Contribution to the phylogeny and taxonomy of the genus Taeniolella, with a focus on lichenicolous taxa

Accepted Manuscript Contribution to the phylogeny and taxonomy of the genus Taeniolella, with a focus on lichenicolous taxa Damien Ertz, Bettina Heuchert, Uwe Braun, Colin E. Freebury, Ralph S. Common, Paul Diederich PII:

S1878-6146(16)30050-2

DOI:

10.1016/j.funbio.2016.05.008

Reference:

FUNBIO 721

To appear in:

Fungal Biology

Received Date: 21 March 2016 Revised Date:

18 May 2016

Accepted Date: 19 May 2016

Please cite this article as: Ertz, D., Heuchert, B., Braun, U., Freebury, C.E., Common, R.S., Diederich, P., Contribution to the phylogeny and taxonomy of the genus Taeniolella, with a focus on lichenicolous taxa, Fungal Biology (2016), doi: 10.1016/j.funbio.2016.05.008. This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.

ACCEPTED MANUSCRIPT

Contribution to the phylogeny and taxonomy of the genus Taeniolella, with a focus on lichenicolous taxa Damien ERTZ1,2*, Bettina HEUCHERT3, Uwe BRAUN3, Colin E. FREEBURY4, Ralph S.

1

Botanic Garden Meise, Department Bryophytes-Thallophytes (BT), Nieuwelaan 38, B–1860 Meise,

Belgium. Email: [email protected] 2

Fédération Wallonie-Bruxelles, Direction Générale de l'Enseignement non obligatoire et de la

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Recherche scientifique, rue A. Lavallée 1, B–1080 Bruxelles, Belgium 3

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COMMON5, Paul DIEDERICH6

Martin-Luther-Universität, Institut für Biologie, Bereich Geobotanik, Herbarium, Neuwerk 21,

[email protected] 4

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06099 Halle (Saale), Germany. Email: [email protected],

Canadian Museum of Nature, PO Box 3443 Stn. “D”, Ottawa, Ontario, Canada, K1P 6P4. Email:

[email protected]

534 Fenton St., Lansing, MI 48910, USA. Email: [email protected]

6

Musée national d'histoire naturelle, 25 rue Munster, L–2160 Luxembourg, Luxembourg. Email:

[email protected]

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5

*Corresponding author: Damien Ertz: Botanic Garden Meise, Department Bryophytes-Thallophytes

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(BT), Nieuwelaan 38, B–1860 Meise, Belgium. Email: [email protected]; tel. +32

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2 2600936; fax +32 2 2600945

Number of figures: 17 Number of tables: 1

Abstract Taeniolella is a genus of asexual ascomycetes with saprophytic, endophytic and lichenicolous life styles. A phylogeny of representative species of the genus is presented, with a focus on lichenicolous taxa. We obtained mtSSU and nuLSU sequence data from culture isolates of Taeniolella and from freshly collected specimens of other species. The genus Taeniolella is recovered as strongly polyphyletic with species distributed between the Dothideomycetes and the Sordariomycetes. The type species, Taeniolella exilis, is placed in the Kirschsteiniotheliaceae within Dothideomycetes. Other saprophytic/endophytic Taeniolella species previously assigned to the

ACCEPTED MANUSCRIPT Sordariomycetes based on sequences were found to represent either contaminants or species that cannot be assigned to Taeniolella for morphological reasons. Lichenicolous species are restricted to the Asterotexiales (Dothideomycetes) where the sequenced species of Taeniolella do not form a monophyletic group, but are related to species of the genera Buelliella s. lat., Karschia, Labrocarpon, Melaspilea s. lat., and Stictographa. Molecular data are, however, not sufficient to reallocate the lichenicolous Taeniolella species to other genera so far. Anamorph-teleomorph relationships between

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these taxa and lichenicolous Taeniolella species are discussed but could not be demonstrated with the current data. Buelliella minimula, the type species of the genus Buelliella, is placed in the

Asterotexiales, and the genus recovered as polyphyletic. Three new lichenicolous Taeniolella species are described, namely T. hawksworthiana, T. pyrenulae, and T. toruloides. Taeniolella rudis is

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transferred to Sterigmatobotrys, as S. rudis. In addition, the taxonomic part comprises detailed

treatments of the generic type T. exilis and of T. punctata, which are both included in the phylogenetic

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trees.

Keywords: Asterotexiales, culture isolation, Dothideomycetes, Kirschsteiniotheliaceae, Melaspilea, Sordariomycetes.

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1. Introduction

Hughes (1958) introduced the new genus Taeniolella for an assemblage of saprobic dematiaceous hyphomycetes characterised by having little differentiated (semi-macronematous), mostly unbranched conidiophores with integrated, terminal, monoblastic, non-cicatrized conidiogenous cells, and

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pigmented, 1- to pluriseptate conidia formed in mostly long acropetal, not easily disarticulating chains. The species reallocated to Taeniolella by Hughes (1958) were originally assigned to the hyphomycete

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genera Dendryphion, Hormiscium, Septonema, and Torula. In his influential treatment of dematiaceous hyphomycetes, Ellis (1971) took up Hughes’s concept of Taeniolella, provided a more detailed generic circumscription, brief descriptions of species, and instructive illustrations. In his second book, Ellis (1976) added the new combination Taeniolella pulvillus, and described the new genus Taeniolina for superficially similar species with usually much branched conidia. The number of Taeniolella species increased over the years to about 53, accompanied by a gradual widening of the morphological concept and circumscription of this genus, inter alia, by the inclusion of aquatic and lichenicolous species. The latter drastic extension of the genus goes back to Hawksworth’s (1979) treatment of lichenicolous hyphomycetes in which several morphologically similar lichen-inhabiting species were assigned to Taeniolella. To this day, the number of lichenicolous Taeniolella species has increased rapidly. The lichenicolous species roughly fit with saprobic Taeniolella spp. in terms of morphology, although most of the saprobic species are characterised by having pluriseptate conidia

ACCEPTED MANUSCRIPT versus amero- to phragmosporous conidia in lichenicolous taxa. These differences are, however, only gradual and barely significant enough to justify the establishment of a separate genus for the lichenicolous species. The striking morphological diversification and wide range of ecological niches within the broad concept of Taeniolella raises the question whether the current morphological circumscription of this genus may withstand phylogenetic approaches. Among ascomycete genera typified by asexual morphs, there are two opposed tendencies. For some genera, traditional

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morphological concepts have been confirmed by molecular methods and this has led to the recognition of larger monophyletic core genera, such as Alternaria (Woudenber et al. 2013) and Cladosporium (Bensch et al. 2012). Other genera, such as Sporidesmium (Shenoy et al. 2006), proved to be totally polyphyletic. Ertz et al. (2015) demonstrated the strong genetic heterogeneity of lichenicolous fungi

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exemplified by former Polycoccum spp. (sexual morphs) and lichenicolous Phoma spp. (asexual morphs) which phylogenetically belong to the genus Didymocyrtis (Phaeosphaeriaceae).

According to the most recent literature (Kirk et al. 2008, Hyde et al. 2013) and without benefit of

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comprehensive phylogenetic analysis, the genus Taeniolella was thought to be part of the family Mytilinidiaceae and to be the anamorph of Glyphium. There are only a few available sequence data retrieved from species of this genus, their reliability has never been established with certainty, and the number of publications containing relevant molecular data is rather limited (Crous et al. 2006, Liang et al. 2011, Réblová et al. 2012). The few published data and the particular positions in the phylogenetic trees indicate a high degree of heterogeneity and polyphyly. But the crux of the problem around the

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phylogeny of Taeniolella and a reassessment of the whole genus lay in the lack of sequence data of lichenicolous species and T. exilis, the type species of the genus. Sequences of the latter species and several lichenicolous taxa are now available, which encouraged us to disentangle the heterogeneous Taeniolella complex by means of a molecular approach and address the following questions: Is the

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genus Taeniolella in its current broad sense mono- or polyphyletic? Are saprobic and lichenicolous species phylogenetically allied? Do the lichenicolous species as an ecological group represent a

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monophyletic assemblage?

2. Material and Methods 2.1. Morphological study – Herbarium specimens are deposited at BR, C, CANL, GLM, H, K(M), M, UPS, TU, WA (abbreviations according to Holmgren et al. 1990), in the herbarium of the Institut für Vegetationskunde und Landschaftsökologie, Hemhofen, Germany (IVL), and in the private collections of P. Diederich and of R. Cezanne / M. Eichler. Microscopical examination (including all microscopical measurements) was carried out using hand-made sections mounted in distilled water and an Olympus BX50 microscope at a magnification of ×1000, without any staining since all structures examined are darkly pigmented. Permanent slides were prepared by sealing the coverglasses with Canada balsam (SERVA, Heidelberg) and dried for 24 hours. Twenty conidiophores,

ACCEPTED MANUSCRIPT conidia and other structures were measured in each collection, and a representative range was depicted. Measurements are given as ranges of values rounded to the nearest half micrometre, except for wall thickness; extreme values are added in brackets. Drawings were done free hand. Microscopic photographs were made using a Zeiss Axioskop 2 equipped with a Zeiss AxioCam HR, occasionally optimised with the software Zeiss AxioVision. Macroscopic photographs were done using a Canon 40D camera with a Nikon BD Plan 10× microscope objective, and with StackShot (Cognisys) and

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Helicon Focus (HeliconSoft) for increasing the depth of field. ESEM examination, conducted at the Interdisciplinary Centre of Materials Science (CMAT) of Martin Luther University Halle-Wittenberg, was carried out to identify details of conidiophores and conidial surface ornamentation. Specimens were excised from the host and attached to aluminium pin stubs. Observations and micrographs were

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made with a Philips XL30 ESEM-FEG environmental electron microscope with digital camera, at 1.2 Torr and 3.0 kV acceleration voltages. The specimens were not coated.

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2.2. Culture isolation and molecular techniques – Cultures of Taeniolella were isolated from conidia of freshly collected material on malt-yeast extract medium (Yoshimura et al. 2002) (Fig 5I). Conidia were spread directly on malt-yeast extract agar. Germination of conidia was observed after several days. Isolated conidia were then immediately transferred to new petri dishes to attain axenic cultures as fast-growing contaminants were often observed in the first step. The cultures were kept at room temperature and exposed to a natural day-light regime in the laboratory of the Botanic Garden Meise.

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No experiments were done to test whether different light or temperature conditions would affect the growth rate. As all the strains were slow-growing, several months were required to obtain sufficient material for DNA extraction. The identity of the cultures was confirmed by phenotypic characters. In the case of Melaspilea s. lat. specimens, hand-made sections of the hymenium were used for direct PCR as described in Ertz et al. (2014). The outer wall of ascomata was removed with a sterile razor

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blade to isolate the hymenium. The material was then added to a tube containing the PCR reaction mixture and amplified directly. We were unable to obtain reliable sequences from the conidia of

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Taeniolella species by direct PCR; e.g. different sequences belonging to Dothideomycetes or Chaetothyriales were obtained from different specimens of T. phaeophysciae for which rich, fresh material was available, suggesting frequent co-occurring fungi in these samples. Unreliable sequences were also obtained from cultures of fungi isolated from thalli fragments infected by lichenicolous fungi, including from several specimens infected by Taeniolella atricerebrina, which suggests that lichen thalli host various endolichenic fungi that are easier to isolate than the lichenicolous taxa visible on the specimens (Muggia et al. 2015). The sequences obtained from the specimens infected by Taeniolella atricerebrina in this latter study were not included in our analyses. Genomic DNA was isolated from cultures using the CTAB extraction protocol (Doyle and Doyle 1990). Amplification reactions were prepared for a 50 µl final volume containing 5 µl 10× DreamTaq Buffer (Fermentas), 1.25 µl of each of the 20 µM primers, 5 µl of 2.5 mg mL-1 bovin serum albumin (Fermentas #B14), 4 µl of 2.5mM each dNTPs (Fermentas), 1.25 U DreamTaq DNA polymerase (Fermentas), and 1 µl of

ACCEPTED MANUSCRIPT template genomic DNA or tiny fragments of fungal material. A targeted fragment of about 1.1 kb at the 5’end of the nuLSU rDNA was amplified using primers LIC15R (Miadlikowska et al. 2002) with LR6 (Vilgalys and Hester 1990), and a fragment of about 0.8 kb of the mtSSU rDNA using primers mrSSU1 and mrSSU3R (Zoller et al. 1999). The yield of the PCRs was verified by running the products on a 1% agarose gel using ethidium bromide. Both strands were sequenced by Macrogen® using amplification primers. Additional primers were used for the sequencing of nuLSU: LR3, more LR3R

and

LR5

(Vilgalys

and

Hester

1990)

(Vilgalys’

website,

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rarely

http://www.botany.duke.edu/fungi/mycolab). Sequence fragments were assembled with Sequencher version 4.6 (Gene Codes Corporation, Ann Arbor, Michigan). Sequences were subjected to MEGABLAST searches to verify their closest relatives and to detect potential contaminations.

