Research Article |
Corresponding author: Martina Réblová ( martina.reblova@ibot.cas.cz ) Academic editor: Thorsten Lumbsch
© 2020 Martina Réblová, Jana Nekvindová, Jacques Fournier, Andrew N. Miller.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Réblová M, Nekvindová J, Fournier J, Miller AN (2020) Delimitation, new species and teleomorph-anamorph relationships in Codinaea, Dendrophoma, Paragaeumannomyces and Striatosphaeria (Chaetosphaeriaceae). MycoKeys 74: 17-74. https://doi.org/10.3897/mycokeys.74.57824
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The Chaetosphaeriaceae are a diverse group of pigmented, predominantly phialidic hyphomycetes comprised of several holomorphic genera including Chaetosphaeria, the most prominent genus of the family. Although the morphology of the teleomorphs of the majority of Chaetosphaeria is rather uniform, their associated anamorphs primarily exhibit the variability and evolutionary change observed in the genus. An exception from the morphological monotony among Chaetosphaeria species is a group characterised by scolecosporous, hyaline to light pink, multiseptate, asymmetrical ascospores and a unique three-layered ascomatal wall. Paragaeumannomyces sphaerocellularis, the type species of the genus, exhibits these morphological traits and is compared with similar Chaetosphaeria with craspedodidymum- and chloridium-like synanamorphs. Morphological comparison and phylogenetic analyses of the combined ITS-28S sequences of 35 isolates and vouchers with these characteristics revealed a strongly-supported, morphologically well-delimited clade in the Chaetosphaeriaceae containing 16 species. The generic name Paragaeumannomyces is applied to this monophyletic clade; eight new combinations and five new species, i.e. P. abietinus sp. nov., P. elegans sp. nov., P. granulatus sp. nov., P. sabinianus sp. nov. and P. smokiensis sp. nov., are proposed. A key to Paragaeumannomyces is provided. Using morphology, cultivation studies and phylogenetic analyses of ITS and 28S rDNA, two additional new species from freshwater and terrestrial habitats, Codinaea paniculata sp. nov. and Striatosphaeria castanea sp. nov., are described in the family. A codinaea-like anamorph of S. castanea forms conidia with setulae at each end in axenic culture; this feature expands the known morphology of Striatosphaeria. A chaetosphaeria-like teleomorph is experimentally linked to Dendrophoma cytisporoides, a sporodochial hyphomycete and type species of Dendrophoma, for the first time.
molecular phylogeny, phialidic conidiogenesis, scolecosporous, systematics, wood-inhabiting fungi, 15 new taxa
The family Chaetosphaeriaceae (
Sexually reproducing fungi encompassed in the Chaetosphaeriaceae are perithecial ascomycetes that share several morphological traits such as similar anatomy of the brittle, melanised ascomatal wall, persistent paraphyses, unitunicate, thin-walled asci with a refractive, non-amyloid apical annulus, transversely septate ascospores that germinate by germ tubes and phialidic conidiogenesis. Several species produce both ascospores and conidia, ascomata are often associated with conspicuous conidiophores arranged in the juxtaposition. Most representatives of the family reproduce only asexually and are known as “anamorphic holomorphs” (
Most of the sexually reproducing fungi in the family were classified in Chaetosphaeria (Tulasne & Tulasne, 1863), a prominent genus of the family. Chloridium botryoideum has long been known to be a part of the life cycle of Ch. innumera, the generic type (
The majority of species accommodated in Chaetosphaeria possess ellipsoidal, fusiform to cylindrical-fusiform, 1–5-septate, hyaline, symmetrical ascospores with their length generally ranging from 6 to 40 μm. Ascomata are brown to black, papillate, often glossy with a two-layered ascomatal wall; the outer layer consisting of several rows of brick-like cells with dark brown, opaque walls. The transfer of a scolecosporous Lasiosphaeria raciborskii (
Our sampling of saprobic lignicolous fungi in terrestrial biotopes in various localities in Europe, New Zealand and North America revealed several species whose morphological characters best match those of the genus Paragaeumannomyces and other scolecosporous Chaetosphaeria, i.e. Ch. albida (
Dendrophoma (
The present study provides new data that improve our understanding of morphological and genetic diversity of the Chaetosphaeriaceae and its pleomorphism. Our longer term goals focus on identification of monophyletic, morphologically well-delimited natural lineages and the life history of species currently assigned to the family. To assess phylogenetic relationships of our isolates, we based the study on morphological and cultivation studies along with the analysis of DNA sequence data from the nuc rDNA internal transcribed spacer region (ITS1-5.8S-ITS2 = ITS) and nuclear large subunit 28S ribosomal DNA gene (28S).
Material for this study was collected in north temperate regions of Europe (France, Germany and Ukraine) and North America (North Carolina, Tennessee), south subtropical and temperate climate zones of New Zealand, and in the neotropical regions of the Caribbean (Puerto Rico) and South America (French Guiana). An additional living culture was obtained from BCCM/
Taxa, isolate information and GenBank accession numbers for new sequences (in bold) determined for this study.
Taxon | Specimen | Status | Country | Host | Substrate | GenBank accessions | Reference | |
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ITS | 28S | |||||||
Codinaea paniculata |
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T | France | unidentified | submerged decaying wood | MT118230 | MT118201 | This study |
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France | Alnus glutinosa | submerged decaying wood | MT118231 | MT118202 | This study | ||
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France | Fraxinus excelsior | submerged decaying wood | MT118232 | MT118203 | This study | ||
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United Kingdom | unidentified | submerged dead leaf | MT118233 | MT118204 | This study | ||
Dendrophoma cytisporoides |
IMI 506817 |
Germany | Buxus sempervivens | decaying periderm of a twig | MT118234 | MT118205 | This study | |
Paragaeumannomyces abietinus |
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T | France | Abies alba | decaying wood | MT118235 | MT118206 | This study |
Paragaeumannomyces albidus | PDD 118738 | New Zealand | unidentified | decaying wood | MT876579 | – | This study | |
Paragaeumannomyces elegans | PDD 118740 | T | New Zealand | unidentified | decaying wood | MT876580 | – | This study |
Paragaeumannomyces granulatus | ICMP 15133 | T | New Zealand | unidentified | decaying wood | MT876575 | MT876577 | This study |
PDD 118745 | New Zealand | unidentified | decaying wood | MT876576 | MT876578 | This study | ||
Paragaeumannomyces lapazianus | S.M.H. 2182 | Costa Rica | unidentified | decaying wood | AY906945 | MT118207 |
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S.M.H. 2900 | Puerto Rico | unidentified | decaying wood | AY906946 | MT118208 |
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S.M.H. 3043 | Puerto Rico | unidentified | decaying wood | AY906947 | MT118209 |
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Paragaeumannomyces longisporus | A.N.M. 1269 | USA, Tennessee | unidentified | decaying wood | MT118239 | MT118210 | This study | |
ILLS00121385 | USA, Tennessee | unidentified | decaying wood | MT118237 | MT118211 | This study | ||
ILLS00121386 | USA, Tennessee | unidentified | decaying wood | MT118238 | MT118212 | This study | ||
S.M.H. 2519 | USA, Indiana | unidentified | decaying wood | AY906939 | MT118213 |
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S.M.H. 2758 | USA, North Carolina | unidentified | decaying wood | AY906940 | MT118214 |
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S.M.H. 3805 | USA, North Carolina | unidentified | decaying wood | MT118236 | MT118215 | This study | ||
S.M.H. 3809 | USA, North Carolina | unidentified | decaying wood | AY906942 | MT118216 |
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S.M.H. 3860 | USA, South Carolina | unidentified | decaying wood | AY906944 | MT118217 |
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Paragaeumannomyces panamensis | S.M.H. 3596 | T | Panama | unidentified | decaying wood | AY906948 | MT118218 |
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Paragaeumannomyces sp. 1 | S.M.H. 2025 | Puerto Rico | unidentified | decaying wood | MT118241 | MT118219 | This study | |
S.M.H. 3014 | Puerto Rico | unidentified | decaying wood | AY906952 | MT118222 |
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Paragaeumannomyces sp. 2 | S.M.H. 2036 | Puerto Rico | unidentified | decaying wood | AY906950 | MT118220 |
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S.M.H. 2132 | Puerto Rico | unidentified | decaying wood | AY906951 | MT118221 |
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Paragaeumannomyces rubicundus | S.M.H. 2881 | PT | Puerto Rico | unidentified | decaying wood | AY906954 | MT118223 |
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S.M.H. 3221 | T | Costa Rica | unidentified | decaying wood | MT118242 | MT118224 | This study | |
Paragaeumannomyces sabinianus | ILLS00121384 | T | USA, Tennessee | unidentified | decaying wood | MT118243 | MT118225 | This study |
S.M.H. 3807 | USA, North Carolina | unidentified | decaying wood | AY906941 | MT118226 |
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S.M.H. 3824 | USA, North Carolina | unidentified | decaying wood | AY906943 | MT118227 |
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Paragaeumannomyces smokiensis | ILLS00121398 | T | USA, Tennessee | unidentified | decaying wood | MT118240 | MT118228 | This study |
Striatosphaeria castanea |
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T | French Guinea | woody liana | decaying periderm | MT118244 | MT118229 | This study |
Striatosphaeria codinaeophora | S.M.H. 1524 | Puerto Rico | Nectandra turbacensis | decaying wood | MT118245 | AF466088 |
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Morphological characteristics were obtained from fungi growing on natural substrate and growth media. Descriptions in the key are based on fungi growing on natural substrate. Herbarium material was rehydrated with tap water and examined with an Olympus SZX12 dissecting microscope (Olympus America, Inc., Melville, USA). Hand-sectioned ascomata and centrum material (asci, ascospores and paraphyses), conidiophores and conidia were mounted in 90 % lactic acid, Melzer’s reagent, and lactophenol with cotton blue. All measurements were in Melzer’s reagent. Means ± standard deviation (SD) based on a minimum of 20–25 measurements are given for dimensions of asci, ascospores and conidia. Micromorphological observations were made using an Olympus BX51 compound microscope with differential interference contrast (DIC) and phase contrast (PC) illumination. Images of microscopic structures were captured with an Olympus DP70 camera operated by Imaging Software Cell^D (Olympus). Macroscopic images of colonies were documented using a Canon EOS 77D digital camera with Canon EF 100mm f/2.8L Macro IS USM objective with daylight spectrum 5500K 16W LED lights (Canon Europe Ltd., Middlesex, United Kingdom). All images were processed with Adobe Photoshop CS6 (Adobe Systems, San Jose, USA).
