MB821552. Facesoffungi number FoF03480. Fig. 20.
Plagiostoma salicicola (MFLU 17–0878). A. Ascomata on substrate. B. Ascomata distribution on bark epidermis (under surface). C. Cross section of ascoma. D. Papilla. E. Peridium. F–H. Asci. I–L. Ascospores. M. Culture on PDA, upper surface view. N. Culture on PDA, lower surface view. Scale bars: A, B = 500 µm, C = 100 µm, D = 50 µm, E–L = 10 µm.
Etymology: Based on the host genus Salix on which this fungus occurs and the Latin “-cola” which means loving.
Saprobic on dead twigs of Salix sp. Sexual morph: Stromata loosely packed comprising pseudoparenchymatous tissues. Ascomata 400–600 μm high, 250–400 μm diam ( = 580 × 300 μm, n = 20), perithecial, aggregated in groups of 3–10, immersed, oblate globose when moist and become convex with irregular dents around base of papilla when dry, coriaceous, black, ostiolate, papillate. Necks 420–700 μm long, 100–150 μm wide at base, 60–150 μm wide at apex, converged or not, eccentric to marginal, slightly curved. Asci 45–70 × 10–20 μm ( = 62 × 16 μm, n = 20), 8-spored, unitunicate, fusiform, apex narrowly obtuse, sessile, with J- apical ring. Ascospores 15–25 × 4–7 μm ( = 17 × 6 μm, n = 20), obliquely biseriate to fasciculate, ellipsoidal to fusiform, medianly 1-septate, constricted or not at the septum, ends rounded to tapering, with upper cell slightly wider than basal cell, hyaline. Asexual morph: Undetermined.
Culture characteristics: Colonies growing on PDA attained 1 cm within 10 d incubated at 18 °C, circular, umbonate, undulate, white median region with ash outer margin, woolly, loosely attached to the substrate.
Specimen examined: Italy, Province of Trento, Val di Sole, near Croviana, on dead branch of Salix sp., 29 Jul. 2013, E. Camporesi, IT 1394 (holotype MFLU 17–0878, isotype BBH 42440, cultures ex-type MFLUCC 13–0656); Province of Trento, Val di Sole, near Croviana, on dead branch of Salix sp., 29 Jul. 2013, E. Camporesi, IT 1394 (paratype MFLU 15–2261, living cultures MFLUCC 17–1666).
Notes: Mejía et al. (2011) revisited the genus Plagiostoma and observed distinct grouping pattern of Plagiostoma species with expanded necks and species with cylindrical necks on Salicaceae. With species with expanded necks, P. salicicola is morphologically similar to Plagiostoma dilatatum. However, P. dilatatum has relatively small ascomata with short necks and long-pedicellate, cylindrical asci. The combined gene sequence analyses herein indicate a relationship of P. salicicola with other species of Plagiostoma separated with moderate support values, but sufficiently distinct of P. dilatatum (Fig. 1, Clade 1).
Plagiostoma jonesii Senan., & K.D. Hyde, sp. nov. MycoBank MB821553. Facesoffungi number FoF03481. Fig. 21.
Plagiostoma jonesii (MFLU 17–0873). A. Ascomata on host surface. B. Cross section of ascoma. C. Peridium. D–G. Asci. H–L. Ascospores. Scale bars: A = 200 µm, B = 100 µm, C = 20 µm, D–L = 10 µm.
Etymology: In honour of Prof. Gareth Jones, an eminent mycologist who collected this species.
Saprobic on umbelliferous stems. Sexual morph: Ascomata 380–420 μm high, 250–280 μm diam ( = 400 × 270 μm, n = 10), solitary or rarely aggregated, erumpent, globose to subglobose, black, coriaceous, ostiolate, papillate. Papilla 165–260 μm high, 70–100 μm wide ( = 200 × 80 μm, n = 10), short, symmetrically or asymmetrically located, narrow at the base, widening towards the top, straight or curved, internally covered by hyaline periphyses. Peridium 15–25 μm wide ( = 18.5 μm, n = 10), comprises thick-walled, brown, compressed cells of textura porrecta. Asci 40–50 × 8.5–9.5 μm ( = 48 × 8.8 μm, n = 10), 8-spored, unitunicate, fusiform to clavate, apex with J-, bilobed, distinct apical ring, short pedicellate. Ascospores 12–14 × 2.6–3.8 μm ( = 13 × 3.2 μm, n = 10), biseriate, fusiform to ellipsoid, hyaline, 1-septate, with two globules in each cell, with small spine-like appendages at both ends. Asexual morph: Undetermined.
Culture characteristics: Colonies growing on PDA attenuated 1 cm within 7 d, incubated at 18 °C, fast-growing, flat, circular, smooth, less in aerial mycelia, white, tightly attached to the medium.
Specimen examined: UK, Sussex Ocidental, Arundel, river bank, on umbelliferous stem, 17 Feb. 2016, E.G.B. Jones, GJ 227 (holotype MFLU 17–0873, isotype BBH 42435, cultures ex-type MFLUCC 16–1189, MFLUCC 17–1654).
Notes: Plagiostoma jonesii is morphologically and phylogenetically distinct from other Plagiostoma species in having long, curved papilla arising out from the substrate appearing as spines and the opening is wider than base, ellipsoid to fusiform, 1-septate, slightly or non-constricted ascospores with small, appendages. Our phylogeny shows that Plagiostoma jonesii is phylogenetically close to P. salicellum and P. populinum, but morphologically distinct from both species.
Gnomoniopsis agrimoniae Senan., Camporesi & K.D. Hyde, sp. nov. MycoBank MB821554. Facesoffungi number FoF03482. Fig. 22.
Gnomoniopsis agrimoniae (MFLU 17–0884). A, B. Ascomata on host substrate. C. Cross section of ascoma. D. Peridium. E–G. Asci. H. Ascospores. I. Culture growing on MEA, upper surface view. J. Culture growing on MEA, lower surface view. Scale bars: A = 100 µm, B = 200 µm, C = 50 µm, D = 10 µm, E–H = 5 µm.
Etymology: Species epithet based on the host genus Agrimonia.
Saprobic on dead stems of Agrimonia eupatoria. Sexual morph: Ascomata 200–320 μm high, 245–400 μm diam ( = 273 × 332 μm, n = 20) solitary, scattered, erumpent, globose, black, coriaceous, ostiolate, papillate. Papilla 100–170 μm high, 70–105 μm diam ( = 160 × 80 μm, n = 20), short, comprising elongate brown cells of textura porrecta. Peridium 35–45 μm ( = 39 μm, n = 10) comprising inner, hyaline, compressed cells of textura angularis and outer, brown, thick-walled, cells of textura globosa. Asci 28–32.5 × 5–5.5 μm ( = 30.5 × 34.9 μm, n = 20), 8-spored, unitunicate, cylindrical to fusiform, short-pedicellate, apex obtuse with bilobed, J- apical ring. Ascospores 7–8 × 1.8–2.2 μm ( = 7.5 × 2.1 μm) overlapping uni- to biseriate, apiosporous, hyaline, uniseptate, smooth-walled. Asexual morph: Undetermined.
Culture characteristics: Colonies growing on MEA becoming 2 cm within 7 d incubated at 18 °C, fast growing, circular, smooth mycelia concentrated at margins making a concave colony, off white, loosely attached to the substrate, wooly.
Specimen examined: Italy, Province of Forlì-Cesena, near Santa Sofia, on dead stem of Agrimonia eupatoria (Rosaceae), 5 Apr. 2014, E. Camporesi, IT 1798 (holotype MFLU 17–0884, isotype BBH 42446, cultures ex-type MFLUCC 14–0844, MFLUCC 17–1662).
Notes: Gnomoniopsis agrimoniae has minute asci and ascospores compared to the other Gnomoniopsis species. Coriaceous, thick-walled ascomata and small apiosporous ascospores are prominent characters in this genus. Our combined gene analyses indicate a moderately supported phylogenetic distinction of Gnomoniopsis agrimoniae from other species with moderate support. Phylogeny analyses based on ITS sequence data following Walker et al. (2010) reported that Gnomoniopsis agrimoniae is distinct from other Gnomoniopsis species. Gnomoniopsis species are considered host specific and only Gnomoniopsis agrimoniae and G. guttulata are reported on Agrimonia species.
Apiognomonia veneta (Sacc. & Speg.) Höhn., Hedwigia 62: 47. 1920. Facesoffungi number FoF03483. Fig. 23.
Apiognomonia veneta (MFLU 15–3710). A–B. Conidiomata on host leaf surface. C. Cross section of conidioma. D–F. Conidiophore and conidiogenous cells attached to conidia. G, H. Conidia. I. Upper surface view of culture. J. Lower surface of culture. Scale bars: A, B = 1 mm, C = 100 µm, D–H = 10 µm.
Basionym: Laestadia veneta Sacc. & Speg., Michelia 1(no. 3): 351. 1878.
Pathogenic on living leaves of Platanus acerifolia. Sexual morph: Undetermined. Asexual morph: Conidiomata 180–200 μm high, 250–265 μm diam ( = 188 × 260 μm, n = 10), acervular, irregularly round or oval, erumpent to immersed, solitary, scattered, conidiogenous layer covering the entire inner surface of acervular chambers and mostly in basal layer, yellowish-brown, initially developing under epidermis, then breaking through epidermis and forming thick whitish amorphous conidial masses. Conidiophores 10–15 × 2.5–4.5 μm ( = 12 × 3.4 μm, n = 20), densely branched, ampulliform, hyaline. Conidiogenous cells 14–21 × 2.7–3.5 μm ( = 18 × 3 μm, n = 10), usually phialidic, rarely annellidic, lageniform to cylindrical, gradually tapering towards the apex, straight or curved, hyaline, smooth. Conidia 12–20 × 4–6 μm ( = 15.6 × 5 μm, n = 10), broadly ellipsoid to oval, sometimes obovoid, occasionally curved or sinuate to slightly angular, hyaline, thick-walled, aseptate, guttulate.
Culture characteristics: Colonies growing on MEA attenuated 1 cm within 7 d, incubated at 18 °C, flat, circular, irregular, with circular ornamentations, margins concentrated with mycelial ends, white, rich in short aerial mycelia, loosely attached to the medium.
Specimen examined: Russia, Rostov region, Krasnosulinsky district, Donskoye forestry, lining-out nursery, on live leaves of Platanus acerifolia (Platanaceae), 27 Oct. 2015, T.S. Bulgakov, R 1048, MFLU 15–3710, living cultures MFLUCC 16–1193, MFLUCC 17–1656.
Notes: Apiognomonia veneta is a common pathogen on Platanaceae. Here we illustrate the asexual morph of Apiognomonia veneta. This is a common epifoliar pathogen. We could not obtain the sexual morph in culture or from the specimen.
Marsupiomyces Senan. & K.D. Hyde, gen. nov. MycoBank MB821555. Facesoffungi number FoF03484.
Etymology: Referring to the ascomata located in mycelial cavity not in stromatic tissues.
Saprobic on leaves of Fagaceae. Sexual morph: Appearing on the surface as black solitary swellings on the leaf surface. Ascomatal cavity pale in colour, tightly packed cells, forming a thin coating around ascomata. Ascomata solitary, scattered, immersed horizontally in the lower and upper leaf epidermis, globose to subglobose, coriaceous, black, ostiolate, papillate. Papilla long, asymmetrically located, slanted or on substrate, curved or erect. Peridium comprising thick-walled, brown, large cells of textura globulosa or textura epidermoidea. Hamathecium aparaphysate. Asci 8-spored, unitunicate, fusiform, with short, pointed pedicel, apex rounded with bi-lobed, distinct, apical ring. Ascospores uni- to tri-seriate, fusiform, cylindrical to elongate fusiform, straight or very slightly curved, 1-septate, hyaline, guttulate, smooth-walled. Asexual morph: Undetermined.
Type species: Marsupiomyces quercina Senan., Camporesi & K.D. Hyde.
Notes: Marsupiomyces is introduced and typified by M. quercina. Members of this genus occur on members of Fagaceae. Marsupiomyces comprises M. quercina and M. epidermoidea. Phylogenetically Marsupiomyces is closely related to Apioplagiostoma (Fig. 1, Clade 1). However, Apioplagiostoma differs from Marsupiomyces in having leaf lesions with dark purple to brown pigmentation, and apiosporous ascospores.
Marsupiomyces quercina Senan., Camporesi & K.D. Hyde, sp. nov. MycoBank MB821556. Facesoffungi number FoF03485. Fig. 24.
Marsupiomyces quercina (MFLU 17–0876). A. Ascomata on substrate. B. Stromatic cavity. C, D. Cross section of ascoma. E. Peridium. F. Apical ring. G–K. Asci. L–N. Ascospores. Scale bars: A, B = 500 µm, C = 200 µm, D = 50 µm, E = 5 µm, F, L–N = 10 µm, G–K = 20 µm.
Etymology: Species epithet based on the host genus Quercus.
Saprobic on leaves of Quercus. Sexual morph: Stromatic cavity pale in colour, tightly packed cells, forming a thin, coating around ascomata. Ascomata 150–250 μm high 160–300 μm diam ( = 175 × 200 μm, n = 10) solitary, scattered, immersed horizontally in the lower and upper leaf epidermis, globose to subglobose, coriaceous, black, ostiolate, papillate. Papilla long, asymmetrically located, slanted or on substrate, curved or erect. Peridium 25–45 μm wide ( = 35 μm, n = 10), comprising thick-walled, brown, large cells of textura globulosa. Hamathecium aparaphysate. Asci 125–150 × 9–11 μm ( = 134 × 9.8 μm, n = 30), unitunicate, 8-spored, fusiform, with short, pointed pedicel, apex rounded with bilobed, distinct, apical ring. Ascospores 15–21 × 6–8 μm ( = 17.4 × 6.6 μm, n = 40), biseriate, cylindrical to elongate fusiform, 1-septate, hyaline, guttulate. Asexual morph: Undetermined.
Specimen examined: Italy, Province of Forlì-Cesena, San Paolo in Alpe, Santa Sofia, dead leaves of Quercus sp. (Fagaceae), 2 May 2013, E. Camporesi, IT 1214 (holotype MFLU 17–0876, isotype BBH 42438, cultures ex-type = MFLUCC 14–0566, MFLUCC 13–0664).
Notes: The combined ITS nrDNA, LSU nrDNA, rpb2 and tef1 sequences analyses of this study shows that Marsupiomyces quercina forms a distinct clade which is sister to Marsupiomyces epidermoidea (Fig. 1, Clade 1).
Marsupiomyces epidermoidea R.H. Perera, Senan., Bulgakov & K.D. Hyde, sp. nov. MycoBank MB821557. Facesoffungi number FoF03486. Fig. 25.
Marsupiomyces epidermoidea (MFLU 15–2921) A. Herbarium specimen. B, C. Appearance of ascoma on the leaf surface. D. Vertical section through ascoma. E. Peridium in surface view. F. Peridium. G–K. Asci. L–O. Ascospores. Scale bars: B = 1 mm, C = 500 µm, D = 100 µm, E = 50 µm, F–K = 20 µm, L–O = 10 µm.