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2.3. Taxon selection and phylogenetic analyses – Although not possible for all taxa, we tried to achieve a sample series with at least two specimens of each newly sequenced species from different

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localities. In addition to Taeniolella sequences, we also provide new sequences of Melaspilea s. lat. and Buelliella that are closely related to the former. Our sample series includes new sequences obtained from the type species of Taeniolella (T. exilis) and the type species of Buelliella (B. minimula). For the nuLSU phylogenetic tree, the closest relatives of the new sequences based on BLAST searches were retrieved from GenBank. NuLSU sequences of Taeniolella already available on GenBank and their closest relatives were also downloaded. The original nuLSU matrix used in Ertz and Diederich (2015) was used as a template to include the newly sequenced species and the taxa

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retrieved from GenBank. According to preliminary phylogenetic analyses, distantly related taxa to our Taeniolella sequences were removed from the dataset. The resulting nuLSU matrix consisted of 104 sequences, mainly from a variety of Dothideomycetes and Sordariomycetes. Three outgroup species were chosen to represent the class Eurotiomycetes (Caliciopsis pinea and Capronia munkii) and the

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class Leotiomycetes (Lachnum virgineum). The alignments were improved manually using Mesquite 3.04 (Maddison and Maddison 2015). Terminal ends of sequences, ambiguous aligned regions, and

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introns were delimited manually and excluded from the nuLSU dataset. The nuLSU data set consisted of 915 unambiguously aligned characters, of which 484 were variable. The best-fit model of DNA evolution GTR+I+G was chosen for the nuLSU data set using the Akaike information criterion (AIC; Akaike 1973) as implemented in Modeltest v. 3.7 (Posada and Crandall 1998). Bayesian analyses were carried out using the Metropolis-coupled Markov chain Monte Carlo method (MCMCMC) in MrBayes v. 3.2.3 (Huelsenbeck and Ronquist 2001; Ronquist and Huelsenbeck 2003) on the CIPRES portal (Miller et al. 2010). Analyses were run under the selected model of nucleotide substitution with six rate categories. Two parallel MCMCMC runs were performed, with each run using four independent chains and 100,000,000 generations, and sampling trees every 1000th generation. Convergence diagnostics were estimated using the PSRF (Potential scale reduction factor) where values closer to one indicated convergence between runs (Gelman and Rubin 1992), and using TRACER v. 1.6 by plotting the log-likelihood values of the sample points against generation time

ACCEPTED MANUSCRIPT (Rambaut and Drummond 2007). Posterior probabilities (PP) were determined by calculating a majority-rule consensus tree generated from the 150,002 post-burnin trees of the 200,002 trees sampled by the two MCMCMC runs using the sumt option of MrBayes. In addition, a Maximum Likelihood (ML) analysis was performed using GARLI (Zwickl 2006, v. 0.951 for OSX) with default settings. One thousand bootstrap pseudoreplicates were used to calculate a majority-rule consensus tree in PAUP* 4.0b10 (Swofford 2002), and to assess the Maximum Likelihood bootstrap values (ML-

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bs).

A combined nuLSU+mtSSU matrix focusing on the Asterotexiales was assembled manually using the matrices in Ertz and Diederich (2015) as a main template. Taxa were selected from Hofmann et al. (2010), Hongsanan et al. (2014), Ertz and Diederich (2015) and Guatimosim et al. (2015) to include

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samples for which at least a nuLSU was available, thus excluding Hemigrapha asteriscus and Melaspileopsis proximella for which only a mtSSU sequence was available. Three outgroup species

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were chosen to represent the orders Capnodiales (Capnodium coffeae), Dothideales (Dothidea insculpta) and Pleosporales (Preussia terricola). As it was not possible to complete the mtSSU sequences for the same set of 58 samples, analyses for topological incongruence among loci were carried out on data sets with 36 samples for which the two genes were available (Table 1). To detect significant conflicts among datasets, Bayesian analyses were carried out on the single-locus data set using the MCMCMC method in MrBayes v. 3.2.6 on the CIPRES Web Portal. The GTR+I+G model was selected for the nuLSU data set while the TVM+I+G model was selected for the mtSSU data set

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using AIC. Two parallel MCMCMC runs were performed each using four independent chains and 20 million generations, and sampling trees every 1000th generation. Posterior probabilities (PP) were determined by calculating a majority-rule consensus tree generated from the 30002 post-burnin trees of the 40002 trees sampled by the two MCMCMC runs using the sumt option of MrBayes. All

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topological bipartitions with PP values ≥ 95 were compared for the two loci. A conflict was assumed to be significant if two different relationships (one being monophyletic and the other being non-

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monophyletic) for the same set of taxa were both supported with PP values ≥ 95. Based on this criterion, no conflict was detected and therefore the nuLSU and mtSSU data sets were concatenated. The combined two-locus data set consisted of 58 taxa and 1387 unambiguously aligned sites, including 877 sites for nuLSU and 510 sites for mtSSU. Bayesian analyses were carried out on the two-locus data set under the selected models for two partitions (nuLSU and mtSSU) and using the same settings as for the analyses for topological incongruence among loci, but using 40 million generations. Posterior probabilities (PP) were determined by calculating a majority-rule consensus tree generated from the 60002 post-burnin trees of the 80002 trees sampled by the two MCMCMC runs using the sumt option of MrBayes. In addition, a Maximum Likelihood (ML) analysis was performed on the two-locus data set using GARLI with default settings. One thousand bootstrap pseudoreplicates were used to calculate a majority-rule consensus tree in PAUP* to assess the ML-bs values.

ACCEPTED MANUSCRIPT The Bayesian tree did not contradict the ML trees topology for the strongly supported branches of the two analyses (nuLSU data set of 104 taxa and the combined nuLSU+mtSSU data set of 58 taxa). Therefore, only the majority-rule consensus tree of the Bayesian analysis is shown with the posterior probabilities of the Bayesian analysis added above the internal branches and the bootstrap support values of the ML analysis added below the internal branches (Figs 1–2). ML-bs ≥ 70 and PP ≥ 95 were considered to be significant. Phylogenetic trees were visualized using FigTree v1.4.2 (Rambaut

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2012).

2.4. Specimens sequenced in addition to those treated in section 3.2 – Buelliella minimula: USA, North Carolina, Dare Co., Buxton Woods Coastal Reserve, SE of jct of Great Ridge Rd. and Water Association Rd., E of West Trail, 35°14’51” N, 75°34’57” W, 5 ft., sand ridge swale complex with

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maritime forest of Pinus and mixed hardwoods (Quercus, Ilex, Carpinus, Persea), on Pertusaria

tetrathalamia on fallen branch, 18 Aug. 2013, J.C. Lendemer 35969 (NY); Cape Hatteras National

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Seashore, Open Ponds Trail, 0.25-0.5 mi W of jct w/ access road, S of Buxton, 35°14’55” N, 75°32’25” W, 9 ft., maritime forest of Quercus-Pinus-Myrica-Ilex vomitoria with occasional sandy openings, on Pertusaria paratuberculifera on Quercus, 19 Mar. 2013, J.C. Lendemer 36238-A (NY); Charleston Co., Edisto Island, Botany Bay Plantation Wildlife Management Area, N of Jason Lake and S of jct of Botany Bay Rd; and Rabbit Rd. 32°32’56” N, 80°16’24” W, 83 ft., upland mixed Pinus and hardwood (Ilex vomitoria, Quercus, Carya, Myrica) forest, on Pertusaria on Quercus, 20 Dec.

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2013, J.C. Lendemer 42273 (NY). Melaspilea s. lat.: USA, Florida, Hernando Co., Baypoint Park at end of CR 50/550, 28°32.166’ N, 82°39.04’ W, on wood, 25 Aug. 2011, R. Common 9134, 9137A (BR). Reunion, Cilaos, Forêt du Grand Matarum, début du sentier vers le Piton des Neiges, 21°07’21” S, 55°29’00” E, alt. 1440 m, tronc en forêt, sur Pyrenula, 11 Dec. 2012, D. Ertz 18012 (BR). Netherlands Antilles, Curaçao, N of Lagun, Playa Abou (Grote Knip), slope north of the beach,

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12°21’09” N, 69°09’06” W, dry forest on a limestone cliff near the sea and a sandy beach, smooth trunk of 3 cm diam., 9 Aug. 2013, D. Ertz 18137 (BR). Democratic Republic of the Congo,

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Yangambi biosphere reserve, 00°47’54” N, 24°29’15” E, alt. 488 m, tropical rain forest, on Pyrenula on liana, 6 Nov. 2012, Van den Broeck 3683, 3684 (BR).

3. Results

3.1. Phylogenetic analyses – We obtained 33 new sequences (21 nuLSU and 12 mtSSU) belonging to 12 species from Azores, Belgium, Canada, Democratic Republic of Congo, Germany, Luxembourg, Rwanda and USA (Table 1, Figs 1–2). The backbone of our nuLSU tree is poorly resolved, with the Dothideomycetes recovered as paraphyletic (Fig 1). Main groups of Dothideomycetes such as Asterotexiales, Asterinaceae,

ACCEPTED MANUSCRIPT Capnodiales, Dothideales, Eremithallales, Hysteriales, Kirschsteiniotheliaceae, Pleosporales, Tubeufiales and Venturiales are strongly supported by both the Bayesian and ML analyses, even though the relationships among these groups are not supported. The Sordariomycetes are strongly supported and divided into three well-supported groups corresponding to Diaporthales, Sordariales (which was only supported by the Bayesian analysis), and Savoryellales + an unnamed clade (the Savoryellales being represented in our tree only by Savoryella lignicola). Taeniolella is resolved as

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polyphyletic, with species being placed in five distantly-related lineages corresponding to Asterotexiales and Kirschsteiniotheliaceae in Dothideomycetes, and to Diaporthales, Sordariales and an unnamed clade related to the Savoryellales in Sordariomycetes. However, the reliability of the sequence placed in the Diaporthales (viz. T. alta) is going to be questioned later (see section 4.2).

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The combined nuLSU+mtSSU tree (Fig 2) includes all lichenicolous species of Taeniolella that were newly sequenced in this study: T. hawksworthiana, T. punctata, T. pyrenulae and T. toruloides, as well

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as two Taeniolella specimens close to T. hawksworthiana and T. punctata. These taxa do not form a monophyletic group but are intermixed with sexual species that were previously assigned to the Asterinales in earlier phylogenies (Hofmann et al. 2010, Hongsanan et al. 2014, Ertz and Diederich 2015). However, the type species of Asterina (and thus the Asterinales) was recently shown to be unrelated and part of a distinct lineage within the Dothideomycetes (Guatimosim et al. 2015). Therefore, the name Asterinales cannot be assigned to this group anymore. Asterotexiales is an order recently described by Guatimosim et al. (2015) for the single species Asterotexis cucurbitacearum that

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is nested in this group. As a consequence, we propose to enlarge here the concept of Asterotexiales to include the group previously assigned to the Asterinales, and thus to also include all ingroup species of our nuLSU+mtSSU tree. Despite the use of two genes, relationships within Asterotexiales are poorly resolved and a generic reappraisal remains problematic. As a consequence, the new species are here

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described in the genus Taeniolella taken in the traditional sense.

3.2. Taxonomy

The following taxonomic treatment of Taeniolella includes the generic type, all sequenced lichenicolous species, and a saprobic species.

Sterigmatobotrys rudis (Sacc.) Heuchert, U.Braun & Ertz comb. nov. Figs 3–4. MycoBank No.: MB 817106 Basionym: Septonema rude Sacc., Michelia 1: 270 (1878). Taeniolella rudis (Sacc.) S. Hughes, Canad. J. Bot. 36: 817 (1958).

ACCEPTED MANUSCRIPT Dendryphion curtipes Ell. & Barth., Erythea 4: 82 (1896) [as ‘Dendryphium’]. [Syntypes: USA, Kansas, Rooks Co., on the underside of an old hog trough, 20 Dec. 1894, E. Bartholomew 1612 (NY 313420, 883692, 883693, 883694; ILLS 34804)]. Brachycladium curtipes (Ell. & Barth.) A.L. Smith, J. Bot. 41: 259 (1903). Misinterpreted name: Septonema hormiscium Sacc. sensu Hughes 1952. Type: [Italy], Hab. in lingo putrescente pyrino a Selva, Sept. 1874 (PAD, not seen).

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Lit.: Hughes (1952: 11, 1980c: 1–2), Matsushima (1975: 132 as Septonema hormiscium), Ellis (1976: 60), Ellis and Ellis (1997: 64), Révay (1988: 98), Kalgutkar (1997: 304), Mel’nik (2000: 309), Jones et al. (2002: 201), Catania and Romero (2009: 46), Wang (2010: 191), Simón (2011: 326).

Ill.: Saccardo (1881: tab. 921), Hughes (1952: 11, fig. 3), Matsushima (1975: F837, P1317 as

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Septonema hormiscium), Ellis (1976: 60, fig. 42C), Hughes (1980c: 1, figs 1–7), Ellis and Ellis (1997, fig. 225), Révay (1988: pl. 3, fig. 2), Caretta et al. (1992: 337, fig. 17), Kalgutkar (1997: 208, pl. 2, fig.

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9), Mel’nik (2000: 308, fig. 216), Jones et al. (2002: 202, figs 1–4; 204, figs 9–15), Catania and Romero (2009: 46, fig. 2A), Esquivel (2009, figs 1–2), Chavarria et al. (2010: 737, figs 11–12), Simón (2011: 327, fig. 116 A–C).

Colonies scattered over the substrate, thin to densely caespitose, effuse, black, slightly shiny, long chains of conidia visible when using a stereomicroscope, erect, in small tufts or procumbent on the substrate. Mycelium immersed, sometimes superficial; hyphae straight to flexuous, branched, 1–4.5

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µm wide, rarely septate, not constricted at the septa, subhyaline to pale brown, smooth, wall unthickened. Stromata lacking. Conidiophores macronematous, mononematous, solitary or aggregated in groups of up to 14, arising from hyphae, terminal or lateral, or from more or less isodiametric hyphal cells, often not very evident, erect, straight, ovoid, obclavate to ellipsoidal with a bulbous or

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rounded base, unbranched, (27–)30−45 × (8–)10−13 µm, 4–6(–8)-septate, not constricted at the septa, smooth, wall thickened, up to 1 µm, dark brown, not enteroblastically proliferating, in morphology and pigmentation the conidiophores are almost indistinguishable from conidia but the conidiophores

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are usually attached to the substrate and only rarely detached. Conidiogenous cells integrated, terminal, monoblastic, monopodial, subcylindrical or doliiform, 4–10 µm long, little differentiated; loci truncate, 4–5(–6) µm diam. Conidia catenate, in unbranched chains, not easily disintegrating, conidia long-adhering, up to 5, chains at least up to 250 µm long, 4–6 µm wide in constricted or narrow segments between single conidia, maximum width (6–)7–13 µm, straight or slightly flexuous, individual conidia obclavate, ellipsoid, fusiform, 28–61 × 7–12(–14.4) µm, 3–12-septate, nonconstricted at the septa, septa conspicuously thickened, 1–2.5 µm, distinctly multilayered, (rarely with a longitudinal septum in one of the cells [Hughes 1980c]), dark brown, somewhat paler at the apex, smooth, wall thickened, 1–1.5 µm, apex rounded in primary conidia, conically truncate in secondary ones, base obconically truncate, hila truncate, unthickened, not darkened, 4–6 µm diam.