For comparative purposes, strains were inoculated in triplicate on cornmeal dextrose agar (CMD) [17 g of cornmeal agar (Oxoid Limited, Hampshire, United Kingdom), 2 g of dextrose, 1 L of distilled water, sterilized for 15 min at 121 C], Modified Leonian’s agar (MLA) (
Methods for the DNA extraction and amplification of samples with A.N.M.,
PCR was carried out in a BioRad C1000 thermal cycler (Bio-Rad Laboratories Inc., USA) as follows: 98 C for 30 s; 40 cycles of denaturation (98 C for 10 s), annealing (62 C for 30 s) and elongation (72 C for 90 s) and a final extension step at 72 C for 5 min. Amplicons were purified from agarose gels using a NucleoSpin® Gel and PCR Clean-up Kit (Macherey-Nagel GmbH & Co. KG, Germany) following the manufacturer’s instructions, with an elution volume of 25 μL. The DNA concentration was assessed fluorimetrically using Quant-iT PicoGreen dsDNA Assay Kit and Qubit fluorometer (Invitrogen / Thermo Fisher Scientific, USA) to assure required sequencing concentrations adjusted for the length of amplicons/ number of reads required.
Each of the amplicons was sequenced in both directions using the PCR primers and nested primers: ITS5, ITS4, JS1, JS7, JS8 and LR7 for ITS-28S (
Two gene markers, ITS and 28S rDNA, were analysed to assess evolutionary relationships of the unknown fungi with members of the Chaetosphaeriaceae. Consensus secondary structure (2D) models for the ITS1 and ITS2 for members of the Chaetosphaeriaceae were built using the Ppfold program v.3.0 (
ITS and 28S sequences were aligned manually in Bioedit v.7.1.8 (
Three analyses were employed to estimate phylogenetic relationships. Bayesian Inference (BI) and Maximum Likelihood (ML) analyses were performed through the CIPRES Science Gateway v.3.3. ML analyses were conducted with RAXML-HPC v.8.2.12 (
Taxa, isolate information and accession numbers for sequences retrieved from GenBank.
Taxon | Strain | Status | Country | Host | Substrate | GenBank accessions | Reference | |
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ITS | 28S | |||||||
Adautomilanezia caesalpiniae | CC-LAMIC 102/12 | T | Brazil | Caesalpina echinata | wood | KX821777 | KU170671 |
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Anacacumisporium appendiculatum | HMAS 245593 | T | China, Hainan | broad-leaved tree | dead stems | KP347129 | KT001553 |
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Brunneodinemasporium brasiliense |
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T | Brazil | unidentified | decaying leaf | JQ889272 | JQ889288 |
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Brunneodinemasporium jonesii | GZCC 16-0050 | T | China | unidentified | decaying wood | KY026058 | KY026055 |
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Cacumisporium capitulatum | FMR 11339 | Spain | unidentified | decaying wood | HF677176 | HF677190 |
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Calvolachnella guaviyunis |
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T | Uruguay | Myrcianthes pungens | bark | KJ834524 | KJ834525 |
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Chaetosphaeria chlorotunicata | S.M.H. 1565 | T | Puerto Rico | unidentified | decaying wood | – | AF466064 |
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Chaetosphaeria innumera | M.R. 1175 | Czech Republic | Fagus sylvatica | decaying wood | AF178551 | AF178551 |
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Chaetosphaeria lignomollis | S.M.H. 3015 | T | Puerto Rico | unidentified | decaying wood | EU037896 | AF466073 |
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Chaetosphaeria myriocarpa |
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The Netherlands | unidentified | decaying wood | AF178552 | AF178552 |
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Chaetosphaeria pygmaea | M.R. 1365 | Czech Republic | Fagus sylvatica | decaying wood | AF178545 | AF178545 |
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Chloridium caesium |
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Austria | Salix cinerea | decaying wood | AF178564 | AF178564 |
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Chloridium gonytrichii |
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South Africa | unidentified | unknown | MH857954 | MH869503 |
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Chloridium virescens |
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France | Acer sp. | unknown | MH857142 | MH868678 |
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Codinaea acaciae |
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T | Malaysia, Sarawak | Acacia mangium | leaf spot | KR476732 | – |
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Codinaea lambertiae |
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T | Australia, N.S. Wales | Lambertia formosa | leaves | MG386052 | MG386105 |
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Codinaea pini |
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T | Uganda | Pinus patula | dead needles | KP004465 | KP004493 |
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Codinaea simplex |
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The Netherlands | Quercus sp. | cupule | AF178559 | AF178559 |
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Codinaeopsis gonytrichodes |
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Japan | unidentified | decaying plant material | AF178556 | AF178556 |
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Conicomyces pseudotransvaalensis | HHUF 29956 | T | Japan | Machilus japonica | dead twig | LC001710 | LC001708 |
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Cryptophiale udagawae | GZCC 18-0047 | China, Guizhou | unidentified | decaying wood | MN104608 | MN104619 |
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Dendrophoma cytisporoides |
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ET | The Netherlands | Rhododendron sp. | branches and twigs | JQ889273 | JQ889289 |
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Dictyochaeta assamica |
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Guadeloupe | Musa sp. | root | MH858788 | MH870426 |
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Dictyochaeta callimorpha | ICMP 15130 | New Zealand | unidentified | decaying wood | MT454483 | MT454498 |
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Dictyochaeta cangshanensis | MFLUCC 17-2214 | T | China, Yunnan | unidentified | submerged decaying wood | MK828632 | MK835832 |
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Dictyochaeta ellipsoidea | MFLUCC 18-1574 | T | China, Yunnan | unidentified | submerged decaying wood | MK828628 | MK835828 |
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Dictyochaeta fuegiana | ICMP 15153 | T | New Zealand | unidentified | decaying wood | MT454487 | EF063574 |
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Dictyochaeta lignicola | DLUCC 0899 | T | China, Yunnan | unidentified | submerged decaying wood | MK828630 | MK835830 |
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Dictyochaeta pandanicola | KUMCC 16-0153 | T | China, Yunnan | Pandanus sp. | decayng leaf | MH388338 | MH376710 |
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Dictyochaeta septata |
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ET | Chile | Eucalyptus grandis × urophylla | leaves | MH107889 | MH107936 |
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Dictyochaeta siamensis | MFLUCC 15-0614 | T | Thailand | unidentified | submerged decaying twig | KX609955 | KX609952 |
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Dictyochaeta terminalis | GZCC 18-0085 | T | China, Guizhou | unidentified | decaying leaves | MN104613 | MN104624 |
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Dinemasporium americanum |
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T | USA, Iowa | n/a | soil of tallgrass prairie | JQ889274 | JQ889290 |
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Dinemasporium pseudoindicum |
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T | USA, Kansas | n/a | soil of tallgrass prairie | JQ889277 | JQ889293 |
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Ellisembia aurea |
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T | France | Sambucus nigra | decaying wood | MH836375 | MH836376 |
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Ellisembia folliculata |
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France | Salix sp. | decaying wood | – | AF261071 | Réblová and Winka (2001) | |
Eucalyptostroma eucalypti |
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T | Malaysia | Eucalyptus pellita | leaf spots | KY173408 | KY173500 |
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Exserticlava vasiformis | TAMA 450 | Japan, Chiba | unidentified | plant debris | – | AB753846 | Tsuchiya et al., unpublished | |
Infundibulomyces cupulatus | BCC 11929 | T | Thailand | Lagerstroemia sp. | dead leaf | EF113976 | EF113979 |
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Infundibulomyces oblongisporus | BCC 13400 | T | Thailand | unidentified, angiosperm | leaf litter | EF113977 | EF113980 |
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Kionochaeta castaneae | GZCC 18-0025 | T | China | Castanea mollissima | decaying seed shell | MN104610 | MN104621 |
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Kionochaeta microspora | GZCC 18-0036 | T | China, Guizhou | unidentified | decaying wood | MN104607 | MN104618 |
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Leptosporella arengae | MFLUCC 15-0330 | T | Thailand | Arenga pinnata | dead rachis | MG272255 | MG272246 |
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Leptosporella bambusae | MFLUCC 12-0846 | T | Thailand | bamboo | dead culms | KU940134 | KU863122 |
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Menispora ciliata |
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T | Czech Republic | Acer campestre | decaying wood | EU488736 | – |
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Menispora tortuosa | DAOM 231154 | unknown | unidentified | unknown | KT225527 | AY544682 |
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Menisporopsis dushanensis | GZCC 18-0084 | T | China, Guizhou | unidentified | decaying leaves | MN104615 | MN104626 |
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Menisporopsis theobromae | MFLUCC 15-0055 | Thailand | unidentified | submerged decaying wood | KX609957 | KX609954 |
