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Department of Plant Taxonomy and Biosystematics, Institute of Botany, Academy of Sciences, 252 43 Pru °honice, Czech Republic
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ABSTRACT |
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 TOP ABSTRACT INTRODUCTIONMATERIALS AND METHODSRESULTSTAXONOMYKEY TO THE GENERA...KEY TO THE SPECIES...KEY TO THE SPECIES...KEY TO THE SPECIES...DISCUSSIONLITERATURE CITED |
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The systematic position and phylogenetic relationships of Ceratostomella sensu lato and phenotypically similar fungi using comparative morphological and culture studies and phylogenetic analyses of the nuclear large- and small-subunit ribosomal DNA were explored. In the light of inferred phylogenies and morphological data the genus Ceratostomella is redescribed, the generic concept is emended and four species are accepted (viz. C. cuspidata, C. pyrenaica, C. rhynchophora and C. rostrata). A new genus Xylomelasma is introduced and delimited from Ceratostomella, with two new species described (viz. X. novaezelandiae and X. sordida). In culture species of both Ceratostomella and Xylomelasma produced sterile mycelium. The genus Lentomitella with a phaeoisaria-like anamorph formed in vitro is reinstated to encompass taxa formerly attributed to the broadly perceived Ceratostomella with three accepted species (viz. L. cirrhosa, L. crinigera and L. tomentosa). Lentomitella and Ceratostomella are clearly distinguishable by the morphology of asci, ascospores and centrum. Lentomitella is compared to phenotypically similar Ceratosphaeria, which formed a harpophora-like anamorph in vitro. In the present phylogenies Ceratostomella, Ceratosphaeria, Lentomitella and Xylomelasma are shown as clearly separate genera belonging to three different groups of perithecial ascomycetes. Ceratostomella, Lentomitella and Xylomelasma reside within a large unsupported clade consisting of members the Ophiostomatales, the freshwater Annulatascaceae and a group of nonstromatic, terrestrial taxa. Ceratosphaeria is well supported within the Magnaporthaceae. The systematic value of morphological characters of ascospores, paraphyses, asci, centrum and conidiogenesis in segregating taxa from Ceratostomella sensu lato and their relatives is discussed.
Key words: Annulatascaceae, Ceratosphaeria, Endoxyla, Harpophora, Lentomitella, LSU and SSU rDNA, Ophiostomatales, Phaeoisaria, Sporothrix, systematics, Wegelina, Xylomelasma
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INTRODUCTION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSTAXONOMYKEY TO THE GENERA...KEY TO THE SPECIES...KEY TO THE SPECIES...KEY TO THE SPECIES...DISCUSSIONLITERATURE CITED |
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simple generic diagnosis of Ceratostomella reads: "Perithecia et asci Ceratostomatis. Sporidia continua, hyalina". Consequently the genus became a large, heterogeneous entity that included more than 100 taxa. Höhnel (1918) removed species with evanescent asci from Ceratostomella to Linostoma Höhn. Sydow and Sydow (1919) recognized Linostoma to be a later homonym for a genus of flowering plants (Thymeleaceae); they proposed the new name Ophiostoma Syd. & P. Syd., where several Ceratostomella species were transferred. Several other species of Ceratostomella with evanescent asci later were recognized congeneric with Ceratocystis Ellis & Halst. in a broad sense (e.g. Elliott 1925, Moreau 1952, Hunt 1956).
Although Ceratostomella should include fungi with hyaline ascospores, nonstromatic perithecia and persistent asci, von Arx and Müller (1954) treated Ceratostomella as a synonym of Endoxyla Fuckel, characterized by ellipsoidal, smoky-brown ascospores with terminal germ pores, asci with dissolving stipe and perithecia with reduced stromatic tissue around the base of a projecting neck. For the next 20 years, Ceratostomella (under Endoxyla) was included in the Diaporthaceae (von Arx 1952, von Arx and Müller 1954, Müller and von Arx 1973, Munk 1952, 1957, Gilman et al 1959). The study of Danish members of Endoxyla by Munk (1965) did not shed light on the systematics of the genus and it remained heterogeneous. Munk (1965) doubted his former placement of Endoxyla in the Rhamphorioideae of the Diaporthaceae and noted that some Endoxyla species may even represent calosphaeriaceous elements. Barr (1978, 1990) excluded Endoxyla from the Diaporthales because of the carbonaceous perithecial wall, asci that do not float freely within the centrum, presence of free-ended paraphyses, occurrence on wood as a secondary saprobe, and she classified the genus in the family Clypeosphaeriaceae. Untereiner (1993) distinguished Ceratostomella from Endoxyla and suggested affinity of the former genus with the Lasiosphaeriaceae while Endoxyla was accepted in the Clypeosphaeriaceae. Eriksson et al (2003) followed Barr’s concept and accepted Ceratostomella as a member in the Clypeosphaeriaceae of the Xylariales.
Ceratostomella sensu Untereiner (1993) was retained for species with dark-colored, immersed, nonstromatic perithecia with opaque, pseudoparenchymatous walls; central, elongate, separately protruding necks; true paraphyses; unitunicate, sessile asci arising from a crozier system with a distinct, refractive, nonamyloid apical annulus and ellipsoidal, hyaline ascospores. The phylogenetic relationships of Ceratostomella have remained unknown and no anamorph connections have been reported.
Höhnel (1905) introduced a monotypic genus, Lentomitella Höhn., for Ceratostomella vestita Sacc. He based the segregation of Lentomitella from Ceratostomella on the ornamented (longitudinally striate) ascospore walls of C. vestita and suggested other species of Ceratostomella to belong to Lentomita Niessl or Ceratosphaeria Niessl. Lentomita brevicollis Niessl the type of the generic name Lentomita (Niessl 1876), was transferred by Müller and von Arx (1962) to Chaetosphaeria, as C. brevicollis (Niessl) E. Müll., and Lentomita was relegated to synonymy of Chaetosphaeria. It is difficult to find a difference in teleomorph morphology between Ceratosphaeria and Ceratostomella except that the ascospores are cylindrical in the type species, Ceratosphaeria lampadophora (Berk. & Broome) Niessl, but can range from ellipsoidal to oblong to subcylindrical in other species currently attributed to the genus (Tsuda and Ueyama 1977). The insufficiently delineated generic concept and life histories continued to hamper the distinction between Ceratostomella and Ceratosphaeria and considerable cross generic and species synonymy can be found in the literature.
When it was proposed originally (Saccardo 1878) two species were included in Ceratostomella Sacc (viz. C. vestita and C. cirrhosa [Pers. : Fr.] Sacc). At the same time Saccardo listed Ceratostoma rostratum (Tode : Fr.) Fuckel under Ceratostomella without comment. Later (Saccardo 1882) transferred Ceratostoma rostratum to Ceratostomella. The examination of type and other herbarium material of the three species cited in the protologue of Ceratostomella (Saccardo 1878), revealed the genus to be heterogeneous from the beginning and the three cited species represent two distinct genera.
Ceratostomella rostrata (Tode : Fr.) Sacc., the lectotype species of Ceratostomella (Clements and Shear 1931), possesses dark, long-beaked perithecia; pale brown, suballantoid to oblong ascospores; branching ascogenous hyphae; broad-celled paraphyses, and clavate asci tapering toward the base from the sporiferous portion with a shallow, indistinct, non-amyloid apical annulus. Ceratostomella rostrata matches the generic concept of Wegelina Berl., as emended by Barr (1998) and it is identical to Wegelina polyporina M.E. Barr. Wegelina was erected by Berlese (1900) to segregate taxa from Calosphaeria Tul. & C. Tul. in the Calosphaeriales. Both genera accommodate species with tiny nonstromatic perithecia; hyaline, aseptate or delicately septate, allantoid ascospores; clavate, long-stipitate asci with thickened ascal apex and persistent paraphyses. Wegelina comprises species with superficial, separately growing perithecia and separately protruding necks, while Calosphaeria encompasses species forming perithecia in ellipsoidal to circinate groups or valsoid configurations with short or elongate, radially converging beaks piercing the periderm. Seven species have been assigned to Wegelina; W. discreta Berl. was designated as lectotype by Clements and Shear (1931). Wegelina was reduced to synonymy of Scoptria Nitschke (Barr 1985) or Calosphaeria (Barr et al 1993), but later it was reinstated (Barr 1998) and one new species and two new combinations were introduced to the genus. The characteristics of W. polyporina and W. subdenudata (Peck) M.E. Barr (Barr 1998), including relatively large perithecia that form a crust on the substratum, thick perithecial wall (50–100 µm thick vs the wall of Wegelina s. str. 20–40 µm), sulcate necks and sub-allantoid to ellipsoidal, pale brown ascospores before discharge, led Barr (1998) to extend Berlese’s concept of Wegelina.
The other original species, C. vestita and C. cirrhosa, are congeneric but not closely related to C. rostrata. A phaeoisaria-like anamorph developed in culture. These fungi match the concept of Ceratostomella sensu Untereiner (1993) and should be accommodated in Lentomitella (Höhnel 1905). Lentomitella is compared here with phenotypically similar species of Ceratosphaeria, based on C. lampadophora, which formed a harpophora-like anamorph in vitro.
During a survey of perithecial lignicolous ascomycetes in temperate regions of both Northern and Southern Hemispheres, several collections of two fungi phenotypically similar to Ceratostomella sensu stricto were encountered. In culture, colonies produced a sterile mycelium. The unknown fungi resemble Ceratostomella sensu stricto in having dark, long-beaked perithecia, densely branched ascogenous hyphae, clavate asci floating freely within the centrum at maturity and pigmented ascospores with terminal pores before discharge. The two taxa differ from the core species of Ceratostomella in thickness of the perithecial wall, morphology of paraphyses, shape of asci and apical annulus, and shape and arrangement of ascospores. One of these fungi has been described as Endoxyla avocetta (Cooke & Ellis) Romero & Samuels (Romero and Samuels 1991) or later as Phaeognomoniella avocetta (Cooke & Ellis) Romero (Romero 1999). The examination of the holotype material of Sphaeria avocetta Cooke & Ellis and herbarium material, on which the cited combinations were based, revealed two distinct perithecial ascomycete genera. Sphaeria avocetta is a species of Endoxyla and it is identical to E. operculata (Fr. : Fr.) Sacc, while one of the unknown fungi is conspecific with E. avocetta sensu Romero and Samuels (1991); its relationship and evolutionary history are examined here. The second unknown fungus differs from E. avocetta sensu Romero and Samuels (1991) in size, shape and color of the ascospores and size of the asci.
To reveal the systematic and phylogenetic relationships of Ceratostomella, Ceratosphaeria, Lentomitella and the unknown fungi, sequences of nuclear small-and large-subunit ribosomal DNA (SSU and LSU nrDNA) of their type species were analyzed in two independent sequence datasets using maximum parsimony and Bayesian analyses.
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MATERIALS AND METHODS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSTAXONOMYKEY TO THE GENERA...KEY TO THE SPECIES...KEY TO THE SPECIES...KEY TO THE SPECIES...DISCUSSIONLITERATURE CITED |
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Single- and mass-ascospore isolates were obtained from fresh material with the aid of a single-spore isolator (Meopta, Czech Republic). Cultures were grown on potato-carrot agar (PCA, Gams et al 1998). Colony characters were taken from cultures grown on PCA for 21 d at room temperature (24 C) under UV light. Cultures are maintained at the Institute of Botany, Academy of Sciences in Pru °honice; Centraalbureau voor Schimmelcultures, Utrecht (CBS); and Landcare Research, Auckland (ICMP). Type material is preserved in Landcare Research, Auckland (PDD), National Museum, Prague (PRM), other herbarium specimens with M.R. numbers are preserved at the Institute of Botany, Pru °honice (PRA). The isolates used in this study and their sources are listed (TABLE I).