ACCEPTED MANUSCRIPT Synanamorph: Terminal conidium frequently extended, long, becoming dichotomously branched at the tip, forming hyaline “metulae” in a penicillate head on which conidiogenous cells are formed; “metulae” erect, straight, subcylindrical, 8–15 × 4–5 µm, aseptate, paler brown than the tips of the conidia, smooth, wall slightly thickened, 2–3 conidiophores at the tip of “metulae”, doliiform, subcylindrical, 5–13 × (1.5–)2–4.5 µm, aseptate, subhyaline or hyaline, smooth, wall unthickened. Conidia solitary, straight, subcylindrical, ellipsoid, 15–25 × 4–5(–6.5) µm, 0–2(–3)-septate, not

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constricted at the septum, subhyaline, smooth, wall unthickened, plasma slightly granulose, apex rounded, base rounded or slightly truncate, hila neither thickened nor darkened.

Host range and distribution: Abies balsamea, Bambusa vulgaris, Chamaecyparis lawsoniana ‘Ellwoodii’, Comarum palustre, Picea glauca, Phragmites australis, Podocarpus parlatotei, Thuja

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occidentalis, on wood of unidentified conifers and on rotten wood. Argentina (Catania and Romero 2009), Canada (Hughes 1980c; Ginns 1986), China (Luo et al. 2004), Estonia (Voronin 1992), Hungary (Révay 1988, 1993), Italy (Caretta et al. 1992), Kazakhstan (Mel’nik 2000), Mexico

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(Chavarria et al. 2010), Panama (Esquivel 2009), Spain (Simón 2011), United Kingdom (Ellis 1976; Jones et al. 2002), USA (Hughes 1980c; Wang 2010).

Additional specimens examined: On conifer plank (on the ground): United Kingdom: Cambridgeshire, Cambridge (Botany Field Station), 16 Jul. 1948, S.J. Hughes (K(M): 180130 = IMI 31196) (as Septonema hormiscium). On hardwood plank: United Kingdom: East Kent, Dungeness, 8 Oct. 1963, B. Sutton & K. Pirozynski (K(M): 180132 = IMI 102586) (as S. hormiscium). On wood:

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United Kingdom: South-west Yorkshire, Sheffield, Taptonville Road [illegible], 7 Feb. 1957, J. Webster [ex herb. Sheffield 1918] (K(M): 180131 = IMI 69401) (as S. hormiscium); South Hampshire, Portsmouth, Dep. Biological Sciences, 28 Jan. 1966, G. Jones (K(M): 180135 = IMI 117214) (as S. hormiscium); South Hampshire, Portsmouth, Dep. Biological Sciences, 28 Jan. 1966, G. Jones (K(M):

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180133 = IMI 117213) (as S. hormiscium). On rotten wood: Hungary: Comit. Pest, pr. pag. Kismaros, ad ripam rivulis Morgó-patak, 14 Jan. 1991, Á. Révay & J. Gönczöl (BP 85790); Comit.

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Borsod-Abauj-Zemplén, pr. pag. Trizs in valley Hidegviz-völgy, 19 Mar. 1991, Á. Révay (BP 87662); Comit. Borsod-Abauj-Zemplén, pr. pag. Perkupa in valley Henc-völgy, 20 Mar. 1991, Á. Révay (BP 87663); montes Börzsöny-hegység, ad ripam rivulis Morgó-patak, 26 Jun. 1991, Á. Révay (BP 87661); Comit. Zala, pr. pag. Budafa, 25 Jul. 2001, Á. Révay (BP 96761); on a pine board lying on the ground, Comit. Pest, in opp. Gödöllő/Hortus Botanicus Univ. Sci. Agr., 28 Dec. 1993, S. Tóth (BP 90896). On Chamaecyparis lawsoniana ‘Ellwoodii’: United Kingdom: Cambridgeshire, Cambridge MAFF, Mar. 1978, P. Gladders PC78/0225 (K(M): 180129 = IMI 225745b). On Pinus sylvestris: United Kingdom: Bedfordshire-Cambridgeshire, Little Barford cooling water power station, on test block, 24 Jan. 1962, R.A. Eaton (K(M): 180128 = IMI 386834). Notes: Taeniolella rudis and Septonema hormiscium are two previously confused names, e.g. by Hughes (1952) and Matsushima (1975). Later, Hughes (1958) reallocated Septonema rude to

ACCEPTED MANUSCRIPT Taeniolella, and Hughes (1980c) reported that an examination of the type collection of S. hormiscium in herb. PAD proved that this species belongs to Sporidesmium. Even though descriptions and illustrations of conidia of T. rudis in Matsushima (1975), Ellis (1976), Ellis and Ellis (1997), Révay (1988), Caretta et al. (1992), Jones et al. (2002), Catania and Romero (2009), Esquivel (2009) and Chavarria et al. (2010) are in agreement with the original description of Saccardo (1878), the conidiophores have been described differently. The illustration of the

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conidiophores in Matsushima (1975) shows the base of the conidiophores with short branches, which is, however, barely interpretable as the base of conidiophores is usually attached to the substrate and not easily discernible. Detached conidiophores have usually a rounded base. Ellis (1976), Mel’nik (2000) and Simón (2011) described the conidiophores as short, pale to mid brown and 1–5 µm thick,

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which may be interpreted as micronematous, whereas Catania and Romero (2009) described the conidiophores as macronematous, usually short, unbranched, dark brown, thick-walled, smooth, 30–40 × 3–5.5 µm, but the given width disagrees with the associated illustration. Re-examination of several

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collections confirmed that the conidiophores are indeed macronematous as described in Matsushima (1975), Hughes (1980c) and Jones et al. (2002).

Kalgutkar (1997) described the fossil taxon Diporicellaesporites taeniolelloides with Taeniolella-like, catenate conidia but generally with coarsely rough wall. Type material of this taxon could not be examined, and its identity and affinity remain unclear.

The occurrence of a synanamorph with penicillately branched heads and colourless conidia was first

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described and illustrated by Hughes (1980c). Strangely, numerous authors who dealt with this species in the following 30 years failed to note this feature. More recently, however, Jones et al. (2002) provided a second detailed description, drawing and micrograph of the synanamorphs that they had frequently observed in the material from England and Hungary (including specimens that were re-

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examined in the course of the present study). The authors discussed the similarity between the synanamorphs of T. rudis and Sterigmatobotrys macrocarpa. Both S. macrocarpa and T. rudis grow in ecologically similar circumstances, on often water-logged wood, and in rather cool temperatures (K.

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Seifert, pers. comm.). Based on the features and dimensions of metulae, conidiogenous cells and conidia (Réblová and Seifert 2011), there are hardly any differences between the two taxa. Taeniolella rudis has somewhat wider metulae (8–15 × 4–5 µm, compared to 6.5–13.5 × (2.5–)3 µm) and conidiogenous cells (5–13 × (1.5–)2–4.5 µm, compared to 5–22 × 1.5–3.5 µm), whereas Sterigmatobotrys macrocarpa has somewhat shorter conidia (17–20.5 × 4.5–5.5, compared to 15–25 × 4–5(–6.5) µm). Distinctly fusiform macroconidia in long-adhering chains, characteristic for T. rudis, are absent in S. macrocarpa, and the conidiophores are straight, stout, up to 325 µm long and 10–13 µm wide. Both species have been included in molecular analyses of Pleurothecium and Pleurotheciella, based on nuclear ribosomal and protein-coding genes (Réblová et al. 2012), in which Pleurotheciella was shown to be sister of a clade containing Sterigmatobotrys, including T. rudis. Pleurotheciella, Pleurothecium, Sterigmatobotrys and T. rudis grouped in phylogenetic trees as a

ACCEPTED MANUSCRIPT sister clade to the Savoryellales (Réblová et al. 2012). T. exilis, the type species of Taeniolella, and T. rudis do not cluster together within our nuLSU phylogenetic tree. They are not closely allied and in any case not congeneric, i.e. the latter species must be excluded from Taeniolella. The molecular data and morphological peculiarities of the synanamorph justify the reallocation of T. rudis to the genus Sterigmatobotrys. Following the new rules of Art. 59 of the ICN (one name for one fungus, i.e. for all

Taeniolella exilis (P. Karst.) S. Hughes, Canad. J. Bot. 36: 817 (1958). Figs 5–6.

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morphs of a single fungus), T. rudis has to be assigned to the genus Sterigmatobotrys.

Basionym: Septonema exile P. Karst., Meddeland. Soc. Fauna Fl. Fennica 14: 98 (1887).

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Type: Finland, Merimasku [Naantali], on bark of living Betula sp., P.A. Karsten (H, not seen).

Lit.: Karsten (1892: 439), Saccardo (1892: 609), Ellis (1971: 93), Migula (1934: 324), Hughes (1980a:

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1–2), Mel’nik (2000: 307).

Ill.: Ellis (1971: 92, fig. 55), Hughes (1980a: 1, figs 1–10), Mel’nik (2000: 307, fig. 214). Colonies scattered on bark, effuse or more or less restricted to lenticels, reticular, caespitose to velvety, sometimes scattered in small tufts or sometimes dense and narrowly oval, somewhat sooty, confluent, black, 1−17 × 0.5−2 mm; bark rarely discoloured, reddish brown. Mycelium immersed and partly superficial, composed of flexuous hyphae, branched, (2–)3−10 µm wide, septate, not constricted

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at the septa in narrow hyphae, sparingly to distinctly constricted at the septa in wider hyphae, subhyaline to dark brown, smooth, wall somewhat thickened, 0.25–0.5 µm. Hyphae sometimes aggregated in scattered, immersed to partly superficial, flattened cell layers 1 to 4 cells thick, forming stromata, 150–320 × 30–60 µm; or penetrating deeply into the tissue and forming a continuous or

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interrupted, superficial or partly immersed, crust-like stroma composed of brown to dark brown irregularly shaped cells, 3–20 × 2–10 µm. Conidiophores seldom micronematous, reduced to

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conidiogenous cells, usually semi-macronematous to macronematous, mononematous, arising from hyphae, terminal or lateral, or from stroma cells, mostly in small caespitose tufts of 3−10 conidiophores, sometimes solitary, erect, straight, unbranched, subcylindrical, conidiophores with attached conidia (11–)30−140(–200) × 8−14 µm, 1–11-septate, slightly or distinctly constricted at the septa, dark brown, paler towards the apex, smooth; wall thickened, up to 0.75 µm wide, often thinner towards the apex; granular cell plasma mostly reduced, with a central vacuole-like cavity, surrounding plasma giving the impression of very thick, three-layered walls, up to 3 µm wide, frequently enteroblastically proliferating with obvious sheath-like wall remnants visible as an irregular collar. Conidiogenous cell integrated, terminal, monoblastic, monopodial, determinate, subcylindrical, doliiform, attenuated at the tip, 9.5−18 µm long; loci truncate to convex, 4−8 µm diam., unthickened, lateral wall thickened, forming a small rim. Distinction between conidiophores and conidial chains difficult. Conidia catenate, usually in unbranched chains, up to five conidia per chain, straight to

ACCEPTED MANUSCRIPT slightly curved, doliiform, subcylindrical to nearly obclavate, (0–)1–7(–13)-euseptate, sometimes also with 1–2 intermixed distosepta: aseptate conidia 19−21 × 11 µm, 1-septate ones 20−32(–45) × 8−15 µm, 2-septate ones 30−42(–56) × 10−15 µm, 3-septate ones 39−49(–68) × 10−17 µm, 4−11-septate ones 50−108(–180) × 10−14(−17) µm, mostly constricted at the septa, brown to dark brown, paler near the apex in secondary conidia, outer wall smooth or seldom roughened, slightly thickened, up to 0.75 µm wide, granular cell plasma mostly reduced, with a central vacuole-like cavity, surrounding plasma

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giving the impression of very thick, three-layered walls, 1.5–3 µm thick, apex rounded in primary conidia, truncate or often slightly obconically truncate in secondary ones, base truncate, sometimes narrowed towards the base, hila truncate to convex, 3−6.5 µm diam., thickened lateral wall sometimes visible as conspicuous rim, in one case microcyclic conidiogenesis observed.

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Host range and distribution: Betula alleghaniensis, B. papyrifera, B. pendula, B. platyphylla subsp. mandshurica, Betula sp., Capinus betulus, Corylus avellana, Quercus robur. Canada (Hughes 1980a; Ginns 1986), Austria (Migula 1934), Finland (Karsten 1887, 1892), Georgia (Svanidze 1984,

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www.cybertruffle.org.uk), Poland (Borowska 1987; Chlebicki and Chmiel 2006), Russia (Mel’nik 2000).