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Nawawia filiformis | MFLUCC 17-2394 | Thailand | unidentified | decaying wood | MH758196 | MH758209 |
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Neopseudolachnella acutispora | MAFF 244358 | T | Japan, Aomori | Pleioblastus chino | dead twigs | AB934065 | AB934041 |
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Neopseudolachnella magnispora | MAFF 244359 | T | Japan, Aomori | Sasa kurilensis | dead twigs | AB934066 | AB934042 |
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Paliphora intermedia |
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IST | Australia, Queensland | unidentified | leaf litter | MH862682 | EF204501 |
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Paragaeumannomyces albidus | PDD 92537 | T | New Zealand | Nothofagus sp. | decaying wood | EU037890 | EU037898 |
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P. albidus | PDD 92540 | New Zealand | Nothofagus sp. | decaying wood | EU037891 | – |
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Paragaeumannomyces bombycinus | PDD 92538 | T | New Zealand | Nothofagus sp. | decaying wood | EU037892 | – |
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Paragaeumannomyces elegans | PDD 92561 | New Zealand | unidentified | decaying wood | EU037895 | – |
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Paragaeumannomyces garethjonesii | MFLUCC 15-1012 | T | Thailand | Fabaceae | seed pod | KY212751 | KY212759 |
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Paragaeumannomyces panamensis | MFLUCC 15-1011 | Thailand | Pinus sp. | decaying wood | KY212752 | KY212760 |
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Paragaeumannomyces sp. 3 | S.M.H. 2017 | Puerto Rico | unidentified | decaying wood | AY906949 | AF466078 |
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Paragaeumannomyces sp. 4 | S.M.H. 3119 | Puerto Rico | unidentified | decaying wood | AY906953 | AY436402 |
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Phialosporostilbe scutiformis | MFLUCC 17-0227 | T | China | unidentified | submerged decaying wood | MH758194 | MH758207 |
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Polynema podocarpi |
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T | New Zealand | Podocarpus totara | unknown | MH327797 | MH327833 |
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Pseudodinemasporium fabiforme |
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Malaysia, Sarawak | Acacia mangium | leaf spots | KR611889 | KR611906 |
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Pseudolachnea fraxini |
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T | Sweden | Fraxinus excelsior | unknown | JQ889287 | JQ889301 |
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Pseudolachnea hispidula | MAFF 244365 | Japan, Aomori | Morus bombycis | dead twig | AB934072 | AB934048 |
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Pseudolachnella asymmetrica | MAFF 244366 | Japan, Fukuoka | Phyllostachys nigra var. henonis | dead twig | AB934073 | AB934049 |
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Pseudolachnella scolecospora | MAFF 244379 | Japan, Gifu | Sasa sp. | dead twigs | AB934086 | AB934062 |
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Pyrigemmula aurantiaca |
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T | Hungary | Vitis vinifera | bark | HM241692 | HM241692 |
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Sporoschisma longicatenatum | MFLUCC 16-0180 | T | Thailand | unidentified | submerged decaying wood | KX505871 | KX358077 |
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Sporoschisma mirabile | FMR 11247 | Spain | unidentified | dead wood | HF677174 | HF677183 |
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Striatosphaeria castanea | monte6.2 | Brazil | Encyclia ghillanyi | root | KC928368 | – | Almeida et al., unpublished | |
Striatosphaeria codinaeophora | M.R. 1230 | Puerto Rico | Dacryodes excelsa | decaying wood | AF178546 | AF178546 |
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Tainosphaeria jonesii | GZCC 16-0065 | PT | China, Guangxi | unidentified | submerged decaying wood | KY026060 | KY026057 |
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Tainosphaeria siamensis | MFLUCC 15-0607 | T | Thailand | unidentified | submerged decaying wood | KX609956 | KX609953 |
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Thozetella nivea | n/a | unknown | unidentified | unknown | EU825201 | EU825200 |
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Thozetella tocklaiensis |
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T | India | Camellia sinensis | decaying flower | MH857817 | MH869349 |
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Tracylla aristata |
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ET | Australia, Victoria | Eucalyptus regnans | leaf | KX306770 | KX306795 |
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Tracylla eucalypti |
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T | Colombia | Eucalyptus urophylla | spots on living leaves | MH327810 | MH327846 |
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Zanclospora iberica |
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T | Spain | unidentified | decaying wood | KY853480 | KY853544 |
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Employing the predicted 2D structure of the variable ITS region and 28S enabled us to construct a reliable multiple sequence alignment of homologous positions at both helices and loops, thus eliminating potential ambiguous regions in the alignments. Initially, we compared trees from ML phylogenetic analyses of the two combined data sets (Chaetosphaeriaceae and scolecosporous species of Chaetosphaeria) after alignments were improved with the 2D structure, with and without applying Gblocks (
Evolutionary relationships of studied fungi were evaluated in the phylogenetic analysis based on the combined ITS and 28S sequences of 87 representative species of the Chaetosphaeriaceae. Leptosporella arengae, L. bambusae (Leptosporellaceae), and Tracylla eucalypti and T. aristata (Tracyllaceae) were used to root the tree. 76 nucleotides (nt) at the 5'-end and 606 nt at the 3'-end of 28S were excluded from the alignment because of missing data in the majority of sequences. The alignment had 1778 characters including gaps and 882 unique character sites (RAxML). In the MP analysis, 1021 characters were constant (proportion = 57.42 %), 134 variable characters were parsimony-uninformative, 623 characters were parsimony-informative (included); two most parsimonious trees were produced (length = 5066 steps, consistency index = 0.0.298, homoplasy index = 0.702, retention index = 0.631). For the BI analysis, GTR+I+G model was selected for ITS and 28S partitions. The ML tree (RAxML) is shown in Fig.
Combined phylogeny using ITS and 28S of selected members of the Chaetosphaeriaceae. Species names given in bold are taxonomic novelties; T, ET, IST and PT indicate ex-type, ex-epitype, ex-isotype and ex-paratype strains. Thickened branches indicate branch support with ML BS = 100%, PP values = 1.0 and MP = 100 %. Branch support of nodes ≥ 75 % ML and MP BS, and ≥ 0.95 PP is indicated above branches.
Phylogenetic relationships within the genus Paragaeumannomyces were assessed in the second analysis of the combined ITS-28S loci. Chaetosphaeria fusiformis and Ch. lignomollis (Chaetosphaeriaceae) were used to root the tree, and thus served as outgroup. The analysis included 35 sequences belonging to 16 species. 28 nt at the 5'-end and 714 nt at the 3'-end of 28S were excluded from the alignment due to missing data in the majority of sequences. The alignment had 1593 characters including gaps and 410 unique character sites (RAxML). In the MP analysis, 1247 characters were constant (proportion = 78.28 %), 78 variable characters were parsimony-uninformative, and 268 characters were parsimony-informative (included); 286 most parsimonious trees were produced (length = 832 steps, consistency index = 0.6118, homoplasy index = 0.3882, retention index = 0.8082). For the BI analysis, SYM+G and GTR+I+G models were selected for ITS and 28S partitions, respectively. The ML tree is shown in Fig.
Combined phylogeny using ITS and 28S of 35 members of Paragaeumannomyces. Species names given in bold are new species; T and PT indicate ex-type and ex-paratype strains. Thickened branches indicate branch support with ML BS = 100%, PP values = 1.0 and MP = 100 %. Branch support of nodes ≥ 75 % ML and MP BS, and ≥ 0.95 PP is indicated above branches.
France – Ariège • Pyrénées Mts., Rimont, La Maille brook; alt. 550 m; 28 May 2018 (incubated in moist chamber for 1 wk); on submerged decaying wood; J. Fournier leg.; M.R. 3950 (holotype:
Panicula (Latin) tuft, referring to the dense groups of setae and conidiophores on the natural substrate.
Colonies on the nature substrate effuse, hairy, greyish-brown. Setae erect, straight or slightly flexuous, smooth-walled, dark brown and thick-walled, becoming pale brown to subhyaline and thin-walled towards the apex, 230–290 μm long, 6–7.5 μm wide above the base, tapering gradually towards the apex which almost always develops into a monophialide. Conidiophores macronematous, mononematous, 62–127 × 3.5–4.5 µm, septate, erect, straight or flexuous, arising singly or in groups of 4–6 from hyphal cells associated with the bases of setae, septate, mid-brown to pale brown becoming gradually paler towards the apex. Conidiogenous cells 16.5–30(–38) × 3.5–5 μm, tapering to 1.5–2 μm just below the collarette, integrated, terminal, monophialidic, cylindrical to cylindrical-lageniform, subhyaline or pale brown at the base becoming hyaline to subhyaline towards the apex, smooth-walled; collarettes funnel-shaped, 3.5–4.5 μm wide, 1.5–2.5 μm deep. Conidia in slimy droplets, hyaline in mass, (11.5–)12–17 × (2–)2.5–3(–3.5) µm (mean ± SD = 14.7 ± 1.5 × 2.5 ± 0.3 µm), of two types, narrower and longer, 13.5–17(–17.5) × 2.5–3.5 μm (mean ± SD = 15.2 ± 1.0 × 2.8 ± 0.3 µm), and shorter and usually wider, 11.5–13.5(–14) × 3–3.5(–4) μm, falcate, asymmetrical, rounded at the apical end, with an inconspicuous scar at the basal end, hyaline, aseptate, smooth-walled, with simple, straight or gently curved setulae at both ends, 5–8 μm long; setulae inserted on the concave sides of the conidia.