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. The LSU and SSU nrDNA sequences of other taxa newly sequenced in this study were generated by methods identical to those described by Réblová and Seifert (2004).
Sequence data analyses.— –
Phylogenetic relationships were examined using 52 LSU nrDNA and 43 SSU nrDNA sequences from 15 or 14 different orders or families of Sordariomycetes, respectively. Members of the Dothideomycetes were used as outgroups in all analyses. New LSU and SSU nrDNA sequences were obtained for these taxa (TABLE I
): the ascospore isolates of Ceratosphaeria lampadophora, Ceratostomella pyrenaica Réblová & Fournier, Lentomitella cirrhosa (Pers. : Fr.) Réblová, L. crinigera (Cooke) Réblová and Xylomelasma sordida Réblová. Homologous LSU and SSU nrDNA sequences from other 88 taxa were retrieved from GenBank; accession numbers are given (FIGS. 1, 2).
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). Predicted models of the secondary structure of the LSU and SSU rRNA molecules of Saccharomyces cerevisiae Meyen ex E.C. Hansen (Gutell 1993, Gutell et al 1993) were used to improve the alignment. The models of the secondary structure of the LSU and SSU rRNA were highly consistent in all taxa. The insertion positions in the SSU nrDNA sequences are named for the 5' flanking nucleotide and correspond to the positions of Escherichia coli (Migula) Castell. & Chalm. (Gargas et al 1995). The alignments are available in TreeBASE as SN2349.
The phylogenetic analyses were performed with PAUP v. 4.0b10 (Swofford 2002) using maximum parsimony; heuristic search with stepwise-addition option with 1000 random taxon addition replicates and branch swapping (tree bisection-recognition, TBR). All characters were unordered and given equal weight during the analysis. For all analyses ambiguously aligned positions were excluded. Gaps were treated as missing data. Branch support was estimated by performing 1000 bootstrap replicates with a full heuristic search consisting of 10 random-addition replicates for each bootstrap replicate.
The model of evolution that fits the data best was determined by MODELTEST 3.5 (Posada and Crandall 1998), showing that the best-fit model for the LSU sequence dataset was GTR+I+G model (Lanave et al 1984) with variable base frequencies (freqA = 0.2221, freqC = 0.2425, freqG = 0.3131, freqT = 0.2223), different rates for transitions and transversions R(a) [A–C] = 1.0263, R(b) [A–G] = 2.3442, R(c) [A–T] = 1.5809, R(d) [C–G] = 0.8755, R(e) [C–T] = 6.7847, R(f) [G–T] = 1.0000, an estimated proportion of invariable sites of 0.3137 and a gamma shape parameter for the rates of variable sites of 0.4790. The best-fit model for the SSU sequence dataset was determined as GTR+I+G model with variable base frequencies (freqA = 0.2420, freqC = 0.2281, freqG = 0.2840, freqT = 0.2459), different rates for transitions and transversions R(a) [A–C] = 1.4452, R(b) [A–G] = 3.5075, R(c) [A–T] = 1.3198, R(d) [C–G] = 0.9220, R(e) [C–T] = 7.2559, R(f) [G–T] = 1.0000, an estimated proportion of invariable sites of 0.5264 and a gamma shape parameter for the rates of variable sites of 0.6233.
Bayesian analyses were performed with MrBayes v. 3.0b4 (Huelsenbeck and Ronquist 2001) to reconstruct phylogenetic trees. The above models of evolution were implemented and Bayesian analyses for both LSU and SSU sequence datasets were conducted with the number of rate categories set to six, rates set to gamma and all remaining parameters estimated from the default prior probabilities. 5 000 000 generations were sampled every 100th generation resulting in 50 000 trees. The first 20 000 trees, which represented the burn-in phase of the analysis (number of trees to be discarded after the likelihoods of the trees have converged on a stable value), were discarded, and the remaining 30 000 trees were used for calculating posterior probabilities in the consensus tree. Posterior probabilities = 95% are indicated as thickened branches in the trees.
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RESULTS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSTAXONOMYKEY TO THE GENERA...KEY TO THE SPECIES...KEY TO THE SPECIES...KEY TO THE SPECIES...DISCUSSIONLITERATURE CITED |
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). The insertions of C. pyrenaica located at 393 and 1424 5' flanking nucleotide positions correspond to the positions 396 and 1086 of E. coli SSU nrDNA (Gargas et al 1995). The first five insertions of X. sordida and both insertions of C. pyrenaica are the group I intron, the sixth insertion of X. sordida represents the spliceosomal intron with the 5'- GT donor and AG - 3' acceptor splice sites. These insertions were excluded from all analyses.
Phylogenetic analyses of the LSU rDNA sequence data.— –
A maximum parsimony analysis (MP1) was performed with 372 phylogenetically informative characters in an alignment including 1256 nt from 52 taxa. The first 75 positions in the alignment that are ambiguous were excluded. Two most parsimonious trees (MPT) were obtained (tree length 2299, consistency index [CI] = 0.375, retention index [RI] = 0.553, homoplasy index [HI] = 0.625) (FIG. 1). The only source of polytomy was a clade containing the Annulatascaceae, the putative terrestrial Trichosphaeriaceae and the fungi newly sequenced within this project. The two trees differed in position of C. pyrenaica and X. sordida; C. pyrenaica was shown either as a sister to Cryptadelphia groenendalensis (Sacc. et al) Réblová & Seifert or to X. sordida with no bootstrap support.
The consensus tree consisted of six major and well supported phylogenetic lineages of the Sordariomycetes (viz. a lineage [75% bootstrap support/100% posterior probability] consisting of four well supported orders or groups), the Hypocreales (77/100) and the Microascales (98/100) clade, which is sister to a clade (87/100) of the Coronophorales (84/100) and Ascotaiwania/Conioscyphascus/Carpoligna group (100/100); a lineage (64/100) consisting of subgroupings of five orders, the Sordariales (91/100), the Chaetosphaeriales (99/100), the Diaporthales (99/100), the Calosphaeriales and the Boliniales (89/100); other major lineages were the well supported Xylariales (89/100) and the Magnaporthaceae (83/100). Ceratosphaeria lampadophora nested on a well supported branch with Ophioceras tenuisporum Shearer et al (96/0) within the Magnaporthaceae. Trichosphaeria pilosa (Pers. : Fr.) Fuckel, representing the Trichosphaeriaceae, is shown basal to all other unitunicate ascomycetes in the tree.
In the sixth major lineage four separate clades were discerned. In one of the two phylogenetic trees (FIG. 1), these clades were the Ophiostomatales (95/100), which were a sister to the larger unsupported clade consisting of X. sordida and C. pyrenaica on one branch and the Annulatascaceae (75/100) subclade and the Cryptadelphia Réblová & Seifert, Lentomitella and Rhamphoria Niessl subclade on the other branch. The two Lentomitella species (100%), Rhamphoria delicatula Niessl and C. groenendalensis represent a terrestrial sister to the freshwater Annulatascaceae. The Annulatascaceae are represented in the phylogeny by Annulusmagnus triseptatus J. Campbell & Shearer, Ascitendus austriacus (Réblová et al) J. Campbell & Shearer, Annulatascus velatisporus K.D. Hyde and Aniptodera chesapeakensis Shearer & M.A. Mill. Xylomelasma sordida and C. pyrenaica reside on a basal branch of this large clade with no branch support.
In the consensus tree inferred from Bayesian analysis the main topology of the six major lineages was retained; the only lineage that did not receive any support was the one containing the Ophiostomatales, the Annulatascaceae, Cryptadelphia, Rhamphoria and the newly sequenced taxa. Within this lineage three clades were discerned (viz. a clade of C. pyrenaica, X. sordida and R. delicatula [98 posterior probability], Cryptadelphia and Lentomitella clade [67] and a clade [97] containing the Annulatascaceae [100] as a sister to the Ophiostomatales [100]).
Phylogenetic analysis of the SSU rDNA sequence data.— –
A maximum parsimony analysis (MP2) was performed with 260 phylogenetically informative characters in an alignment including 1724 nt from 44 taxa. The first 27 positions in the alignment that are ambiguous were excluded. Four MPT were obtained, one of which is shown (FIG. 2) (tree length 1092, CI = 0.503, RI = 0.632, HI = 0.497). The trees differed in grouping of the Sordariales, the Chaetosphaeriales and the Coniochaetales within a single larger clade. Five major lineages of the Sordariomycetes were discerned in the MP2 analysis, (viz. a robust lineage [51/100] consisting of subgroupings of six orders or families, the Coniochaetales [97/100], which are a sister to the unsupported clade containing the Cephalothecaceae on one branch and the Sordariales [97/100], the Boliniales, the Phyllachorales and the Chaetosphaeriales [100/100] on the other branch; a lineage of the Diaporthales [100/100] and the Calosphaeriales; a lineage of the Xylariales [80/100] and a larger lineage of the Hypocreales and Microascales [89/100]; and a lineage consisting of members of the Magnaporthaceae [99/100], which are sister to a clade of the Ophiostomatales [100/100] with Ceratostomella, Lentomitella and Xylomelasma residing on separate basal branches, respectively).
Anamorphteleomorph connection.— – Single ascospores were isolated from L. cirrhosa, L. crinigera, C. lampadophora, C. pyrenaica and X. sordida. The cultures derived from L. crinigera and C. lampadophora sporulated in 21 d on potato-carrot agar at room temperature under UV light. Lentomitella crinigera formed a phaeoisaria-like anamorph and C. lampadophora formed a harpophora-like anamorph in vitro. The cultures derived from C. pyrenaica, L. cirrhosa and X. sordida never sporulated, although the cultural characters of L. cirrhosa otherwise were identical with those of the culture derived from a morphologically similar L. crinigera. Because the cultures of L. crinigera and L. cirrhosa have identical appearance and the LSU sequence data suggest their close relationship within a strongly supported monophyletic clade, it is most likely explanation that a phaeoisaria-like anamorph is also part of the life history of L. cirrhosa.
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TAXONOMY |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSTAXONOMYKEY TO THE GENERA...KEY TO THE SPECIES...KEY TO THE SPECIES...KEY TO THE SPECIES...DISCUSSIONLITERATURE CITED |
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. Although von Arx (1952) considered C. vestita as type species of Ceratostomella the lectotypification proposed by Clements and Shear (1931) must be accepted because even though it was not actually formally combined in Ceratostomella at the time, it is clear that Saccardo (1878) intended to include Ceratostoma rostratum in Ceratostomella (P. Kirk, pers comm).
Tode (1791:14, Tab. IX, Fig. 79.) described Sphaeria rostrata with two varieties (viz. 