Additional specimens examined: On Betula papyrifera: Canada: Quebec, Lake Bernard, Masham Township, Gatineau Co., on felled trunk, 13 Jul. 1958, S.J. Hughes, ex DAOM 59235 (a) (H, K(M) IMI 76361); Gatineau Park, Church Hill Area, 106 m east of Eardley Rd., 45°34’51.7” N, 76°05’25.7”

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W, 217 m, on standing Betula papyrifera, 4 Apr. 2013, C.E. Freebury 1968 (CANL); Ontario, Pembroke, in backyard wood pile, 4 Nov. 1979, G.P. White, ex DAOM 173671 (H). On Carpinus betulus: Poland: Warsaw, Reserve Bielański, on the damaged trunk of a living tree, 5 Jan. 1975, A. Borowska (WA 27758).

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Notes: The type material of Taeniolella exilis, according to Hughes (1958) deposited in Helsinki (H), could unfortunately not be traced recently. Hughes (1958) had previously seen this material and compared it (Hughes 1980a) with collections deposited in DAOM. The descriptions of morphological

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features of this species in Karsten (1887), Migula (1934), Ellis (1971), Hughes (1980a) and Mel’nik (2000) are largely in agreement. The identification of T. exilis recorded on Corylus avellana in Georgia (Svanidze 1984) could not be confirmed. Borowska (1987) published observations of T. exilis in Poland on Betula pendula, Capinus betulus and Quercus robur. Re-examination of a sample on Carpinus betulus (WA 27758) confirmed the identification and the occurrence of this species in Central Europe. Previous authors reported the species from Austria (Migula 1934), Canada (Hughes 1980a) and Finland (original description). Senthilkumar et al. (1993) mentioned a species that he identified as T. exilis during a study about the successional pattern of the mycoflora associated with litter degradation in a Cymbopogon caesiusdominated tropical grassland in South India; the correctness of the identification of this collection is, however, doubtful and not verifiable.

ACCEPTED MANUSCRIPT A sequence erroneously referred to as “Taeniolella exilis” (IP 2199.93) was included in a phylogenetic analysis based on partial LS rRNA sequences by Masclaux et al. (1995). The sequenced material was isolated from a human skin lesion, whereas genuine T. exilis is usually found on bark, which raised doubts about the correct identification of this strain. In the tree published by Masclaux et al. (1995), the sequence based on this culture isolated from human skin clustered adjacent to the type strain (CBS 146.33) Cladosporium elatum (now Ochrocladosporium elatum), which phylogenetically belongs to

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Pleosporales, Incertae sedis (Crous et al. 2007), which is in severe conflict with the recently confirmed phylogenetic position of true T. exilis within the Kirschsteiniotheliaceae (see Fig 1).

In a paper dealing with the morphology of T. rudis, Jones et al. (2002) listed several examined specimens of Taeniolella species, including one collection of T. exilis (IMI 76361). The type material

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of T. exilis (deposited at H, but unfortunately currently not traceable) was not examined. Jones et al. (2002) noted that the examined material of T. exilis ‘did possess a penicillate head’ comparable to similar structures in the aquatic species dealt with in this paper (T. rudis). Several collections of T.

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exilis have been examined in the course of a revision of the genus Taeniolella, but no trace of any synanamorph has been found, which implies that the observations of Jones et al. (2002) are unclear and doubtful. The molecular data and morphological peculiarities of the synanamorphs (penicillately branched heads and colourless conidia) formed by T. rudis justify the reallocation of this species to the genus Sterigmatobotrys.

The colonies of Taeniolella exilis are effuse or more or less restricted to lenticels, usually dense,

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narrowly oval, somewhat sooty and 1−17 × 0.5−2 mm. These distinctive characteristics and the presence of well-developed stromata facilitate the differentiation from other saprophytic Taeniolella species. Taeniolella alta, a similar saprobic species, mainly occurs on bark of branches or roots of Alnus spp., whereas T. exilis mainly inhabits Betula spp., Carpinus betulus and Quercus robur. The

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mycelium of T. alta grows superficially and partly immersed, and the hyphae are narrower (1.5–5 µm wide, vs. (2–)3−10 µm in T. exilis). Taeniolella alta lacks the true stromata as frequently formed in T. exilis (150–320 × 30–60 µm). Furthermore, the conidiophores of T. alta are usually irregularly

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verruculose to verrucose and only rarely smooth; specifically the basal wall of young conidiophores and the walls of supporting hyphal cells are usually distinctly verruculose or verrucose, while the upper part of older conidiophores is less roughened. In contrast, the conidiophores of T. exilis are usually smooth. Conidia in both species are doliiform, subcylindrical to nearly obclavate and of similar size (19–108(–180) × 8–14(−17) µm, 0–13-euseptate in T. exilis, vs. 0–12-septate, 9–107 × 7– 14 µm in T. alta). Taeniolella subsessilis, a similar saprobic species mainly occurring on bark of Smilax hispida, often forms stromatically aggregated cells at the base of conidiophores, but these aggregations are less pronounced than the crust-like stromata, composed of flattened cell layers 1 to 4 cells thick, in T. exilis. Furthermore, the conidiophores of T. subsessilis are usually distinctly shorter (8–28 × 6–8 µm,

ACCEPTED MANUSCRIPT vs. (11–)30−140(–200) × 8−14 µm in T. exilis) and the conidia are usually narrower (15–60 × 7–11 µm, vs. 19–108(–180) × 8-17 µm in T. exilis). Taeniolella hawksworthiana Heuchert, Ertz & Common sp. nov. Figs 7–8. MycoBank No.: MB 817107

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Etym.: The new species is dedicated to the British mycologist and lichenologist David L. Hawksworth on the occasion of his 70th birthday.

Diagn.: Morphologically close to Taeniolella friesii, but conidiophores wider, 5–6 µm; conidia mainly 2-septate, 10−12 × 4–5 µm, adhering in long (up to 60 µm), not easily disintegrating chains.

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Type: USA, Florida, Hillsborough Co., Hillsborough River State Park, 28º08.60’ N, 82º13.79’ W, on Phaeographis cf. brasiliensis, 3 Sept. 2011, R. Common 9199B (BR – holotype; HAL 3031 F, herb. Diederich – isotypes).

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Colonies on lichen thalli, effuse, aggregated in tufts or loose groups, confluent, loosely to densely caespitose, dark brown to black, not causing any discoloration of the thallus. Mycelium inconspicuous, immersed; hyphae flexuous, branched, 3–6 µm wide, septate, slightly constricted at the septa, subhyaline to pale brown, wall thin, up to 0.25 µm, smooth. Stromata lacking; hyphal cells sometimes swollen, subglobose, 4–7 × 5 µm, brown, smooth-walled, rarely aggregated at the base of conidiophores. Conidiophores semi-micronematous, usually reduced to conidiogenous cells,

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mononematous, solitary, arising from hyphae or swollen hyphal cells, loosely to densely aggregated, erect, straight, short, subcylindrical-conical, doliiform, unbranched, 8–15(–20) × 5–6 µm, 0–3(–4)septate, not or slightly constricted at the septa, brown, smooth; wall somewhat thickened, usually 0.5 µm. Conidiogenous cells integrated, terminal, monoblastic, monopodial, doliiform, 4–5 µm long;

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conidiogenous loci truncate, unthickened, 1–3 µm diam. Conidia in long, usually unbranched to sometimes branched chains, chains not easily disarticulating, up to 60 µm long, disintegrating in fragments of different sizes, 3–5-septate, 16–24 × 4–5 µm, conidia straight, ellipsoid, ovoid,

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subcylindrical, (0–)1–2(–3)-septate, aseptate conidia 5–6 × 4–5 µm, 1-septate ones 7−9 × 4−5.5 µm, 2septate ones 10−13 × 4–5 µm, 3-septate ones 13 × 5 µm, constricted at the septa, brown to dark brown, smooth to irregularly verrucose, wall 0.25–1 µm thick, apex rounded in primary conidia, truncate in secondary ones, base truncate, hila truncate, unthickened, not darkened, 1–2 µm diam. Host range and distribution: Phaeographis cf. brasiliensis. USA. Additional specimen examined: On Phaeographis sp.: USA: Florida, Hillsborough Co., Hillsborough River State Park, trail from parking area 2, 28º08.94’ N, 82º13.61’ W, R. Common 9215N (BR, HAL, herb. Diederich). Notes: The new species is very similar to Taeniolella friesii on Strigula stigmatella. The conidiophores are also semi-micronematous but are usually somewhat wider, 8–15(–20) × 5–6 µm, vs.

ACCEPTED MANUSCRIPT 3–12(–15) × (1.5–)3–5 µm in T. friesii. The predominantly 1-septate conidia (5−9(–10) × 3−5 µm) in T. friesii are formed singly or in very short disarticulating chains, whereas in T. hawksworthiana the usually 2-septate conidia (10−12 × 4–5 µm) adhere in long (up to 60 µm), not easily disintegrating chains. The conidial chains are sometimes disarticulating in fragments of different sizes. Unfortunately, molecular data for T. friesii are not available. Based on evident morphological

as a separate species, namely T. hawksworthiana.

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differences and the different hosts of both species, we prefer to describe the species on Phaeographis

The morphological and molecular differences between Taeniolella hawksworthiana and Taeniolella sp. growing on cf. Phaeographis in Rwanda, are discussed in detail under Taeniolella sp.

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Taeniolella punctata M.S. Christ. & D. Hawksw., in Hawksworth, Bull. Brit. Mus. (Nat. Hist.), Bot. 6: 257 (1979).

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Figs 9–11.

Type: Denmark (Lolland), Ryde, W of Maribo, on Carpinus in the wood Kristianssæde Skov, on Graphis scripta, alt. 9–15 m, 24 Jul. 1977, M.S. Christiansen 77.140 (IMI 225002 – holotype, C 6062 – isotype!).

Lit.: Diederich (1989: 253), Clauzade et al. (1989: 121), Montijūnaitė and Anderson (2003: 83). Ill.: Hawksworth (1979: 258, Fig. 37), Diederich et al. (2016).

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Colonies effuse, brown to black, scattered over the host thallus, sometimes in dead or dying portions with grey discolorations, loosely punctiform to densely caespitose, in small tufts, confluent, up to 0.1 mm, usually without any discoloration of the lichen thallus. Mycelium rather sparsely developed; hyphae immersed, flexuous, branched, 3–6 µm wide, septate, constricted at the septa, subhyaline to

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pale brown, smooth; walls slightly thickened, up to 0.5 µm. Stromata lacking, but with some swollen, subglobose rarely aggregated hyphal cells, up to 6 µm diam. Conidiophores macronematous,

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mononematous, sometimes solitary, but usually 3−10 in small caespitose tufts, arising from swollen hyphal cells, straight to slightly flexuous, unbranched or mostly once branched at the base, rarely branched in the upper third, subcylindrical; conidiophores (with adhering conidia) 14−83(–95) × 5−8 µm, 2−25-septate; septa up to 0.75 µm thick, slightly constricted at the septa, brown to dark brown, paler towards the apex; tips of conidiophores and/or the adhering terminal conidium sometimes somewhat swollen, i.e. either entire terminal conium or only conidial tip swollen, up to 9 µm wide; wall light microscopically smooth, occasionally slightly rugose or verruculose, scanning electron microscopically usually verruculose; walls thickened, 1(–1.5) µm, thinner or not thickened toward the apex, enteroblastically proliferating with obvious sheath-like wall remnants visible as irregular collar. Conidiogenous cell integrated, terminal, monoblastic or thalloblastic, monopodial, subcylindrical, doliiform, 3–9 µm long, little differentiated, loci truncate, unthickened, 2.5–5 µm diam. Conidia catenate, in unbranched chains, not easily disintegrating, conidiophores often breaking up at the basis

ACCEPTED MANUSCRIPT or separating into fragments of different sizes, conidia doliiform, broadly subcylindrical, ellipsoid, (0– )1–3-septate, fragments up to 6-septate, aseptate conidia 7−10 × 5−6 µm, 1-septate ones 8−18 × 5−7 µm, 2-septate ones 10−21 × 5–7 µm, 3-septate ones 13−21 × 5−8 µm, 4- to 6-septate fragments 12–28 × 5–7 µm, mostly constricted at the septa, brown to dark brown, wall light microscopically smooth, occasionally slightly rugose or verruculose, scanning electron microscopically usually verruculose, walls thickened, 0.75–1 µm, apex rounded, sometimes swollen in primary conidia, truncate in

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secondary ones, base truncate, sometimes narrowed towards the base, hila truncate, unthickened, not darkened, 2–5 µm diam.

Host range and distribution: Arthonia atra, A. radiata, A. ruana, Fissurina dumastii, Graphis scripta, Graphis sp., Pertusaria leioplaca, Phaeographis dendritica. Austria (Hafellner and Maurer 1994;

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Berger and Türk 1994, 1995; Hafellner 2008), Belgium (Diederich 1986; van den Boom et al. 1998; Diederich and Sérusiaux 2000, Diederich et al. 2016), Czech Republic (Kocourková and van den Boom 2005), Denmark (Hawksworth 1979; Alstrup et al. 2004), Estonia (Suija 2005; Suija et al.