Etymology: Fungal peridium comprising cells of textura epidermoidea.
Saprobic on dead leaves of Quercus robur. Sexual morph: Appearing on the surface as black solitary swellings on the leaf surface. Ascomatal cavity pale in colour, tightly packed cells, forming a thin, coating around ascomata. Ascomata 200–310 μm diam, depressed globose to irregular. Peridium 11–36 μm thick, comprising 3–8 layers of brown to hyaline cells of textura epidermoidea, outer cell layer brown to pale brown, inner cells hyaline, elongate. Asci 54–83 × 11–15 μm ( = 71 × 14 μm, n = 25), 8-spored, unitunicate, clavate, apedicellate, with a J- refractive apical ring, lying without paraphyses. Ascospores 18–21 × 3–3.6 μm ( = 19 × 3.4 μm, n = 30), uni- to tri-seriate, 1-septate, not constricted at the septum, broadly fusiform, rounded at both ends, straight or very slightly curved, hyaline, guttulate, smooth-walled. Asexual morph: Undetermined.
Specimen examined: Russia, Rostov region, Shakhty city, Maisky, Cemetery Park, (47.6922302° E, 40.0925446° N), on dried leaf of Quercus robur (Fagaceae), 21 Jun. 2015, T.S. Bulgakov, T 776 (holotype MFLU 15–2921, isotype BBH 42451).
Notes: Our new taxon, Marsupiomyces epidermoidea is a sister taxon to Marsupiomyces quercina, but sufficiently distinct. In addition, it is different from Marsupiomyces epidermoidea in having a distinct peridium comprising cells of textura epidermoidea.
Ditopella biseptata R.H. Perera, Senan., Camporesi & K.D. Hyde, sp. nov. MycoBank MB821558. Facesoffungi number FoF03487. Fig. 26.
Ditopella biseptata (MFLU 15–2661). A. Herbarium specimen. B, C. Appearance of ascomata on host substrate. D. Vertical section through ascoma. E. Papilla. G–J. Asci (J in Melzer’s reagent). K–M. Ascospores (M in Indian ink). Scale bars: C = 1 mm, D = 200 µm, E = 100 µm, F = 50 µm, G–J = 20 µm, K–M = 10 µm.
Etymology: Species name refers to the ascospores that have two septa.
Saprobic on dead branch of Alnus glutinosa. Sexual morph: Stromata surrounding the perithecial necks, extending outward beneath the host periderm as a distinct clypeus, composed of dark brown thick-walled angular cells. Ascomata 500–900 μm diam, immersed in the ectostroma, situated between the epidermis and the cortex of the host tissue, appearing as solitary swellings of the host epidermis, sometimes epidermis ruptures to expose the rounded apex of the ostiole, perithecial, depressed globose to oval, ostiolate. Ostiolar neck lined with thin-walled hyaline, septate periphyses. Peridium 44 μm thick, 2-layered, outer layer composed of angular, sometimes slightly compressed, dark brown, thick-walled cells of textura angularis, inner layer of elongate, hyaline, thin-walled, compressed cells of textura angularis, wider around the ostiole, composed of dark brown, thick-walled cells of textura angularis. Asci 63–90 × 15–19 μm ( = 79 × 18.3 μm, n = 10), 16- to 32-spored, elongate-ellipsoidal to clavate, apedicellate, with a J- refractive apical ring, lying without paraphyses. Ascospores 18–27 × 3–4 μm ( = 23.8 × 3.6 μm, n = 30), multi-seriate, (1)2(–3)-septate, not constricted at the septum, cylindrical to narrowly ellipsoidal, straight or very slightly curved, tapering slightly to bro2adly rounded ends, hyaline, guttulate, smooth-walled, with 2-polar appendages. Asexual morph: Undetermined.
Specimen examined: Italy, Province of Forlì-Cesena, Bagno di Romagna, near Lago Pontini, on dead branch of Alnus glutinosa (Betulaceae), 26 May 2014, E. Camporesi, IT 1891 (holotype, MFLU 15–2661).
Notes: Here we introduce a new species Ditopella biseptata based on phylogeny. Ditopella biseptata forms a distinct clade which is sister to Ditopella ditopa (Fig. 1, Clade 1). Morphologically Ditopella biseptata has 2-septa and minute appendages at both ends. We could not obtain a culture from this fungus and extracted DNA directly from the sporocarps.
Harknessiaceae Crous, Persoonia 28: 55. 2012. Clade 7.
Saprobic or pathogenic forming leaf spots. Sexual morph: Ascomata perithecial, solitary or aggregated, immersed, globose, coriaceous, brown, papillate. Papilla emergent to depressed, wall comprising 3–5 layers of brown-walled cells of textura angularis. Hamathecium comprising hyaline, septate paraphyses. Asci 8-spored, unitunicate, cylindrical to clavate, short pedicellate, with J- apical ring. Ascospores uni- to biseriate, hyaline, ellipsoid to fusoid, aseptate, thick-walled, guttulate, smooth-walled. Asexual morph: Coelomycetous. Conidiomata eustromatic, pycnidial, scattered or aggregated, immersed, globose, coriaceous, with single or several locules, dark brown to black. Peridium comprising thin-walled, almost hyaline to brown cells of textura angularis. Ostiole wide, central, surrounded by brown cells. Conidiophores lining the inner cavity or reduced to the basal layer, sometimes reduced to conidiogenous cells, sometimes septate, branched. Conidiogenous cells holoblastic, discrete, lageniform, subcylindrical to cylindrical, hyaline to pale yellow, smooth, producing macroconidia and sometimes microconidia from same conidiogenous cell, proliferating sympodially one or several times. Macroconidia with a basal appendage, hyaline when young, brown at maturity, unicellular, although basal appendage separated by a septum thick-walled, smooth-walled, with or without pale and dark longitudinal bands, sometimes longitudinally striate, guttulate, basal appendage cellular, cylindrical to subcylindrical, hyaline, thin-walled, devoid of contents, apical appendage present or absent, if present elongated. Microconidia hyaline, oval to ellipsoid, aseptate, smooth-walled.
Type genus: Harknessia Cooke.
Type species: Harknessia eucalypti Cooke.
Notes: Harknessiaceae (Fig. 1, Clade 7) was introduced to accommodate Harknessia with its wuestneia-like sexual morph. Harknessia species, distributed in both tropical and temperate biomes, are associated with leaves and branches of host trees (Farr & Rossman 2001). Most pathogenic Harknessia species are associated with leaf spots, leaf tip dieback, leaf scorch and stem cankers (Crous et al., 1989, Crous et al., 1993, Yuan et al., 2000), but pathogenicity has not been properly studied (Crous et al. 2012b). Some saprobic species have also been isolated from asymptomatic plant tissues (Marincowitz et al., 2008, Crous et al., 2017). Twenty-one of the 60 species and seven of the 13 wuestneia-like sexual morphs have been linked to Harknessia asexual morphs (Crous et al., 2012b, Crous et al., 2017). Ribosomal DNA sequence analysis of diaporthoid taxa revealed a distinct lineage for Harknessia sensu stricto within Diaporthales (Crous et al. 2012b). Crous et al. (2012b) introduced six novel species of Harknessia from Eucalyptus and phylogenetic relationships based on a multi-gene analysis of ITS nrDNA, calmodulin and beta-tubulin genes were provided for these species. However morphologically, Dwiroopa lythri has similar characters to Harknessia and phylogenetically it is moderately supported here. Hence, Dwiroopa lythri is accommodated within Harknessiaceae for now. Phylogeny herein, indicates support for the establishment of this family.
Harknessia eucalypti Cooke, Grevillea 9 (no. 51): 85. 1881. Facesoffungi number FoF03488. Fig. 27.
Harknessia eucalypti (K (M) 195744). A. Packet of herbarium. B. Herbarium specimen. C. Cross section of conidioma. D–E. Conidia attached to the conidiogenous cells. F–K. Conidia. Scale bars: C = 100 µm, D–E = 20 µm, F–K = 10 µm.
Saprobic on Eucalyptus globulus appearing as nearly circular, black distinct spots. Sexual morph: Undetermined. Asexual morph: Conidiomata 390–550 μm high, 400–600 μm diam, erumpent, scattered, pycnidial, unilocular, globose to subglobose, brown. Peridium 3–4 layers of brown cells of textura angularis. Conidiophores short, cylindrical, almost globose, branched, 1–2 layers, hyaline, mixed with peridium cells. Conidiogenous cells 8–13 × 4–6 μm, ampulliform, cylindrical, hyaline to brown. Conidia 11.5–15 × 8–9.5 μm ( = 13 × 8.5 μm, n = 20), globose to ovoid with a truncate apiculate apex and an obtuse to blunt base, smooth, hyaline when young, brown at maturity, with longitudinal striations along the length of some conidia. Basal appendages 5–15 × 1.5–3 μm ( = 10 × 2.5 μm, n = 20), hyaline, tubular, smooth, thin-walled, often collapsing.
Material examined: USA, California, on leaves of Eucalyptus globulus, Harkness 1280, isotype K (M) 195744.
Notes: Yuan & Mohammed (1997) observed the asexual morph of Wuestneia epispora from culture which was morphologically identical to Harknessia eucalypti, although this has not been proven based on sequences. Harknessia is associated with leaf spots, leaf tip dieback or leaf scorch, stem cankers and is also common on leaf litter (Crous et al., 1989, Marincowitz et al., 2008).
Juglanconidaceae Voglmayr & Jaklitsch, Persoonia 38: 142. 2017. Facesoffungi number FoF03489. Clade 4.
Synonym: Melansporellaceae C.M. Tian et al. Phytotaxa 305: 194. 2017.
Saprobic on dead corticated twigs and branches of Juglandaceae species. Sexual morph: Pseudostromata inconspicuous, ectostromatic disc pale yellow to pale brown, causing a more or less postulate bark surface. Central column more or less conical, beneath the disc. Ascomata surrounding the ectostromatic disc, with long, asymmetrical or symmetrical, lateral ostioles that emerge at the margin or within the ectostromatic disc, globose to subglobose, coriaceous, black. Hamathecium comprising hyaline paraphyses which deliquesce at maturity. Asci 8-spored, unitunicate, with a distinct apical ring, sessile. Ascospores hyaline, bicellular, with or without gelatinous appendages. Asexual morph: melanconium-like. Conidiomata acervular, with ectostromatic disc and central column. Conidiophores aseptate or few-celled, smooth, hyaline to brown. Conidiogenous cells annellidic, cylindrical, base swollen, hyaline to brown. Conidia ellipsoid to oval, brown, with gelatinous sheath. Conidial wall smooth on the outer surface, with inconspicuous to distinct irregular verrucae on the inner surface (description based on Voglmayr et al. 2017).
Type genus: Juglanconis Voglmayr & Jaklitsch.
Type species: Juglanconis juglandina (Kunze) Voglmayr & Jaklitsch.
Notes: Juglanconidaceae (Fig. 1, Clade 4) was introduced by Voglmayr et al. (2017) based on Melanconium juglandinum. This family comprises Juglanconis juglandina, J. oblonga, J. pterocaryae, and J. appendiculata. Juglanconidaceae is morphologically and phylogenetically distinct from other families of Diaporthales. Species in this family are mostly pathogenic on Juglandaceae tree species causing black pustular dieback disease (Graves, 1923, Belisario, 1999). Du et al. (2017) introduced a new family Melansporellaceae for Juglanconis species and here we synonymise Melansporellaceae under Juglanconidaceae.
Juglanconis juglandina (Kunze) Voglmayr & Jaklitsch, Persoonia 38: 144. 2017. Facesoffungi number FoF03490.
Illustration: See Voglmayr et al. (2017).
Saprobic on dead twigs and branches of Juglandaceae. Sexual morph: Pseudostromata 0.8–2 mm diam, typically inconspicuous, sometimes distinct, circular, slightly projecting, without perithecial bumps. Ectostromatic disc 0.5–1.2 mm diam, indistinct, circular or oblong, dark grey, brown or black, often covered by densely arranged ostioles, pulvinate. Central column yellowish to brownish grey. Entostroma indistinct. Ascomata 440–565 μm diam, perithecial, aggregated, immersed, globose to subglobose, coriaceous, black, arrange in various configurations. Asci 140–160 × 17–22 μm, 8-spored, unitunicate, clavate to fusoid, indistinct apical ring, with small narrow stalk. Ascospores 25–30 × 8–11 μm, uni- to irregularly biseriate, hyaline, inequilaterally ellipsoid or broadly fusoid, asymmetric, distinctly constricted at the septum, without appendages, upper cell mostly larger, with rounded to subacute end, lower cell subacute to narrowly rounded, multiguttulate, containing mostly one large and numerous small guttules per cell. Asexual morph: Conidiomata acervular, 1–4 mm diam, black, scattered or occasionally confluent, with central or eccentric stromatic column, at maturity covered by black discharged conidial masses, usually conspicuous. Conidiophores 25–35 μm high, 5–6.5 μm wide, cylindrical to lageniform, simple, rarely branched at the base, smooth, subhyaline to pale brown. Conidiogenous cells annellidic with distinct annellations, integrated. Conidia 20–25 × 12–15 μm, unicellular, hyaline when immature, brown to blackish when mature, broadly ellipsoid to broadly pip-shaped, truncate with distinct scar at the base, multiguttulate, thick-walled, wall ornamented on the inside of the wall with irregular confluent verrucae and with gelatinous sheath.
Notes: Voglmayr et al. (2017) neotypified Melanconium juglandinum based on a freshly collected specimen due to misplacement or loss of the type specimen and poor condition of other authentic specimens. The conidiomata, conidiophores and conidia was nicely illustrated by Corda (1839) and the asexual morph is very common and conspicuous, while the sexual morph is infrequently found in fully-developed condition.
Lamproconiaceae C. Norphanphoun, T.C. Wen & K.D. Hyde, Phytotaxa 270: 94. 2016. Facesoffungi number FoF03491. Clade 22.
Pathogenic and saprobic on dead herbaceous branches. Sexual morph: Stromata prosenchymatous around perithecia, delimited externally by greenish, blackened, dense pseudoparenchymatous zone, interior whitish, composed of interwoven hyphae mixed with substrate cells, 3–5 ascomata in a stroma. Ascomata perithecial, small, aggregated, scattered, globose to subglobose, pale to dark brown, coriaceous, ostiolate, papillate. Papilla converging and erumpent through stroma surface as single, large opening. Peridium comprises pale brown, compressed, cells of textura angularis. Asci 8-spored, unitunicate, cylindrical, short-stalked, J- apical apparatus. Ascospores uniseriate, broadly ellipsoid, 1-septate, not or slightly constricted at the septa, hyaline, smooth. Asexual morph: Conidiomata pycnidial, solitary, partly immersed in host tissue, uni- to multilocular or convoluted, dark blue or dark blackish brown, erumpent in the centre. Pycnidium thick-walled, thin at inner layer, hyaline or dark brown, comprising wall cells of textura angularis or textura intricata. Ostiole absent, dehiscence irregular. Paraphyses interspersed within conidiophores. Conidiophores filiform or cylindrical, pale-bluish or hyaline, septate, branched, smooth-walled, formed at the base of conidiomatal wall. Conidiogenous cells holoblastic, cylindrical to subcylindrical, each forming a single conidium at the apex, or annellidic, colourless to olivaceous, smooth-walled. Conidia fusiform, ellipsoid, thick-walled, contents granular, aseptate, bluish to glistening dark blue or hyaline, smooth-walled, produced in mucilage but without a distinct mucilaginous envelope or appendage.