Codinaea paniculata. A–C setae and conidiophores on nature substrate D–G conidia on nature substrate H–L conidiophores in MLA culture (6 wk) M–O conidia in MLA culture (6 wk) P colonies on CMD, MLA, OA and PCA after 4 wk (from left to right). Images:
Vegetative hyphae hyaline to pale brown. Setae absent. Conidiophores 95–150(–195) μm long, 3.5–4.5 μm wide, conidiogenous cells 25–35 × 3.5–4(–4.5) μm, tapering to 1.5 µm just below the collarette, integrated, terminal, polyphialidic, usually cylindrical, pale brown to subhyaline, smooth-walled; collarette funnel-shaped, 3.5–4(–4.5) μm wide, 1.5–2 μm deep. Conidia in slimy droplets, of two types, narrower and longer (13–)13.5–15.5(–17) × 2.5–3 µm (mean ± SD = 14.4 ± 0.9 × 2.7 ± 0.2 µm), usually slightly wider and shorter 11–13 × 2.5–3.5 µm (mean ± SD = 12.0 ± 0.7 × 3.1 ± 0.3 µm), falcate, asymmetrical, hyaline, with simple setulae 3.5–5.5(–7.5) μm long at both ends.
On CMD colonies 80–85 mm diam, circular, flat, margin fimbriate, aerial mycelium restricted mainly to the centre and margin of the colony, sparsely lanose, floccose centrally becoming mucoid towards the margin, cobwebby at the margin, colony centre whitish, pale brown to creamy towards the margin, pale brown pigment diffusing from the centre of the colony to the agar; reverse creamy. On MLA colonies 65–70 mm diam, circular, slightly raised, margin filiform, lanose, floccose, colony centre whitish becoming brown-grey towards the margin with a brown outer zone of submerged growth, pale brown pigment diffusing to the agar; reverse dark brown. On OA colonies 89–95 mm diam, circular, raised, margin filiform, aerial mycelium occasionally reduced or absent, colonies similar to those on MLA, lanose, floccose, locally mucoid and smooth or cobwebby, whitish becoming dark grey at the margin, a dark brown to burgundy brown pigment diffusing to the agar; reverse dark grey. On PCA colonies 78–89 mm diam, circular, flat to slightly raised, margin entire to weakly filiform, lanose, floccose, occasionally locally mucoid and smooth or with sparse decumbent aerial hyphae, cobwebby at the margin, whitish becoming brown towards the margin; reverse olivaceous brown. Sporulation on MLA, OA, CMD after 8 wk.
France – Ariège • Pyrénées Mts., Rimont, Le Baup stream, ca. 1.5 km from the village along D18 road; alt. 550 m; 12 Jun. 2009; on submerged wood of Fraxinus excelsior; J. Fournier leg.; J.F. 09153 (
All four isolates analysed in this study originated from the freshwater environment and occurred on decaying wood or leaves of Alnus glutinosa, Fraxinus excelsior and other unidentified hosts. Based on the BLASTn search of the ITS sequence of C. paniculata in GenBank, two isolates from roots of Elymus mollis (ITS: KU838460, KU839605,
Among known Codinaea species, C. assamica is similar to C. paniculata, but differs by slightly longer (14.6–16.8 × 2.6–2.8 µm) conidia with longer (9.6–12.8 µm) setulae (
Paragaeumannomyces sphaerocellularis Matsush., Mycol. Mem. 10: 156. (2003) [2001].
Teleomorph: Ascomata perithecial, non-stromatic, superficial, subglobose to conical, solitary, in small groups or aggregated, sometimes collapsing laterally upon drying, ranging from white, yellow-white, light fawn-grey, ginger-brown, reddish-brown, russet to dark brown, papillate, glabrous or setose, setae dark brown, acute, opaque, scattered over entire ascoma and/or clustered around the ostiole, centrum sometimes pink to pale red. Ostiole periphysate. Ascomatal wall three-layered; outer layer composed of thin-walled, globose, subglobose to polyhedral cells, sometimes containing pale purple pigment when fresh; middle layer composed of brick-like, dark brown cells with opaque walls; inner layer of flattened, thin-walled, subhyaline cells. Paraphyses persistent, branching, tapering. Asci unitunicate, 8-spored, cylindrical-fusiform, stipitate, apex with a non-amyloid apical annulus. Ascospores asymmetrical, cylindrical-filiform, slightly tapering towards the basal end, multiseptate, hyaline, occasionally light pink, with negative or positive dextrinoid reaction in Melzer’s reagent. Synanamorphs: Craspedodidymum-like. Conidiophores mononematous, semi-macronematous to micronematous, brown, septate, unbranched or reduced to single conidiogenous cells. Conidiogenous cells phialidic, obclavate or broadly lageniform, brown, with an apical opening; collarettes flared or cup-shaped. Conidia globose, subglobose, subangular to triangular, unicellular, hyaline, with setulae. Chloridium-like. Conidiophores mononematous, macronematous, brown, septate, unbranched. Conidiogenous cells phialidic, cylindrical, subhyaline, elongating percurrently, with an apical opening; collarette indistinct or flared. Conidia globose, ovoid to clavate, unicellular, hyaline, non-setulate, accumulating in slimy droplets. [Characteristics of the synanamorphs adopted from
The holotype of P. sphaerocellularis (Japan, Schimizu-cho, Wakayama Pref., on decaying twig of unknown broadleaf tree, Apr. 2000, MFC-21077), the type species of Paragaeumannomyces (
Members of Paragaeumannomyces display a wide geographical distribution pattern; they have a predominantly pantropical distribution in Central America and Asia but were also encountered in the subtropical and temperate climate zones of Europe, Japan, New Zealand and North America.
1 | Ascomata almost white, yellowish-white or light fawn-grey when fresh, dark yellow, buff, tawny to dark ginger-brown or yellow-grey when dried, purple pigment absent in cells of the outer ascomatal layer, setae absent | 2 |
– | Ascomata reddish, reddish-brown, russet or brown, sometimes with red surface crystals, globose cells in the outer ascomatal layer may contain pale purple pigment when fresh, setae present or absent | 3 |
2 | Ascomata almost white to yellowish-white, translucent, areolate when fresh, with a distinct black papilla, asci 270–295 × 18.5–20.5 μm, ascospores (3–)5–11-septate, (55–)69–86 × 5–6.5(–7) μm | P. albidus |
– | Ascomata light fawn-grey, not areolate when fresh, papilla indistinct or absent, asci 215–270 × 11–14 µm, ascospores (7–)11(–13)-septate, 62–88 × 4.5–6 µm | P. bombycinus |
3 | Ascomatal setae present only at the apex or absent | 4 |
– | Ascomatal setae present at the apex and also scattered over entire surface of the ascoma, or setae only scattered over ascoma, occasionally absent | 8 |
4 | Ascomata with red surface crystals, ascospores 7-septate, 80–100 × 3.5–4.2 μm, anamorph craspedodidymum-like, conidia globose to subglobose in a vertical position, with setulae | P. rubicundus |
– | Ascomata without red surface crystals, ascospores with seven or more septa | 5 |
5 | Ascomata more than 500 μm diam, setae occasionally absent, ascospores 7-septate, 50–100 × 4.5–6 μm, anamorph craspedodidymum-like, conidia oblate to horizontally oblong with a short abscission scar or frill, without setulae, tropical distribution | P. lapazianus |
– | Ascomata less than 500 μm diam, setae present at the apex, ascospores with seven or more septa, temperate and subtropical distribution | 6 |
6 | Ascospores (7–)11–13-septate, (90–)95–123.5 × 4–5(–5.5) μm, asci 210–295 × (16.5–)17–24.5 μm | P. granulatus |
– | Ascospores with up to 11 septa, 87 μm and shorter | 7 |
7 | Ascospores 9–11-septate, (58–)60.5–80.5 × (3–)3.5–4.5(–5) μm, asci (134–)140–174(–189) × 11–13(–14) μm | P. smokiensis |
– | Ascospores (5–)7–9(–11)-septate, (62–)65–87 × (3.5–)4–5.5 μm, asci (185–)195–240 × 12–14.5(–15.5) μm | P. abietinus |
8 | Setae present at the apex and also scattered over entire surface of ascoma, ascospores 7-septate | 9 |
– | Setae scattered over entire surface of ascoma, occasionally absent, ascospores with seven or more septa | 10 |
9 | Ascospores (50.4–)52.5–68 × 3.5–4.5 μm, craspedodidymum- and chloridium-like synanamorphs, conidia without setulae | P. longisporus |
– | Ascospores (64.5–)68.5–86.5(–88.5) × (3–)3.5–4.5 μm, anamorph unknown | P. sabinianus |
10 | Ascospores 7-septate | 11 |
– | Ascospores with more than seven septa | 14 |
11 | Asci up to 152 μm long | 12 |
– | Asci 150 μm and longer | 13 |
12 | Ascomatal setae 60–75 μm long, ascospores 65–75 × 3–4 µm, asci 123–140 × 10–11 µm, craspedodidymum-like anamorph, conidia without setulae, purple-pigmented aleuriospore-like cells present in culture | P. panamensis |
– | Ascomatal setae 38–47 μm long, ascospores 63.3–75 × 2.3–3.7 µm, asci 120–152 × 10.7–13.3 µm, anamorph unknown | P. garethjonesii |
13 | Ascospores (50–)60–100(–150) × 3–3.75(–4.5) µm, asci (150–)180–250(–350) × 10–20(–27) µm, craspedodidymum-like anamorph, conidia with or without setulae, some strains also with a chloridium-like synanamorph |
P. raciborskii s. lat. (fide |
– | Ascospores (57.5–)60–73(–75) × (3.5–)4–4.5(–5) μm, asci (152–)174–221(–227) × 10.5–15(–20) μm, anamorph unknown | P. elegans |
14 | Ascospores 5–10-septate, 65–90 × 3–4 µm, asci 105–125 × 10–12.5 µm | P. sphaerocellularis |
– | Ascospores 13–16-septate, 50–65 × 2–4 μm, asci 70–100 × 10–13 μm |
P. raciborskii s. str. (fide |
France – Ariège • Pyrénées Mts., Ustou, Cirque de Cagateille, path up to the La Hillette lake, mixed Abies forest; alt. 1550 m; 18 Jul. 2018; on decaying wood of a trunk of Abies alba; J. Fournier leg.; J.F. 18057 (holotype:
Referring to the host Abies alba.