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2007), France (Diederich and Roux 1991; Sparrius et al. 2002; Roux et coll. 2014, Diederich et al. 2016), Germany (Diederich 1986; John 1990; Scholz 2000; Bruyn 2001; Triebel and Scholz 2001; Rätzel et al. 2003; Eichler and Cezanne 2003; Cezanne and Eichler 2004, 2015; Bruyn 2005; Kocourková and Brackel 2005; Otte et al. 2006; Cezanne et al. 2008; Bruyn et al. 2008; Brackel 2010; Wirth et al. 2010; John et al. 2011; Schiefelbein 2013), Ireland (Fox 2001), Lithuania (Montijūnaitė and Andersson 2003), Luxembourg (Diederich 1986, 1990; van den Boom et al. 1998; Diederich and

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Sérusiaux 2000; Diederich et al. 2004; Diederich et al. 2016), Netherlands (Sparrius 2000; Aptroot et al. 2004; www.verspreidingsatlas.nl); Poland (Jando and Kukwa 2003; Fałtynowicz 2003; Zalewska and Fałtynowicz 2004; Czyżewska et al. 2005; Kukwa 2005; Kukwa and Czarnota 2006; Czyżewska et al. 2008; Szymczyk and Zalewska 2008; Schiefelbein et al. 2012; Kukwa et al. 2013), Portugal

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(Azores) (Berger and Aptroot 2002; Hafellner 2005, Borges et al. 2010), Russia (Republic Adygeja) (Otte 2004; Kukwa and Jabłońska 2008), Spain (Etayo 2002, 2006, 2010), Sweden (Santesson 1993),

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Ukraine (Bielczyk et al. 2005), United Kingdom (Hawksworth 1990). Additional specimens examined: On Arthonia atra: Belgium: Wellin, à 4 km au sud de Chanly, ruisseau le Glan, alt. 260 m, tronc de Carpinus, 29 Jan. 2012, D. Ertz 17390 (BR, sub Arthonia atra). On Arthonia ruana: Denmark: (Jutland) Yding, WSW of Skanderborg, on the loose bark of a young dead Fraxinus near the brook “Bjergskov Bæk” in the wood “Yding Skov”, alt. 100 m, UTM 32VNH505072, 26 May 1983, M.S. Christiansen 83.020; 83.018 (C 2702; 2704). On Fissurina dumastii: Portugal: Azores, Pico, S of Sao Roque do Pico, forest remnants on the shore of Lagoa Capitao, alt. 780 m, 38°29’9” N, 28°18’58” E, on Juniperus brevifolia, 24 Jul. 2010, P. Diederich 17044 (BR, herb. Diederich); N of Najes do Pico, near Lagoa do Paúl, alt. 790 m, 38°25’58” N, 28°13’58” W, on branches of Juniperus brevifolia, 25 Jul. 2010, P. Diederich 17027 (herb. Diederich). On Graphis scripta: Belgium: (distr. ardennais), Prov. Luxembourg, St-Hubert, valley of the Masblette, near the Point Mauricy, alt. 320 m, ancient mixed woodland by the river, on Fagus, 5

ACCEPTED MANUSCRIPT Jun. 1997, P. Diederich 12647 (herb. Diederich). Denmark: Bornholm, Åkirkeby par., Almindingen, Ekkodalen at Fugelsangsrende, on Carpinus betulus in shaded position, 55°06’ N, 14°53’ E, 30 Jun. 1987, M. Wedin 537 (UPS); EJ, Østjylland, Ry Nørreskov, Ringhoved, 13 May 1995, V. Alstrup (C 1962); NJ, Himmerland, Ravnkilde, on beech in the wood Nörlund skov (Rold Skov), 31 Oct. 1971, M.S. Christiansen (C 2751); NMJ, Salling, Brigshøj, Krat, UTM 32VMH8175, 22 Oct. 2002, V. Alstrup (C 5739); Zealand, Nordrupöster, east of Ringsted, on the smooth bark of young oak in the

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woods around the manor house Giesegaard, 14 Sep. 1966, M.S. Christiansen 66.750a (C 2886); Vemmetofte, in the wood “Vemmetofte Dyrehave”, alt. 5–10 m, UTM 33UUB240270-3, on Corylus, 6 Aug. 1982, M.S. Christiansen 82.077 (H, C 2470, C 2471, M-0043798, duplicates distributed in A. Vĕzda: Lich. Sel. Exs. nr. 1925). Estonia: Ida-Viru county, Puhatu Nature reserve, Kivinõmme

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forestry, deciduous forest, 59°10’31” N, 27°38’26” E, on Alnus incana, 2 Sep. 2006, A. Suija 123 (TU-45018). France: Pyrénées-Atlantiques, au sud de Tardets-Sorholus, Ste-Engrâce, vers Pierre-StMartin, on Fagus, 17 Jul. 1991, P. Diederich 9521 & J. Etayo (herb. Diederich); Pas-de-Calais, forêt

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domaniale de Desvres, parc. 22, Lamb1 562.8/1332.3, alt. 50 m, on Fraxinus, 10 Aug. 2000, P. Diederich 14297 & J. Signoret (herb. Diederich). Germany: Baden-Württemberg, Nördliches Oberrheintiefland, Hainbuchen-Eschen-Bestand im Kastenwört WSW von Karlsruhe, 107 m, TK 7015-2, 20 Feb. 2007, R. Cezanne & M. Eichler (Cezanne-Eichler 7278); Bavaria, Menterschwaige, bei München, auf Rinde von Hainbuche, A. v. Krempelhuber, 1859 (M-0043800); Schwaben, Günzburg/Donau, Donauauen “Leibi” auf Tilia, 446 m, 29 Mar. 1963, Doppelbauer (M-0043799);

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Schwaben, Kreis Oberallgäu, Kürnacher Wald W Kempten, Ulmertal, on Fagus sylvatica im Mischwald am Bach, 8226/4, 820 m, 9 Sep. 2004, J. Kocourková & W. v. Brackel (IVL 2983); Kreis Oberallgäu, Weißbachtal, SW Oberstaufen, 700 m SW Steinebach, am Stamm von Corylus avellana, 8425/4, 700 m, 10 Sep. 2004, J. Kocourková & W. v. Brackel (IVL 2862); Unterfranken, Kreis

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Haßberge, ehem. Standortübungsplatz Ebern, S Unterpreppach, an Hainbuche im Eichen-HainbuchenWald, 5930/2, 300 m, 19 Oct. 2008, W. v. Brackel (IVL 4803); Rheinland-Pfalz, S Manderscheid, vallée de la Kleine Kyll, 19 Jun. 1984, on Carpinus, P. Diederich 5524 (herb. Diederich).

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Luxembourg: entre Mersch et Angelsberg, vallon du Bënzelterbaach, 1 Jul. 1999, sur Carpinus, P. Diederich 13824 (herb. Diederich); S of Capellen, Jongebësch, 26 Mar. 2005, on Carpinus, P. Diederich 16040 (herb. Diederich, also Santesson Fungi Lich. Exs. 371); Vogelsmühle, vallon du Halerbaach, rive gauche, on Fagus, 9 Dec. 2007, P. Diederich 16718 (herb. Diederich). Spain: Catuluña, Prov. Barcelona, km 32 de la carretera de Cantonigros a Olot, alt. 900 m, 12 Feb. 1991, P. Diederich 9855 (herb. Diederich). United Kingdom: Isle of Skye, S Broadford, Kilmore, churchyard and rocks near the sea, NG 66 07, on Corylus, 30 May 1987, P. Diederich 8818 (herb. Diederich). On Pertusaria cf. leioplaca: Germany: Baden-Württemberg, Nördliches Oberrheintiefland, alter Hainbuchen-Bestand im Kastenwört WSW von Karlsruhe, 107 m, TK 7015-2, 20 Feb. 2007, R. Cezanne & M. Eichler (Cezanne-Eichler 7279).

ACCEPTED MANUSCRIPT Notes: Hawksworth (1979) and Diederich (1989) considered Taeniolella punctata as a true lichenicolous species, which can be confirmed on the basis of the examination of numerous specimens. The colonies of this species do not penetrate into the adjacent cortical tissue. The principal host is Graphis scripta, but there are additional collections on other hosts (see above), which were confirmed by our phylogenetic analyses for Arthonia atra and Pertusaria leioplaca (Fig. 2). Taeniolella punctata is the only Taeniolella species known to occur on Graphis scripta. The

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collections on Thelotrema lepadinum reported by Diederich (1986) and Clauzade et al. (1989) were reexamined and identified as belonging to a similar new species, Taeniolella toruloides. Compared to Taeniolella punctata, T. toruloides has unbranched and shorter conidiophores (6–34 × 4–7 µm, 0−5septate), lacks conspicuously thickened conidial septa, and has conidial chains that are not easily

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disintegrating, but usually toruloid. In our phylogenetic analyses (Fig. 2), Taeniolella punctata clusters with Buelliella minimula and Karschia cezannei in an unsupported polytomy. The species is clearly distinct from two strains of Taeniolella toruloides isolated from Thelotrema antoninii. Unfortunately,

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we were unable to obtain sequences from material on Thelotrema lepadinum for analysis. We examined two collections on Fissurina dumastii from the Azores with conidiophores and conidia measurements within the range of variability of Taeniolella punctata. The transition between hyphae and conidiophores is sometimes not very evident in this material. We observed brown to dark brown hyphae, in addition to the subhyaline to pale brown hyphae typical for T. punctata. Hyphal cells are are densely aggregated, branched, and up to 8 µm wide; the septa are up to 1 µm thick and rarely

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oblique; the wall is smooth and thickened, up to 0.5 µm. The colonies sometimes develop on damaged portions of the thallus with a grey discoloration. As Fissurina is a recent segregate of Graphis, according to Lücking (2009) and Dal-Forno (2009), older records of T. punctata on Graphis sp. from the Azores (Berger and Aptroot 2002; Hafellner 2005, Borges et al. 2010) might also refer to material

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on Fissurina. A specimen on Fissurina dumastii from the Azores (P. Diederich 17044) was successfully cultured and does not cluster with other sequences from Taeniolella punctata in our phylogenetic analyses. However, this specimen is placed close to the cluster of T. punctata in a lineage

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for which the internal relationships are poorly supported (Fig. 2). For that reason, and owing to morphological similarities, the collections on Fissurina dumastii from the Azores are refered to Taeniolella cf. punctata.

The wall of conidiophores and conidia in Taeniolella punctata appears to be smooth by light microscopy, as described in Hawksworth (1979), Diederich (1989) and Montijūnaitė and Andersson (2003), but we occasionally observed them to be slightly rugose or verrucose. Examination by scanning electron microscopy showed that the wall is usually finely verruculose. Taeniolella caespitosa is very similar to T. punctata; the two species are barely distinguishable using dimensions of conidiophores and conidia. Conidiophores in T. punctata are slightly longer (14−83(– 95) × 5−8 µm, vs. 7–71(–81) × (4.5–)5–8 µm). In T. punctata the tips of the conidiophores and/or the adhering terminal conidium are occasionally swollen up to 9 µm wide (see arrows in Fig 9F). The wall

ACCEPTED MANUSCRIPT of the conidiophores in the examined material of T. caespitosa is mostly smooth, occasionaly somewhat irregularly rough, but without cracks and squamules, whereas in T. punctata the conidiophores wall can be smooth, slightly rugose or verruculose. The dimorphic mycelium of T. caespitosa can be composed on the one hand of pale brown, flexuous to tortuous, 1.5–3 µm wide, smooth and thick-walled hyphae that penetrate the host thallus, or composed of subhyaline to pale brown, densely aggregated, 3-10 µm wide, irregularly shaped, thin-walled, hyphae with a granular

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surface entering the underlying cortex cells. In the latter case, the hyphae are developed only around the base of conidiophores. Such dimorphic hyphae are unknown in Taeniolella punctata. In addition to the discussed differences, T. punctata grows preferably on Graphis scripta and only rarely on other hosts such as Pertusaria leioplaca, whereas T. caespitosa is undoubtedly confined to Pertusaria spp.

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In the phylogenetic analysis, a single specimen from Germany on Pertusaria leioplaca (CPS 14809 = Cezanne-Eichler 7279) proved to be T. punctata, which is in accordance with the morphology of this

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sample, above all the lacking formation of immersed dimorphic hyphae. Taeniolella pyrenulae Heuchert & Diederich sp. nov. Figs 12–13. MycoBank No.: MB 817108

Etym.: Epithet derived from the host genus, Pyrenula.

Diagn.: Morphologically close to Taeniolella friesii, but with distinctly longer and wider conidiophores and conidia. Distinguished from T. toruloides by having wider, clearly structured,

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verrucose-rugose and sometimes rimulose conidia.

Type: Portugal, Azores, Pico, between Lajes do Pico and Sao Roque do Pico, Bosque da Junqueira, 1 km S of crossing with road going to the east, 38°27’56” N, 28°17’57” W, on Pyrenula cf. hibernica, on Vaccinium cylindraceum, in laurisilva, 26 Jul. 2010, P. Diederich 17075 (BR – holotype; HAL

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3032 F, herb. Diederich – isotypes)

Colonies on the surface of thalli and perithecia, in slight infections forming small tufts, effuse,

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confluent, loosely caespitose, in severe infections densely caespitose over the entire lichen thallus, black; thallus without any discoloration. Mycelium immersed; hyphae flexuous, branched, 2–7µm wide, septate, mostly constricted at the septa, pale brown to brown, more intensively pigmented below the conidiophores, smooth, walls thickened, 0.25–0.5 µm wide. Stromata lacking, but with swollen, rarely aggregated hyphal cells, subglobose, ellipsoid or square, 3–6 µm diam. Conidiophores semimicronematous, usually reduced to conidiogenous cells, not easily distinguishable from swollen hyphal cells, mononematous, solitary or in small groups, erect, straight, unbranched, subcylindrical, ovoid, doliiform, 3–7 µm long and wide, aseptate, brown, thick-walled, 0.25–0.5 µm wide, smooth. Conidiophores aseptate, i.e. reduced to conidiogenous cells, monoblastic, monopodial, conidiogenous loci truncate, unthickened, 2−4 µm diam. Conidia catenate, long-adhering in unbranched chains, not easily disarticulating, chains up to 100 µm long, but later disintegrating into fragments of different sizes, conidia usually easily discernible within the chain by obvious constrictions between individual

ACCEPTED MANUSCRIPT conidia, enteroblastically proliferating with obvious sheath-like wall remnants visible as irregular collar, straight, ellipsoid, subcylindrical to broadly subcylindrical, doliiform, pyriform, ovoid, 1–4septate, septa often conspicuously thickened, up to 1.25 µm, 1-septate ones 6–14 × 5–8 µm, 2-septate ones 11–20 × 5–8 µm, 3-septate ones 15–25 × 6–7 µm, 4-septate ones 18–33 × 6–8(–9) µm, slightly or not constricted at the septa, brown to dark brown, wall thickened, 0.5–2 µm wide, sometimes distinctly multi-layered, usually clearly structured, verrucose-rugose, sometimes rimulose, apex rounded in

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primary conidia, truncate in secondary ones, base truncate, sometimes narrowed, hila truncate, unthickened, not darkened, 1.5–4(–5) µm diam.