Type: Lamproconium (Grove) Grove.
Type species: Lamproconium desmazieri (Berk. & Broome) Grove.
Lamproconium desmazieri (Berk. & Broome) Grove [as ‘desmazieri’], British Stem- and Leaf-Fungi (Coelomycetes) (Cambridge) 2: 321. 1937. Facesoffungi number FoF03492. Fig. 28.
Lamproconium desmazieresi (MFLU 14–0780).A–C. Conidiomata on host. D. Vertical section of conidioma. E. Peridium and raised host. F. Apex of conidioma. G, H. Conidiogenous cells with attached conidia. I–M. Immature conidia. N. Mature conidium. Scale bars: A = 2 mm, B = 1 mm, C = 500 µm, D = 300 µm, E, F = 200 µm, G, H = 40 µm, I–N = 20 µm.
Pathogenic and saprobic on dead twigs and branches of lime trees (Tilia sp.). Sexual morph: Undetermined. Asexual morph: Conidiomata 0.8–1 × 0.4–0.55 mm, pycnidial, solitary, partly immersed in host tissue, uniloculate, dark blue, with a raised centre. Pycnidium 50–70 μm, with multi-layered wall, thin at inner layer, hyaline, wall cells of textura angularis. Paraphyses interspersed with conidiophores. Conidiophores 30–120 μm high, arising from the outermost wall layer at the basal of pycnidium, filiform or cylindrical, pale-bluish to hyaline, septate, branched, smooth-walled. Conidiogenous cells cylindrical to sub-cylindrical, annellidic, with flared periclinal thickenings in the collarette zone, colourless to olivaceous, smooth-walled. Conidia 22–28 × 8–10 μm ( = 25.25 × 9 μm, n = 30), fusiform, ellipsoid, infrequently slightly curved, aseptate, initially hyaline, bluish to glistening dark blue at maturity, narrowly rounded at ends, smooth-walled.
Material examined: Russia, Rostov region, Krasnosulinsky district, Donskoye forestry, artificial forest, on dead branches of Tilia cordata (Tiliaceae), 21 May 2014, T. Bulgakov, MFLU 14–0780.
Notes: Melanconium desmazieri was reported as the asexual morph of Melanconis desmazieri from Tilia species (Petrak 1938). Grove (1937) re-circumscribed the species of Melanconium and postulated that Melanconium desmazieri differed from the type species of Melanconium in having 1-septate, bluish to glistening dark blue conidia. Therefore, Grove introduced a new genus Lamproconium to accommodate this taxon (Grove, 1937, Sutton, 1980), and Lamproconium desmazieri was placed in Diaporthales genera incertae sedis by Cannon & Minter (2014). Based on phylogenetic study, Norphanphoun et al. (2016) synonymised Melanconis desmazieri under Lamproconium desmazieri and introduced a new family Lamproconiaceae to accommodate Lamproconium and Hercospora. Morphologically Lamproconiaceae is distinct from other families of Diaporthales in having dark blue or dark blackish brown pycnidial conidiomata and fusiform to ellipsoid, aseptate, bluish to glistening dark blue or hyaline conidia. The sexual morph is reported only for Hercospora. Combined gene analysis of LSU nrDNA, ITS nrDNA, rpb2, and tef1 shows that Lamproconiaceae is a distinct family that is sister to Sydowiellaceae (Fig. 1, Clade 22).
Macrohilaceae Crous, IMA Fungus 6: 180. 2015. Clade 15.
Pathogenic forming leaf spots. Sexual morph: Undetermined. Asexual morph: Coelomycetous. Conidiomata pycnidial, immersed, becoming erumpent, medium brown, globose. Conidiophore reduced to conidiogenous cells. Conidiogenous cells lining the inner cavity, pale brown, cylindrical, proliferating percurrently near the apex. Conidia solitary, medium to dark brown, ovoid, smooth, guttulate, medianly septate, apex obtuse, base truncate with a visible scar.
Type genus: Macrohilum H.J. Swart.
Type species: Macrohilum eucalypti H.J. Swart.
Notes: The family Macrohilaceae was introduced and typified by Macrohilum (Crous et al. 2015) and its taxonomic placement in Diaporthales has been reported based on LSU nrDNA sequence data. In this study, our concatenated analysis on LSU nrDNA, ITS nrDNA, rpb2 and tef1 also indicates that the Macrohilum eucalypti strains cluster together with high support and belong to the Macrohilaceae (Fig. 1, Clade 15). Macrohilaceae differs from other families of Diaporthales in having single, dark brown, guttulate, thick-walled, medianly septate, oval conidia with obtuse apex and truncate base (Crous et al. 2015). This monotypic family comprises only a single species commonly associated with leaf spots of Eucalyptus.
Macrohilum eucalypti H.J. Swart, Trans. Br. mycol. Soc. 90: 288. 1988. Facesoffungi number FoF03493.
Illustration: See Crous et al. (2015).
Pathogenic forming leaf spots. Sexual morph: Undetermined. Asexual morph: Conidiomata immersed, becoming erumpent, medium brown, globose, to 300 μm diam. Conidiogenous cells lining the inner cavity, pale brown, cylindrical, finely roughened, proliferating percurrently near the apex, 10–15 × 3–5 μm. Conidia solitary, medium to dark brown, ovoid, smooth, guttulate, developing a single, dark brown, supra-median septum, thick-walled, frequently constricted at the septum, apex obtuse, base truncate and protruding, with a visible scar, 2–3 μm wide, 15–20 × 10–12 μm (description based on Crous et al. 2015).
Notes: Crous et al. (2015) epitypified Macrohilum eucalypti using an Australian specimen collected from Eucalyptus piperita. Although a New Zealand isolate (CPC 10945) differed from the Australian ex-epitype isolate (CPC 19421) by four base pairs in the ITS nrDNA, Crous et al. (2015) did not propose this isolate as a new species pending collection of more material.
Melanconidaceae G. Winter [as ‘Melanconideae’], Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1.2: 764. 1886. Clade 2.
Saprobic or pathogenic on plants. Sexual morph: Pseudostromata well-developed, obvious, erumpent. Ectostromatic disc surrounded by bark or not, yellowish white, ostiolar canal opening around the disc. Ascomata perithecial, arranged as circles around the ectostromatic disc, oblique or horizontal, globose to subglobose, coriaceous, black, with long, periphysate, lateral ostiolar canals. Peridium comprising outer, thick-walled, brown cells of textura globosa to textura angularis and inner, thick-walled, flat, hyaline cells of textura angularis. Hamathecium comprising wide, hyphae-like, paraphyses, deliquescent at maturity. Asci 8-spored, unitunicate, oblong to fusiform, short pedicellate, with distinct, J- apical ring. Ascospores overlapping uni- to biseriate, hyaline, ellipsoid, 1-septate. Asexual morph: coelomycetous, melanconium-like. Conidiomata acervular, scattered, solitary, superficial, black, coriaceous. Conidiophores branched at the base, septate, Conidiogenous cells annellidic, cylindrical. Conidia hyaline to brown, ellipsoid or subglobose, smooth-walled, thick-walled.
Type genus: Melanconis Tul. & C. Tul.
Type species: Melanconis stilbostoma (Fr.) Tul. & C. Tul.
Notes: The family Melanconidaceae was introduced by Winter (1886) to accommodate species having yellowish-white ectostromatic discs surrounding ascomata arranged in a circle. Members of this family are plant pathogens causing disease on economic plant species, as well as saprobes. Maharachchikumbura et al. (2016) listed 24 genera under this family based on morphology, following Lumbsch & Huhndorf (2010). However, most genera do not have any DNA sequence data, except Dicarpella, Melanconiella, Melanconis, Melanconium, and Prosthecium. Voglmayr & Jaklitsch (2014) synonymised Prosthecium under Stilbospora and included it in Stilbosporaceae. Crous et al. (2012b) have linked more than half of known wuestneia-like species to Harknessia species accommodating it in Harknessiaceae. Based on morphological and phylogenetic evidence, Castlebury et al. (2002) and Rossman et al. (2007) reported that this family is monogeneric with Melanconis and its asexual morph Melanconium. However, Rossman et al. (2015) synonymised Melanconium under Melanconis based on the poor phylogenetic resolution of Melanconium species and poor host-specificity. Phylogenies generated in this study position Dicarpella and Melanconiella (both in Melanconiellaceae; Fig. 1, Clade 10) outside the Melanconidaceae (Fig. 1, Clade 2). Considering the lack of molecular data, diverse ecological strategies and variable morphology, the family Melanconidaceae is restricted to Melanconis sensu stricto. Hence, we exclude all genera listed in Maharachchikumbura et al. (2016) from this family except Melanconis.
Melanconis apiocarpum and M. marginale have been reported from leaf spots of Alnus species in Canada, England, and Switzerland (Sieber et al. 1991). Melanconium juglandinum causes black pustular dieback of Juglans species in Europe and was consistently isolated from diseased twigs and branches of Persian walnut trees (Juglans regia), proving to be a virulent pathogen (Belisario 1999).
Melanconis italica Senan., Camporesi & K.D. Hyde, sp. nov. MycoBank MB821560. Facesoffungi number FoF03494. Fig. 29, Fig. 30.
Melanconis italica (MFLU 17–0879). A. Stromata on substrate. B–D. Horizontal cross section of stroma. E. Vertical section of ascoma. F. Peridium. G. Paraphyses. H–K. Asci. L–O. Ascospores. Scale bars: A = 500 µm, B-D = 200 µm, E = 100 µm, F = 50 µm, G–O = 10 µm.
Asexual morph of Melanconis italica (MFLUCC 16–1199).A, B. Conidiomata on PDA. C–F. Conidia attached to conidiogenous cells, and conidiophores. G. Conidia. Scale bars: A = 500 µm, B = 200 µm, C–F = 10 µm, G = 5 µm.
Etymology: Species epithet based on the country where the fungus was collected, Italy.
Saprobic on Alnus cordata. Sexual morph: Pseudostromata poorly-developed, erumpent. Ectostromatic disc 500–600 μm diam, surrounded by bark or not, yellowish-white, causing a coarse bark surface, inverted conical, ostioles open into margin and rarely middle of the disc. Ascomata 0.90–1 mm high, 0.4–0.5 mm diam ( = 0.98 × 0.47 mm, n = 20), perithecial, oblique, globose to subglobose, coriaceous, black, with long periphysate, lateral ostiolar canal. Hamathecium comprising wide, hyphae-like, hyaline, septate 4–12 μm wide ( = 9 μm, n = 20) paraphyses. Peridium 15–28 μm diam ( = 21 μm, n = 10), comprising thick-walled, brown cells of textura angularis. Asci 80–92 × 11–14 μm ( = 13 × 8.5 μm, n = 20), 8-spored, unitunicate, cylindrical, short pedicellate, with distinct, J- apical ring. Ascospores 18–21 × 2.8–4 μm ( = 19 × 3.5 μm), biseriate, hyaline, fusiform, 1-septate, slightly constricted at the septum, smooth-walled. Asexual morph: Conidiomata on MEA solitary, superficial, globose, appears as slimy bubbles of conidia mass, black. Conidiophores cylindrical, branched, thick-walled, hyaline. Conidiogenous cells blastic, terminal or intercalary, bottle-shaped, narrowing towards the apex, hyaline, thick-walled. Conidia fusiform to ellipsoidal, aseptate, thick-walled, basal end pointed, apical end blunt, olivaceous.
Culture characteristics: Colonies growing on MEA attained 1 cm within 7 d incubation at 18 °C, flat, circular, smooth margin, white, tightly attached to the substrate, little aerial mycelia.
Specimen examined: Italy, Province of Forlì-Cesena, Fiumicello di Premilcuore, on dead branch of Alnus cordata (Betulaceae), 4 Dec. 2013, E. Camporesi, IT 1557 (holotype MFLU 17–0879, isotype BBH 42441, cultures ex-type, MFLUCC 16–1199, MFLUCC 17–1659).
Notes: Melanconis italica clusters in a clade with M. alni with high support (Fig. 1, Clade 2). Both M. italica and M. alni are associated with Alnus species. Melanconis alnicola is also reported from Alnus species. However, there are no DNA sequence data in GenBank for Melanconis alnicola. Morphologically, M. alni differs from M. italica in having short to long, hyaline, filiform appendages at both ends, and oval to ellipsoid ascospores. In contrast, M. alnicola has large, oval to ellipsoid ascospores (25–45 × 9–12 μm) and small asci (50–60 × 10–15 μm). Hence, a new species, Melanconis italica, is introduced to accommodate this taxon.
Melanconiellaceae Senan. & Maharachch., K.D. Hyde, fam. nov. MycoBank MB821561. Facesoffungi number FoF03495. Clade 10.
Synonym: Melanconiellaceae Locq., Mycol. gén. struct. (Paris): 210. 1984. (nom. inval., Art 39.1, Melbourne Code).
Phytopathogenic or saprobic. Sexual morph: Stromata present or absent. If present; Pseudostromata inconspicuous, erumpent, pale or dark coloured ectostromatic disc or pulvillus causing a more or less pustulate bark surface. Ectostromatic disc convex, flat to concave, surrounded by bark or not. Central column beneath the disc more or less conical, comprising hyaline or pigmented hyphae mixed with a pigmented, cream, yellow, olive, brownish or grey, powdery amorphous substance. Perithecia sometimes epiphyllous without stromatic tissues and immersed in host substrate, inconspicuous or appearing as rounded bumps beneath the bark surrounding the ectostromatic disc, oblique or horizontal, scattered or often arranged in a circle around the central column, with long lateral ostioles that converge at the margin of the central column. Ostioles emerging in various positions in the ectostromatic disc, cylindrical. Peridium comprising dark, thick-walled cells of textura angularis. Hamathecium aparaphysate or comprising broad, hyaline paraphyses. Asci 2–8-spored, unitunicate, cylindrical-clavate, oblong or fusoid, with a J- distinct apical ring, tapering below to a short, narrow pedicel. Ascospores hyaline, yellowish or brown, oblong, fusoid or ellipsoid, 0–1-septate, septa central or slightly eccentric, slightly constricted or not, smooth, with or without short, blunt appendages and sometimes with a narrow gelatinous sheath. Asexual morph: Conidiomata acervular or pycnidia, punctiform, subcuticular, immersed or erumpent, sometimes with a central, well-developed, pale brown, pseudoparenchymatous layer becoming thinner or absent at the margin of the conidiomata, multiloculate, sometimes papillate, sometimes with pale coloured, ectostromatic disc and central column or with radiate scutella. Scutella convex, membranous, brown, somewhat translucent, with a central hyaline or pale disc, giving rise to radiating hyphae, thick-walled cells radiating from a central point, rounded to pointed at the tips. Conidiophores reduced to conidiogenous cells or branched, sometimes septate only at the base, few-celled, smooth, hyaline to pale brown, sometimes short, forming under the developing scutellum. Pseudoparaphyses filiform. Conidiogenous cells annellidic or phialidic. Conidia initially hyaline becoming brown, ellipsoid, obovoid, subglobose, ovoid or oblong, thick-walled, smooth to finely verrucose, with or without distinct hyaline sheath, each with a truncate base and obtuse to bluntly pointed apex, sometimes somewhat granular, sometimes with inconspicuous to conspicuous basal hilum, with or without distinct hyaline sheath or frill.