. atra [perithecia black, tuberculate] and ß. nigrofusca [perithecia brown to black, glabrous]). No type or other authenticated material of S. rostrata is available because none of Tode’s original collections are preserved. The illustration accompanying the original description is the only surviving original element (FIG. 22). The protologue does not include critical features and the illustrations of var. and ß show only dark tuberculate or glabrous perithecia with long central necks, respectively. Persoon (1801:58) cited the name S. rostrata var. . and referred to the Tode’s description. Fries (1823:473) based the description of S. rostrata Tode : Fr. on material distributed in his exsiccate collection Scleromyceti Sueciae 116 with a reference to Tode (1791) and Persoon (1801). Later Saccardo (1878) accepted S. rostrata in Ceratostomella. The examination of the specimens of Scleromyceti Sueciae 116 by the present author revealed that S. rostrata is not congeneric with C. cirrhosa, C. vestita or other species currently attributed to Ceratostomella sensu Untereiner (1993). Sphaeria rostrata differs from C. cirrhosa and C. vestita in having suballantoid to ellipsoid to reniform ascospores that are pale brown before discharge. Ascogenous hyphae are branching, producing terminal and lateral, aseptate cells, from each of which one ascus arises. The asci are clavate, short-stipitate, tapering toward the base from the sporiferous portion, floating freely within the centrum at maturity. The apical annulus is shallow, indistinct and nonamyloid. Paraphyses are broad-celled, constricted at the septa and tapering.
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and the lectotype species C. rostrata are accepted by the present author as valid. Because it never will be possible to know what Tode actually studied, I accept Fries’ treatment of S. rostrata with a reference to Tode (1791) and Persoon (1801) based on material distributed in Scleromyceti Sueciae 116. Several exsiccate collections of S. rostrata under Scleromyceti Sueciae 116 were studied (C, G, K, PRM, UPS) but only the G and PRM specimens contained many intact mature perithecia in good condition. To avoid confusion I herewith lectotypify the name S. rostrata with Tode’s original illustration and epitypify with Fries’ material distributed in Scleromyceti Sueciae 116 (PRM 666367!). In this study Ceratostomella is redescribed based on the lectotype species C. rostrata, and three other species are accepted (viz. C. cuspidata, C. pyrenaica and C. rhynchophora). Wegelina polyporina is relegated to synonymy of C. rostrata. Other species currently attributed to Ceratostomella sensu lato that do not match the emended generic concept are under revision by the author.
The generic concept of Wegelina based on the lectotype species W. discreta (Exsiccate. Mycotheca Veneta 1450: ITALY. Belluno, on decorticated branch of Acer pseudoplatanus, autumn 1879, P.A. Saccardo (as Calosphaeria wahlenbergii; isotype of W. discreta, NY!) should follow Berlese’s treatment (Berlese 1900).
The distinction between the core species of Ceratostomella and the phenotypically similar unknown fungi are based on these features: thick perithecial wall (ca. [60–]80–110 µm), clavate asci; broad-celled paraphyses; shallow, indistinct apical annulus, and suballantoid to reniform to irregularly ellipsoidal ascospores 2–3-seriate or in a fascicle within the ascus of Ceratostomella sensu stricto, vs thinner perithecial wall (ca. 50–60[–80] µm); cylindrical asci; cylindrical, slightly constricted paraphyses, distinct apical annulus, and ellipsoidal ascospores, usually obliquely 1-seriate within the ascus of the unknown fungi. The LSU and SSU molecular data do not support closer relationship between Ceratostomella, represented by C. pyrenaica, and the unknown fungi. In the consensus tree of the two most parsimonious trees C. pyrenaica resides either on a branch together with C. groenendalensis as sister to Lentomitella and Rhamphoria subclade, which is sister to the Annulatascaceae subclade, while the unknown fungus is shown on a basal branch of the whole clade, or both C. pyrenaica and the unknown fungus are nested on a basal branch of this large clade (FIG. 1). Based on the morphological characters and results from molecular analyses, both taxa are distinct fungi at the genus level. Because no ascomycete genus could be found to match these characteristics, a new genus, Xylomelasma, is introduced for the two unknown fungi and the new species X. novaezelandiae and X. sordida are described. Xylomelasma sordida includes taxa formerly known as E. avocetta or P. avocetta (Romero and Samuels 1991, Romero 1999).
Taxonomic affinities of Ceratostomella cirrhosa and Ceratostomella vestita.— –
Sphaeria cirrhosa Pers. was described and illustrated by Persoon (1800: Tab. 24, Fig. 3; 1801:59) for fungi with immersed to semiimmersed, dark perithecia sparsely covered with short hairs and with sulcate necks. No type has ever been designed for S. cirrhosa. Von Arx (1952) reported four collections of S. cirrhosa from the Persoon herbarium (L) and said that only one of them was a fungus that matched the description of S. cirrhosa sensu Persoon (1801). That collection, according to von Arx (1952), contained few ascospores (8–10 x 3.5 µm). Unfortunately von Arx (1952) did not note which specimen contained ascospores; according to Untereiner (1993) and in my own experience, none of the four collections can be located in Persoon’s herbarium.
Von Arx (1952) considered C. cirrhosa and C. vestita conspecific and cited other 13 synonymous names for C. cirrhosa. The examination of the type material of C. vestita by the present author revealed a fungus identical to C. cirrhosa sensu Persoon (1801) and von Arx (1952). Though the name S. cirrhosa was sanctioned by Fries, no suitable herbarium material matching the concept of the species could be located in Fries’ herbarium. The only material of S. cirrhosa in Fries’ herbarium was an exsiccate collection Scleromyceti Sueciae 346, which represents another fungus, Lentomitella crinigera (this study). Because no type or other authenticated material of S. cirrhosa could be found, I herewith lectotypify the name S. cirrhosa with Persoon’s original illustration of this species (Persoon 1800: Tab 24, Fig. 3). The recent collection of this fungus, from which the culture and LSU and SSU sequences were derived, is designated here as an epitype (PDD 81434!).
Lentomitella (Höhnel 1905) was segregated from Ceratostomella for the single species C. vestita Sacc. as L. vestita (Sacc.) Höhn. The name Lentomitella is available to accommodate taxa that do not match the generic concept of Ceratostomella based on C. rostrata and that possess hyaline, ellipsoidal ascospores; unitunicate, shortstipitate, sessile asci formed on croziers and with a conspicuous, refractive, nonamyloid apical ring, cylindrical paraphyses and the phaeoisarialike anamorph. Therefore Lentomitella is reinstated and three species are accepted (viz. L. cirrhosa with L. vestita relegated to its synonymy, L. crinigera and L. tomentosa). Lentomitella cirrhosa and L. crinigera formed a strongly supported monophyletic clade in the MP1 analysis. They were clearly separated from the phenotypically similar species of Ceratosphaeria, which resides on a well supported branch with Ophioceras Sacc. (96/100) within the Magnaporthaceae. The differences between the sequence data, conidiogenesis (phaeoisaria-like anamorph of Lentomitella vs harpophora-like anamorph of Ceratosphaeria) and ascospore shape warrant the delimitation of the two holomorph genera, Lentomitella and Ceratosphaeria.
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KEY TO THE GENERA CERATOSPHAERIA, CERATOSTOMELLA, ENDOXYLA, LENTOMITELLA AND XYLOMELASMA |
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Ceratostomella Sacc., Michelia 1:370. 1878, emend. Réblová
Perithecia solitary to densely crowded, nonstromatic, venter globose to subglobose, superficial, semiimmersed or immersed, glabrous or tuberculate, dark brown to black, surrounded by sparse mycelium; hyphae growing out of the bottom part of the outer perithecial wall. Necks protruding separately, central, elongate, cylindrical, straight to slightly flexuous, perpendicular, oblique to almost decumbent toward the substratum, sulcate or glabrous, with periphysate ostiole. Perithecial wall leathery to fragile, two-layered. Outer wall of brown, thick-walled cells, textura prismatica to textura angularis to textura epidermoidea, and textura prismatica to porrecta in the neck; often with a distinct, external crustose layer of heavily pigmented, dark brown cells with opaque walls. Inner layer of thinner-walled, subhyaline to hyaline, elongated and compressed cells. Ascogenous hyphae branching, discrete, with croziers, each producing several lateral and terminal, dehiscent cells sequentially and simultaneously, from each of which one ascus arises as an outgrowth. Paraphyses abundant, unbranched, septate, hyaline, broad-celled and heavily constricted at the septa, wider near the base, tapering toward the tip, apically free, longer than the asci, dissolving with age. Asci unitunicate, clavate to cylindricalclavate, short-stipitate, truncate to broadly rounded at the apex, tapering toward the base from the sporiferous portion, floating freely within the centrum at maturity, with a shallow, indistinct, nonamyloid apical annulus, 8-spored. Ascus stipe usually containing non-refractive material deposited at the bottom part, visible after ascus dehiscence from the ascogenous hypha. Ascospores ranging from suballantoid to irregularly ellipsoid to globose to reniform, straight or curved, often flattened on one side, hyaline when young, pale brown before discharge, aseptate, smooth, sometimes with terminal pores, arranged in a fascicle in the upper part of the ascus or 2–3-seriate in the sporiferous part.
Typus. –
Ceratostomella rostrata (Tode : Fr.) Sacc., Syll. Fung. 1:409. 1882. (Lectotype designated by Clements and Shear [1931].)
Anamorph. – Hyphomycetous, dematiaceous; in culture only sterile mycelium seen.
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Ceratostomella cuspidata (Fr. : Fr.) Réblová, comb. nov. FIGS. 3–6, 23A, B.
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= Ceratostoma cuspidatum (Fr. : Fr.) Sacc., Syll. Fung. 1:474. 1882.
Anamorph. – Unknown.
Perithecia solitary to densely crowded forming a crust, superficial with base of the venter slightly immersed, subglobose to globose, 400–500 µm high, 380–500 µm diam, glabrous, slightly tuberculate, dark brown, surrounded by dense mycelium, especially at the bottom; hyphae reddish brown, septate, 4.5–6 µm wide; neck central, elongate, straight or slightly flexuous, rounded at the top, sulcate with 4–5 ridges, (250–)400–1000 µm long, 110–130 µm diam. Perithecial wall leathery to fragile, textura prismatica to textura angularis, 55–80 µm thick, up to 90–112 µm thick in places with tubercules, external crustose layer 10–17 µm thick. Some cells in the outer layer with "Munk pores". Paraphyses 6–11 µm wide near the base, tapering to 1.5–2.0 µm at the top. Asci cylindricalclavate, 21–30 x (5–)6–7 (mean ± SE = 24.7 ± 0.4 x 6.3 ± 0.1) µm, truncate at the apex, tapering basally, short-stipitate, with an apical annulus, 3–3.5 µm diam, ca. 1 µm high. Ascospores suballantoid to reniform, curved, flattened on one side, 4–5 x 2–3 (mean ± SE = 4.4 ± 0.1 x 2.7 ± 0.1) µm, pale brown, aseptate, smooth, in a fascicle in the upper part of the ascus or 2–3-seriate in the ascus.
ISOTYPE. – Fries’ Scleromyceti Sueciae 117, decayed wood (PRM 666203).
Additional specimens examined. – CZECH REPUBLIC. SOUTHERN MORAVIA: Valtice, Randevous, decayed wood of Quercus sp., 4 Jun 1999, A. Vágner, M.R. 1813 (PRA). NEW ZEALAND. WESTLAND: Mount Aspiring National Park, Makarora Bush Walk, 500 m N of NP Headquaters in Makarora, decayed wood of Nothofagus sp., 30 Mar 2005, M. Réblová (PRA).
Habitat. – Saprobe on decayed wood of deciduous trees.
Distribution. – Czech Republic, New Zealand, Sweden.
Commentary. – Ceratostomella cuspidata is the most similar to C. rostrata, which differs by larger perithecia and narrower suballantoid to somewhat oblong ascospores.