Teleomorph: Ascomata perithecial, non-stromatic, superficial, solitary or in small groups, 350–450 μm diam, 360–500 μm high, broadly conical, collapsing laterally upon drying, finely roughened, dark reddish-brown, glabrous except for the black conical papilla, with dark brown, stiff, acute setae, 32–40 × 3–4 μm, densely clustered around the ostiole; centrum pink. Ostiole periphysate. Ascomatal wall leathery, three-layered. Outer layer of textura angularis, 33–58 μm thick, consisting of thin-walled, globose, subglobose to polyhedral, dark ginger-brown to reddish-brown cells, 6.5–11 μm diam, grading into smaller cells towards the exterior. Middle layer of textura prismatica, 18–25 μm thick, composed of thick-walled, polyhedral, dark brown, melanised cells. Inner layer of textura prismatica, 10–15 μm thick, composed of thin-walled, flattened and elongated hyaline cells. Paraphyses abundant, hyaline, sparsely branched, septate, 4.5–7 μm wide, tapering to 2–2.5 μm, longer than the asci. Asci (185–)195–240 × 12–14.5(–15.5) μm (mean ± SD = 209.2 ± 12.0 × 14.1 ± 0.8 μm), (145–)155–205 µm (mean ± SD = 172.6 ± 14.3 μm) long in the sporiferous part, cylindrical-fusiform, stipitate, apically rounded, ascal apex non-amyloid with a distinct apical annulus 3–3.5 μm wide, 2–3 μm high. Ascospores (62–)65–87 × (3.5–)4–5.5 μm, filiform to cylindrical, straight or slightly curved to sigmoid, hyaline, light pink in mass, with dextrinoid reaction in Melzer’s reagent turning reddish-brown except for the end cells which remain hyaline, (5–)7–9(–11)-septate, septa often unevenly distributed, not constricted or slightly constricted at the septa, especially at the septa above and below the middle, asymmetrical, rounded at the apical end, tapering towards the basal end, with one or two guttules in each cell, 2–3-seriate or 4-seriate and partially overlapping or 4-seriate forming two fascicles end to end. Anamorph: Unknown.
Paragaeumannomyces abietinus. A, B ascomata C, D, F vertical section of ascomal wall E vertical section of the ascomal wall and papilla with apical setae G, H ascospores I, J asci K, L ascal apex with apical ring M paraphyses N colonies on CMD, MLA, OA and PCA after 4 wk (from left to right). Images:
On CMD colonies 10–11 mm diam, circular, slightly convex, margin entire to weakly fimbriate, lanose, beige-brown with a dark brown outer zone of submerged growth, dark brown pigment diffusing from the colony margin to agar; reverse dark brown to black. On MLA colonies 12–15 mm diam, circular, slightly convex, margin entire, lanose, floccose, cobwebby at the margin, beige-brown with a dark brown outer zone of submerged growth, brown pigment diffusing from the colony margin to agar; reverse dark brown. On OA colonies 8–9 mm diam, circular, convex, margin entire, lanose, beige-brown, with a paler outer ring; reverse brown. On PCA colonies 14–15 mm diam, circular, convex, margin entire, lanose, floccose, cobwebby towards the margin, beige, pale brown towards the margin; reverse brown. Sporulation absent on all media, even after prolonged incubation (> 3 mo).
Ukraine • Carpathian Mts., Kvasi, Bliznica near Rachiv, right bank of the upper flow of the Tisa river; alt. 1000 m; 28 Jun. 1997; on decaying wood of Abies alba; M. Réblová leg.; M.R. 946 (
All specimens of P. abietinus occur on decaying wood of Abies alba. The species has been collected in mountain areas and is known in Europe in France and Ukraine.
Attempts to cultivate this species were unsuccessful for the Ukrainian specimens; the ascospores germinated over five days with long inflated germ tubes from both ends but did not grow after isolation on agar medium. The axenic culture derived from the ascospore isolate of the French material yielded sterile mycelium only.
Paragaeumannomyces abietinus is similar to P. rubicundus and P. lapazianus in reddish-brown ascomata, the arrangement of setae around the ostiole and distribution in the north temperate region. Paragaeumannomyces rubicundus (
Chaetosphaeria albida T.J. Atk., A.N. Mill. & Huhndorf, New Zealand J. Bot. 45: 688. 2007.
New Zealand – Tasman • Tasman District, Abel Tasman National Park, Pigeon Saddle point, unpaved road between Tata Beach and Totaranui ca. 10 km NW of Totaranui; 24 Feb. 2003; on decaying wood of Nothofagus sp. buried in soil; M. Réblová leg.; M.R. 2605/NZ 76 (PDD 118737). – West Coast • Westland District, Arthur’s Pass National Park, Kelly Shelter ca. 5 km W of Otira, Cockayane Nature Walk, a podocarp-broadleaf forest; 16 Mar. 2003; on decaying wood of a trunk; M. Réblová leg.; M.R. 2840/NZ 351 (PDD 118738). • Ibid.; 16 Mar. 2003; on decaying wood and bark of a branch; M. Réblová leg.; M.R. 2843/NZ 355 (PDD 118739).
Paragaeumannomyces albidus has been collected on Metrosideros robusta, Metrosideros sp., Nothofagus sp. and other unidentified hosts and is known from New Zealand (
For additional illustrations and description, see
Paragaeumannomyces albidus is the only species of the genus characterised by a wide range of ascoma colours that change when ascomata are young and fresh or mature and dried. Different colours were used by
Attempts to isolate our specimens of P. albidus in living culture were not successful. Therefore, the DNA was extracted from herbarium material of all three collections, but only ITS1 of PDD 118738 could be amplified and sequenced. Comparison of the ITS1 sequences of our specimen and the holotype of P. albidus revealed 100 % similarity (Fig.
Chaetosphaeria bombycina T.J. Atk., A.N. Mill. & Huhndorf, New Zealand J. Bot. 45: 691. 2007.
The species has been collected on decaying wood of Nothofagus sp. and is known from New Zealand (
For description, illustration and holotype information, see
New Zealand – West Coast • Westland District, Mount Aspiring National Park, Haast, Roaring Billy track; 22 Mar. 2005; on decaying wood; M. Réblová leg.; M.R. 3295/NZ 566A (holotype: PDD 118740!).
Elegans (L) elegant, referring to elegant and lovely ascomata adorned with setae.
Teleomorph: Ascomata perithecial, non-stromatic, superficial, in small groups, often gregarious, 290–350 μm diam, 280–350 μm high, subglobose to slightly conical, dully glossy, brown with a light grey tinge except for the tiny black papilla composed of thick-walled, dark brown cells, ascomata densely setose, setae 28–60 × 4.5–6 μm, stiff, acute, dark brown, thick-walled, opaque. Ostiole periphysate. Ascomatal wall leathery, three-layered. Outer layer of textura angularis, 33–41 μm thick, consisting of thin-walled, globose to subglobose to polyhedral, reddish-brown cells ca. 5–12 μm diam. Middle layer of textura prismatica, 9.5–18 μm thick, composed of thick-walled, polyhedral, elongated, dark brown, melanised cells. Inner layer of textura prismatica, 5–8 μm thick, composed of thin-walled, flattened and elongated hyaline cells. Paraphyses abundant, hyaline, sparsely branched, septate, 3.5–5 μm wide, tapering to ca. 2 μm, longer than the asci. Asci (152–)174–221(–227) × 10.5–15(–20) μm (mean ± SD = 204.8 ± 13.7 × 12.3 ± 1.5 μm), (129–)141–197(–204) μm (mean ± SD = 168.2 ± 17.2 μm) long in the sporiferous part, cylindrical-fusiform, stipitate, apically rounded, ascal apex non-amyloid with a distinct apical annulus 2.5–3 μm wide, 2–2.5 μm high. Ascospores (57.5–)60–73(–75) × (3.5–)4–4.5(–5) μm (mean ± SD = 65.5 ± 3.2 × 4.1 ± 0.2 μm), filiform to cylindrical, straight or slightly curved to sigmoid, hyaline, with negative or very weak dextrinoid reaction in Melzer’s reagent, 7–septate, septa usually obscured by large guttules, not constricted at the septa, asymmetrical, rounded at the apical end, slightly tapering towards the basal end, with one or two guttules in each cell, 2–3-seriate or 3–4-seriate, partially overlapping. Anamorph: Unknown.