Type genus: Melanconiella Sacc.
Type species: Melanconiella spodiaea (Tul. & C. Tul.) Sacc.
Notes: The phylogenetic analyses of this study showed that Greeneria, Melanconiella, and Dicarpella (previously placed in Melanconidaceae), Tubakia (previously placed as Diaporthales incertae sedis), Sphaeronaemella fragariae (previously placed in Microascales incertae sedis) and Microascospora gen. nov. forms a distinct clade with moderate support, which we recognise as Melanconiellaceae (Fig. 1, Clade 10).
The genus Greeneria was introduced based on G. fuliginea (Scribner & Viala 1887) and was synonymised under Melanconium (Cavara 1889) as Melanconium fuligineum. Later van der Aa (1973) accommodated this genus in Phyllosticta as P. ampelicida. Punithalingam (1974) renamed this taxon as Greeneria uvicola providing a detailed description and illustration. A LSU nrDNA sequence analysis by Farr & Rossman (2001) showed the phylogenetic placement of Greeneria uvicola outside of Melanconidaceae, but within Diaporthales. Analyses in this current study showed the phylogenetic placement of G. uvicola and G. saprophytica within Melanconiellaceae (Fig. 1, Clade 10). However, G. saprophytica does not show a very close affinity to G. uvicola. Greeneria uvicola is one of the most common pathogens causing various diseases in grapes (Navarrete et al. 2009). Greeneria lacks a known sexual morph (Zhang & Blackwell 2001) and it differs from other diaporthalean asexual morphs in having holoblastic conidiogenesis, producing phialidic conidiogenous cells in acervuli or pycnidia, and pale brown conidia (Barr 1978). The fungus overwinters on stem lesions, mummified berries, leaves, and tendrils. It is known to attack several species of Vitis including V. aestivalis, V. labrusca, V. rotundifolia, and V. vinifera.
The genus Dicarpella is based on Dicarpella bina and the asexual morph of this genus was reported as Tubakia (Belisario 1991). Tubakia is typified by Tubakia japonica. The type species of these two genera are not linked to each other. However molecular data linked Tubakia and Diplacella together and a few Diplacella species have Tubakia asexual morphs (Sogonov et al. 2008). Tubakia is more commonly encountered compared with Dicarpella and it is also a more widely used name than Dicarpella. The phylogenetic analyses in this study indicate a plausible relationship of Dicarpella dryina and Tubakia seoraksanensis as a holomorphic genus (Fig. 1, Clade 10). However, it is hard to confirm that Dicarpella and Tubakia are congeneric without analysing sequence data of the type species. Thus, here we retain Dicarpella and Tubakia as two separate genera until sequence data becomes available.
Voglmayr et al. (2012) reviewed the genus Melanconiella based on herbarium material and recently collected specimens. The morphological and phylogenetic distinctness of Melanconiella from Melanconis was discussed. The generic type of Melanconiella was confirmed as M. spodiaea. Phylogenetic analysis in this study showed the distinct placement of Melanconiella within this new clade (Fig. 1, Clade 10).
A new genus Microascospora is introduced to this family based on Microascospora rubi. Phylogenetically Sphaeronaemella fragariae did not cluster with other Sphaeronaemella species and it forms a clade with Microascospora rubi. Hence Sphaeronaemella fragariae is excluded from Sphaeronaemella and accommodated in Microascospora as M. fragariae. However, Melanconiellaceae was originally invalidly published (Art. 39.1, Melbourne) by Locquin (1984). Hence Melanconiellaceae is herewith validated to accommodate Dicarpella, Greeneria, Melanconiella, Microascospora and Tubakia.
Melanconiella chrysodiscosporina Voglmayr & Jaklitsch, Fungal Diversity 57: 14. 2012. Facesoffungi number FoF03496. Fig. 31.
Melanconiella chrysodiscosporina (MFLU 17–0893). A. Herbarium specimen. B. Vertical cross section of ascoma. C. Peridium. D–F. Asci. G–J. Ascospores. K. Conidiomata on substrate. L. Vertical cross section of conidioma. M. Conidiophores, conidiogenous cells and conidia. N–Q. Conidia. Scale bars: A = 1 mm, B, L = 100 µm, C = 20 µm, D–J, M = 10 µm, K = 200 µm, N–Q = 5 µm.
Saprobic on dead branch of Fagus sylvatica. Sexual morph: Pseudostromata indistinct, irregular or circular outline. Ectostromatic disc minute, circular, narrowly fusoid to oblong, yellow or greyish brown, central column yellow. Entostroma comprising subhyaline to yellowish hyphae. Ascomata 1.2–1.3 mm high, 0.3–0.5 mm diam ( = 1.25 × 0.45 mm, n = 20), immersed, aggregated, globose to subglobose, coriaceous, black, arranged in valsoid configuration. Papilla 600–950 μm high, 90–130 μm diam ( = 800 × 117 μm, n = 10), long, asymmetrical or symmetrical, black, converging at upper region and make a common canal to open out, internally covered by hyaline periphyses. Peridium 14–17 μm diam ( = 15 μm, n = 10), comprising outer few layers of thick-walled, brown, compressed cells of textura angularis and inner thick-walled, hyaline, compressed cells of textura angularis. Asci 85–100 × 13–17 μm ( = 95 × 15 μm, n = 20), 8-spored, unitunicate, cylindrical to fusoid, with J- distinct apical ring, sessile or with short pedicel. Ascospores 17–20 × 6–9 μm ( = 17.6 × 7.5 μm, n = 20), uni- or biseriate, broadly ellipsoid, not constricted at the septum, ends broadly rounded, hyaline, medianly 1-septate, multi-guttulate with one large and numerous small guttules per cell, wall swelling and sometimes thicken and stuffed at the septum. Asexual morph: discosporina-like. Conidiomata 140–180 μm high, 490–600 μm diam ( = 150 × 507 μm, n = 20), visible as darker spots marginated by a distinct dark brown to blackish marginal zone, with a central stromatic column, at maturity covered by whitish discharged conidial masses. Conidiophores 6–10 × 5–7.5 μm ( = 7.4 × 6 μm, n = 20), few layers, cubic, thick-walled, hyaline. Conidiogenous cells 9–12 × 1–2 μm ( = 11 × 1.8 μm, n = 20), phialidic, conical, base enlarged, narrowing towards the apex, thick-walled, hyaline. Conidia 10–12 × 4–6 μm ( = 10.8 × 5.2 μm, n = 20), ellipsoid, oblong or cylindrical, with two large and numerous small guttules, hyaline, with gelatinous sheath.
Specimen examined: Italy, Province of Forlì-Cesena, Bagno di Romagna, near Riofreddo, on dead branches of Fagus sylvatica (Fagaceae), 14 Aug. 2016, E. Camporesi, IT 3066, MFLU 17–0893, living culture MFLUCC 17–1671.
Notes: Melanconiella chrysodiscosporina was introduced by Voglmayr et al. (2012). This fungus was mostly found in the summer season from late spring to autumn (Voglmayr et al. 2012). The holotype and other authentic specimens were collected from dead branches of Carpinus betulus (Betulaceae). However, we collected this specimen from dead branches of Fagus sylvatica (Fagaceae). This is the first host record of Melanconiella chrysodiscosporina on Fagus sylvatica.
Melanconiella chrysomelanconium Voglmayr & Jaklitsch, Fungal Diversity 57: 16. 2012. Facesoffungi number FoF03497. Fig. 32.
Melanconiella chrysomelanconium (MFLU 17–0966).A, B. Conidiomata on substrate. C. Cross section of conidioma. D–F. Conidiophores, conidiogenous cells and attached conidia. G–J. Conidia. Scale bars: B = 200 µm, C = 500 µm, D–F = 20 µm, G–J = 10 µm.
Saprobic on branches of Carpinus betulus. Sexual morph: Not observed. Asexual morph: melanconium-like. Conidiomata 0.4–1 mm diam, visible as blackish spots with central or eccentric ostiolar opening, pycnidial, epidermal to subepidermal, globose to subglobose, black, coriaceous, at maturity covered by black discharged conidial masses. Ostiole present, pointed. Conidiomatal wall composed of thin-walled, brown cells of textura angularis. Conidiophores reduced to conidiogenous cells, arising from the uppermost layer of cells of the basal stromatic pycnidial wall. Conidiogenous cells 7–18 μm high, 2–6 μm diam ( = 14.4 × 4 μm, n = 20), annellidic, hyaline, cylindrical, thick-walled, determinate, integrated, with flared collarette and periclinal wall-thickening. Conidia 13–20 × 7–11 μm ( = 15 × 8.5 μm, n = 20), dark brown, broadly ellipsoid to globose, circular in outline, slightly truncate at base, aseptate, multiguttulate with 1–2 large and numerous small guttules, thick-walled, with distinct gelatinous sheath, smooth.
Specimen examined: Italy, Province of Forlì-Cesena, Via Nenni, Forlì, on dead aerial branches of Carpinus betulus (Betulaceae), 2 Jan. 2015, E. Camporesi, IT 1622, MFLU 17-0966.
Notes: Melanconiella chrysomelanconium is morphologically similar and phylogenetically related to M. chrysodiscosporina (Voglmayr et al. 2012). The combined gene analysis of this study illustrates its phylogenetic relationship to other Melanconiella species (Fig. 1, Clade 10).
Microascospora Senan. & K.D. Hyde, gen. nov. MycoBank MB821562. Facesoffungi number FoF03498.
Etymology: Name based on small ascospores (<20 μm in length).
Saprobic on dead stems. Sexual morph: Ascomata scattered, solitary, immersed, globose to subglobose, brown, coriaceous, papillate, ostiolate. Papilla narrow, long, straight or curved, comprising thick-walled, brown, compressed cells of textura angularis, internally covered by hyaline periphyses. Peridium comprising thick-walled, brown, somewhat compressed cells of textura angularis. Hamathecium aparaphysate. Asci 8-spored, unitunicate, clavate to fusiform, J- apical ring, attached to base without a pedicel. Ascospores overlapping biseriate, ellipsoid to fusiform, hyaline, aseptate, with two large fat globules at ends, appendages long, filiform to wavy, hyaline. Asexual morph: Undetermined.
Type species: Microascospora rubi Senan., Maharachch. & K.D. Hyde.
Microascospora rubi Senan., Camporesi & K.D. Hyde, sp. nov. MycoBank MB821563. Facesoffungi number FoF03499. Fig. 33.
Microascospora rubi (MFLU 15–1112). A. Herbarium specimen. B. Vertical section of ascoma. C. Peridium. D–F. Asci. G–J. Ascospores. K. Upper side of culture growing on MEA. L. Lower side of culture growing on MEA. Scale bars: A = 200 µm, B = 100 µm, C = 25 µm, D–F = 10 µm, G–J = 5 µm.
Etymology: Name based on host genus Rubus.
Saprobic on dead stems of Rubus ulmifolia. Sexual morph: Ascomata 250–290 × 205–255 μm ( = 269 × 230 μm), scattered, solitary, immersed, globose to subglobose, brown, coriaceous, papillate, ostiolate. Papilla 115–155 μm high, 55–67 μm diam ( = 139 × 65 μm, n = 10), narrow, long, straight or curved, comprising thick-walled, brown, compressed cells of textura angularis, internally covered by hyaline periphyses. Peridium 10–25 μm ( = 18 μm), comprising thick-walled, brown, large, somewhat compressed cells of textura angularis. Hamathecium aparaphysate. Asci 68–70 × 15–18 μm ( = 69 × 16 μm, n = 20), 8-spored, unitunicate, clavate to fusiform, J- apical ring, attached to base without a pedicel. Ascospores 14–19 × 5–7 μm ( = 17 × 6 μm, n = 20), overlapping biseriate, ellipsoid to fusiform, hyaline, aseptate, with two large fat globules at ends, appendages long, filiform to wavy, hyaline. Asexual morph: Undetermined.
Culture characteristics: Colonies growing on MEA slow growing, becoming 1 cm within 10 d at 18 °C, circular, umbonate, irregular margin, cream to olivaceous, cotton-like, loosely attached to the substrate.
Specimen(s) examined: Italy, Province of Forlì-Cesena, Bagno di Romagna, Ridracoli, on dead branch of Rubus ulmifolia (Rosaceae), 24 Jan. 2014, E. Camporesi, IT 1675 (holotype MFLU 15–1112, isotype BBH 42445).
Notes: A new genus Microascospora is introduced based on M. rubi. This genus is morphologically and phylogenetically distinct from other genera in Melanconiellaceae having small ascospores (<20 μm length) with wavy, filiform long appendages, and immersed, solitary ascomata with wavy papilla.
Microascospora fragariae (F. Stevens & Peterson) Senan., Maharachch. & K.D. Hyde, comb. nov. MycoBank MB821631. Facesoffungi number FoF03500.
Basionym: Sphaeronaemella fragariae F. Stevens & Peterson, Phytopathology 6: 258. 1916.
Notes: The multi-gene sequence analysis in this study shows that Sphaeronaemella fragariae (Fig. 1, Clade 10) does not have any affinities to the type species of Sphaeronaemella, S. helvellae (incertae sedis in Microascales). Sphaeronaemella fragariae forms a well-supported clade that is sister to Microascospora rubi in Melanconiellaceae. Hence, we exclude Sphaeronaemella fragariae from Sphaeronaemella and accommodate this taxon in Microascospora and propose a new combination as Microascospora fragariae.
Tubakia thailandensis Senan., Tangthir. & K.D. Hyde, sp. nov. MycoBank MB821564. Facesoffungi number FoF03501. Fig. 34.
Tubakia thailandensis (MFLU 13–0260).A. Herbarium specimen. B. Conidiomata on the host surface. C. Vertical section of pycnothyrium. D. Top view of radiate scutellum and conidiogenous cells with developing conidia. E–H. Conidiogenous cells with developing conidia stained with lactophenol cotton blue. I–K. Conidia. L. Conidia stained with lactophenol cotton blue. M. Germinating conidium. N–O. Colonies on PDA from top. P–Q. Colonies on PDA from reverse. Scale bar: C = 50 μm, D, F–M = 10 μm, E = 5 μm.
Etymology: Name based on the country from which this species was collected, Thailand.
Saprobic on dead leaves. Conidiomata 40–50 μm high, 50–75 μm diam, pycnothyria with radiate scutella, scattered to gregarious, superficial on the substratum. Scutella convex, brown to dark brown, thick-walled cells, radiating from a central point. Conidiophores short, forming under the developing scutella. Conidiogenous cells 5–10 × 2–4 μm, phialidic, with a minute collarette and wide periclinal thickening. Conidia 10–12.4 × 7.4–8.7 μm ( = 11.3 × 8.1 μm, n = 20), globose to subglobose, smooth, hyaline, thick-walled.