Ceratostomella pyrenaica Réblová & Fournier, sp. nov. FIGS. 7–11, 23C, D
Perithecia gregaria, immersa vel semiimmersa, globosa vel subglobosa, brunnea, 400–550 µm diam, 350–550 µm alta, ostiolo centrali elongato, cylindraceo, 600–1130 µm longo, 95–110 µm diam. Paries perithecii 45–65 µm crassus, bistratosus. Paraphyses copiosae, hyalinae. Asci cylindraceo-clavati, (30–)33–40 x 5.5–7 µm, breviter stipitati, 8-spori. Ascosporae ellipsoideae vel oblongae, apiculatae, 7–9 x 3–4 µm, unicellulares, brunneae.
Anamorph. – Hyphomycetous, dematiaceous; in culture only sterile mycelium seen.
Perithecia gregarious, venter immersed to semiim-mersed, globose to subglobose, 350–550 µm high, 400–550 µm diam, slightly roughened, dark brown, surrounded by sparse mycelium; hyphae dark brown, septate, flexuous, 4–5 µm wide; neck central, elongate, straight or slightly flexuous, rounded at the apex, sulcate with 4–5 ridges, 600–1130 µm long, 95–110 µm diam. Perithecial wall leathery to fragile, textura prismatica, 45–65 µm thick, external crustose layer 10–15 µm thick. Paraphyses 10–12 µm wide near the base, tapering to 2–2.5 µm at the top. Asci cylindricalclavate, (30–)33–40 x 5.5–7 (mean ± SE = 36.2 ± 0.7 x 7.3 ± 0.3) µm, truncate to broadly rounded at the apex, short-stipitate, with an apical annulus ca. 3 µm diam, 0.5–1 µm high. Ascospores ellipsoidal to oblong, slightly curved and apiculate at both ends, flattened on one side, 7–9 x 3–4 (mean ± SE = 8.2 ± 0.1 x 3.1 ± 0.1) µm, pale brown, aseptate, smooth, often with two large drops, 2–3-seriate in the ascus or in a fascicle in the upper part of the ascus.
Characteristics in culture. – Colonies reaching 10–11 mm diam on PCA, grayish-brown. Aerial mycelium well developed, denser in the center of colony; hyphae subhyaline, 3–3.5 µm wide, septate, smooth. Substrate mycelium of brown, septate hyphae, 3–4 µm wide, smooth. Sporulation not seen. Margin gnawed. Reverse dark brown.
HOLOTYPE. – FRANCE. PYRENEES ATLANTIQUES: Rimont, Las Muros, Ruisseau de Peyran, decayed wood of a trunk of Alnus glutinosa, 8 Sep 2000, J. Fournier J.F. 00195 (PRM 902273).
Additional specimens examined. –
CZECH REPUBLIC. SOUTHERN MORAVIA: Hodonín district, Ska
iny Nature Reserve near Mikul
ice, decayed wood of a trunk of Acer campestre, 24 Oct 2004, M. Réblová, M.R. 2912 (PRA, culture CBS 117116). FRANCE. PYRENEES ATLANTIQUES: Rimont, Las Muros, Ruisseau de Peyran, decayed wood of a trunk of A. glutinosa, 18 Jun 2001, J. Fournier J.F. 01114 (PRA); ibid., Charente Maritime, La Villedieu, Forêt d'Aulnay, decayed basidioma of Trametes gibbosa on a dead stump of Fagus sylvatica, 30 Apr 2004, J. Fournier J.F. 04071 (PRA).
Etymology. – Pyrenaica referring to the locality of origin, Pyrénées Atlantiques.
Culture. – CBS 117116 ex M.R. 2912.
Habitat. – Saprobe on decayed deciduous wood. Distribution. Czech Republic, France.
Commentary. – Ceratostomella pyrenaica is characterized by pale brown, smooth, ellipsoidal to oblong ascospores, slightly apiculate at the ends, a shape that is well distinguishable within other Ceratostomella species.
Ceratostomella rhynchophora (De Not.) Réblová, comb. nov. FIGS. 12–15, 23E–F
Basionym. – Sordaria rhynchophora De Not., Comm. Crittog. Ital. 2:480. 1867.
= Ceratostoma rhynchophorum (De Not.) W. Kirschst., Krypt. Fl. Mark Brandenb. 7:249. 1911.
= Ceratostoma notarisii Sacc., Fung Ven. Ser. II, p. 308, 1874. (Illegit. name, Art. 52)
Anamorph. – Unknown.
Perithecia solitary to gregarious, venter superficial to completely immersed, globose to subglobose, 480–660 µm high, 500–650 µm diam, slightly roughened, dark brown, surrounded by dense mycelium; hyphae dark brown, septate, branching, flexuous, 4–5 µm wide; neck central, elongate, straight or slightly flexuous, sulcate at the apex with 3–4 ridges, 400–2000 µm long, 140–190 µm diam. Perithecial wall leathery, textura prismatica to textura epidermoidea, 75–100 µm thick, external crustose layer 12–17.5 µm thick. Paraphyses 8–11 µm wide near the base, tapering to 2–2.5 µm at the top. Asci cylindrical-clavate, (33–)35–44 x 7–8.5(–10) (mean ± SE = 38.8 ± 0.5 x 8.6 ± 0.2) µm, truncate to broadly rounded at the apex, short-stipitate, with an apical annulus 2.5–3 µm diam, ca. 1 µm high. Ascospores ellipsoidal, sometimes flattened on one side, slightly apiculate at both ends, 6–7 x (3.5–)4–5 (mean ± SE = 6.4 ± 0.1 x 4.3 ± 0.1) µm, midbrown, aseptate, smooth, with a minute pore at each end, 1–2-seriate in the ascus or in a fascicle in the ascus.
NEOTYPE. – ITALY. Decayed wood of Prunus domestica, P.A. Saccardo (PAD, as Ceratostoma notarisii; neotype designated here).
Additional specimens examined. – FRANCE. PYRENEES ATLANTIQUES: Rimont, Las Muros, decayed wood of P. domestica, 3 Feb 2002, J. Fournier J.F. 02022 (PRA); ibid. 21 Apr 2002, J. Fournier J.F. 02070 (PRA).
Habitat. – Saprobe on decayed deciduous wood.
Distribution. – France, Italy.
Commentary. –
Saccardo (1874:308) created an obligate synonym Ceratostoma notarisii based on Sordaria rhynchophora with explanation that the name "Ceratostoma rhynchophorum" would have an identical meaning as the already existing name Ceratostoma rostratum.
Two specimens of C. rhynchophora were found in PAD under the name Ceratostoma notarisii. One contained old and empty, minute, brown perithecia with long, central filiform necks strongly similar to species of Melanospora. The second specimen was collected by Saccardo and contained mature perithecia with asci, ascospores and paraphyses that fully match the original description of S. rhynchophora. Because no type or authenticated material of S. rhynchophora could be found (PAD, RO, MI), the collection from Saccardo’s herbarium labeled as C. notarisii is chosen as a neotype.
Ceratostomella rostrata (Tode : Fr.) Sacc., Syll. Fung. 1:409. 1882. FIGS. 16–18, 23G, H
Basionym. – Sphaeria rostrata Tode, Fungi Mecklenb. Sel. 2:14. 1791 : Fries, Syst. Mycol. 2:473. 1823.
= Ceratostoma rostratum (Tode : Fr.) Fuckel, Jahrb. Nassau. Ver. Naturk. 23–24:127. 1870.
= Ceratosphaeria rostrata (Fr.) Sacc., Syll. Fung. 2:227. 1883. (as "[Kickx] Sacc.")
= Endoxyla rostrata (Tode : Fr.) Munk, Dansk Bot. Ark. 17(1):196. 1957.
= Wegelina polyporina M.E. Barr, Cryptogamie, Bryol. Lichénol. 19:170. 1998.
= Ceratostoma grumsinianum W. Kirschst., Ann. Mycol. 34:199. 1936.
Anamorph. – Unknown.
Perithecia densely aggregated forming a crust on the substratum, venter superficial to semiimmersed, globose to subglobose, 700–750 µm high, 650–750 µm diam, tuberculate, dark brown to black, surrounded by sparse mycelium; hyphae dark brown, septate, flexuous, branching, 6–7 µm wide; neck central, elongate, straight or slightly flexuous, tapering toward the apex and narrowly rounded, sulcate with 4–5 ridges, 800–2500 µm long, 130–170 µm diam. Perithecial wall leathery to fragile, textura prismatica to textura angularis, 62–85 µm thick, 110–125 µm thick in places with tubercules, external crustose layer 12–25 µm thick. Some cells in the outer layer with "Munk pores". Paraphyses 8–12 µm wide near the base, tapering to 2.5–3 µm at the top. Asci cylindricalclavate, (26–)30–39 x 5–6 (mean ± SE = 33.4 ± 0.4 x 5.9 ± 0.1) µm, truncate to broadly rounded at the apex, short-stipitate, with an apical annulus 1.5–2 µm diam, ca. 1 µm high. Ascospores allantoid to suballantoid, 4.5–6 x 1.5–2 (mean ± SE = 5.3 ± 0.1 x 1.9 ± 0.1) µm, pale brown, aseptate, smooth-walled, usually in a fascicle in the upper part of the ascus or 2–3-seriate within the ascus.
LECTOTYPE illustration: Tode, Fungi Mecklenbur-genses Selecti, Fasc. 2: Tab. IX, Fig. 79. 1791 (lectotype illustration designated here).
EPITYPE. – Fries’ Scleromyceti Sueciae 116, decayed wood (PRM 666367, epitype designated here).
Additional specimens examined. –
Fries’ Scleromyceti Sueciae 116, decayed wood (C, G, K 114933, UPS, as Sphaeria rostrata). CZECH REPUBLIC. SOUTHERN BOHEMIA: 
umava Mountains National Park, Volary, Medv
dice Nature Reserve, decayed wood of Ulmus glabra, 26 Oct 2001, M. Réblová M.R. 1887, 1925 (PRA). FINLAND. Wehmersalmi, on hymenium of Fomes fomentarius, 12 Aug 1992, U. Söderholm, L. Kasoner 2052 (DAOM 227785). GERMANY. Bredowen Forst, Orthavelland, in hymenium of the decayed basidioma of F. fomentarius on Betula sp., 6 Oct 1916, W. Kirschstein (B700006552, holotype of Ceratostoma grumsinianum). POLAND. W. Siemaszko: Fungi Bialowiezenses exsiccati no. 158, Bialystok, Bialowieza, National Park Puzcza Bialowieszka, in hymenium of the decayed basi-dioma of Fomes sp., Aug 1922, W. Siemaszko (PRM 655803, as Ceratostoma rhynchophorum). SWEDEN. Närke, Glanshammar Parish, Skölv, on hymenium of F. fomentarius, 20 May 1993, K.G. Nilsson (DAOM 227786, isotype of Wegelina polyporina); Öland, Vickleby Parish, Nature Reserve Vickleby ängar 2 km NW of Vickleby, decayed bracket basidiomata of F. fomentarius on Betula pendulosa, 4 Jun 2001, N. Lundqvist N.L. 21416.
Habitat. – Saprobe on decayed deciduous wood and decayed aphyllophoraceous basidiomata.
