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Cryptodiscus Muriformis and Schizoxylon Gilenstamii, Two New Species of Stictidaceae (Ascomycota)

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Cryptodiscus Muriformis and Schizoxylon Gilenstamii, Two New Species of Stictidaceae (Ascomycota) Mycological Progress (2018) 17:295–305 https://doi.org/10.1007/s11557-017-1363-4 ORIGINAL ARTICLE Cryptodiscus muriformis and Schizoxylon gilenstamii, two new species of Stictidaceae (Ascomycota) Samantha Fernández-Brime1 & Ibai Olariaga1 & Hans-Otto Baral2 & Gernot Friebes3 & Walter Jaklitsch4,5 & Beatrice Senn-Irlet6 & Mats Wedin1 Received: 14 August 2017 /Revised: 8 November 2017 /Accepted: 13 November 2017 /Published online: 18 December 2017 # The Author(s) 2017. This article is an open access publication Abstract Cryptodicus muriformis and Schizoxylon gilenstamii (Stictidaceae, Ostropales, Lecanoromycetes, Ascomycota) are described as new to science from collections made in Europe (Sweden, Austria, Switzerland) and North America (Arizona), based on phylogenetic analyses of three loci. Both species show unique morphological characters that distinguish them from closely related species; Cryptodiscus muriformis, growing on decorticated or sometimes corticated branches of Picea abies and Pseudotsuga menziesii,thatresemblesC. foveolaris and C. tabularum in the yellowish-orange ascomatal disc, but is clearly distinguished by the muriform ascospores. Schizoxylon gilenstamii grows on dead stems of Globularia vulgaris and has a characteristic unique red-purple pigment in the epithecium and upper part of the excipulum that changes to bluish-green with KOH. We also report for the first time the anamorph of Stictis brunnescens. Keywords Ostropales . Phylogeny . Saprotrophic fungi . Taxonomy Introduction The family Stictidaceae (Ostropales, Lecanoromycetes, discomycetes, but also lichens, parasites, and endophytes Ascomycota) contains predominantly saprotrophic (e.g., Gilenstam 1969; Fernández-Brime et al. 2011; Baloch et al. 2013; Aptroot et al. 2014;Jahnetal.2017). Sherwood This article is part of the BSpecial Issue on ascomycete systematics in (1977a, 1977b) produced a detailed and still very useful honour of Richard P. Korf who died in August 2016^. worldwide taxonomic and nomenclatural revision of the Section Editor: Gerhard Rambold family with main focus on taxa from the USA. With the advent of DNA technologies, molecular phylogenetic studies * Samantha Fernández-Brime have been crucial to revise established generic limits and [email protected] species concepts in the family, although these studies are 1 Department of Botany, Swedish Museum of Natural History, still in their infancy. Wedin et al. (2004, 2005, 2006)investi- P.O. Box 50007, SE-10405 Stockholm, Sweden gated taxa living on Populus and Salix in Northern 2 Blaihofstraße 42, D-72074 Tübingen, Germany Scandinavia. They found that several species of Stictis and 3 Centre of Natural History, Botany, Universalmuseum Joanneum, Schizoxylon can live either as lichens or as saprotrophs de- Weinzöttlstraße 16, 8045 Graz, Austria pending on the substrate they inhabit, a phenomenon they B ^ 4 Institute of Forest Entomology Forest Pathology and Forest called optional lichenization . These results led to a widened Protection, Department of Forest and Soil Sciences, concept of Stictis and Schizoxylon, which until then only in- BOKU-University of Natural Resources and Life Sciences, cluded saprotrophic species. Wedin et al. (2006) pointed out Hasenauerstraße 38, 1190 Vienna, Austria that these two genera are potentially heterogeneous, but as 5 Division of Systematic and Evolutionary Botany, Department of molecular data was available only for a small number of spe- Botany and Biodiversity Research, University of Vienna, Rennweg cies they could not pursue this further at that time. Finally, 14, 1030 Wien, Austria Cryptodiscus was studied in detail in Scandinavia by Baloch 6 Swiss Federal Institute for Forest, Snow, and Landscape Research et al. (2009), resulting in a revision of the generic concept and WSL, Biodiversity and Conservation Biology, Zürcherstr. 111, 8903 Birmensdorf, Switzerland the description of a new species. 