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Molecular systematics of Ceratostomella sensu lato and morphologically similar fungi

     Department of Plant Taxonomy and Biosystematics, Institute of Botany, Academy of Sciences, 252 43 Pru °honice, Czech Republic

	ABSTRACT

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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

	INTRODUCTION

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Ceratostomella Sacc. is a cosmopolitan genus of perithecial nonstromatic ascomycetes that colonize wood of angiosperms and gymnosperms in late stages of decay. Saccardo’s (1878) 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.

	MATERIALS AND METHODS

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Herbarium material and fungal strains.— – Dried herbarium specimens were rehydrated in water and studied in Melzer’s reagent or 90% lactic acid. All measurements were made in Melzer’s reagent. Means ± standard errors (SE) for asci and ascospores are given, based on 20–25 measurements. The length/width ratios (L/W) for asci are given. Images were captured in Melzer’s reagent with differential interference microscopy (DIC) and phase contrast (PC) and processed using Adobe Photoshop 6.0 CE.

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).

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).

View larger version (33K): [in this window] [in a new window]   FIG. 1. Phylogram of the most parsimonious tree from a heuristic analysis (MP1) of LSU rDNA sequences from 14 ascomycetous orders. Bootstrap values = 50% from 1000 replicates are included at the nodes. Thickened branches indicate posterior probabilities = 95%. Branch lengths are drawn to scale.

View larger version (30K): [in this window] [in a new window]   FIG. 2. Phylogram of one of the four equally parsimonious trees from a heuristic analysis (MP2) of SSU rDNA sequences from 15 ascomycetous orders. Bootstrap values = 50% from 1000 replicates are included at the nodes. Thickened branches indicate posterior probabilities = 95%. Branch lengths are drawn to scale.

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.

	RESULTS

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Insertions in the SSU rDNA.— – In the SSU six insertions from Xylomelasma sordida were identified (i.e. 64-nt, 369-nt, 492-nt, 71-nt, 135-nt and 64-nt) along with two insertions of Ceratostomella pyrenaica (i.e. 196-nt and 129-nt). The insertions are located at 330, 925, 1052, 1220, and 1384 5' flanking nucleotide positions of X. sordida, corresponding to the positions 330, 786, 904, 1072, 1184 and 1230 of E. coli SSU nrDNA (Gargas et al 1995). 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.

	TAXONOMY

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Taxonomic affinities of Ceratostomella rostrata.— – The lectotype species Ceratostomella rostrata was cited by Clements and Shear (1931). 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.

View larger version (36K): [in this window] [in a new window]   FIG. 22. Ceratostomella rostrata. A, B. Sphaeria rostrata . atra. A. Habit sketch of perithecia on wood. B. Perithecia with central elongate neck. C. Sphaeria rostrata ß. nigrofusca. Perithecia with central elongate neck. A–C copied from Tode (1791).

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.

	KEY TO THE GENERA CERATOSPHAERIA, CERATOSTOMELLA, ENDOXYLA, LENTOMITELLA AND XYLOMELASMA

TOP ABSTRACT INTRODUCTION MATERIALS AND METHODS RESULTS TAXONOMY KEY TO THE GENERA... KEY TO THE SPECIES... KEY TO THE SPECIES... KEY TO THE SPECIES... DISCUSSION LITERATURE CITED

 

1.Ascospores hyaline 2 1.Ascospores dull brown 3 2.Ascospores ellipsoid; phaeoisaria-like anamorph Lentomitella 2.Ascospores cylindrical to fusiform; harpophora-like anamorph Ceratosphaeria 3.Ascogenous hyphae ramifying; asci short-stipitate; ascomata without stromatic tissue 4 3.Ascogenous hyphae no ramifying; asci long-stipitate; ascomata surrounded with reduced stromatic tissue around neck Endoxyla 4.Ascospores suballantoid, globose, reniform to irregularly ellipsoidal; 2–3-seriate or in a fascicle in the ascus; paraphyses broad-celled; apical annulus indistinct Ceratostomella 4.Ascospores ellipsoid to oblong, obliquely uniseriate in the ascus; paraphyses cylindrical, slightly constricted at the distant septa; apical annulus distinct Xylomelasma

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.

	KEY TO THE SPECIES OF CERATOSTOMELLA

TOP ABSTRACT INTRODUCTION MATERIALS AND METHODS RESULTS TAXONOMY KEY TO THE GENERA... KEY TO THE SPECIES... KEY TO THE SPECIES... KEY TO THE SPECIES... DISCUSSION LITERATURE CITED

 

1.Asci shorter than 30 µm 2 1.Asci longer than 30 µm 3 2.Ascospores suballantoid to reniform, 4–5 x 2–3 µm C. cuspidata 2.Ascospores ellipsoidal to globose, 3–4 x (2.5–) 3–4 µm Ceratostomella sp. 3.Ascospores suballantoid, non-apiculate at the ends, 4.5–6 x 1.5–2 µm C. rostrata 3.Ascospores ellipsoidal to oblong, slightly apiculate at the ends, usually flattened at one side 4 4.Ascospores ellipsoidal, 6–7 x (3.5–)4–5 µm C. rhynchophora 4. Ascospores ellipsoidal to oblong, 7–9 x 3–4 µm C. pyrenaica

Ceratostomella cuspidata (Fr. : Fr.) Réblová, comb. nov. FIGS. 3–6, 23A, B.