Nomenclatural note. –
Saccardo (1883:227) proposed a new combination Ceratosphaeria rostrata (Kickx) Sacc. based on Sphaeria rostrata Kickx, Fl. Crypt. Flandres 1:338. 1867, where: "Sphaeria rostrata Fr., Kx. Rech. Cent. III. p. 17; West. Herb. Crypt. fasc. 11, No. 552; Dryinosphaera rostrata Dmtr.—Desm. Pl. Crypt. fasc. 20, No. 973" is given. In Kickx’s earlier work cited above (Kickx 1847: 17) "Sphaeria rostrata Fr., Desm. Crypt., fasc. 20, no. 973" is given. It is obvious that in both publications, Kickx referred to Fries’ concept of S. rostrata. It is probable that Saccardo (1883) misinterpreted the name Sphaeria rostrata and incorrectly ascribed the name to J. Kickx.
Kirschstein (1911) proposed a new combination for Ceratostoma rhynchophorum, however according to its description and comparison to the material of C. rhynchophora and C. rostrata this species was mis-identified; it is C. rostrata. No herbarium material of C. rhynchophorum could be found in Kirschstein herbarium (B).
Commentary. – Ceratostomella rostrata is similar to C. cuspidata in having tuberculate perithecia with sulcate neck. However size of perithecia, shape and width of ascospores and length of asci clearly distinguish the two taxa; in C. cuspidata asci are usually shorter than 30 µm, and ascospores are suballantoid to reniform, 2–3 µm wide.
Untereiner (1993) cited two other taxa in the synonymy of S. rostrata (viz. Endoxyla laevirostris Munk [Denmark. Jylland, Staksrode near Juelsminde, distr. 24, rotten wood of a branch of Fagus sylvatica, 12 Oct 1963, A. Munk, C!] and Ceratostomella ampullasca [Cooke] Sacc. [U.K. Shere, on rotten wood of Quercus sp., Feb 1869, K 84424!]). According to the study of their type material by the present author these two taxa are neither conspecific nor are they species of Ceratostomella. Additional culture and molecular data are needed to confirm their systematic position.
Ceratostomella sp. FIGS. 19–21
Anamorph. – Unknown.
Perithecia superficial to semiimmersed, sitting densely on wood around the margin of a resupinate basidiomycete, globose to subglobose 530–570 µm high, 500–550 µm diam, tuberculate, dark brown, surrounded by dense mycelium; hyphae dark brown, septate, branching, flexuous, 2.5–3 µm diam; neck central, elongate, straight or slightly flexuous, sulcate with four ridges, 600–1000 µm long, 110–130 µm diam. Perithecial wall fragile, textura prismatica to textura angularis, 87–92 µm thick, 97–125 µm thick in places with tubercules, external crustose layer 10–12 µm thick. Some cells in the outer layer with "Munk pores". Paraphyses 6–8 µm wide near the base, tapering to 2.5–3 µm at the top. Asci clavate, 25– 32 x 5–6 (mean ± SE = 28.7 ± 0.9 x 5.5 ± 0.1) µm, truncate to broadly rounded at the apex, short-stipitate, with an apical annulus 2–2.5 µm diam, 0.5–1 µm tall. Ascospores ellipsoidal to globose, 3–4 x (2.5–)3–4 (mean ± SE = 3.6 ± 0.1 x 3.2 ± 0.1) µm, pale brown to stramineous, aseptate, smooth-walled, 1–2-seriate in the ascus.
Specimen examined. – SWEDEN. SCANIA: Skurup, Svaneholm, near the west shore of the lake, decayed wood of Fagus sylvatica in association with Steccherinum ochraceum, 10 Nov 2000, S.-Å. Hanson, M.R. 2592.
Habitat. – Saprobe on decayed deciduous wood in association with old basidiomata.
Distribution. – Sweden.
Commentary. – The shape of the ascospores is distinctive and unique within the reported ascospore variability of Ceratostomella. However, the material of this fungus is too poor to designate a holotype and additional material is needed for a more detailed study to describe a new species.
Ceratosphaeria lampadophora (Berk. & Broome) Niessl, Verh. Naturf. Vereins Brünn 14:203. 1876. FIG. 24
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For synonymy, detailed description and illustration refer to Hyde et al (1997).
Characteristics in culture. Colonies reaching 9–11 mm diam on PCA, yellowish, whitish in the center. Aerial mycelium scarcely developed in the center of the colony and on the inoculating block; hyphae subhyaline to hyaline 2–3 µm wide, septate, smooth. Sporulation copious, widespread throughout the colony in 1 mo. Phialides or short conidiophores arising on aerial hyphae, conidial heads slimy, inconspicuous, and transparent. Reverse dark yellow. Conidiogenous cells phialidic, terminal or intercalary, hyaline to subhyaline, cylindrical, 14–18 x 2–2.5 µm, tapering below the collarette to 1.5 µm. Conidia cylindrical with curvature, hyaline, narrowly rounded at both ends, 9–11(12) x 1–1.5 µm, aseptate, smooth.
Culture. – CBS 117555 ex J.F. 01065.
HOLOTYPE. – UNITED KINGDOM. ENGLAND: Bath, Combe Hay, decayed wood, Apr 1855, C.E. Broome (holotype of Sphaeria lampadophora, K).
Additional specimens examined. –
CZECH REPUBLIC. SOUTHERN BOHEMIA: umava Mountains National Park, Mount Spáleni
t near Sto
ec, decayed wood of F. sylvatica, 12 Nov 2000, M. Réblová M.R. 1734 (PRA). FRANCE. PYRENEES ATLANTIQUES: Rimont, Ariège, Las Muros, on decorticated wood of Populus tremula, 11 Apr 2001, J. Fournier J.F. 01065 (PRA; culture CBS 117555); ibid. 18 Jun 2001, J.F. 01115 (PRA). SWITZERLAND. Grange canal near Genève, decayed wood of a trunk, 8 Feb 1859, J. Mü ller (Herb. Barbey-Boissier 605, G). Lentomitella Höhnel, Ann. Mycol. 3:552. 1905.
Perithecia solitary to gregarious, nonstromatic, venter superficial, semiimmersed or immersed, globose, subglobose to conical, glabrous or roughened, dark brown to black, surrounded by sparse mycelium; hyphae growing out of the bottom part of the outer perithecial wall. Necks dark, protruding separately, central, perpendicular or oblique to the substratum, elongate, cylindrical, straight to slightly flexuous, sulcate or glabrous, ostiolum periphysate. Perithecial wall leathery, 2-layered. Outer wall of brown, thick-walled cells, textura prismatica to textura angularis; external layer of heavily melanized cells with opaque walls. Inner layer of thinner-walled, subhyaline to hyaline, elongated and compressed cells. Ascogenous hyphae not extensive, proliferating through croziers. Paraphyses abundant, cylindrical, tapering toward the tip, septate, slightly constricted at the septa or nonconstricted, hyaline, apically free, longer than the asci, dissolving partly at maturity. Asci unitunicate, cylindrical-clavate, short-stipitate, sessile, truncate to broadly rounded at the apex, with a distinct, refractive, nonamyloid apical annulus, 8-spored. Ascospores ellipsoidal, hyaline, aseptate or several-septate, smooth or with few longitudinal ridges, obliquely 1-seriate or 1–2-seriate in the ascus.
Typus. – Lentomitella vestita (Sacc.) Höhn.
Anamorph. – Phaeoisaria-like.
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Lentomitella cirrhosa (Pers. : Fr.) Réblová, comb. nov. FIGS. 25–28, 44A–C
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= Ceratostoma cirrhosum (Pers. : Fr.) Fuckel, Jahr. Nassau. Ver. Naturk. 23–24:127. 1870.
= Ceratostomella cirrhosa (Pers. : Fr.) Sacc., Michelia 1:370. 1878.
= Endoxyla cirrhosa (Pers. : Fr.) Arx & E. Müll., Beitr. Krypt-Fl. Schweiz 11(1):355. 1954.
= Sphaeria investita Schw., Trans. Amer. Phil. Soc. 2, Vol. 4:216. 1834.
= Ceratostoma investitum (Schw.) Ellis & Everh., North Amer. Pyrenom. p 193. 1892.
= Ceratostomella investita (Schw.) Starbäck, Bih. Kongl. Svenska Vet.-Akad. Handl. 19(2):26. 1894.
= Ceratostomella vestita Sacc., Michelia 1:370. 1878.
= Lentomitella vestita (Sacc.) Höhn., Ann. Mycol. 3:548. 1905.
= Cerastomis vestita (Sacc.) Clem., Gen. Fungi p. 259. 1931.
= Endoxyla vestita (Sacc.) Munk, Bot. Tidsskr. 61:64. 1965.
= Ceratostomella vestita Sacc. var. varvicensis Grove, J. Bot. 23:131. 1885.
= Eriosphaeria conoidea Feltg., Pilz Flora Luxemb. 3:282. 1903.
Anamorph. – Hyphomycetous, dematiaceous; in culture only sterile mycelium seen.
Perithecia solitary or gregarious, venter immersed to superficial, globose to subglobose, 300–410 µm high, 300–400 µm diam, slightly roughened, dark brown, surrounded by sparse mycelium; hyphae brown, 4.5–5 µm wide; neck dark brown, central, elongate, cylindrical, straight to slightly flexuous, sulcate with four ridges at the apex, 300–900 µm long, 100–120 µm diam. Perithecial wall leathery, 30–50 µm thick, textura prismatica to textura angularis; externally with a row of heavily melanized cells with opaque walls. Paraphyses 6–8 µm wide near the base, tapering to 2.5–3 µm. Asci cylindrical-clavate, truncate to broadly rounded at the apex, (42–)50–74 x 6–7(–8) (mean ± SE = 62.3 ± 1.1 x 7.1 ± 0.1) µm, sessile, with an apical annulus 2.5–3 µm diam, 1–1.5 m high, 8-spored. Ascospores ellipsoidal, (5.5–)6–9(–10) x 3–4.5(–5) (mean ± SE = 7.9 ± 0.2 x 3.8 ± 0.1) µm, hyaline, 1(–3)-septate, not constricted at the septa, with few longitudinal ridges, obliquely 1-seriate or 1–2-seriate.
Characteristics in culture. – Colonies 8–12 mm diam on PCA, dark brown, cocoa brown in the center. Aerial mycelium well developed, densest in the center of the colony, hyphae subhyaline to pale brown, 3–4 µm wide, septate, smooth. Substrate mycelium of brown, septate hyphae, 3.5–4 µm wide, septate, smooth. Sporulation not observed. Reverse dark brown.
LECTOTYPE illustration: Persoon, Icones et Descriptiones Fungorum minus Cognitorum. Fasc. 2, Tab. 24, Fig. 3. 1800 (lectotype illustration designated here).
EPITYPE. – NEW ZEALAND. WESTLAND: Hokitika, 40 km S of Greymouth, Mananui Point, Lake Mahinapua, Swimmers Beach walks, decayed wood of Podocarpus totara, 5 Mar 2003, M. Réblová M.R. 2705 (PDD 81434, culture ICMP 15131; epitype designated here).