Host range and distribution: Pyrenula cf. hibernica, P. laevigata. Portugal (Azores), Russia. N.B.: Taeniolella pyrenulae does not colonize Pyrenula dermatodes, although this species is present in the

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type specimen.

Additional specimens examined: On Pyrenula laevigata: Russia: Republik Adygeja, Majkopskij Rajon, Gebiet des Berges Bol. Tcvač, Talgrund beim Lagerplatz am oberen Nal-Sachrai, 44°06’ N,

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40°24’ E, ca. 950 m, on Alnus incana, 24 Aug. 2003, V. Otte (GLM 23601); Gebiet des Berges Bol. Tcvač, Talgrund beim Bache Bol. Sachrai oberhalb der letzten Furt vor dem Zusammenfluss mit dem Nal-Sachrai, 44°05’30” N, 40°23’ E, ca. 920 m, an jungen noch glattrindigen Acer cf. trautvetteri/pseudoplatanus, 30 Aug. 2003, V. Otte (GLM 23720); ibid., an totem jungen Ulmus, 30 Aug. 2003, V. Otte (GLM 23609). On Pyrenula sp.: Portugal: Azores, Pico, NW of Lages, near Cabeço do Farrobo, 38°26’29” N, 28°16’17” W, ca. 600 m, trunk in remnants of laurisilva along a

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road, 25 Aug. 2011, D. Ertz 16807 (BR).

Notes: The specimen from the Azores is the best developed sample of this species, with abundant conidiophores, and is therefore selected as the holotype. All collections from Russia are less welldeveloped: the conidial chains are often shorter and the walls are less thickened and less structured. In

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specimen GLM 23609 from Russia, aseptate, smooth conidia (7–8 × 4.5–5.5 µm) were occasionally observed. All other features agree with those of the other specimens. Therefore, the collections from

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Russia and from the Azores are assigned to a single new species. Taeniolella friesii, currently known only on Strigula stigmatella, is similar to the new species. The conidiophores are also micronematous and the conidia are solitary or in short disarticulating chains. However, T. friesii is easily distinguishable from T. pyrenulae by having distinctly shorter and narrower conidia (e.g. 2-septate ones 10−12 × 4–5 µm, vs. 11–20 × 5–8 µm in T. pyrenulae). The new species T. toruloides on Thelotrema is also similar to T. pyrenulae by having conidia that adhere in long firm chains up to 100 µm, but the conidia are usually distinctly narrower (e.g. 2-septate ones 14–22 × 4.5–6 µm, vs. 11–20 × 5–8 µm in T. pyrenulae). Additionally, T. toruloides conidia are mostly smooth, except older ones that may be verrucose and with single cracks, compared to the textured conidia of T. pyrenulae.

ACCEPTED MANUSCRIPT The holotype of T. pyrenulae was successfully cultured and sequenced. It is the sister species to two unidentified lichenicolous Melaspilea s. lat. specimens from D.R. Congo and growing on Pyrenula. However, this relationship is only supported by the Bayesian analysis.

Taeniolella sp.

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Figs 14–15.

Specimen examined: Rwanda, Province de l’Ouest, forêt de Nyungwe, Uwinka, sentier vers les

chutes d’eau, forêt dense, sure tronc, alt. 2460 m, 2°28’ S, 29°12’ E, on cf. Phaeographis sp., 9 Aug. 2007, D. Ertz 11026 (BR-LICH 2029-89).

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Colonies on lichen thalli, effuse, aggregated in tufts, colonies punctiform, confluent, up to 1 mm diam., caespitose, dark brown to black, not causing any discoloration of the thallus. Mycelium

inconspicuous, immersed; hyphae flexuous, branched, 3–6 µm wide, septate, slightly constricted at the

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septa, subhyaline to pale brown; wall thin, up to 0.25 µm, smooth. Conidiophores semimacronematous to macronematous, mononematous, solitary, arising from hyphae, loosely to densely aggregated, erect, straight or slightly flexuous, subcylindrical, doliiform, unbranched, 12–70 × 5–6(–7) µm, 1–10(–13)-septate; septa conspicuous, 1–2 µm thick, slightly to distinctly constricted at the septa; conidiophores dark brown, smooth or rarely irregularly rough; wall thickened, 0.5–2 µm, cell plasma mostly reduced, with a central vacuole-like cavity; surrounding plasma giving the impression of a very

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thick wall, rarely enteroblastically proliferating with obvious sheath-like wall remnants visible as irregular collar. Conidiogenous cells integrated, terminal, monoblastic, monopodial, doliiform, 5–7 µm long; conidiogenous loci truncate, unthickened, 3–4(–5) µm diam. Conidia in long, usually unbranched chains; chains not easily disarticulating, up to 70 µm long, disintegrating in fragments of

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different sizes, 3–9-septate, 19–46 × 6–7 µm; conidia straight or slightly sinuous, ellipsoid, ovoid, subcylindrical, 1–5-septate, 1-septate conidia 9–10 × 5–6 µm, 2-septate ones 10−18 × 6 µm, 3-septate

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ones 15–20 × 6–6.5 µm, 4 and 5-septate 21–31 × 5.5–7 µm, slightly to distinctly constricted at the septa, septa conspicuously thickened and darkened, 1–1.5 µm thick, brown, young conidia at the apex paler, smooth, rarely irregularly rough, often rough at the apex, wall 0.5–2 µm thick, cell plasma mostly reduced, with a central vacuole-like cavity, surrounding plasma giving the impression of very thick wall, vacuoles usually with 1–2 oil-like droplets, wall thinner at the apex, apex rounded in primary conidia, truncate in secondary ones, base truncate and often narrowed, hila truncate, unthickened, not darkened, 2–4 µm diam. Host range and distribution: cf. Phaeographis sp. Rwanda. Notes: The sample on Phaeographis sp. from Rwanda is morphologically similar to Taeniolella hawksworthiana, a new species on Phaeographis sp. from Florida. The conidiophores in both species are not easily distinguishable from conidia adhering in long chains. However, in contrast to semi-

ACCEPTED MANUSCRIPT macronematous [12–70 × 5–6(–7) µm] conidiophores in Taeniolella sp. from Rwanda, the conidiophores in T. hawksworthiana are much shorter and semi-micronematous [8–15(–20) × 5–6 µm]. The conidia in T. hawksworthiana are distinctly smaller and narrower, (0–)1–2(–3)-septate, 6–13 × 4–5 µm, vs. 1–5-septate, 9–31 × 5–7 µm in Taeniolella sp.; and the hila in T. hawksworthiana are accordingly narrower (1–2 µm, vs. 2–4 µm in Taeniolella sp.). The position of both species in the mtSSU+nuLSU-tree indicates a close relationship. The small number of specimens known from

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Phaeographis does not allow us understanding if they represent two distinct species, T.

hawksworthiana from Azores and an unnamed species from Rwanda, or if they belong to a single, morphologically variable species.

The septa in the conidiophores and conidia in Taeniolella sp. from Rwanda are conspicuously

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thickened, up to 2 µm, which is similar to other Taeniolella species, e.g. T. pyrenulae known on

Pyrenula cf. hibernica from Azores and P. laevigata (Pyrenulales, Pyrenulaceae) from Russia. The material from the Azores and Russia differs in having slightly wider, 1–4-septate conidia, 6–33 × 5–

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8(–9) µm (vs. 1–5-septate, 9–31 × 5–7 µm in the specimens from Rwanda), and usually clearly sculptured, verrucose-rugose, sometimes rimulose walls (vs. smooth, rarely irregularly rough in the specimen from Rwanda). Taeniolella pyrenulae is phylogenetically clearly distinct from Taeniolella sp. from Rwanda (Fig. 2).

Taeniolella punctata is also very similar to the specimen from Rwanda. The most common host of T. punctata, Graphis scripta, and the host of Taeniolella sp. belong the same family (Graphidaceae), The

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morphological features of conidiophores and conidia in both species of Taeniolella are very similar. The conidiophores of T. punctata are unbranched or only once branched at the base, rarely branched in the upper third. The tips of conidiophores are often swollen up to 9 µm, and the cell plasma is not conspicuously reduced and without oil-like droplets. Taeniolella punctata and both Taeniolella

separate species.

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populations on Phaeographis sp. are phylogenetically clearly distinct (Fig. 2) supporting at least two

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In the specimen examined, the host lichen Phaeographis sp. is additionally colonized by Tremella phaeographinae, which is a rare lichenicolous basidiomycete previously known only from Florida (Diederich 1996). Interestingly, Taeniolella sp. grows both on the thallus of Phaeographis and as a hyperparasite on the basidiomata of Tremella.

Taeniolella toruloides Heuchert & Diederich sp. nov. Figs 16–17. MycoBank No.: MB 817109 Etym.: Epithet referring to the superficial similarity of the conidial chains to those of Torula species with distinct constrictions.

ACCEPTED MANUSCRIPT Diagn.: Morphologically close to Taeniolella friesii, but with distinctly longer and wider conidiophores and conidia. Distinguished from T. pyrenulae by having narrower, smooth conidia. Type: Portugal, Azores, Pico, S of Sao Roque do Pico, forest remnants on the shore of Lagoa Capitao, alt. 780 m, 38°29’9” N, 28°18’58” E, on Thelotrema antoninii, on Juniperus brevifolia, 24 Jul. 2010, P. Diederich 17047 (BR – holotype; herb Diederich – isotype). Colonies on the surface of thalli and apothecia, punctiform, up to 0.1 mm diam., effuse, confluent,

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loosely to densely caespitose, black; thallus without any discoloration or sometimes with grey

discoloration. Mycelium immersed, inconspicuous; hyphae flexuous, branched, 2–6 µm wide, septate, sometimes constricted at the septa; cells ellipsoid to irregularly shaped, subhyaline to pale brown, rarely brown, smooth, walls slightly to distinctly thickened, up to 1.5 µm wide; cell lumen reduced.

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Stromata poorly formed with swollen, rarely aggregated hyphal cells, subglobose, 5–8 µm diam.

Conidiophores micronematous or semi-macronematous, mononematous, solitary or in dense fascicles, arising from hyphae, lateral and terminal, or arising from swollen hyphal cells, erect, straight,

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unbranched, subcylindrical, doliiform, 6–34 × 4–7 µm, 0−5-septate, non-constricted or slightly constricted at the septa, brown to dark brown, paler at the base, thick-walled, 0.25–0.75(–1) µm wide, smooth, with age becoming rimulose-rugose or irregularly verrucose, rarely enteroblastically proliferating with obvious sheath-like wall remnants visible as irregular collar. Conidiogenous cell integrated, terminal, monoblastic, monopodial, short cylindrical, narrowed at the apex, 5−7 µm long; conidiogenous loci truncate, unthickened, 2−3 µm diam. Conidia catenate, mostly in unbranched

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chains, adhering in long firm, toruloid chains, i.e. with distinct constrictions, not easily disarticulating, chains up to 100 µm long, straight, subcylindrical, doliiform, obovoid, 0–2-septate, aseptate conidia 4– 11 × 3–6 µm, 1-septate ones 9–14(–20) × 4–7 µm, 2-septate ones 14–22 × 4.5–6 µm, chains later disintegrating in fragments of different sizes, mostly constricted at the septa, brown to olivaceous-

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brown or grey, wall thickened, 0.25–1 µm wide, cell-lumen frequently with oil-like droplets, smooth, occasionally irregularly rugose-verrucose, older conidia verrucose or with single cracks in the outer

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wall, apex rounded to attenuated in primary conidia, truncate in secondary ones, base truncate, sometimes narrowed, hila truncate, unthickened, not darkened, 2–3.5 µm diam. Host range and distribution: Thelotrema antoninii, T. lepadinum. France, Luxembourg, Portugal (Azores), Spain.