296 Mycol Progress (2018) 17:295–305 Our knowledge of the Stictidaceae is obviously still very DNA Kit following an improved protocol (Stielow et al. limited. The family contains small and generally drought- 2013). tolerant fungi, which are easily overlooked and rarely collect- The selected markers for this study were the internal tran- ed. For this reason, in recent years we have continued explor- scribed spacer complete repeat (ITS), the first part (D1–D2 ing plant debris and, as a result, we found specimens of region, ca. 600 bp) of the large subunit (nLSU) of the nuclear stictidaceous fungi that did not fit any described taxa. This ribosomal DNA, and the small subunit of the mitochondrial paper introduces two new species collected in Sweden, ribosomal DNA (mtSSU). Primer combinations for amplifica- Austria, Switzerland and USA, and provides detailed descrip- tion of the three loci were: ITS1F (Gardes and Bruns 1993) tions and a three-locus phylogeny that places them in the here and ITS2, ITS3, and ITS4 (White et al. 1990)for≈ 0.6 kb ITS, adopted genera within the context of the Stictidaceae. LR0R and LR3 (Vilgalys and Hester 1990)for≈ 0.6 kb nLSU, Anamorphs have been reported from several Stictidaceae and mrSSU1 and mrSSU3R (Zoller et al. 1999) for the ≈ species. Pycnidial anamorphs have been found in 0.8 kb mtSSU. Symmetric PCR amplifications were per- Acarosporina microspora, Conotrema urceolatum, formed using IllustraTM Hot Start PCR beads, according to Cryptodiscus gloeocapsa, Cyanodermella oleoligni,and the manufacturer’s instructions, with the following settings for Stictis radiata (e.g., Gilenstam 1969; Sherwood 1977a; the three loci: initial denaturation 94 °C for 5 min, followed by Johnston 1983; Baloch et al. 2009; Nieuwenhuijzen et al. five cycles (94 °C for 45 s, 54 °C for 30 s, and 72 °C for 60 s), 2016). Johnston (1983) established the connection between 30 cycles (94 °C for 45 s, 52 °C for 30 s, and 72 °C for 60 s), anamorphs and teleomorphs of several Stictis species using and a final extension 72 °C for 8 min. After examination by cultures. In all cases, the anamorphs described were pycnidia. gel electrophoresis, amplification products were purified As a result from recent field work in Sweden, we found sev- using ExoSAP-IT (USB Corp., U.S.A.). Sequencing of both eral conidiomata, which are presumably the anamorph of strands was performed with the Big Dye Terminator technol- Stictis brunnescens. We have also used the three-locus phy- ogy kit v3.1 (ABI PRISM, U.S.A.) using the PCR primers. logeny to test the connection between the anamorph and teleomorph. Sequence alignment and phylogenetic analyses Sequence fragments were edited and assembled with Materials and methods Sequencher 4.9 (Gene Codes Corp., USA) and consensus se- quences subjected to BLAST searches to rule out amplifica- tion of contaminants. The newly produced sequences were Morphological and anatomical studies included in individual alignments for each locus, together with sequences retrieved from GenBank representing morphologi- Specimens were examined morphologically and anatomically cally similar species from the family Stictidaceae. The taxon and preserved in the herbaria GJO, S, UPS (herbarium codes Karstenia rhopaloides was selected as an outgroup as it is according to Thiers [2017, continuously updated]) and H. O. closely related to Stictidaceae, but Baloch et al. (2009) Baral’s private herbarium (H.B.). Hand-cut sections mounted showed that it does not belong to the family. Sequences were in water were used to measure margin, hymenium, asci, pa- aligned with the online version of MAFFT v7 (http://mafft. raphyses, and ascospores. Freezing microtome sections, 20– cbrc.jp/alignment/server/) using the algorithms L-INS-I for 30 μm thick, mounted in water or stained with lactic blue or ITS and nLSU, and E-INS-I for mtSSU. The ambiguously Congo red, were produced for a more detailed study of the aligned regions were identified using the G-blocks v0.91b anatomy. Lugol’s iodine solution, before and after treatment (http://molevol.cmima.csic.es/castresana/Gblocks_server. with 10% potassium hydroxide (KOH), was used to detect html) and removed before analysis. Alignments were amyloid structures. Thereby, Btype RR^ means that the ascus submitted to TreeBase (http://www.treebase.org;IDnumber wall reacts red at both low and high concentration before 21812). applying KOH. For the anatomical features measurements, Phylogenetic analyses were performed for each locus indi- the extreme values are shown after rejecting 10% of the vidually with RAxML v 8.2.0 (Stamatakis 2014), using the highest and the lowest values; the highest and lowest absolute rapid bootstrap analysis and search for best-scoring maximum values are given in parentheses. likelihood algorithm (−f a), the GTRGAMMA model and 1000 bootstrap replicates (ML-BS). Then, the individual phy- DNA extraction, PCR amplification, and sequencing logenies were compared to assess congruence, considering there to be a significant conflict when two alternative relation- Total DNA was extracted from a selected number of samples ships corresponding to different loci were both supported with with the DNeasy Plant Mini Kit (Qiagen, Germany) following bootstrap values ≥70% (Mason-Gamer and Kellog 1996). As the manufacturer’s instructions or with the E.Z.N.A. Forensic no significant conflicts were detected, the three alignments Mycol Progress (2018) 17:295–305 297 were concatenated into a single dataset with five partitions structure clustered unequivocally with the specimens of (ITS1, 5.8S, ITS2, nLSU and mtSSU). Phylogenetic relation-
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