Additional specimens examined. –
CZECH REPUBLIC. SOUTHERN BOHEMIA: umava Mountains. National Park, Volary, Spálenit Mount near Stoec, decayed wood of Fraxinus excelsior, 16 Aug 1999, M. Réblová M.R. 1545; ibid., Povydí National Nature Reserve, 
e
kova pila, decayed wood of a trunk of U. glabra, 27 Aug 2000, M. Réblová M.R. 1677; ibid., 
elezná Ruda, glacial cirque of the ertovo jezero lake, decayed wood of Picea abies, 22 Oct 1996, M. Réblová M.R. 858/96; ibid., Práily, Mount danidla, 22 Aug 2000, M. Réblová M.R. 2234; Novohradské hory Mountains, Nové Hrady, ofínsk
prales National Nature Reserve, decayed wood of F. sylvatica, 20 May 1997, M. Réblová M.R. 917 (PRA). Moravia: Flora Bohem. et Morav. exsicc. Lfg. 14, No. 675, Hranice na Morav, Podhoí, decayed wood, 24 Mar 1913, F. Petrak (PRM 195089). BELGIUM. Decayed wood of Pyrus communis, 25 May 1902, Feltgen (LUX 043455!, holotype of Eriosphaeria conoidea). DENMARK. Snarup Mose SW of Kvaerndrup, inner side of the bark of Betula sp., 1 Apr 1999, J. Vesterholt J.V. 99-015 (C 45296). SJALLAND: Charlottenlund skov, rotten wood, 10 Nov 1964, A. Munk (C); ibid., Dyrehaven, rotten wood, 26 Nov 1964, A. Munk (C); ibid., Dyrehaven, rotten wood of a stump of F. sylvatica, 2 Dec 1963, A. Munk (C). FRANCE. PYRENEES ATLANTIQUES: Rimont, Ariège, Las Muros, decayed wood, 9 May 2004, J. Fournier J.F. 04084 (PRA). GERMANY. Nassau, decayed wood, autumn 1894, L. Fuckel (Herb. Barbey-Boissier 603, as Ceratostomella rhenana, G). ITALY. On decayed undetermined wood, Sep 1878. P.A. Saccardo (PAD!, holotype of C. vestita). SWEDEN. SCANIA: Kattarp distr., Kattarps mosse, on decorticated wood of a branch of Betula sp., 21 Apr 2001, S.- Å. Hanson. USA. PENNSYLVANIA: Northhampton County, Bethlehem, L.D. Schweinitz, S.F. 1621.476 (PH 01016198!, holotype of Sphaeria investita).
Culture. – ICMP 15131 exepitype PDD 81434.
Habitat. – Saprobe on decayed wood.
Distribution. – Austria, Czech Republic, Belgium, Denmark, France, Italy, New Zealand, Sweden, USA.
Commentary. – Lentomitella cirrhosa is a common saprobe growing on decayed wood. It is characterized by ellipsoidal, longitudinally striate ascospores with one, rarely three septa and long-necked perithecia with the sulcate top of the neck.
Höhnel (1906) in his revision of Feltgen’s original herbarium material distinguished between C. cirrhosa and L. debaryana and considered them separate taxa. Höhnel (1906) considered E. conoidea conspecific with Lentomita debaryana (Auersw.) Höhn. (as de Baryana) and listed four other species closely related to L. debaryana. Von Arx (1952) accepted L. debaryana and the four species cited by Höhnel (1906) as synonyms of C. cirrhosa and listed other nine synonymous names but without studying their type material. Re-examination of the respective types by the present author revealed: C. vestita (= L. cirrhosa, PAD!), C. investita (= L. cirrhosa, PH 01016198!), E. conoidea (= L. cirrhosa, LUX 043455!), and Ceratostomella similis W. Kirschst. (= C. ampullasca, B 700009209!). The examination of C. ampullasca (K 84424!) revealed that it is not a species of Lentomitella and requires additional study. Ceratostomella rostrata is not congeneric with L. cirrhosa and it is redescribed under Ceratostomella sensu stricto in this study. The type material of Ceratostomella capilliformis Bomm. et al, Ceratostomella albocoronata (Ellis) Sacc. and Ceratostomella multirostrata (Fuckel) Sacc. contained only empty perithecia. The type specimen of Ceratostomella subpilosa (Fuckel) Sacc. (Fungi rhenani 2251, G!) contained several perithecia with only few ascospores attached to the outer perithecial wall; ascospores ellipsoidal, slightly tapering toward the ends, 8–9 x 4–5 µm, finely verruculose. The fungus is not conspecific with L. cirrhosa, but it is likely a member of Lentomitella based on the ascospore characters; however, fresh material is needed to clarify its systematics. The type material of Ceratostomella unedonis H. Fabre, Ceratostomella debaryana (Auersw.) Sacc. and Ceratostomella lejocarpa Sacc., which were cited by von Arx (1952) among the synonyms of C. cirrhosa, could not be located.
Lentomitella crinigera (Cooke) Réblová, comb. nov. FIGS. 29–33, 44D–H
Basionym. – Sphaeria crinigera Cooke, Grevillea 1:156. 1873.
Ceratosphaeria crinigera (Cooke) Sacc. Syll. Fung. 2:227. 1883.
Anamorph. – Phaeoisaria-like.
Perithecia solitary or gregarious, venter immersed to superficial, globose to subglobose, 510–700 µm high, 500–600 µm diam, slightly roughened, dark brown, surrounded by sparse mycelium; hyphae brown, 3.5–5 µm wide; neck dark brown, central, elongate, cylindrical, straight to slightly flexuous, sulcate with 4 ridges, 300–1000 µm long, 120–140 µm diam. Perithecial wall leathery, 37–50 µm thick, textura prismatica to textura angularis; externally with a row of heavily melanized cells with opaque walls. Paraphyses 6–8 µm wide near the base, tapering to 2.5–3 µm. Asci cylindrical-clavate, truncate to broadly rounded at the apex, (65–)78–98(–103) x 7–9(–11) (mean ± SE = 86.7 ± 0.8 x 8.3 ± 0.1) µm, sessile, with an apical annulus 2.5–3.5 µm diam, ca. 1–1.5 µm high. Ascospores ellipsoidal, 10–14 x (4–)5–6 (mean ± SE = 12 ± 0.1 x 4.6 ± 0.1) µm, hyaline, 1–3-septate, not constricted at the septa, with few longitudinal ridges, obliquely 1-seriate or 1–2-seriate.
Characteristics in culture. – Colonies reaching 8–10 mm diam on PCA, dark brown, cocoa-brown in the center of the colony. Aerial mycelium well developed, denser in the center of colony; hyphae subhyaline, 3–4 µm wide, septate, smooth. Substrate mycelium of brown, septate hyphae, 3.5–4 µm wide, smooth. Sporulation copious, widespread throughout the colony. Short conidiophores arising from aerial hyphae, conidial heads slimy, inconspicuous, transparent. Reverse dark brown. Conidiophores macronematous, mononematous, arising terminally or laterally, hyaline, 24–45 x 2–2.5 µm. Conidiogenous cells terminal or intercalary, hyaline, cylindrical, tapering toward the apex, 9–20(–25) x 2–2.5 µm, bearing 2–6 hyaline denticles 0.5–1 µm wide, 0.5–1 µm long. Conidia ellipsoidal to globose, apiculate at the base, 4–6 x 2–2.5 (mean ± SE = 5.2 ± 0.2 x 2.2 ± 0.3) µm, hyaline, aseptate, smooth.
HOLOTYPE. – UNITED KINGDOM. ENGLAND: Norfolk, King’s Lynn, decayed wood, C.B. Plowright (holotype of Sphaeria crinigera, K 84422!).
Additional specimens examined. –
CZECH REPUBLIC. SOUTHERN BOHEMIA: umava Mountains National Park, elezná Ruda, glacial cirque of the ertovo jezero Lake Nature Reserve, decayed wood of Abies alba, 28 Aug 1997, M. Réblová M.R. 997, M.R. 1034; ibid., 12 Aug 1999, M. Réblová M.R. 1544, M.R. 1585; ibid., 21 Aug 2000, M. Réblová M.R. 1672; ibid., glacial cirque of the erné jezero Lake Nature Reserve, decayed wood of A. alba, 13 Aug 1999, M. Réblová M.R. 1526, M.R. 1546; ibid., Práily, Mount danidla, 24 Aug 2000, decayed wood of F. sylvatica, M. Réblová M.R. 1671; ibid., Boubínsk prales Nature Reserve, decayed wood of P. abies, 17 Aug 1999, M. Réblová M.R. 1611; ibid., Modrava, Ptaí nádr, decayed wood of P. abies, 14 Aug 1999, M. Réblová M.R. 1457; ibid., Modrava, Modravské slat, Pytláck roh, decayed wood of a trunk of P. abies, 14 Aug 1999, M. Réblová M.R. 1652 (PRA). DENMARK. Silkeborg, Vesterskov, decayed wood of F. sylvatica, 24 Mar 1954, A. Munk (C). NEW ZEALAND. WESTLAND: Victoria National Park, Reefton, 90 km NE of Greymouth, Lake Stream track, 30 km SE of Reefton, decayed wood of a trunk of Nothofagus sp., 27 Feb 2003, M. Réblová M.R. 2659 (PDD 81435; cultures CBS 113655, ICMP 15124). SWEDEN. Fries’ Scleromyceti Sueciae No. 346, decayed wood (UPS).
Cultures. – CBS 113655, ICMP 15124, both isolates ex PDD 81435.
Habitat. – Saprobe on decayed wood.
Distribution. – Czech Republic, Denmark, New Zealand, Sweden, U.K.
Commentary. – Lentomitella crinigera is characterized by ellipsoidal, longitudinally striate, 1–3-septate ascospores. The related L. cirrhosa is distinguished from L. crinigera by shorter ascospores and asci and smaller perithecia.
Lentomitella tomentosa Réblová & J. Fournier, sp. nov. FIGS. 34, 35, 45A, B
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Anamorph. – Unknown.
Perithecia gregarious, venter immersed to entirely superficial, globose to subglobose, 400–500(–600) µm high, 350–600 µm diam, dark brown, collapsed laterally on drying, surrounded by sparse hyphae 4–5 µm wide, growing out of the outer perithecial wall; neck dark brown, central, perpendicular to slightly decumbent toward the substratum, elongate, cylindrical, straight to slightly flexuous, sulcate with 3–4 ridges, 200–450 µm long, 120–180 µm diam. Perithecial wall leathery, laterally 40–50 µm thick, 35–37 µm thick at the bottom, textura prismatica to textura angularis; externally with a row of heavily melanized cells with opaque walls. On the outer perithecial wall a red waxy layer, 8–18 µm thick attached to the surface of melanized cells; granules of red pigment also attached to the surface of the perithecial hairs and also staining the surrounding substratum. Outer perithecial wall of brown, thick-walled cells, inner layer of thinner-walled, subhyaline to hyaline, elongated and compressed cells. Paraphyses 4–6 µm wide near the base, tapering to 3–3.5 µm, longer than the asci. Asci cylindrical-clavate, truncate to broadly rounded at the apex, (75–)80–98 x (9–)10–11 (mean ± SE = 84.8 ± 1.4 x 10.6 ± 0.1) µm, sessile, with an apical annulus 2.5–3.5 µm diam, ca. 1.5 µm high. Ascospores ellipsoidal, 13–16(–17) x (5–)6–7 (mean ± SE = 14.4 ± 0.2 x 5.9 ± 0.1) µm, hyaline, aseptate, smooth, obliquely 1-seriate or 1–2-seriate.
HOLOTYPE. – FRANCE. Finistère, Plohars, Forêt de Carnoët, Pont Douar, 40 m a.s.l., decorticated wood of F. sylvatica, 26 Oct 2002, J. Fournier J.F. 02196 (PRM 902274).