Specimen examined: Thailand, Chiang Rai, Doi Mae Salong, on dead leaf, 2 May 2012, K. Wisitrassameewong, NTCL059 (holotype MFLU 13–0260, culture ex-type MFLUCC 12–0303).
Culture characteristics: Mycelium white when young, dark green, pale grey to black from above and reverse when aged, with medium mycelium, flat, rhizoid to irregular form, labate margin, and attaining a diam of 46 mm on PDA in 7 d at 27 °C.
Notes: Tubakia comprises seven species (Index Fungorum, 2017, MycoBank, 2017). Braun et al. (2014) presented a taxonomic key to the genus Tubakia and according to that key, this species is morphologically quite similar to “Tubakia sp.” which has a small scutellum (40–80 μm diam.) and hyaline or subhyaline conidia (9–11 × 7–9 μm) collected from Castanea henryi in China. Therefore, we introduce this species as Tubakia thailandensis. Tubakia thailandensis differs from other Tubakia species in having small (length < 15 μm), globose or subglobose, hyaline conidia, without microconidial development. Tubakia shares close phylogenetic affinities to Greeneria saprophytica (Fig. 1, Clade 10).
Prosopidicolaceae Senan. & K.D. Hyde, fam. nov. MycoBank MB821565. Facesoffungi number FoF03502. Clade 17.
Pathogenic on species of Fabaceae. Conidiomata pycnidial, rarely acervular, solitary or aggregated in a eustromatic stroma with one to several ostioles or astromatic, grey to black, erumpent to immersed. Peridium comprising grey-brown cells of textura angularis. Conidiophores reduced to conidiogenous cells or lining the whole inner layer of the wall, subcylindrical, branched, septate, straight to irregularly curved, base pale brown, becoming medium green-brown at apex. Conidiogenous cells mono- to polyphialidic, tightly aggregated, hyaline, smooth, ampulliform, subcylindrical to lageniform, prominent periclinal thickening, at times with percurrent proliferation. Conidia solitary, subhyaline to brown, smooth, guttulate, straight to variously curved, ellipsoid to fusoid-ellipsoid, apex obtuse, base truncate to bluntly round.
Type genus: Prosopidicola Crous & C.L. Lennox.
Type species: Prosopidicola mexicana Crous & C.L. Lennox.
Notes: Prosopidicolaceae is a monotypic family introduced here to accommodate Prosopidicola species. Prosopidicolaceae (Fig. 1, Clade 17) is phylogenetically not associated with any support to known families of Diaporthales, but is morphologically well-delineated. Species in this family are pathogens on Fabaceae host plants. This family comprises Prosopidicola albizziae and P. mexicana (Lennox et al., 2004, Crous et al., 2016).
Prosopidicola mexicana Crous & C.L. Lennox, Stud. Mycol. 50: 191. 2004. Facesoffungi number FoF03503.
Illustration: See Lennox et al. (2004).
Pathogenic causing pod rot disease on Prosopidis glandulosae. Lesions 2–3 mm wide and up to 7 mm long, covering the pod, irregular, extending across the width of the pod, pale brown with a raised, dark brown margin. Conidiomata up to 250 μm diam, amphigenous, pycnidial, rarely acervular, scattered, immersed to erumpent, globose to subglobose, unilocular, black. Peridium up to 15 μm thick, consisting of 3–4 layers of brown cells of textura angularis. Conidiophores 5–50 μm high, 3–4 μm diam, lining the whole inner layer of the wall, subcylindrical, branched, 0–3-septate, straight to irregularly curved, base pale brown, becoming medium greenbrown at apex. Conidiogenous cells 5–16 × 3–4 μm, phialidic when young, with prominent periclinal thickening and proliferating percurrently with age, subcylindrical to lageniform, green-brown, smooth when young, becoming medium to dark green-brown and warty with maturity, apex obtaining flared collarettes, rarely with two loci per conidiogenous cell. Conidia 10–15 × 4.5–5.5 μm, solitary, broadly ellipsoidal, medium brown, straight to slightly curved, rounded at the apex, tapering to a subtruncate base, with an inconspicuous dehiscence scar, smooth, thin-walled, aseptate (description based on Lennox et al. 2004).
Notes: Prosopidicola mexicana is the cause of a severe pod rot disease on Prosopidis glandulosa. It appears as black lesions surrounded by a dark brown margin. Lennox et al. (2004) revealed it to group closely to Cryphonectriaceae. However in the phylogenetic analyses generated in this study, it forms a distinct clade which is basal to Cytosporaceae.
Pseudoplagiostomataceae Cheew. et al., as “Pseudoplagiostomaceae”, Fungal Diversity 44: 95. 2010. Clade 12.
Pathogenic on leaves, forming spots. Sexual morph: Ascomata solitary, scattered, immersed, slanted to horizontal on host tissue, globose or elliptical, black, coriaceous, papillate, ostiolate. Papilla short, internally covered with hyaline, filamentous periphyses. Peridium comprising a few layers of thick-walled, brown cells of textura angularis. Hamathecium lacking paraphyses. Asci 8-spored, unitunicate, cylindrical, sessile, with J-, subapical ring. Ascospores overlapping uni- to biseriate, hyaline, fusiform to ellipsoid, 1-septate, with terminal, elongate, hyaline appendages. Asexual morph: Coelomycetous. Conidiomata acervular or pycnidial, brown. Peridium comprising small, brown cells of textura angularis. Conidiophores absent. Conidiogenous cells cylindrical to ampulliform, enteroblastic, percurrently proliferating with periclinal thickening and collarette. Conidia holoblastic, hyaline to brown, ellipsoid, unicellular, subglobose to broadly allantoid, with obtuse apex and a flat protruding scar at the base.
Type genus: Pseudoplagiostoma Cheew. M.J. Wingf. & Crous.
Type species: Pseudoplagiostoma eucalypti Cheew., M.J. Wingf. & Crous.
Notes: Pseudoplagiostomaceae was introduced by Cheewangkoon et al. (2010). Pseudoplagiostomaceae is similar to Gnomoniaceae (Fig. 1, Clade 1) based on morphological characters of its sexual morph, such as solitary, immersed, non stromatic ascomata with lateral beaks, asci with a distinct apical ring and 1-septate ascospores (Sogonov et al. 2008). However, in our phylogenetic analyses it formed a fully-supported clade (Fig. 1, Clade 12) sister to Apoharknessiaceae.
Pseudoplagiostoma eucalypti Cheew., M.J. Wingf. & Crous, Fungal Diversity 44: 98. 2010. Facesoffungi number FoF03504.
Illustration: See Cheewangkoon et al. (2010).
Pathogenic on leaves forming leaf spots. Sexual morph: Ascomata 130–150 μm high, 100–130 μm diam, perithecia, immersed in host tissue, slanted to horizontal, globose to elliptical, coriaceous, brown to black, papillate, ostiolate. Papilla 60–65 μm diam, erumpent, internal wall lined by hyaline periphyses. Peridium comprising few layers of thick-walled, brown cells of textura angularis. Hamathecium aparaphysate. Asci 65–70 × 11–13 μm, 8-spored, unitunicate, subcylindrical to long obovoid, with wedge-shaped, J- subapical ring, apex blunt and without a distinct pedicel. Ascospores 17–19 × 5–7 μm, overlapping uni- to biseriate, ellipsoid, tapering towards rounded ends, hyaline, median 1-septate, widest at septum, with terminal, elongate, hyaline appendages. Asexual morph: Conidiomata 180–200 μm high, 170–190 μm diam, acervular to pycnidial, subcutical to epidermal. Peridium comprising small, brown cells of textura angularis. Conidiophores absent. Conidiogenous cells 8–12 × 2–4 μm, cylindrical to ampulliform, enteroblastic proliferation with periclinal thickening. Conidia 17–19 × 7–8 μm, holoblastic, ellipsoid, unicellular, with obtuse apex and a flat protruding scar at the base (description based on Cheewangkoon et al. 2010).
Notes: The monotypic family Pseudoplagiostomataceae was introduced by Cheewangkoon et al. (2010) to accommodate a cryptosporiopsis-like fungus isolated from Eucalyptus. The type species, P. eucalypti (as Cryptosporiopsis eucalypti) and two other new species, P. oldii and P. variabile, were isolated as foliar pathogens from Eucalyptus leaf spots. However, Cryptosporiopsis eucalypti was not closely related to the generic type of Cryptosporiopsis, C. nigra (Dermateaceae, Helotiales), and hence Cheewangkoon et al. (2010) introduced Pseudoplagiostoma to accommodate this taxon. Pseudoplagiostoma corymbiae (Crous et al. 2012c) and P. dipterocarpi (Suwannarach et al. 2016) were introduced to this genus from Corymbia sp. and Dipterocarpus tuberculatus respectively. The ascospore morphology, in particular, is distinct and morphologically this family differs from other families in the order in having astromatic, slanted to horizontal, globose ascomata with aparaphysate hamathecium, ascospores with terminal, elongate, hyaline appendages and a cryptosporiopsis-like asexual morph. Cheewangkoon et al. (2010) analysed LSU nrDNA sequence data of the order Diaporthales, to show the distinct placement of Pseudoplagiostomaceae with 100 % bootstrap support.
Schizoparmaceae Rossman, Mycoscience 48: 137. 2007. Clade 8.
Saprobic, parasitic or pathogenic on woody, herbaceous plants. Sexual morph: Ascomata perithecial, solitary, scattered, subepidermal, immersed to erumpent, becoming superficial, globose, coriaceous, brown to black, short papillate, ostiole with hyaline periphyses, plate-like ornamentation around ostiole. Peridium comprising thick-walled, brown-cells of textura angularis. Hamathecium aparaphysate. Asci 8-spored, unitunicate, ellipsoid to fusiform, sessile, with a J- apical ring. Ascospores biseriate, hyaline to becoming pale brown at maturity, ellipsoidal, aseptate, with or without mucoid caps. Asexual morph: Coelomycetous. Conidiomata pycnidial, subepidermal, immersed to erumpent, unilocular, globose, slightly depressed globose to subglobose. Conidiophores densely aggregated, slender, subulate, simple or branched, hyaline, smooth, occasionally septate and branched at base, invested in mucus, developing from basal pad. Conidiogenous cells discrete, simple, subcylindrical, obclavate or lageniform, smooth, proliferating percurrently or with prominent periclinal thickening. Conidia ellipsoid, globose, napiform, fusiform or naviculate with a truncate base and an obtuse to apiculate apex, hyaline or olivaceous brown to brown, unicellular, broadly or narrowly ellipsoidal, apices tapering, with or without a longitudinal germ slit, with or without a mucoid appendage.
Type genus: Coniella Höhn.
Type species: Coniella pulchella Höhn.
Notes: The monogeneric family Schizoparmaceae (Fig. 1, Clade 8) was introduced to accommodate Coniella (= Pilidiella, Schizoparme). Species of the asexual Pilidiella have been more widely reported than Schizoparme (Farr & Rossman 2017) and thus, Schizoparme was synonymised under Pilidiella giving priority to the older name (Rossman et al. 2015). Although van Niekerk et al. (2004) treated Coniella and Pilidiella as two distinct genera, the generic boundaries of the former were recently expanded to include “hyaline to dark brown conidia”, as Alvarez et al. (2016) reported that conidial pigmentation was lost or gained several times during the evolution of species within Coniella.
Coniella pseudokoreana Senan., Tangthir. & K.D. Hyde, sp. nov. MycoBank MB821542. Facesoffungi number FoF03505. Fig. 35.
Coniella pseudokoreana (MFLU 13–0282). A. Specimen on dead leaf. B. Conidiomata on host surface. C–D. Longitudinal section of a conidioma. E–H. Conidiogenous cells with developing conidia. I. Conidiogenous cells with developing conidia stained in lactophenol cotton blue. J. Conidia. K. Conidia stained with lactophenol cotton blue. L. Germinating conidium. M. Colonies on PDA from top. N. Colonies on PDA from reverse. Scale bar: C = 100 μm, D = 50 μm, E–L = 10 μm.
Etymology: Somewhat similar to Coniella koreana, however phylogenetically distant from this species.
Saprobic on dead leaves. Conidiomata pycnidial, solitary to gregarious, globose, brown, unilocular, ostiolate, 85–130 μm high, 78–106 μm diam ( = 108 × 92 μm, n = 10), immersed, with a central short ostiolar canal on each conidioma. Conidiomata wall 2–4-layered, 6–15 μm wide ( = 10 μm), with outer brown to dark brown layers composed of thick-walled cells of textura angularis, with inner pale brown layer composed of thin-walled cells of textura prismatica, except at the base which has a pulvinate convex giving rise to conidiophores or conidiogenous cells. Conidiophores 4–8 μm high, 1.5–4 μm wide, short, branched at the base, hyaline, smooth. Conidiogenous cells 5–10 μm high, 1.5–2 μm wide, holoblastic to enteroblastic, phialidic. Conidia 18–26 × 3–4 ( = 23 × 3.6 μm) fusiform, navicular, with one side slightly curved and another straight, smooth, hyaline, conidium length/width ratio = 6.5:1.
Culture characteristics: Colonies attaining a diam of 4 cm on PDA after 5 d at 27 ºC; surface white with medium to sparse mycelium, flat, irregular, undulate or wavy margin.
Specimen examined: Thailand, Phitsanullok, Tung Salang Luang, on decaying leaf, 18 Jun. 2012, N. Tangthirasunun, NTCL093 (holotype MFLU 13–0282, culture ex-type MFLUCC 12–0427).
Notes: Coniella pseudokoreana displays somewhat similar morphological characters to C. koreana and C. castaneicola in having linear, falcate, pale brown conidia (Alvarez et al. 2016). The colony morphology of Coniella koreana described in Alvarez et al. (2016) on PDA is similar to the colony morphology of our strain. However, conidiomatal morphology and size of the conidia are different. We collected Coniella pseudokoreana on a decaying leaf in Thailand. Our phylogeny reveals, C. pseudokoreana is distant from Coniella koreana and shares a sister taxon relationship to C. straminea (Fig. 1, Clade 8).
Stilbosporaceae Link [as ‘Stilbosporei’], Abh. Königl. Akad.Wiss. Berlin 1824: 180. 1826, emend. Clade 19.
Saprobic on bark of trees and shrubs. Sexual morph: Pseudostromata inconspicuous, immersed. Ectostromatic disc absent or present, if present inconspicuous, pale brown, rarely dark brown. Entostroma prosenchymatous, pale coloured, slightly differentiated from the surrounding bark tissue. Ascomata loosely arranged as valsoid groups in a single layer, immersed, aggregated, globose to subglobose, coriaceous, black, ostiolate, papillate. Ostiole not obvious, convergent in groups. Hamathecium comprising filiform, aseptate, hyaline paraphyses. Asci 8-spored, unitunicate, cylindrical, initially attached to the base, later floating in centrum, with J- refractive, apical ring. Ascospores overlapping uni- to biseriate, brown, ellipsoid to oblong, distoseptate. Asexual morph: Coelomycetous. Conidiomata stromatic, acervular with circular outline, epidermal, immersed to semi-immersed, brown, basal stroma of textura angularis to textura globulosa, with simple, septate, hyaline paraphyses and hyaline, unbranched cylindrical conidiophores. Conidiophores arising from the uppermost cells of basal and parietal tissue, unbranched, cylindrical, septate at only the base, hyaline, smooth, invested in mucus. Conidiogenous cells annellidic, discrete or integrated, cylindrical to lageniform, hyaline, smooth-walled, proliferating several times percurrently at apex. Conidia ellipsoid or oblong, with an obtuse apex and broad truncate base, sometimes 3-euseptate or distoseptate, with a hyaline sheath, hyaline to brown, thick-walled, smooth, sometimes with several, tubular, unbranched, filiform, flexuous, apical appendages.