New Zealand – Otago • Clutha District, The Catlins, Catlins Coastal Rain Forest Park, MacLennan Range, Lake Wilkie Walk; 17 Mar. 2005; on decaying wood of a branch; M. Réblová leg.; M.R. 3289/NZ 549 (PDD 118742). – West Coast • Westland District, Ship Creek Point, Kahikatea Swamp Forest walk; 8 Mar. 2003; on decaying wood; M. Réblová leg.; M.R. 2819/NZ 329 (PDD 118741). – West Coast • Westland District, Ross, Totara Valley Road, 12 Apr. 2005; on decaying wood; M. Réblová leg.; M.R. 3486/NZ 775 (PDD 118743).
The present species is a saprobe on decaying wood of Nothofagus sp. and other unidentified hosts, known from New Zealand (
Paragaeumannomyces elegans is distinguishable from other members of the genus by densely setose, dull glistening brown ascomata with a light grey tinge, which gives them an almost grey appearance when dried. The new species resembles P. garethjonesii (
Comparison of the ITS sequence of the holotype of P. elegans with available Paragaeumannomyces sequences revealed 100 % sequence similarity with a specimen PDD 92561 (New Zealand, Taupo, Ohakune, ITS: EUO37895) tentatively identified as P. raciborskii (
Chaetosphaeria garethjonesii R.H. Perera, Maharachch. & K.D. Hyde, Mycosphere 7: 1308. 2016.
Paragaeumannomyces garethjonesii was collected on a Fabaceae seed pod and is known from Asia in Thailand (
For description, illustration and holotype information see
New Zealand – West Coast • Westland District, Hokitika, Mananui Point, Lake Mahinapua, Swimmers Beach walks; 5 Mar. 2003; on decaying wood; M. Réblová leg.; M.R. 2715/NZ 216 (holotype: PDD 118744!, ex-type culture ICMP 15133).
Granulum (L), granule, small grain, diminutive of granum, referring to the roughened surface of the ascomatal wall composed of globose cells, which appears granulose in the surface view.
Teleomorph: Ascomata perithecial, non-stromatic, superficial, solitary or in small groups, 380–495 μm diam, 415–530 μm high, subglobose to conical, finely roughened, dark brown to dark reddish-brown, sometimes with irregular reddish colour except for the black papilla; papilla composed of dark brown, thick-walled, cylindrical to subulate, apically narrowly rounded soft setae; centrum pink. Ostiole periphysate. Ascomatal wall leathery, three-layered. Outer layer of textura angularis, 95–115 μm thick, consisting of thin-walled, globose to subglobose ginger-brown cells ca. 27–33 μm diam, grading into smaller cells 8–16 μm diam. Middle layer of textura prismatica, 14–21 μm thick, composed of thick-walled, polyhedral, elongated, dark brown, melanised cells. Inner layer of textura prismatica, 7–12 μm thick, composed of thin-walled, flattened and elongated hyaline cells. Paraphyses abundant, hyaline, sparsely branched, septate, 3.5–5 μm wide, tapering to 2–2.5 μm, longer than the asci. Asci 210–295 × (16.5–)17–24.5 μm (mean ± SD = 239.7 ± 15.5 × 20.3 ± 2.1 μm), 165–200(–250) μm (mean ± SD = 184.7 ± 10.3 μm) long in the sporiferous part, cylindrical-fusiform, stipitate, apically rounded, ascal apex non-amyloid with a distinct apical annulus 3.5–4 μm wide, 2.5–3(–3.5) μm high. Ascospores (90–)95–123.5 × 4–5(–5.5) μm (mean ± SD = 101.4 ± 10.2 × 4.8 ± 0.4 μm), filiform to cylindrical, straight or slightly curved to sigmoid, hyaline to very light pink, light pink-brown in mass, with dextrinoid reaction in Melzer’s reagent turning reddish-brown except for the end cells which remain hyaline, (7–)11–13-septate, septa often unevenly distributed, not constricted or slightly constricted at the septa, asymmetrical, broadly rounded at the apical end, tapering and narrowly rounded at the basal end, with one or two guttules in each cell, 2–3-seriate or 4-seriate and partially overlapping. Anamorph: Unknown.
Paragaeumannomyces granulatus (PDD 118744). A, B ascomata C, D vertical section of ascomal wall E papilla and the upper part of ascomal wall in surface view F, G asci H ascal apex with apical annulus I paraphyses J ascospores K colonies on CMD, MLA, OA and PCA after 4 wk (from left to right). Scale bars: 250 μm (A, B); 100 μm (C); 25 μm (G); 20 μm (D–F, I); 10 μm (H, J); 1 cm (K).
On CMD colonies 14–16 mm diam, circular, convex, margin fimbriate, lanose, grey-brown, reverse dark brown to almost black. On MLA colonies 19–20 mm diam, circular, raised, margin entire to weakly fimbriate, lanose, beige-brown, with a dark brown outer zone, reverse dark brown to almost black. On OA colonies 13–16mm diam, circular, raised, margin weakly fimbriate, lanose, beige-grey becoming grey towards the periphery, reverse dark brown to almost black. On PCA colonies 15–17 mm diam, circular, slightly convex, margin weakly fimbriate, lanose, beige, pale brown at the margin, reverse black. Sporulation absent on all media.
New Zealand – Auckland • Auckland district, Waitakere Ranges Nature Reserve, Anawhata Road; 24 Apr. 2005; on decaying wood; M. Réblová leg.; M.R. 3543/NZ 838 (PDD 118745).
A saprobe on decaying wood, known from New Zealand.
Paragaeumannomyces granulatus most closely resembles P. abietinus in the ascoma appearance, pink content of the ascoma centrum, ascospores with usually more than seven septa and positive dextrinoid reaction in Melzer’s reagent but both species are separated by size of asci and ascospores. The ascospores of P. abietinus are (5–)7–9(–11)-septate and shorter [(62–)65–87 μm] and asci are shorter and narrower [(185–)195–240 × 12–14.5(–15.5) μm].
≡ Chaetosphaeria lapaziana (G.C. Carroll & Munk) F.A. Fernández & Huhndorf, Fung. Diver. 18: 49. 2005.
Lasiosphaeria lapaziana G.C. Carroll & Munk, Mycologia 56: 90. 1964.
Paragaeumannomyces lapazianus is common on decaying wood in the neotropics and is known from the Caribbean in Puerto Rico and Jamaica, from Central America in Costa Rica, and from South America in French Guiana (
For description, illustration and holotype information see
≡ Sphaeria longispora Ellis, Bull. Torrey Bot. Club 6: 135. 1877 non
≡ Ceratostomella longispora (Sacc.) Cooke, Grevillea 17: 50. 1889.
≡ Chaetosphaeria longispora (Sacc.) P.M. Kirk, Index Fung. 120: 1. 2014.
= Lasiosphaeria ellisii M.E. Barr, Mycotaxon 46: 48. 1993.
≡ Chaetosphaeria ellisii (M.E. Barr) Huhndorf & F.A. Fernández, Fung. Diver. 19: 27. 2005.
Ophioceras longisporum Sacc., Syll. fung. 2: 360. 1883.
USA – Tennessee • Cocke Co., Great Smoky Mountains National Park, Cosby, Cosby Nature Trail; alt. 716 m; 23 Mar. 2007; on decaying wood; A.N. Miller, P. Chaudhary & H.A. Raja leg.; A.N.M. 1134 (ILLS00121385). • Ibid.; 19 Jul. 2007; T.J. Atkinson & P. Chaudhary leg.; A.N.M. 1250 (ILLS00121386).
The species occurs on decaying wood and is known from the north temperate region in the USA (Indiana, New Jersey, North Carolina, South Carolina, Tennessee) (
For description and illustration, refer to
Sphaeria longispora (
Paragaeumannomyces longisporus. A, B ascomata C, D vertical section of ascomal wall E vertical section of ascomal wall and papilla with apical of setae F ascomal wall with setae G globose cells of the outer layer of the ascomal wall H, I asci J paraphyses K–N ascospores. Images: ILLS00121385 (A, B, G); S.M.H. 3860 (C, E, M); S.M.H. 2519 (D); ILLS00121386 (F, H–J, K); S.M.H. 2758 (L); S.M.H. 3809 (N). Scale bars: 250 μm (A–D); 50 μm (E–G); 20 μm (H–J); 10 μm (K–N).
Chaetosphaeria panamensis Huhndorf & F.A. Fernández, Fung. Diver. 19:33. 2005.
Paragaeumannomyces panamensis has been collected on decaying wood of Pinus sp. and an unidentified host, and it is known from Asia in Thailand and Central America in Panama (
For description, illustrations and holotype information see
≡ Lasiosphaeria raciborskii (Penz. & Sacc.) G.C. Carroll & Munk, Mycologia 56: 91. 1964.
≡ Chaetosphaeria raciborskii (Penz. & Sacc.) F.A. Fernández & Huhndorf, Mycol. Res. 108: 29. 2004.
Ophiochaeta raciborskii Penz. & Sacc., Malpighia 11: 406. 1897.
Paragaeumannomyces raciborskii has been collected on culms of Chusquea bamboo and other unidentified bamboo species, on palm wood and fruit, and decaying wood of unknown trees. The species has a pantropical geographical distribution and is probably the most commonly encountered species of the genus; it is known from Indonesia in Java and Central America in Costa Rica (
For descriptions and illustrations, see
Paragaeumannomyces raciborskii fide
Chaetosphaeria rubicunda Huhndorf & F.A. Fernández, Fung. Diver. 19: 39. 2005.