Additional specimen examined. – DENMARK. SJALAND: Dyrehaven, decayed wood of a stump, 3 Oct 1964, A. Munk (C); ibid., 20 Mar 1965, A. Munk (C); ibid., Silkeborg, Spring area near Almindsø, decayed wood of A. glutinosa, 4 Sep 1953, A. Munk (C).
Etymology. – Tomentosus (L), tomentose, refers to the felted appearance of the surface of the perithecia.
Habitat. – Saprobe on decayed wood.
Distribution. – Denmark, France.
Commentary. – Lentomitella tomentosa is characterized by aseptate, ellipsoidal, smooth ascospores and perithecia with stout, sulcate, black neck in contrast to the finely tomentose venter with a red waxy layer covering the surface of the wall and hairs; the red pigment dissolves in KOH and stains the nearby wood reddish.
Xylomelasma Réblová, gen. nov.
Perithecia solitaria usque gregaria, superficialia usque immersa, globosa vel subglobosa, atro-brunnea, ostiolo centrali elongato, cylindraceo. Paries perithecii bistratosus. Paraphyses copiosae, hyalinae, cylindraceae. Asci cylindraceo-clavati, breviter stipitati, 8-spori. Ascosporae ellipsoideae usque oblongae, apiculatae, unicellulares vel raro 1-septatae, brunneae dum in asco.
Perithecia solitary to gregarious, nonstromatic, venter globose to subglobose, superficial, semiim-mersed or immersed, glabrous or roughened, dark brown to black, surrounded by sparse mycelium; hyphae growing out of the bottom part of the outer perithecial wall. Necks protruding separately, central, elongate, cylindrical, straight to slightly flexuous, perpendicular, oblique to almost decumbent toward the substratum, sulcate or glabrous, ostiolum periphysate. Perithecial wall leathery to fragile, two-layered. Outer wall of brown, thick-walled cells, textura prismatica; externally with dark brown cells with opaque walls but without any distinct crustose layer. Inner layer of thinner-walled, subhyaline to hyaline, elongated and compressed cells. Ascogenous hyphae sparsely branching, discrete, each sequentially and simultaneously producing several lateral and terminal dehiscent cells, each forming one ascus arises. Paraphyses abundant, branching, septate, hyaline, cylindrical, slightly constricted, slightly tapering toward the tip, apically free, longer than the asci. Asci unitunicate, cylindrical, short-stipitate, truncate to broadly rounded at the apex, narrowly rounded at the base, floating freely within the centrum at maturity, with a distinct, refractive, nonamyloid apical annulus, 8-spored. Ascus stipe usually containing a non-refractive material deposited at the bottom part, visible after ascal dehiscence from the ascogenous hyphae. Ascospores ellipsoidal to oblong, hyaline when young, pale brown before discharge, aseptate or some developing a median septum, smooth or finely verruculose, sometimes with terminal pores, obliquely 1-seriate in the ascus.
Typus. – Xylomelasma sordida Réblová.
Anamorph. – Hyphomycetous, dematiaceous; in culture only sterile mycelium seen.
Etymology. – Xylon (Gr) = wood; melas (Gr) black, referring to the dark, opaque perithecia.
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Xylomelasma novaezelandiae Réblová, sp. nov. FIGS. 36–38, 46A, B
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Anamorph. – Unknown.
Perithecia solitary to gregarious, venter immersed or partly erumpent, globose to subglobose, 350–400 µm high, 310–340 µm diam, smooth to slightly roughened, dark brown, surrounded by sparse mycelium; hyphae brown, septate, sparsely branched, 3–4 µm wide; neck central, elongate, straight or slightly flexuous, rounded at the top, not sulcate, 225–375 µm long, 95–115 µm diam. Perithecial wall leathery to fragile, textura prismatica, 52–62 µm thick; external crustose layer 11–14 µm thick. Paraphyses 6–7 µm wide near the base, tapering to 2.5–3 µm at the top. Asci cylindrical-clavate, 50–60(–65) x 7–8(–9) (mean ± SE = 53.2 ± 0.9 x 7.7 ± 0.2) µm, truncate at the apex, short-stipitate, with an apical annulus 2.5–3 µm diam, 1–1.5 µm high. Ascospores ellipsoidal, tapering toward both ends, narrowly rounded and slightly apiculate, 7–8 x (3.5–)4–5 (mean ± SE = 6.7 ± 0.3 x 4.1 ± 0.1) µm, pale brown, aseptate or rarely with one median septum at maturity, smooth-walled, with two large drops; mature ascospores become finely verruculose and dark brown, with a minute pore at each end, 1-, rarely 1–2-seriate in the ascus.
HOLOTYPE. – NEW ZEALAND. WESTLAND: Haast, 300 km SW of Greymouth, Jackson River valley, track to the Lake Ellery, S 43°52.9559', E 169°02.6211', rotten wood of a stump of Nothofagus sp., 10 Mar 2003, M. Réblová M.R. 2787 (PDD 81433).
Etymology. – Novaezelandiae refers to country of origin, New Zealand.
Habitat. – Saprobe on decayed deciduous wood.
Commentary. – Attempts to germinate ascospores of the cited specimen were unsuccessful.
Although Xylomelasma and Ceratostomella are readily distinguishable in the main characteristics of asci, apical annulus, ascospores and perithecial wall, X. novaezelandiae could be confused with C. rhynchophora because of the similar ascospore shape; but the ascospores are regularly ellipsoidal, not flattened on a side and shorter in X. novaezelandiae. Ceratostomella rhynchophora differs from X. novaezelandiae also in having shorter asci, ascospores 2–3-seriately arranged in the asci and larger perithecia with a thicker wall.
Xylomelasma sordida Réblová, sp. nov. FIGS. 40–43, 46C, D
Perithecia solitaria usque gregaria, superficialia usque immersa, globosa vel subglobosa, brunnea, 490–550 µm diam, 450–600 µm alta, ostiolo centrali elongato, cylindraceo, 120–150 µm diam, 950–1500 µm longo. Paries perithecii 55–62(–80) µm crassus, bistratosus. Paraphyses copiosae, hyalinae. Asci cylindraceo-clavati, (58–)60–76(–81) x 7–10(–13) µm, breviter stipitati, 8-spori. Ascosporae ellipsoi-deae, apiculatae, 9–12 x 4–6 µm, unicellulares, brunneae.
Anamorph. – Hyphomycetous, dematiaceous; in culture only sterile mycelium seen.
Perithecia solitary to gregarious, venter superficial to immersed, globose to subglobose, 490–550 µm diam, 450–600 µm high, slightly roughened, dark brown, surrounded by sparse mycelium; hyphae dark reddish brown, septate, slightly flexuous, 5–6 µm wide; neck central, elongate, slightly roughened, sometimes unevenly thickened, rounded at the top, sulcate with four ridges, 950–1500 µm long, 120–150 µm diam. Perithecial wall leathery to fragile, textura prismatica to textura angularis, 55–62(–80) µm thick, external crustose layer 7–12 µm thick. Some cells in the outer layer with "Munk pores". Paraphyses 6–10 µm wide near the base, tapering to 2– 3 µm at the top. Asci cylindrical-clavate, (58–)60–76 (–81) x 7–10(–13) (mean ± SE = 72.4 ± 1.1 x 8.2 ± 0.1) µm, truncate to broadly rounded at the apex, short-stipitate, with an apical annulus 2–2.5 µm diam, ca. 1 µm high. Ascospores ellipsoidal, sometimes slightly apiculate at both ends, 9–12 x 4–6 (mean ± SE = 10 ± 0.1 x 5 ± 0.1) µm, pale brown, aseptate, smooth, with a minute germ pore at each end, with 1(–2) oil drops, 1–2-seriate in the ascus.
Characteristics in culture. – Colonies on PCA reaching 12–14 mm diam in 14 d, dark olivaceous brown, grayish in the center. Aerial mycelium scarcely developed, densest in the center of colony; hyphae subhyaline to pale brown, 3–3.5 µm wide, septate, smooth. Substrate mycelium of dark brown, septate hyphae, 3–4 µm. Sporulation absent. Margin discrete, somewhat gnawed. Reverse dark brown.
HOLOTYPE: FRANCE. PYRENEES ATLANTIQUES: Ariège, Lescure, Bois du Pas du Baup, 500 m a.s.l., rotten wood of A. glutinosa, 24 Feb 2004, J. Fournier 04020 (PRM 902275, culture CBS 116000).
Additional specimens examined. –
ARGENTINA. BUENOS AIRES PROVINCE: Pdo. San Pedro, Gdor. Castro, Eucalyptus viminalis, May 1982, A. Romero 9, 19 (BAFC 32028, BAFC 32030, BAFC 32031); ibid., Aug 1982, A. Romero 10 (BAFC 51316); ibid., decayed wood, Aug 1981, A. Romero 6/22-8 (BAFC 32027); ibid. decayed wood, Feb 1982, A. Romero 38 (BAFC 32029). CZECH REPUBLIC. SOUTHERN MORAVIA: Beclav district, obora Soutok near Lanhot, on decayed deciduous wood. 23 Oct 2004, M. Réblová M.R. 2904, 2911 (PRA). DENMARK. SJALLAND: Dyrehaven, Distr. 45a, decayed wood, 5 Apr 1963, G. Carroll (C, as Endoxyla parallela); ibid. Rude Skov, decayed wood of a stump of F. sylvatica, 4 Feb 1964 (C, as Endoxyla parallela). HUNGARY. Bükk National Park in NE Hungary, decayed wood of F. sylvatica, Oct 2000, K. Kovács K.K. 215, K.K. 236 (C).
Culture. – CBS 116000 ex-type PRM 902275.
Etymology. – Sordidus (L), dull, refers to the dull brown ascospores.
Habitat. – Saprobe on decayed wood.
Distribution. – Argentina, Denmark, France, Hungary.
Commentary. –
Romero and Samuels (1991:43, Pl. 3m–p, Pl. 6g–j) reported a fungus identical with X. sordida from Argentina, but they considered it to be conspecific with Sphaeria avocetta Cooke & Ellis and proposed a new combination Endoxyla avocetta. They based their identification on the comparison with isotype material of S. avocetta (BPI, as Ceratostoma avocettum). Untereiner (1993) concluded that E. avocetta could not be classified in Endoxyla based on the description and illustration by Romero and Samuels (1991) and suggested its placement in the Diaporthales. Romero (1999) introduced a new genus Phaeognomoniella Romero for S. avocetta in this sense.
The revision of the holotype material of S. avocetta (FIGS. 47–50; USA. NEW JERSEY: Newfield, undetermined decayed wood, Jan 1876, J.B. Ellis, K 114931!, as Ceratostoma avocettum) by the present author revealed a fungus that possesses immersed, dark, conical, glabrous, monostichous perithecia with short, stout, cohering necks protruding just above the substrate surface, with remnants of stromatic tissue surrounding the projecting necks; paraphyses filiform, hyaline, septate, apically free, ca. 2 µm wide, longer than the asci; asci unitunicate, cylindrical, long-stipitate, rounded apically, floating free within the centrum, 66–88 µm long in pars sporifera, 6–7 µm wide, stipe 35–52 µm long, dissolving at maturity; with a nonamyloid indistinct apical annulus; ascospores ellipsoidal to subcylindrical, 11–13 x (3–)3.5–4 µm, aseptate, dull brown before discharge, with a minute pore at one end and slightly tapering toward this end, smooth, 1-seriate in the ascus. These characteristics of S. avocetta as seen on the holotype material are typical of members of Endoxyla (Untereiner 1993). Sphaeria avocetta is conspecific with Endoxyla operculata and should be relegated to its synonymy. Phaeognomoniella, although based on misidentified material, becomes a generic synonym of Endoxyla. For full synonymy of Endoxyla and E. operculata I refer to Untereiner (1993). The sizes of ascospores and asci in pars sporifera of X. sordida and S. avocetta are similar and might be confusing but in all other respects both fungi represent two distinct genera.