Type genus: Stilbospora Pers.
Type species: Stilbospora macrosperma Pers.
Notes: The family Stilbosporaceae was introduced by Link (1826) to accommodate Prosthecium and its asexual morph. However, it is not a phylogenetically well-supported family and hence, Stilbosporaceae has been synonymised under several different families. Voglmayr & Jaklitsch (2014) resurrected the family Stilbosporaceae in Diaporthales based on a phylogenetic analysis of LSU nrDNA sequence data and accommodated the genera Stegonsporium and Stilbospora within the family, synonymising Prosthecium under Stilbospora. This decision is also supported by our multi-gene phylogeny (Fig. 1, Clade 19). The type species of Stilbospora, S. macrosperma has been linked to its asexual morph Prosthecium ellipsosporum, the generic type of Prosthecium (Voglmayr & Jaklitsch 2008). This genus comprises opportunistic or moderately phytopathogenic fungal species that cause branch dieback or twig blight of various plants. Maharachchikumbura et al. (2015) included Natarajania in Stilbosporaceae based on LSU nrDNA, SSU nrDNA, tef1 and rpb2 sequence data. However, in other analyses (not shown here), phylogenies also indicated a close association to the genera Crinitospora, Stilbospora and Stegonsporium. This is rather interesting as up to date, this is the only hyphomycetous taxon affiliated to the diaporthean taxa which are known to have coelomyceteous asexual morphs. The reliability of the deposited sequences as well as the identification of that taxon needs further investigation.
Stilbospora macrosperma Pers., Syn. meth. fung. (Göttingen) 1: 96 (1801). Facesoffungi number FoF03506. Fig. 36.
Stilbospora macrosperma (WU 28068). A. Herbarium specimen. B. Stromata on host substrate. C. Cross section of ascoma. D. Peridium. E–H. Asci. I–M. Ascospores. N. Paraphyses. O. Cross section of conidioma. P. Conidia attached to conidiogenous cells. Q–S. Conidia. Scale bars: C–D = 500 µm, E, O, P = 100 µm, F = 50 µm, H–K = 20 µm, G, L–M, Q–S = 10 µm.
Saprobic on branches of Acer pseudoplatanus. Sexual morph: Pseudostroma comprising white, greyish to yellowish hyphae. Ascomata 300–350 μm high, 350–465 μm diam ( = 325 × 420 μm, n = 20), immersed, aggregated, globose to subglobose, coriaceous, ostiolate, papillate. Papilla cylindrical, pale brown, emerging from perithecial apices and merging separately with the stromatal disc, inconspicuous, often invisible on the bark surface. Peridium 20–40 μm diam ( = 32 μm, n = 20), comprising thick-walled, brown, large, cells of textura angularis and hyaline, thick-walled, compressed cells of textura angularis around the base of papilla. Hamathecium comprising multiguttulate, hyaline, septate paraphyses. Asci 165–200 × 35–50 μm ( = 182 × 42 μm, n = 20), 8-spored, unitunicate, clavate to ellipsoidal, thick-walled, very short pedicellate, apex containing a J- refractive canal usually wider towards its base. Ascospores 40–50 × 20–26 μm ( = 46 × 22 μm, n = 20), biseriate, ellipsoidal, oblong or rarely pyriform, with (3–)5-distosepta and sometimes 1, longitudinal, distoseptum, appendages on both ends projecting, subglobose, outer margin becoming diffuse. Asexual morph: Conidiomata 340–450 μm high, 450–460 μm diam ( = 410 × 453 μm, n = 20), immersed, acervular, solitary, with circular outline, dark brown to black. Paraphyses 2.5–4 μm diam ( = 3.2 μm, n = 10), unbranched, aseptate, hyaline. Conidiophores reduced to conidiogenous cells. Conidiogenous cells 25–35 × 7–10 μm ( = 31 × 9 μm, n = 20), holoblastic, cylindrical, septate, hyaline. Conidia 40–45 × 20–25 μm ( = 43 × 23 μm, n = 20), pyriform, oval, ellipsoid or oblong, base truncate and hyaline, brown, with several distosepta, 1(–2)-longitudinal distosepta, with hyaline sheath.
Material examined: Austria, Wien, Landstraße, 3rd District, Botanical Garden of the University of Vienna (HBV), grid square 7864/1, on dead corticated branches of Acer pseudoplatanus (Sapindaceae), holomorph, 4 Feb. 2006, H. Voglmayr, D39 (epitype WU 28068).
Notes: Voglmayr & Jaklitsch (2014) epitypified the type species of Stilbospora, Stilbospora macrosperma and S. macrosperma was confirmed as the asexual morph of Prosthecium ellipsosporum, the generic type of Prosthecium (Voglmayr & Jaklitsch 2008). Stilbospora (1801) is older than Prosthecium (1852) and therefore Stilbospora has priority (Voglmayr & Jaklitsch 2014).
Sydowiellaceae Lar.N. Vassiljeva, Pirenomits. Lokuloaskomits. Severa Dal'nego Vostoka (Leningrad): 210. 1987. Clade 21.
Saprobic or parasitic on plant matter. Sexual morph: Stromata well- or poorly developed, prosenchymatous, scattered, immersed to erumpent, appearing as an aggregation of ostioles, rounded or elliptic in shape, dark brown to black, composed of compact pseudoparenchymatous tissues, several ascoma in a stromata, some species turn umber in 5 % KOH. Ascomata solitary or aggregated, immersed or erumpent, globose to sub-globose, sometimes circinate, coriaceous, central or asymmetrically located ostiolar canal opens through an individual or converged ostiole, internally covered by filamentous, hyaline periphyses, sometime ostiolar opening wider than canal, black to brown. Peridium comprising a few layers of brown, thick-walled cells of textura angularis. Hamathecium comprising cellular, septate, branched, hyaline paraphyses. Asci 8-spored, unitunicate, cylindrical to sub-globose, short pedicellate, apex blunt with J- apical ring. Ascospores uni- to multi-seriate, filamentous, ellipsoid or long fusoid-cylindrical, 1–11-septate, hyaline, pale brown to dark brown, sometimes with apical and basal appendages, wall smooth. Asexual morph: Conidiomata sometimes stromatic, pycnidia, uniloculate, superficial, aggregated 3–5 in one group, globose, orange to brown. Conidiomatal wall comprising thick-walled, orange cells of textura angularis. Conidiophores elongate, branched, hyaline, few conidiogenous cells arising from one conidiophore, attached to conidiomatal wall. Conidiogenous cells cylindrical, hyaline, ampulliform, septate, ends pointed, phialidic. Conidia ovoid to ellipsoid, unicellular, hyaline, smooth-walled.
Type genus: Sydowiella Petr.
Type species: Sydowiella fenestrans (Duby) Petr.
Notes: The family Sydowiellaceae (Fig. 1, Clade 21) was established to accommodate the genus Sydowiella, which is typified by S. fenestrans. Members of this family occur on herbaceous plants, dicotyledonous and hardwood trees as saprobes, parasites and pathogens. Initially, most genera in this family were placed in Diaporthales incertae sedis (Rossman et al. 2006). However, DNA sequence data analyses of different gene regions of taxa in the family Sydowiellaceae proved it to be a well-supported, and its relationships to other families have been clarified (Rossman et al., 2007, Maharachchikumbura et al., 2015, Senanayake et al., 2017). Sydowiellaceae comprises the genera Alborbis, Breviappendix, Cainiella, Calosporella, Chapeckia, Italiomyces, Hapalocystis, Lambro, Paragnomonia, Ranulospora, Rossmania, Sillia, Sydowiella, Tenuiappendicula and Tortilispora (Senanayake et al. 2017). Here we introduce a new Sydowiella species as S. urticicola. Sydowiella urticicola produce solitary ascomata and ascospores containing a large guttule in each cell which clearly demarcates it from other species. Phylogenetically it is also distinct from other Sydowiella species.
Sydowiella urticicola Senan., Camporesi & K.D. Hyde, sp. nov. MycoBank MB821566. Facesoffungi number FoF03507. Fig. 37.
Sydowiella urticicola (MFLU 17–0877). A–C. Ascomata on host surface. D. Cross section of ascoma. E. Peridium. F. Papilla. G–I. Asci. J–N. Ascospores. Scale bars: A = 500 µm, B, C = 200 µm, D, E = 100 µm, F–I = 50 µm, J–N = 20 µm.
Etymology: Named after the host genus Urtica.
Saprobic on dead branches of deciduous plants. Sexual morph: Ascomata 290–325 μm high, 290–395 μm diam ( = 309 × 314 μm, n = 20), perithecia, astromatic, scattered, solitary, superficial to erumpent, globose to sub-globose, coriaceous, black, papillate, ostiolate. Papilla 115–150 μm, 155–205 μm diam ( = 133 × 173 μm, n = 20), short, wide, internally covered by hyaline periphyses. Peridium 15–25 μm diam ( = 19 μm, n = 20) comprising inner, hyaline, thick-walled, compressed, 1–3 layers of cells of textura angularis and outer, dark brown, thick-walled, rigid, 3–7 layers of cells of textura angularis. Hamathecium comprising wide, cellular, septate paraphyses. Asci 125–145 × 10–15 μm ( = 136 × 14 μm, n = 20) 8-spored, unitunicate, cylindrical, short pedicellate, distinct, J- apical ring. Ascospores 20–25 × 10–15 μm ( = 22 × 12 μm, n = 20), overlapping uniseriate, ends blunted, hyaline, 1-septate, slightly constricted at the septum, with a large fat globule in each cell. Asexual morph: Undetermined.
Culture characteristics: Colonies growing on MEA becoming 2 cm within 5 d at 18 °C, fast growing, circular, umbonate, margin irregular, white, tightly attached to the substrate without spreading aerial mycelium.
Specimen examined: Italy, Province of Forlì-Cesena, Monte Fumaiolo, dead stem of Urtica dioica (Urticaceae), 16 May 2013, N. Camporesi, IT 1268 (holotype MFLU 17–0877, isotype BBH 42439, cultures ex-type MFLUCC 13–0665, MFLUCC 17–1665).
Notes: Here we introduce a new Sydowiella species as S. urticicola. Sydowiella urticicola produces solitary ascomata and ascospores containing a large guttule in each cell. Phylogenetically this is distinct from other Sydowiella species (Fig. 1, Clade 21).
Diaporthales genera incertae sedis
Phaeoappendicospora Senan., Q.R. Li & K.D. Hyde, gen. nov. MycoBank MB821572. Facesoffungi number FoF03508. Clade 18.
Etymology: Name based on three Latin words “phaeo”, “appendicem” and “spora” referring to the brown spores with appendages.
Saprobic on dead stems of Fagaceae species. Sexual morph: Stromata forming thin weft of pale brown hyphae around upper part of perithecia. Ascomata immersed, aggregated, subglobose to globose, coriaceous, black, ostiolate, papillate. Papilla black, cylindrical, lateral, periphysate. Periphyses hyaline, filamentous. Peridium comprising few layers of black, thick-walled, cells of textura angularis. Hamathecium comprising few, septate, hyaline paraphyses attached to the base, longer than asci. Asci 8-spored, unitunicate, ellipsoid, with a short pedicel, inconspicuous, flat, J- refractive ring at the lower end of the thickened apical wall, apex narrow and blunted. Ascospores ellipsoidal to ovoid, with broadly rounded ends, pale brown, 1-septate, not constricted at the septa, thick and smooth-walled, guttulate, with short, hyaline, rounded appendages at both ends. Asexual morph: Undetermined.
Type species: Phaeoappendicospora thailandensis Q.R. Li, Senan. & K.D. Hyde.
Phaeoappendicospora thailandensis Senan., Q.R. Li & K.D. Hyde, sp. nov. MycoBank MB821573. Facesoffungi number FoF03509. Fig. 38.
Phaeoappendicospora thailandensis (MFLU 12–2131). A. Appearance of fungus on host substrate. B. Horizontal cross section of stroma. C. Vertical cross section of stromata. D. Peridium. E. Apical ring in Melzer’s reagent. F–I. Asci. J–M. Ascospores. Scale bars: A = 1 mm, B = 500 μm, C = 100 μm, D = 25 μm, F–M = 10 μm.
Etymology: The species epithet is based on the collection locality.
Saprobic on dead stems of Quercus. Sexual morph: Stromata 2–4 mm diam ( = 3 mm, n = 15), forming thin weft of pale brown hyphae around upper part of perithecia. Ascomata 180–230 μm high, 170–220 μm diam ( = 208 × 198 μm, n = 25), immersed, aggregated, subglobose to globose, coriaceous, black, ostiolate, papillate. Papilla black, cylindrical, lateral, periphysate. Periphyses hyaline, filamentous. Peridium 20–35 μm wide ( = 28 μm, n = 20), comprising few layers of black, thick-walled cells of textura angularis. Hamathecium comprising few, septate, hyaline paraphyses attached to the base, longer than asci. Asci 195–265 × 18.5–27 μm ( = 223 × 22.5 μm, n = 30), 8-spored, unitunicate, ellipsoid, with a short pedicel, inconspicuous, flat, J- refractive ring at the lower end of the thickened apical wall, apex narrow and blunted. Ascospores 26–34.5 × 11–13 μm ( = 30 × 12.5 μm, n = 30), ellipsoidal to ovoid, with broadly rounded ends, pale brown, 1-septate, not constricted at the septa, thick and smooth-walled, guttulate, with short, hyaline, rounded appendages at both ends. Asexual morph: Undetermined.
Culture characteristics: Colonies on PDA reaching 5 cm in 7 d at 25 °C, fast growing, circular, flat, smooth, whitish cream in upper, pale yellow in reverse.
Specimen examined: Thailand, Chiang Rai, near Khun korn water fall, on branch of Quercus sp. (Fagaceae), 25 Dec. 2012, Q.R. Li, TL19 (holotype MFLU 12–2131, culture ex-type MFLUCC 13–0161).
Notes: The newly introduced monotypic genus Phaeoappendicospora is typified by P. thailandensis and it comprises brown ascospores with guttules and appendages. Phaeoappendicospora thailandensis is associated with dead plant parts may be as saprobes. This fungus is morphologically similar to Hapalocystis berkeleyi in Sydowiellaceae. However, phylogenetically it does not show an affinity to any of the families in Diaporthales (Fig. 1, Clade 18). Hence, we accommodate this species in Diaporthales genera incertae sedis.