View larger version (143K): [in this window] [in a new window]   FIGS. 3–21. Ceratostomella spp. 3–6. C. cuspidata. 3, 4. Asci. 5. Longitudinal section of perithecial wall. 6. Ascogenous hyphae and paraphyses. 7–11. C. pyrenaica. 7, 8. Asci. 9. Young ascus on an ascogenous hypha. 10. Ascogenous hyphae and paraphyses. 11. Longitudinal section of perithecial wall. 12–15. C. rhynchophora. 12, 13. Asci. 14. Ascogenous hyphae and paraphyses. 15. Longitudinal section of perithecial wall. 16–18. C. rostrata. 16. Ascogenous hyphae and paraphyses. 17. Asci. 18. Longitudinal section of perithecial wall. 19–21. Ceratostomella sp. 19, 20. Asci. 21. Longitudinal section of perithecial wall. DIC: 4, 5, 7, 8, 11–13, 15, 17–19, 21; PC: 3, 6, 9, 10, 14, 16, 20. FIGS. 3–6 from M.R. 1813; 7–11 from J.F. 04071; 12, 15 from PAD; 13, 14 from J.F. 02022; 16, 18 from DAOM 227786; 17 from M.R. 1887; 19–21 from M.R. 2592. Bars: 3, 4, 6–10, 12–14, 16, 17, 19–21 = 10 µm; 5, 11, 15, 18 = 50 µm.

View larger version (58K): [in this window] [in a new window]   FIG. 23. Ceratostomella spp. A, B. C. cuspidata. Asci with ascospores, ascogenous hyphae, paraphyses and habit sketches of perithecia. C, D. C. pyrenaica. Asci with ascospores, ascogenous hyphae, paraphyses and habit sketches of perithecia. E, F. C. rhynchophora. Asci with ascospores, ascogenous hyphae, paraphyses and habit sketches of perithecia. G, H. C. rostrata. Asci with ascospores, ascogenous hyphae, paraphyses and habit sketch of perithecia. A, B from M.R. 1813; C, D from PRM 902273; E, F from J.F. 02022; G, H from DAOM 227780. Bars: A, C, E, G = 10 µm; B, D, F, H = 500 µm.

= 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, Skainy Nature Reserve near Mikulice, 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, Medvdice 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

View larger version (11K): [in this window] [in a new window]   FIG. 24. Ceratosphaeria lampadophora. Harpophora-like anamorph in vitro. A. Conidiophores. B. Conidia. From CBS 117555 ex J.F. 01065 (PCA, 1 mo). Bar = 10 µm.

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 Spálenit near Stoec, 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.

	KEY TO THE SPECIES OF LENTOMITELLA

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1.Ascospores longitudinally striate. 1.Ascospores smooth L. tomentosa 2.Ascospores shorter than 10 µm, (5.5–)6–9 (–10) x 3–4.5(–5) µm L. cirrhosa 2.Ascospores longer than 10 µm, 10–14 x (4–) 5–6 µm L. crinigera

Lentomitella cirrhosa (Pers. : Fr.) Réblová, comb. nov. FIGS. 25–28, 44A–C

View larger version (139K): [in this window] [in a new window]   FIGS. 25–43. Lentomitella spp., Xylomelasma spp. 25–28. L. cirrhosa. 25, 26. Asci. 27. Ascospores. 28. Longitudinal section of perithecial wall. 29–33. L. crinigera. 29. Ascospores. 30. Asci. 31, 32. Conidiophores and conidia of phaeoisaria-like anamorph of L. crinigera. Conidiophores and conidia in vitro. 33. Longitudinal section of perithecial wall. 34, 35. L. tomentosa. 34. Ascus. 35. Longitudinal section of perithecial wall. 36–38. X. novaezelandiae. 36. Asci. 37, 38. Ascogenous hyphae and paraphyses. 39. Longitudinal section of perithecial wall. 40–43. X. sordida. 40–42. Asci. 43. Longitudinal section of perithecial wall. DIC: 25–31, 33–36, 39, 41–43; PC: 32, 37, 38, 40. FIGS. 25, 27 from PDD 81434; 26 from M.R. 2234; 29, 30, 33 from PDD 81435; 31, 32 from CBS 113655 ex PDD 81435 (PCA, 14 d); 34, 35 from PRM 902274; 36–39 from PDD 81433; 40–42 from PRM 902275; 43 from K.K. 236 (C). Bars: 25–27, 29–32, 34, 36–38, 40–42 = 10 µm 28, 33, 35, 39, 43 = 50 µm.

View larger version (59K): [in this window] [in a new window]   FIG. 44. Lentomitella spp. A-C. L. cirrhosa. Asci, ascogenous hyphae, paraphyses, transverse section of ascospore and habit sketches of perithecia. D-H. L. crinigera. Asci, ascogenous hyphae, paraphyses, transverse section of ascospore and habit sketches of perithecia. G, H. Conidiophores and conidia of the phaeoisaria-like anamorph of L. crinigera in vitro. A–C from PDD 81434. D–F from PDD 81435, G, H from CBS 113655 ex PDD 81435 (PCA, 14 d). Bars: A, B, D, E, G, H = 10 µm; C, F = 500 µm.

= 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 rid