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DISCUSSION |
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Among comparatively well known ascomycetes, species of Ceratostomella and Xylomelasma can be compared with some phenotypically similar taxa of the Ceratostomataceae (e.g. Melanospora Corda or Arxiomyces P. Cannon & D. Hawksw.) especially because of the long-beaked perithecia and dull brown, aseptate ascospores with terminal pores. However, based on morphology of the asci, ascogenous system and interthecial filaments, none of the species attributed to Ceratostomella or Xylomelasma are melanosporaceous. According to molecular analyses of ribosomal DNA and RPB2, the phylogenetic affinity of Melanospora and other members of the Ceratosto-mataceae lie with the Hypocreales (Zhang and Blackwell 2002). In all recent analyses Melanospora takes this position. Ceratostomella bears also some resemblance to Cannonia Taylor & K.D. Hyde with the single species C. australis (Speg.) J. Taylor & K.D. Hyde in dark perithecia with immersed subglobose venter, separately projecting necks and brown ellipsoidal ascospores. Cannonia differs from Ceratostomella in ascospores that have a full-length germ slit, filiform paraphyses, presence of rudimentary stroma and a dark clypeus around the base of the neck (Taylor and Hyde 1999).
Lentomitella encompasses segregates from Ceratostomella sensu lato with hyaline, ellipsoidal, septate or aseptate ascospores; unitunicate, short-stipitate, sessile asci formed on croziers with a conspicuous, refractive, nonamyloid apical ring, cylindrical paraphyses and the phaeoisaria-like anamorph. Based on the LSU and SSU nrDNA sequence data (FIGS. 1, 2), Lentomitella has no obvious affinity to the Diaporthales, which originally were suggested to accommodate the genus (as Ceratostomella) (von Arx 1952; von Arx and Müller 1954; Müller and von Arx 1973; Munk 1952, 1957, 1965). Members of the Diaporthales have perithecia embedded in a stroma or only partly surrounded by reduced stromatic tissue; short-stipitate, basally rounded asci floating free within the centrum; the centrum is paraphysate, the tissue deliquesce early in development; anamorphs are phialidic coelomycetous with occasionally annellate conidiogenous cells indicating percurrent proliferation, and most of the taxa have phytopathogenic lives. Also another preliminary placement of Lentomitella in the Lasiosphaeriaceae (Untereiner 1993) or the Clypeosphaeriaceae (Barr 1990, Eriksson et al 2003) could not be confirmed in either MP1 or MP2 analyses. The Lasiosphaeriaceae were shown to form a well supported clade within the Sordariales (Huhndorf et al 2004) and Clypeosphaeria phillyreae Sousa da Câmara representing the Clypeosphaeriaceae was nested within the Hypocreales in MP1.
In the MP1 analysis, Lentomitella resides in a larger unsupported clade containing the Ophiostomatales and members of the Annulatascaceae (viz. A. triseptatus, A. austriacus, A. chesapeakensis and A. velatisporus) and a group of terrestrial perithecial ascomycetes (viz. C. groenendalensis and R. delicatula) tentatively placed in the Trichosphaeriaceae (Réblová and Seifert 2004). In the MP2 analysis Lentomitella together with Ceratostomella and Xylomelasma takes a basal position in relation to the Ophiostomatales, but on three separate branches within a single clade. While the anamorphs of the Annulatascaceae are generally unknown, the anamorphs of Cryptadelphia and Rhamphoria are hyphomycetous, dematiaceous with holoblastic denticulate conidiogenesis and have been linked to Brachysporium Sacc. and Phaeoisaria Höhn., respectively (Müller and Samuels 1982, Réblová and Winka 2001, Réblová and Seifert 2004). Lentomitella shares with Ceratostomella only dark, long-beaked perithecia with the top of the neck sulcate, and with Xylomelasma ellipsoidal ascospores arranged obliquely 1-seriately in the ascus; cylindrical, short-stipitate asci and cylindrical paraphyses; Xylomelasma differs from Lentomitella by ascospores with terminal pores, pigmented before discharge, asci floating freely within the centrum and characteristic ramification of ascogenous hyphae. Although the cultures of X. sordida and C. pyrenaica remained sterile, their grouping near Lentomitella, the Ophiostomatales and other taxa with holoblastic denticulate conidiogenesis in both phylogenies is suggestive of a similar conidiogenous pattern.
The anamorph of Lentomitella crinigera obtained in vitro is hyphomycetous, dematiaceous, with subhya-line to pale brown, denticulate conidiogenous cells. It is difficult to satisfactorily place the anamorph of L. crinigera in any of the known hyphomycetous genera. Several dematiaceous hyphomycetous genera such as Sporothrix Hektoen & Perkins, Rhinocladiella Nannf., Ramichloridium Stahel ex de Hoog or Tritirachium Limber produce similar holoblastic denticulate conidia, of which the latter has strongly geniculate proliferations, but the three latter have been linked to phylogenetically unrelated ascomycetes. Tritirachium has been linked to Trichosphaerella ceratophora (Höhn.) E. Müll. (Tritirachium oryzae [Vincens] de Hoog; Müller and Samuels 1982) of the Niessliaceae (Hypocreales). Rhinocladiella is known as an anamorph of Capronia Sacc. of the Herpotrichiellaceae (Chaetothyriales), while most of the Ramichloridium species (particularly the common R. schulzeri [Sacc.] de Hoog) are asexual and do not belong there. However phenotypically similar anamorphs are reported also for the perithecial ascomycete Calosphaeria fagi Samuels & Candoussau (ramichloridium-like and sporothrix-like synanamorphs, Samuels and Candoussau 1993), Rhamphoria (phaeoisaria-like anamorph, Müller and Samuels 1982) and some Ophiostoma species that also possess sporothrix-like anamorphs (Hoog 1974, Samuels and Müller 1979). The Lentomitella anamorph is most similar to the anamorphs of C. fagi, Rhamphoria or some Ophiostoma species; the two latter genera were close relatives to Lentomitella in both phylogenetic analyses; unfortunately, no molecular data are available for C. fagi. The Ophiostomatales are clearly distinguishable from Lentomitella by minute ascomata with filiform beaks, evanescent asci without any discharge mechanism and minute, reniform to crescent-shaped ascospores arranged in a fascicle. The teleomorphs of Lentomitella and C. fagi differ significantly in the morphology of ascospores (ellipsoidal vs allantoid), asci (cylindricalclavate, short-stipitate, sessile vs clavate, long-stipitate, floating free within the centrum), ascogenous system (ascogenous hyphae not branched, proliferating shortly through formation of croziers vs ascogenous hyphae branching in several levels), and habitat of perithecia (separately growing with separately projecting vertical necks vs forming circinate groups with long, cohering necks often perpendicular to the substratum). The perithecia of Rhamphoria are conical and superficial, asci are long cylindricalclavate and ascospores are irregularly ellipsoidal-fusiform, muriform, producing secondary minute ellipsoidal spores as outgrows from cells of primary ascospores within the ascus.
Ceratosphaeria lampadophora, the type species of the genus, is a lignicolous perithecial ascomycete possessing immersed nonstromatic perithecia with globose semi- or entirely immersed venter and elongate, separately protruding necks; long-cylindrical, short-stipitate asci arising from croziers, containing a distinct, refractive, nonamyloid apical annulus and eight hyaline, 5–8-septate, cylindrical to cylindrical-fusiform 2–3-seriate ascospores. According to the present phylogenies C. lampadophora resides on a strongly supported branch within the Magnaporthaceae (96/100 in MP1 and 100/100 in MP2) without close relationship with Lentomitella. The placement of the exclusively saprobic C. lampadophora in the Magnaporthaceae is of interest because the family was erected (Cannon 1994) for important plant pathogens specialized as necrotrophic parasites with a preference for Gramineae and Cyperaceae. The ordinal affiliation of the Magnaporthaceae has not yet been resolved within the Sordariomycetes (Cannon 1994, Kirk et al 2001, Eriksson et al 2003). In MP1 and MP2 phylogenies the Magnaporthaceae were shown as a well supported clade. The anamorphs that have been linked to the Magnaporthaceae form either pycnothyrial conidiomata producing phialidic conidia (Mycoleptodiscus Ostazeski, Pseudotracylla B. Sutton & Hodges), or they are hyphomycetous, with conidiogenesis being either holoblastic denticulate with rhexolytic seccession (Nakataea Hara, Pyricularia Sacc.), or holoblastic with schizolytic secession (Clasterosporium Schwein.), or hyphomycetous with phialidic conidiogenesis (Harpophora W. Gams).
The genus Harpophora was erected for phialophora-like fungi associated with Gaeumannomyces Arx & D.L. Olivier and Magnaporthe R.A. Krause & R.K. Webster species (Scott and Deacon 1983, Landschoot and Jackson 1989, Gams 2000) producing more or less pigmented phialides and cylindrical, hyaline, strongly curved conidia aggregated in slimy heads. Despite the similarity in the anamorphs, the teleomorphs of C. lampadophora and Magnaporthe are rather different. Magnaporthe species have erumpent perithecia with elongate necks and produce lightly pigmented, allantoid, 3-septate ascospores (Barr 1978). The two different types of conidiogenesis associated with Magnaporthe (Harpophora, Nakataea and Pyricularia) are suggestive of some heterogeneity in the generic concept. The anamorphs of Gaeumannomyces and Ophioceras of the Magnaporthaceae clade are also phialophora-like. Their overall teleomorphic similarity with C. lampadophora lies in dark, long-beaked perithecia with cylindrical to scolecosporous, septate, hyaline ascospores.
Trichosphaeria pilosa, representing the Tricho-sphaeriaceae in the phylogeny, takes a basal position in relation to all other unitunicate ascomycetes in the MP1 analysis (FIG. 1). The BLAST search of the T. pilosa LSU nrDNA sequence (Campbell and Shearer 2004) revealed that the closest relatives are members of the Dothideomycetes (viz. Raciborskiomyces Siemaszko [Capnodiales], Cladosporium Link, Dothidea Fr. or Mycosphaerella Johanson [Dothideales]). The delimitation of the Trichosphaeriales has not yet been resolved because of high degree of phenotype convergence and because of the absence of molecular data of T. pilosa, the representative taxon of the order. Trichosphaeria pilosa traditionally was accepted as a unitunicate perithecial ascomycete (von Arx and Müller 1954, Réblová et al 1999) but placement of the sequence in the tree is suggestive of its affinity to the ascolocular ascomycetes. The examination and comparison of the type and herbarium material, from which the sequence was derived, will be necessary to reveal the true systematic position of the fungus.
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ACKNOWLEDGMENTS |
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FOOTNOTES |
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1 E-mail: reblova{at}ibot.cas.cz
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LITERATURE CITED |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSTAXONOMYKEY TO THE GENERA...KEY TO THE SPECIES...KEY TO THE SPECIES...KEY TO THE SPECIES...DISCUSSIONLITERATURE CITED |
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