Key to families and genera of Diaporthales
| 1. | Coelomycetous……………………………………………………………..2 |
| 1. | Ascomycetous………………………………………………………………7 |
| 2. | Conidia hyaline to olivaceous or bluish to glistening dark blue……...3 |
| 2. | Conidia brown to dark brown…………………………………………….6 |
| 3. | Conidial wall thick, mostly specific on Tiliaceae host …………………………………………………………………………….Lamproconiaceae |
| 3. | Conidial wall thin, mostly on Myrtaceae, Fabaceae host……………..4 |
| 4. | Conidia aseptata…………………………………………………………...5 |
| 4. | Conidia 1-septate……………………………….Auratiopycnidiellaceae |
| 5. | Conidiophores reduced to conidiogenous cells, conidia with acute apex…………………………………………………….Erythrogloeaceae |
| 5a. | Conidia olivaceous…………………………………………….Disculoides |
| 5a. | Conidia hyaline……………………………………………………………3b |
| 5b. | Conidiomata acervular, sides of conidiomatal wall appearing dark brown to black cells of textura angularis, conidiogenous cells lageniform to cylindrical, conidia monomorphic………………………………………………………………………………………….Erythrogloeum |
| 5b. | Conidiomata subglobose, conidiomatal wall uniformly comprises orange-brown cells of textura angularis conidiogenous cells ampulliform, conidia dimorphic…………………………….Chrysocrypta |
| 5. | Conidiophores subcylindrical, branched, 0–3-septate, conidia with obtuse apex…………………………………………..Prosopidicolaceae |
| 6. | Conidia ovoid, obclavate to conical……………………………………...7 |
| 6. | Conidia transversely distoseptate, consisting of four arms………………………………………………………………………..Asterosporiaceae |
| 7. | Conidial wall smooth on the outer surface, with inconspicuous to distinct irregular verrucae on the inner surface…..Juglanconidaceae |
| 7. | Conidial wall without ornamentation……………………………………..8 |
| 8. | Conidia ovoid, 1-septate………………………………….Macrohilaceae |
| 8. | Conidia obclavate to conical, aseptata…………..Apoharknessiaceae |
| 8a. | Conidia obclavate, with a scar at the base…………………..Lasmenia |
| 8a. | Conidia conical, with small appendage at base and apical apiculus………………………………………………………………….Apoharknessia |
| 9. | Stromata well-developed………………………………………………...10 |
| 9. | Stromata absent or poorly developed………………………………….17 |
| 10. | Stromatic tissues orange, becoming purple in KOH………………………………………………………………………………Cryphonectriaceae |
| 10a. | Asexual morph not reported, hamathecium paraphysate, ascospores brown……………………………………………………….Chromendothia |
| 10a. | Asexual morph reported, hamathecium aparaphysate, ascospores hyaline……………………………………………………………………10b |
| 10b. | Coelomycetes……………………………………………………………10c |
| 10b. | Ascomycetes……………………………………………………………..10i |
| 10c. | Conidia sigmoid with obtuse to subobtuse apex and swollen, obtuse base……………………………………………………….Aurantiosacculus |
| 10c. | Conidia cylindrical, ellipsoid to fusoid, occasionally allantoid……………………………………………………………………………………….10d |
| 10d. | Conidiomata pulvinate………………………………………………….10e |
| 10d. | Conidiomata rostrate, pyriform to globose…………………………...10g |
| 10e. | Neck present…………………………………………………..Luteocirrhus |
| 10e. | Neck absent………………………………………………………………10f |
| 10f. | Conidiomata superficial, pale to medium brown, conidiophores consisting of basal subglobular to angular cells, that branch irregularly, becoming cylindrical, transversely septate………………….Foliocryphia |
| 10f. | Conidiomata immersed to erumpent, orange, conidiophores aseptate, occasionally with separating septa and branching……………………..………………………………………………………………..Immersiporthe |
| 10g. | Conidiophores hyaline, cylindrical, delimited by septa or not, conidia cylindrical………………………………………………………...Ursicollum |
| 10g. | Conidiophores reduced to conidiogenous cells, conidia fusoid-ellipsoid or allantoid……………………………………………..10h |
| 10h. | Conidia with apical appendage………………………….Mastigosporella |
| 10h. | Conidia without apical appendage…………………………...Chrysofolia |
| 10i. | Conidiomata uniformly orange………………………………………….10j |
| 10i. | Conidiomata uniformly brown to black, with or without orange necks………………………………………………………………………10t |
| 10j. | Conidiomata pulvinate to globose……………………………………..10k |
| 10j. | Conidiomata conical, rostrate, pyriform or convex………………….10q |
| 10k. | Ascospores septate……………………………………………………...10l |
| 10k. | Ascospores aseptate……………………………………………………10p |
| 10l. | Ascostromata superficial, conidiomata paraphysate……………….10m |
| 10l. | Ascostromata immersed to erumpent, conidiomata aparaphysate………………………………………………………………………………….10n |
| 10m. | Perithecia valsoid…………………………………………...Diversimorbus |
| 10m. | Perithecia diatrypoid………………………………………………Microthia |
| 10n. | Conidiomata usually more than 350 μm diam, ascospores with median septum………………………………………………Cryphonectria |
| 10n. | Conidiomata usually less than 350 μm diam, ascospores with median to submedian septum…………………………………………10o |
| 10o. | Ascospores oval to ellipsoid, papilla parallel to each other and open individually……………………………………………………Cryptometrion |
| 10o. | Ascospores fusiform, sometimes curved, papilla close to each other and converge at the apex…………………………………Aurantioporthe |
| 10p. | Stromata strongly developed, large, erumpent, mostly superficial, numerous conidial locules, no paraphyses…………………….Endothia |
| 10p. | Stromata small to medium, semi-immersed, few conidial locules or one convoluted locule, paraphyses present…………….Holocryphia |
| 10q. | Conidiomata with necks…………………………………………………10r |
| 10q. | Conidiomata without necks…………………………………………….10s |
| 10r. | Conidiomata with prominent, delimited neck………………Latruncellus |
| 10r. | Conidiomata with neck continuous with base, rostrate, white sheath of tissue surrounding perithecial necks……………………Rostraureum |
| 10s. | Conidiomata conical, uniformly orange………………………Amphilogia |
| 10s. | Conidiomata convex, with blackened ostiolar openings……...Aurifilum |
| 10t. | Conidiomata uniformly black when mature…………………………..10u |
| 10t. | Conidiomata black with orange neck……………………………Aurapex |
| 10u. | Conidiomata base tissue of textura globulosa when sectioned longitudinally, perithecial necks long and covered with dark tissue, emerging from orange stroma……………………………..Chrysoporthe |
| 10u. | Conidiomata base tissue prosenchymatous, apices of conidiomata can be orange to scarlet when young, perithecial necks short, orange to umber stroma………………………………………Celoporthe |
| 10. | Stromatic tissues dark brown to black, not becoming purple in KOH…………………………………………………………………………11 |
| 11. | Perithecia with very long, narrow, wavy ostiolar neck opening to or around ectostromatic disc………………………………………………..12 |
| 11. | Perithecia with medium to short, somewhat wide, straight ostiolar neck opening to host surface……………………………………………14 |
| 12. | Ectostroma conspicuous………………………………Melanconidaceae |
| 12. | Ectostroma inconspicuous………………………………………………13 |
| 13. | Ascospores distoseptata……………………………….Stilbosporaceae |
| 13a. | Conidia hyaline with several tubular, unbranched, filiform apical appendages…………………………………………………….Crinitospora |
| 13a. | Conidia brown without any appendages……………………………..13b |
| 13b. | Ascospores and conidia with three transverse eusepta, ellipsoid to oblong; asci without a refractive canal in the apex………..Stilbospora |
| 13b. | Ascospores and conidia with more than three transverse distosepta, ascospores sometimes and conidia always with additional longitudinal distosepta, ascospores ellipsoid to oblong, conidia mostly pyriform; asci with a cylindrical, slightly refractive canal in the apex…………………………………………………………………………….Stegonsporium |
| 13. | Ascospores not distoseptata………………………..Melanconiellaceae |
| 13a. | Coelomycetous…………………………………………………………..13b |
| 13a. | Ascomycetous……………………………………………………………13c |
| 13b. | Conidiomata subcuticular, acervular…………………………..Greeneria |
| 13b. | Conidiomata epiphyllous or hypophyllous with radiate scutella……………………………………………………………………………….Tubakia |
| 13c. | Astromatic with solitary ascomata…………………………………….13d |
| 13c. | Stromatic with aggregated ascomata…………………….Mellanconiella |
| 13d. | Ascospores ellipsoid without appendages…………………….Dicarpella |
| 13d. | Ascospores fusiform with appendages…………………Microascospora |
| 14. | Perithecia arranged in valsoid configuration………………………….15 |
| 14. | Perithecia not arranged in valsoid configuration……………………..16 |
| 15. | Conidiomata non-loculate and forming both alpha and beta conidia…………………………………………………………………Diaporthaceae |
| 15a. | Coelomycetes……………………………………………………………15b |
| 15a. | Ascomycetes…………………………………………………………….15e |
| 15b. | Conidia olivaceous………………………………………………………15c |
| 15b. | Conidia brown……………………………………………………………15d |
| 15c. | Conidia aseptate, guttulate, elongate fusiform to sigmoid……………..………………………………………………………………...Pustulomyces |
| 15c. | Conidia 2(-3)-septate, ovoid with filiform apical cell and obtuse base………………………………………………………………….Chaetoconis |
| 15d. | Conidia dimorphic, aseptate, ellipsoid to pyriform…………………………………………………………………………………….Phaeocytostroma |
| 15d. | Conidia monomorphic, uniseptate, subcylindrical to narrowly ellipsoid…………………………………………………………………Stenocarpella |
| 15e. | Ascospores brown………………………………………..Phaeodiaporthe |
| 15e. | Ascospores hyaline………………………………………………………15f |
| 15f. | Ascospores aseptata……………………………………………Mazzantia |
| 15f. | Ascospores septate……………………………………………………..15g |
| 15g. | Septa submedian, large cell usually 2-guttulate, small cell usually 1-guttulate……………………………………………………..Apioporthella |
| 15g. | Septa median, with or without guttules………………………………15h |
| 15h. | Ascospores with long slender, thread-like appendage at both ends……………………………………………………………..Hyaliappendispora |
| 15h. | Ascospores without appendages………………………………………15i |
| 15i. | Papilla long-cylindrical, conidia globose to subglobose, multiguttulata………………………………………………………………Ophiodiaporthe |
| 15i. | Papilla short, conidia absent or if present; ovate to ellipsoidal, biguttulata…………………………………………………………………15j |
| 15j. | Ascospores ovoid, not constricted at the septa……………………..15k |
| 15j. | Ascospores fusiform to elongate-ellipsoid, constricted at the septa….……………………………………………………………………………..15l |
| 15k. | Asci form long, pointed apex by narrowing towards the apical ring….…………………………………………………………….Chiangraiomyces |
| 15k. | Asci form blunt apex………………………………………Leucodiaporthe |
| 15l. | Ascospores overlapping uniseriate, often with 4 guttules, larger guttules at the center and smaller ones at the ends………………15m |
| 15l. | Ascospores biseriate, without guttules…………………….Allantoporthe |
| 15m. | Ascospores fusiform, ends pointed, papilla short and wide……………………………………………………………………………...Paradiaporthe |
| 15m. | Ascospores elongate to elliptical, ends round, papilla long and narrow……………………………………………………………………..Diaporthe |
| 15. | Conidiomata loculate forming numerous interconnecting chambers arranged radially or irregularly with in ectostromatic tissues and without forming beta conidia…………………………….Cytosporaceae |
| 15a. | Coelomycetes……………………………………………………..Waydora |
| 15a. | Ascomycetes…………………………………………………………….15b |
| 15b. | Stromata inconspicuous, ascospores allantoids…………………….15c |
| 15b. | Stromata conspicuous, well-developed, ascospores fusiform…………………………………………………………………………………….…15d |
| 15c. | Ascomata solitary………………………………………………..Paravalsa |
| 15c. | Ascomata aggregated……………………………………………Xenotypa |
| 15d. | Perithecia in groups with convergent beaks; asci clavate to fusoid………………………………………………………………………..Cytospora |
| 15d. | Perithecia in groups with non-convergent beaks; asci more or less rectangular………………………………………………………Pachytrype |
| 16. | Conidia dark brown, broadly fusiform to cylindrical or clavate, 3-5-cellular, distoseptata……………………………………Coryneaceae |
| 16. | Conidia hyaline, ellipsoid, unicellular…………………..Sydowiellaceae |
| 16a. | Stromata conspicuous, well-developed………………………………16b |
| 16a. | Stromata absent, inconspicuous, poorly developed………………..16d |
| 16b. | Ascomata valsoid, stromatic tissues do not turn any colour with 10 % KOH………………………………………………………………………16c |
| 16b. | Ascomata diatrypoid, stromatic tissues form dull red with 10 % KOH……………………………………………………………………………Sillia |
| 16c. | Ascospores oval to fusoid-oval, 1-septate, hyaline or hyaline to brown…………………………………………………………………….Chapeckia |
| 16c. | Ascospores fusiform to ellipsoid with long filiform basal cell, 2-septate, hyaline…………………………………………………………..Ranulospora |
| 16d. | Ascomata solitary……………………………………………………….16e |
| 16d. | Ascomata aggregated…………………………………………………...16i |
| 16e. | Asci 4-spored……………………………………………….Breviappendix |
| 16e. | Asci 8-spored…………………………………………………………….16f |
| 16f. | Ascospores apiosporous…………………………………………..Lambro |
| 16f. | Ascospores non-apiosporous………………………………………….16g |
| 16g. | Ascospores oval, hyaline to brown, not constricted at the septa……..………………………………………………………………………Cainiella |
| 16g. | Ascospores fusoid, hyaline, constricted at the septa………………16h |
| 16h. | Ascospores non-appendaged, globules at the center of each cell…………………………………………………………………………Sydowiella |
| 16h. | Ascospores appendaged, globules at the septa of the spore………………………………………………………………………………Italiomyces |
| 16i. | Ascospores oval, short fusoid to ellipsoidal…………………………..16j |