Additional specimens examined: On Thelotrema antoninii: Portugal: Azores, Pico, S of Sao Roque do Pico, forest remnants on the shore of Lagoa Capitao, alt. 780 m, 38°29’9” N, 28°18’58” E, on Juniperus brevifolia, 24 Jul. 2010, P. Diederich 17048 (herb. Diederich); ibid., 22 Aug. 2011, D. Ertz 16593 (BR). On Thelotrema lepadinum: France: Pyrénées-Atlantiques, south of Tardets-Sorholus, Ste-Engrace, gorges de Kakouetta, on Buxus, 17 Jul. 1991, P. Diederich 9576 (herb. Diederich); 15 km south-southeast of Saint-Jean-de-Luz, south of Sare, forêt communale de Sare, near road D306 to Col de Lizarrieta, alt. 310–360 m, 43.26115° N, 1.6089° W, on very old trunks of Quercus, aged 300– 400 years, 26 Aug. 2015, P. Diederich 18158 (HAL 3033 F); 20 km southeast of Saint-Jean-Pied-de-

ACCEPTED MANUSCRIPT Port, forêt d’Iraty, 500 m south of Chalet Pedro, alt. 1000–1030 m, 43.03126° N, 1.07961° W, on Fagus, 3 Sep. 2015, P. Diederich 18139 (herb. Diederich). Luxembourg: Berdorf, Binzeltschlëff et Predigtstuhl, 49.81° N, 6.33° E, on Acer, 15 Aug. 1981, P. Diederich 3870 (herb. Diederich); Berdorf, Binzeltschlëff, on Acer, 11 Jun. 1984, P. Diederich 5739 (herb. Diederich). Portugal: Azores, Sao Miquel, N of Vila Franca do Campo, forest around Congro lake, alt. 470 m, 37°45’17” N, 25°24’30” W, on bark of tree, 29 Jul. 2010, P. Diederich 17015 (herb. Diederich). Spain: Navarra, au nord de

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Orbaiceta, au nord de la Fabrica de Orbaiceta, 19 Jul. 1991, P. Diederich 9626 (herb. Diederich). Notes: The morphological traits of Taeniolella punctata, with Graphis scripta (Graphidaceae) as principal host, are very similar to those of T. toruloides on Thelotrema lepadinum, which also belongs to the Graphidaceae. However, T. punctata has conidiophores that are often branched at the base or

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rarely branched in the upper third, longer (14−83(–95) × 5−8 µm), and usually have numerous septa that can be up to 0.75 µm thick. The tips of conidiophores and/or the adhering terminal conidium in T. punctata are sometimes somewhat swollen up to 9 µm (see arrows in Fig 9F). The conidial chains,

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while not easily disintegrating, as in T. toruloides, are not conspicuously toruloid. The lichenicolous Taeniolella friesii is similar to T. toruloides by forming conidia in chains with conspicuous constrictions between individual conidia, but T. friesii, known only on Strigula stigmatella, is distinguishable from T. toruloides by its shorter and narrower conidiophores (3–12(–15) × (1.5–)3–5 µm, vs. 6–34 × 4–7 µm in T. toruloides). Furthermore, the conidia, developed in very short disarticulating chains, are usually much shorter and often narrower (1-septate ones 5−9(–10) × 3−5

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µm, vs. 1-septate ones 9–14(–20) × 4–7 µm in T. toruloides). Collections of T. punctata and T. toruloides were successfully cultured. T. toruloides is phylogenetically clearly distinct from T. punctata in our mtSSU+nuLSU tree (Fig 2). It is the sister species to ‘Melaspilea sp. 18012’, a lichenicolous fungus belonging to Melaspilea s. lat., and forming

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4. Discussion

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black lirellae on the thallus of a Pyrenula.

4.1. Phylogenetic placement of the type species of Taeniolella – Our phylogenetic inference based on nuLSU sequences show for the first time the placement of the type species of Taeniolella (T. exilis, Figs 5–6) within the family Kirschsteiniotheliaceae (type: K. aethiops) and as the sister species to Kirschsteiniothelia thujina (Fig 1). Kirschsteiniotheliaceae was recently proposed by Boonmee et al. (2012) for the single genus Kirschsteiniothelia that includes saprobic fungi occurring on dead wood and forming globose, unilocular ascomata, and pigmented, septate ascospores. The family was placed as Dothideomycetes incertae sedis by Hyde et al. (2013), and thus lacks an ordinal rank. Dendryphiopsis was shown to be the asexual state of Kirschsteiniothelia aethiops and characterized by mononematous, erect, branched, septate, coloured conidiophores, and broadly ellipsoid-obovoid,

ACCEPTED MANUSCRIPT septate, light to dark brown conidia (Hughes 1953, Hawksworth 1985), and thus to be different from Taeniolella s. str. Wijayawardene et al. (2014) proposed synonymizing Dendryphiopsis with Kirschsteiniothelia. Based on our analysis, Kirschsteiniothelia thujina probably needs to be transferred to Taeniolella, but we refrain from making that taxonomic change and prefer to wait until more taxa of these genera have been sequenced. The species has also been included in the genus Splanchnonema and referred to the family Pleomassariaceae (Barr 1993). While that family is clearly part of the

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Pleosporales and thus distinct from the Kirschsteiniotheliaceae (Zhang et al. 2012, Hyde et al. 2013), sequences of the type species of Splanchnonema (S. pustulatum) are needed to establish the

phylogenetic placement of Splanchnonema. As this genus is older than Taeniolella, it will be

important to establish its phylogenetic placement before proposing taxonomic changes for the clade

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Taeniolella exilis-Kirschsteiniothelia thujina.

4.2. Polyphyly of the genus Taeniolella – The genus Taeniolella is strongly polyphyletic, with species

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being placed in two different classes, Dothideomycetes and Sordariomycetes (Fig 1). Saprobic and/or endophytic species are not allied and are distributed between both classes, while hitherto phylogenetically examined lichenicolous species are all included in the Asterotexiales within Dothideomycetes.

Dothideomycetes — Taeniolella exilis is the only Taeniolella species currently included within the

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Kirschsteiniotheliaceae. Taeniolella typhoides, isolated from submerged freshwater wood, was placed in the family Lindgomycetaceae (Pleosporales) as the sister taxon to Massariosphaeria typhicola using an 18S ribosomal RNA sequence (Shearer et al. 2009; although Taeniolella typhoides is not included in our phylogenetic tree because the nuLSU sequence is not available, note the placement of

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M. typhicola in Fig 1). The lichenicolous species that we sequenced are surprisingly placed in the Asterotexiales as newly defined (see section 3.1), which means that this order now includes plant pathogens, saprobic and lichenicolous fungi. The five species of Taeniolella do not form a

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monophyletic group within the Asterotexiales, but are intermixed with species of the genera Buelliella s. lat., Karschia, Labrocarpon, Melaspilea s. lat. and Stictographa (Fig 2). Speciation by host switching might have occurred here as has been assumed in mycoparasites (Chaverri and Samuels 2013) including lichenicolous fungi (Millanes et al. 2014, Ertz et al. 2015) because many of these species are highly host-specific. This is obvious for Taeniolella toruloides, a species that is confined to the lichen host genus Thelotrema and is the sister species to ‘Melaspilea 18012’ growing on Pyrenula. However, Taeniolella punctata, thought to have a preference for the lichen host Graphis scripta, also grows on other lichen hosts, as confirmed by our molecular results for Arthonia atra and Pertusaria leioplaca (Fig 2). These crustose lichens often grow intermixed with Graphis scripta in the same habitat, usually on the smooth bark of various trees in deciduous forests. Speciation might have recently occurred on closely related lichen hosts, as suggested by the lineage including Taeniolella

ACCEPTED MANUSCRIPT hawksworthiana and Taeniolella sp. from Rwanda (both on Phaeographis spp.) and ‘Melaspilea’ lekae, which all grow on hosts belonging to the family Graphidaceae. We hypothesize that lichenicolous Taeniolella species might represent asexual stages of other lichenicolous teleomorphic genera of Asterotexiales. However, no obvious anamorph-teleomorph relationships could be demonstrated with the molecular data available. It should be noted that a fossil record of a lichenassociated Taeniolella-like fungus, resembling extant species of Taeniolella s.lat., has recently been

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reported from Paleogene amber (Kettunen et al. 2015) dating back to at least 24 million years, suggesting that ‘the evolutionary associations between lichen-associated microfungi to their substrate must extend back much further, most probably to the Mesozoic’. The type species of Buelliella, B. minimula, is included in the Asterotexiales but does not cluster with the two other sequenced species

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of the genus, B. physciicola and B. poetschii (Fig 2). We were not able to find morphological evidence to explain this polyphyly.

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Sordariomycetes — Taeniolella species are distributed in three different orders (Fig 1), namely Diaporthales, Sordariales, and a clade lacking familial and ordinal affiliation but related to the Savoryellales (Réblová et al. 2012):

1) Diaporthales: The phylogenetic placement of Taeniolella alta in a lineage with Phomopsis sp. and species of Diaporthe was already shown in previous studies (Crous et al. 2006, Damm et al. 2007, Crous et al. 2011) using the same nuLSU sequence originally published by Masclaux et al. (1995).

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Taeniolella alta is known from corticated branches of various tree genera (e.g. Alnus, Quercus) and from decaying wood of conifers (Hughes 1980b). Species of Diaporthe-Phomopsis represent a large group of plant-inhabiting fungi, which are commonly encountered as endophytes of woody plants and are often responsible for plants diseases (e.g. Uecker 1988, Rossman et al. 2007). The published

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nuLSU sequence of T. alta was obtained from a specimen growing on Carpinus betulus in Switzerland. Its reliability is questioned here because the related culture was non-sporulating and

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could thus not be identified by one of us (B.H.). Moreover, morphological features of Taeniolella alta agree with common characteristics of the genus Taeniolella and strongly differ from species of Diaporthe (sexual) and Phomopsis (asexual), the asexual state Phomopsis being characterized by pycnidia producing hyaline, simple conidia.

2) Sordariales: Taeniolella phialosperma was isolated from strawberry rhizosphere soil in Japan (Watanabe 1992), but the nuLSU sequence does not seem to have been included in a phylogenetic tree in the past. The species was included in the Sordariales based on an ITS sequence in the framework of a phylogenetic study of thermotolerant fungi (Liang et al. 2011). It was placed in a polytomy with Sordaria fimicola, Thielavia intermedia and a clade including Corynascus div. spp., Chaetomium div. spp., Thielavia div. spp. and Humicola fuscoatra. A similar ITS sequence was obtained from an isolate of orchids roots in southwestern China (Huang and Zhang 2015). The descriptions and illustrations in

ACCEPTED MANUSCRIPT Watanabe (2002, 2010) are consistent with the original description in Watanabe (1992). The type material from TFM (Watanabe TW 73-466, culture) was not available for a re-examination. The combination of morphological features (e.g. Phialophora-like synanamorph; clavate, ellipsoidal or cylindrical large phragmospores often with an apical hyaline papillate or blactis cell) is unusual for Taeniolella s. str. and, along with the phylogenetic results, justifies the exclusion of this species from Taeniolella. However, the true generic affinity remains unclear and a re-examination of type material

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with more detailed molecular analyses and using a larger dataset of Sordariales is required.

3) Savoryellales s. lat.: Taeniolella rudis (Figs 3–4) is a species mainly confined to dead wood and bark of various conifers, and producing erect, straight, unbranched conidiophores that are almost

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indistinguishable from conidia that are ellipsoid and truncate at the ends (Hughes 1980c; see section 3.2 for more details). The strain included in our nuLSU phylogenetic tree was isolated from wood submerged in freshwater, and it groups with Sterigmatobotrys macrocarpa growing in similar habitats.

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Taeniolella rudis is also very close phylogenetically to Sterigmatobotrys macrocarpa in a phylogram inferred from ITS1-5.8S-ITS2 sequences (Réblová et al. 2012), which suggests that both taxa are closely related. The anamorph of Sterigmatobotrys macrocarpa appears to differ from Taeniolella rudis by having conidiophores with penicillate heads producing conidia (Réblová and Seifert 2011), but similar structures were recently found and illustrated for Taeniolella rudis by Jones et al. (2002). And Hughes (1980c) had previously reported terminal branching structures on some conidia of T.

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rudis and suggested the presence of a synanamorph. Despite these observations, Jones et al. (2002) still consider that Sterigmatobotrys macrocarpa differs from T. rudis by the lack of macroconidia and the presence of much longer conidiophores and shorter conidia, and this has been confirmed in the course of monographic studies on Taeniolella spp. (Heuchert et al. in prep.). Based on molecular and

taxonomic section).

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morphological evidence Taeniolella rudis belongs undoubtedly to the genus Sterigmatobotrys (see

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4.3. Conclusions – Taeniolella s. str. is placed in the Kirschsteiniotheliaceae within Dothideomycetes, and the lichenicolous species sequenced so far are confined to the Asterotexiales (Dothideomycetes). Other saprophytic/endophytic Taeniolella species previously assigned to the Sordariomycetes based on sequences were found to represent either contaminants or species that cannot be assigned to Taeniolella for morphological reasons. The present study provides phylogenetic insights into the evolution of a genus of anamorphic fungi with highly diversified life-styles. The relationship of Taeniolella species to other saprobic, endophytic or lichenicolous genera deserves further investigation. Current molecular data were not sufficient to demonstrate anamorph-teleomorph relationships. This is especially the case of lichenicolous taxa belonging to the Asterotexiales, where a considerable molecular work remains to be done before the genera can be re-appraised. Moreover, a revision of Taeniolella based on phenotypic characters suggests that the genus can be regarded as

ACCEPTED MANUSCRIPT more heterogeneous than our molecular data could highlight (Heuchert et al. in prep.). Therefore, we currently refrain from proposing any generic changes and redispositions for lichenicolous species, and prefer to maintain Taeniolella s.lat. until a much larger sample is available for reassessment. We hope that the results obtained from the present work will trigger further molecular studies on Taeniolella

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and related groups.

Acknowledgements

We are much obliged to the curators of BR, C, CANL, GLM, H, IVL, K(M), M, UPS, TU and WA as well as R. Cezanne and M. Eichler for the opportunity to examine collections from their herbaria.

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Richard C. Harris and James Lendemer are especially thanked for allowing us to sequence specimens of Buelliella minimula from NY. We are very grateful to Frank Syrowatka from Interdisciplinary Centre of Materials Science (CMAT) of Martin Luther University Halle-Wittenberg for allowing us to

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prepare ESEM examinations and for his kind technical support. The CBS-KNAW Fungal Biodiversity Centre is warmly thanked for cultures and sequences of Taeniolella exilis (CBS 122902) and of T. punctata (CPC12201 and CPC14809) used in the present study. DE is grateful to the Belgian Fonds National de la Recherche Scientifique for financial support.

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of Texas at Austin. pp 1–115.