Paragaeumannomyces rubicundus occurs on decaying wood and is known from the Caribbean in Puerto Rico and from Central America in Costa Rica (
For description, illustration and holotype information see
USA – Tennessee • Sevier Co., Great Smoky Mountains National Park, Twin Creeks, Twin Creeks Nature Trail, near ATBI plot; alt. 549 m; 11 Oct. 2006; on decaying wood; A.N. Miller & P. Chaudhary leg.; A.N.M. 1011 (holotype: ILLS00121384!).
The species epithet is proposed in honour of Sabine M. Huhndorf for her contribution to mycology and studies in Chaetosphaeria.
Teleomorph: Ascomata perithecial, non-stromatic, superficial, usually solitary or in small groups, 250–300(–400) μm diam, 280–320 μm high, subglobose to broadly conical, rarely collapsing laterally upon drying, finely roughened, dark reddish-brown except for a black indistinct papilla, setose, setae 30–41.5 × 4–5 μm, dark brown, stiff, acute, scattered over entire ascoma, shorter and narrower setae 16.5–35 × 2.5–3 μm densely aggregated around the ostiole. Ostiole periphysate. Ascomatal wall leathery, three-layered. Outer layer of textura angularis, 32–53 μm thick, consisting of thin-walled, globose to subglobose, dark orange-brown to reddish-brown cells, 11.5–28 μm diam. Middle layer of textura prismatica, 12–22 μm thick, composed of thick-walled, polyhedral, dark brown, melanised cells. Inner layer of textura prismatica, 3–5 μm thick, composed of thin-walled, flattened and elongated hyaline to subhyaline cells. Paraphyses abundant, hyaline, sparsely branched, septate, 3.5–4.5(–6) μm wide, tapering to 2–2.5 μm, longer than the asci. Asci (154–)161–189 × (11–)12.5–14.5(–15.5) μm (mean ± SD = 174.2 ± 8.7 × 13.0 ± 0.8 μm), (130–)144–165 µm (mean ± SD = 155.2 ± 8.3 μm) long in the sporiferous part, cylindrical-fusiform, stipitate, apically broadly rounded to obtuse, ascal apex non-amyloid with a distinct apical annulus 2.5–3 μm wide, 1.5–2 μm high. Ascospores (64.5–)68.5–86.5(–88.5) × (3–)3.5–4.5 μm, (mean ± SD = 79.1 ± 5.3 × 4.0 ± 0.3 μm), filiform to cylindrical, straight or slightly curved to sigmoid, hyaline, 7-septate, septa often unevenly distributed, not constricted at the septa, asymmetrical, broadly rounded at the apical end and tapering towards the narrowly rounded basal end, with one or two guttules in each cell, 2–3-seriate, rarely 4-seriate, partially overlapping, with a negative or weak dextrinoid reaction in Melzer’s reagent. Anamorph: Unknown.
Paragaeumannomyces sabinianus. A ascomata. B ascomatal setae C vertical section of ascomal wall D, E ascomal wall F, G upper part of the ascoma with ostiole surrounded by setae H setae from the ostiolar region I paraphyses J ascal apex with apical ring K asci L, M ascospores. Images: ILLS00121384 (A, B, D, E, G–K); S.M.H. 3824 (C, F, L); S.M.H. 3807 (M). Scale bars: 500 μm (A); 20 μm (B, E, G, H); 100 μm (C, D); 25 μm (F); 5 μm (J); 10 μm (I, K–M).
USA – North Carolina • Macon Co., Coweeta Hydrological Laboratory; 27 Jun. 1998, on decorticated wood; F.A. Fernández leg.; S.M.H. 3807. • Ibid., Horse Cove Drive & Bull Pen Road, alt. 1000 m; 27 Jun. 1998; on decorticated wood; F.A. Fernández leg.; S.M.H. 3824.
A saprobe on decaying wood, so far known from North America in the USA (North Carolina, Tennessee) (Ellis 1887;
In our ITS-28S phylogeny, P. longisporus fide
USA – Tennessee • Sevier Co., Great Smoky Mountains National Park, Greenbrier, alternative side trail to Whaley Cemetery; alt. 549 m; 10 Jul. 2005; on decaying wood; A.N. Miller & A.M. Stchigel leg.; A.N.M. 466 (holotype: ILLS00121398!).
Named after the Great Smoky Mountains National Park from where it was collected.
Teleomorph: Ascomata perithecial, non-stromatic, superficial, solitary or in small groups, 270–390 μm diam, 320–400 μm high, subglobose to broadly conical, finely roughened, dark reddish-brown, glabrous except for the black papilla with dark brown, stiff, acute setae, 9.5–13.5 × 2–2.5 μm, densely aggregated around the ostiole. Ostiole periphysate. Ascomatal wall leathery, three-layered; outer layer of textura angularis consisting of globose to subglobose, reddish-brown cells; middle layer composed of brick-like, brown cells; inner layer of flattened, thin-walled, subhyaline cells. Paraphyses abundant, hyaline, longer than the asci, tapering. Asci (134–)140–174(–189) × 11–13(–14) μm (mean ± SD = 158.2 ± 19.4 × 12.3 ± 0.9 μm), cylindrical-fusiform, stipitate, apically narrowly rounded, ascal apex non-amyloid with a distinct apical annulus 2–2.5 μm wide, 1–1.5 μm high. Ascospores (58–)60.5–80.5 × (3–)3.5–4.5(–5) μm (mean ± SD = 69.8 ± 6.3 × 4.0 ± 0.5 μm), filiform to cylindrical, straight or slightly curved, hyaline, 9–11-septate, septa often unevenly distributed, not constricted at the septa, asymmetrical, rounded at the apical end, tapering towards the basal end, with one or two guttules in each cell, 2–3-seriate or 4-seriate and partially overlapping, seldom in a single fascicle. Anamorph: Unknown.
A saprobe on decaying wood, known only from the USA.
The present species is most similar to P. abietinus, the only member of the genus known from Europe. They share dark reddish-brown, glabrous ascomata with short setae surrounding the ostiole. Although the size of ascospores of both species overlap, P. abietinus differs from P. smokiensis by longer [(185–)195–240 × 12–14.5(–15.5) μm] asci and slightly longer and wider ascospores [(62–)65–87 × (3.5–)4–5.5 μm] with usually less septa [(5–)7–9(–11)].
The species was described from dead twigs of an unknown broadleaf tree and is so far known only from the subtropical climate zone of the northern hemisphere in Japan, Wakayama Prefecture (
Paragaeumannomyces sphaerocellularis is similar to P. panamensis (
French Guiana • Maripasoula, Saül, sentier des gros arbres, disturbed secondary rainforest; alt. 200 m; 25 Aug. 2018; on the bark of decaying woody liana on the ground associated with Xylaria papillatoides; C. Lechat leg.; GY.J.F. 18140-1 (holotype:
Castanea (Latin) chestnut-coloured, referring to the colour of conidia.
Teleomorph: Ascomata perithecial, non-stromatic or formed on rudimentary basal stroma, superficial, solitary or in small groups or dense clusters, 160–200 μm diam, 170–220 μm high, subglobose to broadly conical, dark brown, glabrous, papillate. Ostiole periphysate. Ascomatal wall fragile, carbonaceous, 20–27 μm thick, two-layered. Outer layer of textura prismatica, consisting of brown, polyhedral cells with opaque walls. Inner layer of textura prismatica, consisting of several rows of thin-walled, hyaline, flattened cells. Paraphyses sparse, partially disintegrating at maturity, septate, 3–5 μm wide, tapering to ca. 2.5 μm, longer than the asci. Asci (75–)78–97(–102) × (10.5–)11–14.5 μm (mean ± SD = 87.3 ± 5.0 × 12.5 ± 1.1 µm), (56.5–)65–77(–82.5) μm (mean ± SD = 70.0 ± 4.9 µm) long in the sporiferous part, cylindrical-fusiform, stipitate, apically obtuse, ascal apex with a shallow, non-amyloid apical annulus 3.5–4.5 μm wide, 1–1.5 μm high. Ascospores (10.5–)11–13.5(–14.5) × (5.5–)6–7.5 μm (mean ± SD = 12.2 ± 0.5 × 6.7 ± 0.5 µm), ellipsoidal-fusiform, dark brown to chestnut brown, 1-septate; septum median, dark brown, with a central pore, not constricted or slightly constricted at the septum, with longitudinally arranged darker brown ridges alternating with lighter brown furrows, ascospores uniseriate or obliquely uniseriate in the ascus. Anamorph: Codinaea-like, not present on the nature substrate.
Striatosphaeria castanea (
Colonies effuse, vegetative hyphae hyaline, branched, 2.5–3.5 μm wide. Conidiophores macronematous, mononematous, 22–66 μm long, 3.5–5 μm wide near the base, erect, straight, cylindrical, several-septate, brown, paler towards the apex, unbranched, smooth-walled, or reduced to single conidiogenous cells. Conidiogenous cells (9.5–)12.5–25(–33) × 4–4.5 μm, tapering to 2.5–3.5 μm just below the collarette, monophialidic, integrated, terminal, cylindrical to cylindrical-lageniform, pale brown, subhyaline towards the apex, smooth-walled; collarettes funnel-shaped, 4–6.5 μm wide, (1.5–)2–2.5 μm deep. Conidia (10–)11–13.5 × 4–5.5 μm (mean ± SD = 11.9 ± 0.8 × 4.8 ± 0.4 µm), reniform to ellipsoidal, straight or slightly curved, asymmetrical, narrowly rounded at the apical end, truncate at the basal end, brown, 1-septate, not constricted or slightly constricted at the septum, with 1–2.5 μm long, hyaline setulae at each end, smooth-walled, in slimy droplets, dark brown in mass.