| 16i. | Ascospores long fusoid cylindrical…………………………………….16n |
| 16j. | Ascospores 1-septate……………………………………………………16k |
| 16j. | Ascospores multi-septate………………………………………………16m |
| 16k. | Ascospores with long strip-like appendages…………Tenuiappendicula |
| 16k. | Ascospores with short, appendages……………………………………16l |
| 16l. | Ascospores ellipsoid to cylindrical, usually 4-guttulate…………Alborbis |
| 16l. | Ascospores fusiform, usually multi-guttulate…………….Paragnomonia |
| 16m. | Ascospores hyaline to brown, ellipsoidal with broadly rounded ends, 1–3-septate, constricted at the septa………………………Hapalocystis |
| 16m. | Ascospores hyaline, fusoid to oblong, 3–4-eusepta, not constricted at the septa…………………………………………………….Calosporella |
| 16n. | Ascospores 0–5-septate……………………………………….Tortilispora |
| 16n. | Ascospores 6–11-septate……………………………………..Rossmania |
| 17. | Ascospores unicellular……………………………………………………18 |
| 17. | Ascospores multicellular………………………………………………….19 |
| 18. | Ascomata superficial to erumpent, conidia elongate ellipsoidal to fusiform without appendages…………………………Schizoparmaceae |
| 18. | Ascomata immersed, conidia oval to globose with basal appendage………………………………………………………………..Harknessiaceae |
| 19. | Perithecia with 2–3 necks opening on both sides of the substrate/leaf blade…………………………………………………Apiosporopsidaceae |
| 19. | Perithecia with one neck opening to upper side of the substrate/leaf blade………………………………………………………………………...20 |
| 20. | Papilla short, conidia with microcyclic conidiation……………………………………………………………………..…Pseudoplagiostomataceae |
| 20. | Papilla long, conidia do not have microcyclic conidiation………………………………………………………………………………Gnomoniaceae |
| 20a. | Coelomycetes…………………………………………………………….20b |
| 20a. | Ascomycetes………………………………………………………………20f |
| 20b. | Conidia hyaline……………………………………………………………20c |
| 20b. | Conidia brown……………………………………………………….Uniseta |
| 20c. | Conidiomata pycnidia……………………………………………………20e |
| 20c. | Conidiomata acervuli…………………………………………………….20d |
| 20d. | Conidia 1-septate………………………………………………..Sirococcus |
| 20d. | Conidia aseptate……………………………………………...Millerburtonia |
| 20e. | Conidia filiform to fusiform……………………………………….Asteroma |
| 20e. | Conidia ellipsoidal…………………………………………Cylindrosporella |
| 20f. | Pseudostromata/ stromata absent; perithecia immersed in host tissues……………………………………………………………………..20g |
| 20f. | Pseudostromata/ stromata present; perithecia immersed in stromatic tissues…………………………………………………………………….20u |
| 20g. | Perithecial necks not parallel to substrate……………………………20h |
| 20g. | Perithecial necks parallel to substrate and not fused…………………..………………………………………………………………..Cryptosporella |
| 20h. | Infected lesions distinct with dark purple to brown pigmentation or blackish area with pale brown sharp margin…………………………20i |
| 20h. | Infected lesions indistinct………………………………………………..20j |
| 20i. | Ascospores apiosporous………………………………...Apioplagiostoma |
| 20i. | Ascospores non-apiosporous……………………………………Diplacella |
| 20j. | Ascospores slightly isthmoid with a median septum, often readily separate as part spores……………………………………….Pleuroceras |
| 20j. | Ascospores non-isthmoid, do not separate into part spores……….20k |
| 20k. | Appendages generally present…………………………………………20l |
| 20k. | Appendages generally absent…………………………………………20o |
| 20l. | Perithecia mostly epiphyllous…………………………………………20m |
| 20l. | Perithecia mostly hypophyllous………………………………………..20n |
| 20m. | Appendages ovoid to subulate………………………...Ambarignomonia |
| 20m. | Appendages cuneiform with diffuse ends or ovoid, subulate acicular……………………………………………………………Gnomonia |
| 20n. | Ascospores fusiform; arranged irregularly fasciculate or, obliquely in one longitudinal row…………………………………….Apiognomonia |
| 20n. | Ascospores oval to filiform; arranged unevenly parallel, irregularly multiseriate or obliquely uniseriate, occasionally evenly parallel……..………………………………………………………………Ophiognomonia |
| 20o. | Perithecia occurring on both sides of the host leaf…………………20q |
| 20o. | Perithecia occurring on only upper or lower side of host leaf…….20p |
| 20p. | Necks present……………………………………………………………20r |
| 20p. | Necks absent………………………………………………Gloeosporidina |
| 20q. | Bell-shaped to hemispherical chamber at base of neck present……..…………………………………………………………………..Spataporthe |
| 20q. | Bell-shaped to hemispherical chamber at base of neck absent………………………………………………………………………….Gnomoniella |
| 20r. | Asci 32-spored……………………………………………………;Ditopella |
| 20r. | Asci 8-spored……………………………………………………………20s |
| 20s. | Ascomata immersed in pale brownish, parenchymatous cellular cavity and ascomata easily separate from them……..Marsupiomyces |
| 20s. | Ascomata not immersed in any, parenchymatous cellular pockets………………………………………………………………………………..20t |
| 20t. | Stromata black………………………………………………Gnomoniopsis |
| 20t. | Stromata grey, brownish, cream, yellowish white………...Plagiostoma |
| 20u. | Ectostromatic disc present………………………………..…Ditopellopsis |
| 20u. | Ectostromatic disc absent………………………………………………20v |
| 20v. | Stromatic tissues grey to pale brown; on top of perithecia and cream yellow mycelium at bottom of perithecia………..Occultocarpon |
| 20v. | Stromatic tissues black; do not form any mycelium at bottom of perithecia…………………………………………………………………20w |
| 20w. | Ascospores septate……………………………………………………..20x |
| 20w. | Ascospores aseptate…………………………………………………..20ac |
| 20x. | Cytoplasm of ascospores granular and divide into two parts with wide vacuous space forming diplastic polarity………………Bagcheea |
| 20x. | Cytoplasm of ascospores does not divide into any parts………….20y |
| 20y. | Ascospores apiosporous……………………………………………….20z |
| 20y. | Ascospores non-apiosporous………………………………………...20aa |
| 20z. | Ascospores form brown large cell and small hyaline cell at maturity……………………………………………………………………..Anisomyces |
| 20z. | Ascospores form both cells hyaline at maturity…………...Mamianiella |
| 20aa. | Ascospores 3-septate…………………………………….Phragmoporthe |
| 20aa. | Ascospores 1-euseptate………………………………………………20ab |
| 20ab. | Stromata immersed in host tissues in bark, ascospores fusiform…….………………………………………………………………………Alnecium |
| 20ab. | Stromata erumpent to superficial on leaves, ascospores oval……………………………………………………………………………..Phylloporthe |
| 20ac. | Ascospores oval to ellipsoid…………………………………Amphiporthe |
| 20ac. | Ascospores allantoid………………………………………….Valsalnicola |
List of accepted families and genera in Diaporthales
| Apiosporopsidaceae Senan. et al. |
| Apiosporopsis (Traverso) Mariani |
| Apoharknessiaceae Senan. et al. |
| Apoharknessia Crous & S.J. Lee |
| Lasmenia Speg. |
| Asterosporiaceae Senan. et al. |
| Asterosporium Kunze |
| Auratiopycnidiellaceae Senan. et al. |
| Auratiopycnidiella Crous & Summerell |
| Coryneaceae Corda |
| = Pseudovalsaceae M.E. Barr |
| Coryneum Nees |
| Cryphonectriaceae Gryzenh. & M.J. Wingf. |
| Amphilogia Gryzenh. et al. |
| Aurantiosacculus Dyko & B. Sutton |
| Aurapex Gryzenh. & M.J. Wingf. |
| Aurifilum Begoude et al. |
| Celoporthe Nakab. et al. |
| Chromendothia Lar.N. Vassiljeva |
| Chrysofolia Crous & M.J. Wingf. |
| Chrysoporthe Gryzenh. & M.J. Wingf. |
| =Chrysoporthella Gryzenh. & M.J. Wingf. |
| Cryphonectria (Sacc.) Sacc. & D. Sacc. |
| Cryptometrion Gryzenh. & M.J. Wingf. |
| Diversimorbus S.F. Chen & J. Roux |
| Endothia Fr. |
| Foliocryphia Cheewangkoon & Crous |
| Holocryphia Gryzenh. & M.J. Wingf. |
| Immersiporthe S.F. Chen et al. |
| Latruncellus M. Verm. et al. |
| Luteocirrhus C.F. Crane & T.I. Burgess |
| Mastigosporella Höhn. |
| Microthia Gryzenh. & M.J. Wingf. |
| Rostraureum Gryzenh. & M.J. Wingf. |
| Ursicollum Gryzenh. & M.J. Wingf. |
| Cytosporaceae Fr. |
| =Valsaceae Tul. & C. Tul. |
| Cytospora Ehrenb. |
| Pachytrype Berl. ex M.E. Barr et al. |
| Paravalsa Ananthap |
| Waydora B. Sutton |
| Xenotypa Petr. |
| Diaporthaceae Höhn. ex Wehm. |
| Allantoporthe Petr. |
| Apioporthella Petr. |
| Chaetoconis Clem. |
| Chiangraiomyces Senan. & K.D. Hyde |
| Diaporthe Nitschke |
| Hyaliappendispora Senan. et al. |
| Leucodiaporthe M.E. Barr et al. |
| Mazzantia Mont. |
| Ophiodiaporthe Y.M. Ju et al. |
| Paradiaporthe Senan. et al. |
| Phaeocytostroma Petr. |
| Phaeodiaporthe Petr. |
| Pustulomyces D.Q. Dai et al. |
| Stenocarpella Syd. & P. Syd. |
| Erythrogloeaceae Senan. et al. |
| Chrysocrypta Crous & Summerell |
| Disculoides Crous et al. |
| Erythrogloeum Petr. |
| Gnomoniaceae G. Winter |
| Alnecium Voglmayr & Jaklitsch |
| Ambarignomonia Sogonov |
| Amphiporthe Petr. |
| Anisomyces Theiss. & Syd. |
| Apiognomonia Höhn. |
| Apioplagiostoma M.E. Barr |
| Asteroma DC |
| Bagcheea E. Müll. & R. Menon |
| Clypeoporthe Höhn. |
| Cryptosporella Sacc. |
| Cylindrosporella Höhn. |
| Diplacella Syd. |
| Ditopella De Not. |
| Ditopellopsis J. Reid & C. Booth |
| Gloeosporidina Petr. |
| Gnomonia Ces. & De Not. |
| Gnomoniella Sacc. |
| Gnomoniopsis Berl. |
| Marsupiomyces Senan. & K.D. Hyde |
| Millerburtonia Cif. |
| Occultocarpon L.C. Mejía et al. |
| Ophiognomonia (Sacc.) Sacc. |
| Phragmoporthe Petr. |
| Phylloporthe Syd. |
| Plagiostoma Fuckel |
| Pleuroceras Riess. |
| Sirococcus Preuss |
| Spataporthe Bronson et al. |
| Uniseta Ciccar |
| Valsalnicola D.M. Walker & Rossman |
| Harknessiaceae Crous |
| Dwiroopa Subram. & Muthumary |
| Harknessia Cooke |
| Juglanconidaceae Voglmayr & Jaklitsch |
| Juglanconis Voglmayr & Jaklitsch |
| Lamproconiaceae C. Norphanphoun et al. |
| Hercospora Fr. |
| Lamproconium (Grove) Grove |
| Macrohilaceae Crous |
| Macrohilum H.J. Swart |
| Melanconidaceae G. Winter |
| Melanconis Tul. & C. Tul. |
| Melanconiellaceae Senan. et al. |
| Dicarpella Syd. |
| Greeneria Scribn. & Viala |
| Melanconiella Sacc. |
| Microascospora Senan. & K.D. Hyde |
| Tubakia B. Sutton |
| Prosopidicolaceae Senan. & K.D. Hyde |
| Prosopidicola Crous & C.L. Lennox |
| Pseudoplagiostomataceae Cheew. et al. |
| Pseudoplagiostoma Cheew. et al. |
| Schizoparmaceae Rossman DF et al. |
| Coniella Höhn. |
| Stilbosporaceae Link |
| Crinitospora B. Sutton & Alcorn |
| Stegonsporium Corda |
| Stilbospora Pers. |
| Sydowiellaceae Lar.N. Vassiljeva |
| Alborbis Senan. & K.D. Hyde |
| Breviappendix Senan. & K.D. Hyde |
| Cainiella E. Müll |
| Calosporella J. Schröt |
| Chapeckia M.E. Barr |
| Italiomyces Senan. et al. |
| Hapalocystis Auersw. ex Fuckel |
| Lambro Racib. |
| Paragnomonia Senan. & K.D. Hyde |
| Ranulospora Senan. et al. |
| Rossmania Lar.N. Vassiljeva |
| Sillia P. Karst. |
| Sydowiella Petr. |
| Tenuiappendicula Senan. et al. |
| Tortilispora (Sacc.) Senan. & K.D. Hyde |
| Diaporthales genera incertae sedis |
| Anisomycopsis I. Hino & Katum. |
| Caudospora Starbäck |
| Chadefaudiomyces Kamat et al. |
| Cryptascoma Ananthap. |
| Cryptoleptosphaeria Petr. |
| Cytomelanconis Naumov |
| Dictyoporthe Petr. |
| Ditopellina J. Reid & C. Booth |
| Durispora K.D. Hyde |
| Fremineavia Nieuwl. |
| Hypodermina Höhn. |
| Hypophloeda K.D. Hyde & E.B.G. Jones |
| Kapooria J. Reid & C. Booth |
| Keinstirschia J. Reid & C. Booth |
| Lollipopaia Inderbitzin |
| Macrodiaporthe Petr. |
| Maculatipalma J. Fröhlich & K.D. Hyde |
| Massariovalsa Sacc. |
| Mebarria J. Reid & C. Booth |
| Melanamphora Lafl. |
| Melanconiopsis Ellis & Everh. |
| Natarajania Pratibha & Bhat |
| Phaeoappendicospora Senan. et al. |
| Phragmodiaporthe Wehm. |
| Plagiophiale Petr. |
| Plagiostigme Syd. |
| Prostratus Sivan. et al. |
| Pseudocryptosporella J. Reid & C. Booth |
| Pseudothis Theiss. & Syd. |
| Pseudovalsella Höhn. |
| Rabenhorstia Fr. |
| Savulescua Petr. |
| Skottsbergiella Petr. |
| Stioclettia Dennis |
| Trematovalsa Jacobesco |
| Uleoporthe Petr. |
| Vismaya V.V. Sarma & K.D. Hyde |
| Wehmeyera J. Reid & C. Booth |
| Wuestneia Auersw. ex Fuckel |
| Wuestneiopsis J. Reid & Dowsett |
Acknowledgements
We thank Elias Jonk, Westerdijk Fungal Biodiversity Institute, for DNA isolation and sequencing of some of the included CBS cultures. Alan J.L. Phillips acknowledges the support from Biosystems and Integrative Sciences Institute (BioISI, FCT/UID/ Multi/04046/2013). Kevin D. Hyde thanks to National Research Council of Thailand (Mae Fah Luang University) for the grant “Biodiversity, phylogeny and role of fungal endophytes of Pandanaceae” (Grant No: 592010200112) and Thailand Research Fund (TRF) grant no RSA5980068 entitled “Biodiversity, phylogeny and role of fungal endophytes on above parts of Rhizophora apiculata and Nypa fruticans”, the National Research Council of Thailand (Mae Fah Luang University) grant no 60201000201 entitled “Diseases of mangrove trees and maintenance of good forestry practice”, and the Chinese Academy of Sciences, project number 2013T2S0030, for the award of Visiting Professorship for Senior International Scientists at Kunming Institute of Botany. Samantha C. Karunarathna thanks to Yunnan Provincial Department of Human Resources and Social Security funded postdoctoral project (number 179122).
Footnotes
Peer review under responsibility of Westerdijk Fungal Biodiversity Institute.
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