Fig 1 – Phylogenetic relationships among 104 samples within Dothideomycetes and

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Sordariomycetes (with three outgroup taxa) based on a data set of nuLSU sequences that resulted from a Bayesian analysis using MrBayes. Posterior probabilities ≥ 95 are shown above internal branches, and Maximum Likelihood bootstrap values ≥ 70 obtained from a Garli analysis are shown below internal branches. Internal branches considered strongly supported by both analyses are represented by thicker lines. The newly sequenced specimens are in bold. Taeniolella s. lat. species are in red in the online version. Collecting numbers of the authors following species names act as specimen and sequence identifiers. Main lineages including Taeniolella species are highlighted in colour in the online version. The length of the branches represented by dashed lines was reduced by 50% for editing reason.

ACCEPTED MANUSCRIPT Fig 2 – Phylogenetic relationships among 55 samples of Asterotexiales (with three outgroup taxa) based on a data set of nuLSU and mtSSU sequences that resulted from a Bayesian analysis using MrBayes. Posterior probabilities ≥ 95 are shown above internal branches and Maximum Likelihood bootstrap values ≥ 70 obtained from a Garli analysis are shown below internal branches. Internal branches considered strongly supported by both analyses are represented by thicker lines. The newly sequenced specimens are in bold. Taeniolella species are in red in the

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online version. Collecting numbers of the authors following species names act as specimen and sequence identifiers. For the main lineage including Taeniolella species, the collecting numbers are followed by the country and by the substrate or host lichen.

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Fig 3 – Sterigmatobotrys rudis [A: BP 96761, B–D: BP 87661]. (A) conidiophores with adhering conidia; (B–D) synanamorph on extension of conidium. Scale bars: 50 µm (A), 20 µm (B, C), 10

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µm (D).

Fig 4 – Sterigmatobotrys rudis [BP 96761, BP 87661]. (A) conidiophores with adhering conidia; (B) synanamorph on extension of conidium. Scale bar = 10 µm (B. Heuchert del.). Fig 5 – Taeniolella exilis [A, I: Freebury 1968; B, C, E, G, H: ex DAOM 59235; D, F: ex DAOM 173671]. (A) macroscopical overview of colonies; (B, C, E, F) conidiophores with adhering conidia arising from hyphae or crust-like stroma; (D) conidial chain; (G, H) conidia; (I)

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germinating conidia in culture after three days. Scale bars: 1 mm (A), 200 µm (I), 50 µm (B–C), 20 µm (F), 10 µm (D–E, G–H).

Fig 6 – Taeniolella exilis [A–B: ex DAOM 59235; C–E ex DAOM 173671]. (A) conidiophores arising from hyphae or crust-like stroma; (B) conidial chain; (C) conidia; (D) conidiophores

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arising from hyphae; (E) conidiophores with enteroblastical proliferations with obvious sheathlike wall remnants visible as an irregular collar. Scale bars = 10 µm (B. Heuchert del.).

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Fig 7 – Taeniolella hawksworthiana [holotype]. (A) macroscopical overview of colonies; (B) conidiophores with adhering conidial chains; (C, E) conidia; (D, F–H) conidia adhering in long chains; (I) semi-micronematous conidiophores arising from aggregated swollen hyphal cells. Scale bars: 200 µm (A), 10 µm (B–I). Fig 8 – Taeniolella hawksworthiana [holotype]. (A) semi-micronematous conidiophores with adhering conidial chains; (B) branched conidial chain; (C) conidia and fragments of conidial chains. Scale bar = 10 µm (B. Heuchert del.). Fig 9 – Taeniolella punctata [A: Diederich 16714; B, E, H, K: Diederich 12647; C, D, F, G, I, J: C 5739,]. (A) macroscopical overview of colonies; (B) SEM overview of colonies; (C, F, H)

ACCEPTED MANUSCRIPT conidiophores, (arrows) tips of conidiophores and/or the adhering terminal conidium somewhat swollen; (G, I, K) conidia. Scale bars: 1 mm (A), 50 µm (B), 10 µm (C–K). Fig 10 – Taeniolella punctata (C 6062). (A) hyphae; (B) conidiophores arising from hyphae or swollen hyphal cells; (C) conidia. Scale bar = 10 µm (B. Heuchert del.). Fig 11 – Taeniolella cf. punctata on Fissurina dumastii (Diederich 17044). (A) hyphae; (B)

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conidiophores arising from hyphae or swollen hyphal cells; (C) conidia. Scale bar = 10 µm (B. Heuchert del.).

Fig 12 – Taeniolella pyrenulae [holotype]. (A) macroscopical overview of colonies; (B) SEM overview of colonies; (C, D, G–J) conidia and conidial chains; (E–F) semi-micronematous

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conidiophores (swollen hyphal cells) with adhering conidia. Scale bars: 1 mm (A), 70 µm (B), 20 µm (D, J), 10 µm (C, E–I).

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Fig 13 – Taeniolella pyrenulae [holotype]. (A) hyphae with semi-micronematous conidiophores and adhering conidia; (B) conidial chains; (C) conidia and fragments of conidial chains. Scale bar = 10 µm (B. Heuchert del.).

Fig 14 – Taeniolella sp. from Rwanda [Ertz 11026]. (A) macroscopical overview of colonies; (B) conidiophores with adhering conidial chains; (C) branched conidial chain; (D) tips of conidial chains; (E) fragment of conidial chain with germination tube; (F–I) conidia and chain fragments

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of different sizes. Scale bars: 200 µm (A), 20 µm (B), 10 µm (C–I). Fig 15 – Taeniolella sp. from Rwanda [Ertz 11026]. (A) conidiophores arising from hyphae with adhering conidial chains; (B) branched conidial chain; (C) conidia and fragments of conidial

del.).

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chains; (D) fragment of conidial chain with germinating tube. Scale bar = 10 µm (B. Heuchert

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Fig 16 – Taeniolella toruloides [A: Diederich 18158; B–K: holotype]. (A) macroscopical overview of colonies; (B) SEM overview of colonies; (C–F) conidiophores with adhering long, unbranched toruloid conidial chains; (G, I–K) parts of conidial chains with a smooth, occasionally irregularly rugose-verrucose surface; (H) conidia. Scale bars: 1 mm (A), 50 µm (B, C), 20 µm (E, F), 10 µm (D, H), 6 µm (G, J, K), 3 µm (I). Fig 17 – Taeniolella toruloides (A–C: holotype; D: Diederich 18158). (A) hyphae with micronematous or semi-macronematous conidiophores; (B) conidiophores with adhering long toruloid chains; (C–D) conidia and fragments of conidial chains. Scale bar = 10 µm (B. Heuchert del.).

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Lichenothelia convexa

Aulographum hederae Laurera megasperma 1 Polycoccum pulvinatum 1 Friedmanniomyces endolithicus Xanthoriicola physciae 100 Capnodium coffeae 1 1 Davidiella tassiana Verrucocladosporium dirinae 89 99 Mycosphaerella punctiformis 0.99 Delphinella strobiligena 1 97 Sydowia polyspora Dothidea insculpta 96 Plowrightia abietis Myriangium duriaei 1 Asterina melastomatis 0.96 Lembosia abaxialis 99 1 Asterina chrysophylli 82 Batistinula gallesiae 92 Prillieuxina baccharidincola Parmularia styracis Coniosporium uncinatum 1 Encephalographa elisae Melaspilea enteroleuca 98 Abrothallus parmotrematis Jahnula aquatica Buelliella minimula 42273 1 Buelliella minimula 36238A 100 Buelliella minimula 35969 Karschia cezannei 19186 Taeniolella punctata 16714 1 Taeniolella punctata 17390 100 Taeniolella punctata 16040 Taeniolella punctata 7279 1 Karschia talcophila Taeniolella cf. punctata 17044 Melaspilea sp. 9134 0.95 Melaspilea sp. 18137 83 Melaspilea sp. 9137A 0.97 Melaspilea lekae 17325 0.97 Taeniolella sp. 11026 0.97 Taeniolella hawksworthiana 9199B Buelliella physciicola 18113 Taeniolella pyrenulae 17075 0.98 Labrocarpon canariense 10878 0.98 Stictographa lentiginosa 17570 Taeniolella toruloides 17047 1 85 Taeniolella toruloides 17048 0.99 84 Melaspilea sp. 18012 1 Melaspilea sp. 3684 Melaspilea sp. 3683 100 1 Melaspileopsis cf. diplasiospora 16247 Melaspileopsis sp. 17904 95 1 Asterina phenacis 1 Asterina weinmanniae 97 80 Inocyclus angularis 1 Lembosia albersii Asterotexis cucurbitacearum 1 1 Hemigrapha atlantica Buelliella poetschii 18116 96 Mycosphaerella pneumatophorae Geastrumia polystigmatis Acrospermum compressum 1 Acanthostigma minutum Tubeufia cerea 86 1 Arthopyrenia salicis 1 Didymocyrtis cladoniicola 0.96 87 85 Preussia terricola Lepidosphaeria nicotiae 78 Massariosphaeria typhicola 1 Hysterium angustatum 0.99 Psiloglonium araucanum 88 Glonium stellatum Taeniolella exilis 1 1 74 Taeniolella exilis 2 100 1 Kirschsteiniothelia thujina 1 Dendryphiopsis atra 100 1 0.99 89 Kirschsteiniothelia aethiops 90 Kirschsteiniothelia lignicola Phaeotrichum benjaminii 1 Sympoventuria capensis Venturia chlorospora 84 1 Diaporthe arctii 0.97 91 Phomopsis sp. Taeniolella alta (contaminant?) S 1 94 Diaporthe phaseolorum TE E Phaeocytostroma ambiguum 100 YC Stenocarpella maydis OM 1 Triangularia mangenotii I R Cercophora mirabilis A D Arnium mendax R SO Taeniolella phialosperma 1 Sordaria fimicola 1 Thielavia subthermophila 0.98 Carpoligna pleurothecii 99 0.97 75 Pleurotheciella centenaria 1 1 Sterigmatobotrys macrocarpa 1 98 Sterigmatobotrys (Taeniolella) rudis 89 Helicoon farinosum 100 Savoryella lignicola Lachnum virgineum Caliciopsis pinea 0.09 Capronia munkii

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ACCEPTED MANUSCRIPT Melaspilea sp. 3683, D.R. Congo/Pyrenula Melaspilea sp. 3684, D.R. Congo/Pyrenula Taeniolella pyrenulae 17075, Azores/Pyrenula Taeniolella toruloides 17047, Azores/Thelotrema antoninii 1 96 Taeniolella toruloides 17048, Azores/Thelotrema antoninii 0.99 100 Melaspilea sp. 18012, Reunion/Pyrenula Stictographa lentiginosa 17447, France/Phaeographis dendritica 1 Stictographa lentiginosa 17570, France/Phaeographis dendritica 100 Stictographa lentiginosa 47621, Madeira/Phaeographis dendritica 0.98 Taeniolella hawksworthiana 9199B, Florida/Phaeographis 1 84 Taeniolella sp. 11026, Rwanda/cf. Phaeographis 100 Melaspilea lekae 17325, Thailand/Sarcographa 1 Buelliella physciicola 18113, Belgium/Phaeophyscia orbicularis 100 Buelliella physciicola 19173, Belgium/Phaeophyscia orbicularis 1 Labrocarpon canariense 16308, Canary Islands/Pertusaria 1 100 Labrocarpon canariense 16907, Canary Islands/Pertusaria 97 Labrocarpon canariense 10878, Canary Islands/Pertusaria Taeniolella punctata 16714, Luxembourg/Graphis scripta 1 Taeniolella punctata 17390, Belgium/Arthonia atra 100 Taeniolella punctata 16040, Luxembourg/Graphis scripta Taeniolella punctata 7279, Germany/Pertusaria leioplaca Buelliella minimula 42273, USA/Pertusaria 1 Buelliella minimula 36238A, USA/Pertusaria paratuberculifera 100 Buelliella minimula 35969, USA/Pertusaria tetrathalamia 0.97 Karschia cezannei B26, Luxembourg/bark 100 Karschia cezannei 19186, Belgium/bark 81 Karschia cezannei 7453, Luxembourg/bark 0.97 Karschia talcophila 16749, Switzerland/Diploschistes 1 Taeniolella cf. punctata 17044, Azores/Fissurina dumastii 82 Melaspilea sp. 9134, Florida/wood 1 Melaspilea sp. 18137, Curaçao/bark 95 Melaspilea sp. 9137A, Florida/wood Melaspileopsis cf. diplasiospora 16247, Canary Islands/bark 1 Melaspileopsis cf. diplasiospora 16624, Azores/bark 100 1 Melaspileopsis cf. diplasiospora 16625, Azores/bark 98 1 Melaspileopsis sp. 17904, Reunion/bark 100 Melaspileopsis sp. 17913, Reunion/bark 1 Asterotexis cucurbitacearum 1 100 Asterotexis cucurbitacearum 2 1 ASTEROTEXIALES Inocyclus angularis 0.95 Lembosia albersii 1 Buelliella poetschii 18115 100 Buelliella poetschii 18116 1 Hemigrapha atlantica 1 100 Mycosphaerella pneumatophorae 92 ‘Asterinales’ SH2014 Asterina zanthoxyli 1 Mahanteshomyces sp 98 Asterina SH2014 Asterina weinmanniae Asterina fuchsiae 1 Asterina phenacis 93 1 81 Asterina siphocampyli 89 Asterina cestricola Geastrumia polystigmatis 1 Dothidea insculpta 97 Capnodium coffeae Preussia terricola 1 100

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ACCEPTED MANUSCRIPT Research Highlights Taeniolella manuscript Ertz et al.

The type species of Taeniolella is placed in the Kirschsteiniotheliaceae.

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The phylogeny shows Taeniolella to be strongly polyphyletic.

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Lichenicolous species of Taeniolella are polyphyletic within the Asterotexiales.

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Anamorph-teleomorph relationships between Taeniolella and other genera are discussed.

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Three new species of Taeniolella are described.

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