On CMD colonies 23–25 mm diam, circular, flat, margin entire, lanose, floccose, funiculose at the centre, cobwebby towards the periphery, whitish with irregular pale brown spots due to pigmented funiculose mycelium, with an isabelline outer zone of submerged growth; reverse beige. On MLA colonies 22–25 mm diam, circular, raised, margin entire, lanose, floccose, zonate, with grey, brown and white zones, with an isabelline outer zone of submerged growth; reverse dark grey. On OA colonies 31–33 mm diam, circular, flat, margin entire, sparsely lanose, floccose, cobwebby at the margin, zonate, whitish, colony centre with irregular dark brown spots due to pigmented submerged mycelium, pale brown towards the margin, with an olivaceous outer zone of submerged growth, dark brown pigment diffusing to agar at the colony centre; reverse olivaceous grey. On PCA colonies 17–19 mm diam, circular, slightly convex centrally, margin entire, lanose, floccose becoming cobwebby towards the periphery, isabelline to light beige with irregular brown spots due to pigmented mycelium; reverse light beige. Sporulation abundant on CMD, CMA with Urtica stems and PCA, sparse on MLA and OA.
Brazil • Bahia; isolated from roots of Encyclia ghillanyi; isolate monte6.2; GenBank (ITS): KC928368, unpublished. (Specimen not available).
Striatosphaeria castanea occurs on the bark of woody liana and as an endophyte of Encyclia ghillanyi. It is known from South America in Brazil and French Guiana.
Striatosphaeria codinaeophora closely resembles S. castanea, but differs in having larger [(130–)140–160(–170) × 25–35(– 40) μm] asci, [(17–)19–23(–26) × (6–)7–9(–10) μm] ascospores and (15–20 × 4.5–6 μm) conidia (
≡ Phoma cytisporoides Sacc., Michelia 1: 522. 1879.
≡ Phoma cytisporoides subsp. punicina Sacc., Michelia 2: 273. 1881.
≡ Dendrophoma cytisporoides var. punicina (Sacc.) Sacc., Syll. fung. 3: 180. 1884.
≡ Dendrophoma cytisporoides var. pruni-virginianae Sacc., Riv. Accad. di Padova 33: 169. 1917.
= Dendrophoma punicina (Sacc.) Sacc., Rabenh. Krypt.-Fl., Edn. 2, 1(6): 409. 1901.
Teleomorph: Ascomata perithecial, non-stromatic, immersed becoming erumpent, in small groups or dense caespitose clusters on the bark of the host, 120–150 μm diam, 180–200 μm high, subglobose to broadly conical, dark brown, glabrous, papillate. Ostiole periphysate. Ascomatal wall fragile, carbonaceous, 22–28 μm thick, two-layered. Outer layer of textura prismatica, consisting of brown, polyhedral cells with opaque walls. Inner layer of textura prismatica, consisting of several rows of thin-walled, hyaline, flattened cells. Paraphyses sparse, persistent, septate, anastomosing, 2–2.5 μm wide, tapering to ca. 1–1.5 μm, longer than the asci. Asci (66–)69–88.5(–92) × 6.5–8 μm (mean ± SD = 77.5 ± 5.9 × 7.3 ± 0.5 µm), (35.5–)43.5–63(–69) μm (mean ± SD = 53.9 ± 5.7 µm) long in the sporiferous part, unitunicate, arising from densely branched, short ascogenous hyphae, 8-spored, cylindrical to cylindrical-clavate, stipitate, apically broadly rounded, ascal apex with an indistinct, non-amyloid apical annulus visible only with the PC illumination, ca. 1.5 μm wide, 1.5–2 μm high. Ascospores 8–10.5(–11) × 2.5–3.5 μm (mean ± SD = 9.5 ± 0.6 × 3.0 ± 0.3 µm), fusiform, hyaline, 1-septate, not constricted at the median septum, uniseriate, obliquely uniseriate or partially biseriate in the ascus. Anamorph: Not observed.
Dendrophoma cytisporoides (
Colonies effuse, vegetative hyphae hyaline, 2–3 μm wide. Conidiomata stromatic, globose becoming cupulate, up to 350 μm diam. Setae absent. Conidiophores macronematous, septate, branched or unbranched, up to 60 μm long, hyaline. Conidiogenous cells 7–13 × 1.5 μm, monophialidic, integrated and terminal or discrete and lateral, subcylindrical, single or in terminal whorls, hyaline, tapering to ca. 1 μm; collarettes indistinct. Conidia 2.5–3.5 × 1–1.5 (mean ± SD = 3.1 ± 0.3 × 1.2 ± 0.1 µm), naviculate to botuliform, with 0.5–1 μm long setulae at each end, aseptate, smooth-walled, in slimy droplets, hyaline in mass.
On CMD colonies 18–20 mm diam, circular, flat, margin entire, velvety-lanose, mucoid at the margin, white, isabelline towards the periphery, reverse white. On MLA colonies 18–21 mm diam, slightly convex, circular, margin entire to fimbriate, lanose, floccose, cobwebby at the margin, white, with an isabelline outer zone of submerged growth, reverse isabelline. On OA colonies 19–21 mm diam, circular, flat, raised margin, margin entire, velvety becoming lanose towards the margin, mucoid at the margin, zonate, colony centre grey to whitish-grey with intermediate white zone becoming brown, beige at the margin, reverse grey. On PCA colonies 13–15 mm diam, circular, flat, slightly convex centrally, margin entire, velvety-lanose, floccose becoming mucoid towards the periphery, white, pale brown centrally, with an isabelline outer zone of submerged growth, reverse isabelline. Sporulation absent on all media after 4 wk; abundant on OA and PCA after 10 wk or 4 mo, respectively, on MLA sterile primordia of conidiomata were formed.
Specimen examined. Germany • 16 Jan. 2012; on bark and wood of a dead twig of Buxus sempervivens; R. Schumacher leg.; (
Saprobic on decaying wood and bark of Buxus sempervivens, Deutzia scabra, Rhododendron sp., and Ulmus sp. The species is known from Europe in France, Germany and the Netherlands (
Our strain sporulated in vitro only after prolonged incubation. On OA it formed globose to pulvinate fertile conidiomata, while on PCA the conidiomata often became confluent. The comparison of ITS and 28S sequences of our strain with those of the epitype strain of D. cytisporoides
Based on morphology, cultivation studies and phylogenetic analysis of the combined ITS-28S loci, seven new species and eight new combinations are introduced in the Chaetosphaeriaceae. Paragaeumannomyces (
Examination of our material of Paragaeumannomyces revealed that ascospores of four species exhibit a strong dextrinoid reaction in Melzer’s reagent, namely P. abietinus, P. albidus, P. granulatus and P. smokiensis. The ascospores turned reddish-brown except for the end cells, which remained partially hyaline, especially at the tips. Interestingly, these species share glabrous, non-setose ascomata or only minute setae are arranged around the ostiole. The chemical reaction is visible in ascospores without guttules, which otherwise fill the cells and obscure the colour. The ascospores of P. elegans, P. longisporus and P. sabinianus exhibit a negative or weak dextrinoid reaction; some mature ascospores turned light pink-brown. These species share setose ascomata, sometimes with ostioles surrounded by minute setae. Although more species need to be examined to evaluate this character, we hope it is not premature to argue that the dextrinoid reaction of ascospores is species-specific and has the potential to become another diagnostic feature facilitating species identification. Because we did not examine all known species of Paragaeumannomyces, this character has not been used in the key, but it is mentioned in the species descriptions, if known.
Paragaeumannomyces, typified by P. sphaerocellularis, encompasses 18 species. The present phylogenetic tree (Fig.
Members of Paragaeumannomyces are not easily cultivated and only seldom sporulate in vitro.
The original P. longisporus clade with two strongly supported subclades was recognized as two species, the short-spored P. longisporus and the long-spored P. sabinianus. In general, a high degree of ITS sequence variability and more or less uniform teleomorphic phenotype pose special problems in species identification, especially in P. raciborskii, which was resolved as polyphyletic (
Codinaea (
Based on published records, Striatosphaeria is an uncommon lignicolous genus with a known distribution in the neotropics. It was introduced by
The genus Dendrophoma with a single species, D. cytisporoides, was proposed by
We thank Sabine Huhndorf for the use of her collections, images and DNA extracts and Alberto Stchigel and Preeti Chaudhary for assistance in our collection. Peter Johnston is thanked for his assistance to M.R. in obtaining the Manaaki Whenua Fellowship and collecting permits for New Zealand. We thank J. Leland Crane for grammatical review of some of the new names. J.F. gratefully acknowledges the Parc Naturel Amazonien de Guyane for initiating, financing and organizing fieldwork in Saül in 2018 and 2019 in connection with the inventorial ABC project, during which a new species of Striatosphaeria castanea was collected. We thank René Schumacher for a collection of Dendrophoma cytisporoides. We thank both reviewers for their valuable comments and suggestions.
This study was supported by the project of the Czech Science Foundation (GAČR 20-14840S), and as long-term research development projects of the Czech Academy of Sciences, Institute of Botany (RVO 67985939) (M.R.) and the University Hospital Hradec Králové MH CZ – DRO (UHHK, 00179906) (J.N.). This study was also supported by a National Science Foundation award (DEB-0515558) to A.N.M. The WM Keck Center at the University of Illinois Urbana-Champaign is thanked for sequencing services. The field work of M.R. in New Zealand was partly supported by Studienstiftung für mykologische Systematik und Ökologie (2003) and Manaaki Whenua Fellowship, Landcare Research Auckland (2005).The authors have declared that no competing interests exist.