Ceratostomella Molecular Systematics M98 2006

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Molecular systematics of Ceratostomella sensu lato and

morphologically similar fungi

Martina Reblova1

Department of Plant Taxonomy and Biosystematics,Institute of Botany, Academy of Sciences, 252 43Pruhonice, Czech Republic

Abstract: The systematic position and phylogeneticrelationships of Ceratostomella sensu lato and pheno-typically similar fungi using comparative morpholog-ical and culture studies and phylogenetic analyses ofthe nuclear large- and small-subunit ribosomal DNA

were explored. In the light of inferred phylogeniesand morphological data the genus Ceratostomella isredescribed, the generic concept is emended andfour species are accepted (viz. C. cuspidata, C.

pyrenaica, C. rhynchophora and C. rostrata). A newgenus Xylomelasmais introduced and delimited fromCeratostomella, with two new species described (viz. X.novaezelandiae and X. sordida). In culture species ofboth Ceratostomellaand Xylomelasmaproduced sterilemycelium. The genus Lentomitellawith a phaeoisaria-like anamorph formed in vitro is reinstated toencompass taxa formerly attributed to the broadlyperceived Ceratostomella with three accepted species(viz. L. cirrhosa, L. crinigera and L. tomentosa).Lentomitellaand Ceratostomellaare clearly distinguish-

able by the morphology of asci, ascospores andcentrum. Lentomitella is compared to phenotypicallysimilar Ceratosphaeria, which formed a harpophora-like anamorph in vitro. In the present phylogeniesCeratostomella, Ceratosphaeria, Lentomitella and Xylo-melasmaare shown as clearly separate genera belong-ing to three different groups of perithecial ascomy-cetes. Ceratostomella, Lentomitella and Xylomelasmareside within a large unsupported clade consisting ofmembers the Ophiostomatales, the freshwater Annu-latascaceae and a group of nonstromatic, terrestrialtaxa. Ceratosphaeria is well supported within theMagnaporthaceae. The systematic value of morpho-

logical characters of ascospores, paraphyses, asci,centrum and conidiogenesis in segregating taxa fromCeratostomella sensu lato and their relatives is dis-cussed.

Key words: Annulatascaceae, Ceratosphaeria, En-doxyla, Harpophora, Lentomitella, LSU and SSUrDNA, Ophiostomatales, Phaeoisaria, Sporothrix, sys-tematics, Wegelina, Xylomelasma

INTRODUCTION

Ceratostomella Sacc. is a cosmopolitan genus ofperithecial nonstromatic ascomycetes that colonize

wood of angiosperms and gymnosperms in late stagesof decay. Saccardos (1878) simple generic diagnosisof Ceratostomella reads: Perithecia et asci Ceratosto-matis. Sporidia continua, hyalina. Consequently thegenus became a large, heterogeneous entity thatincluded more than 100 taxa. Hohnel (1918) re-moved species with evanescent asci from Ceratosto-mella to Linostoma Hohn. Sydow and Sydow (1919)recognized Linostoma to be a later homonym fora genus of flowering plants (Thymeleaceae); they

proposed the new name Ophiostoma Syd. & P. Syd.,where several Ceratostomellaspecies were transferred.Several other species ofCeratostomellawith evanescentasci later were recognized congeneric with CeratocystisEllis & Halst. in a broad sense (e.g. Elliott 1925,Moreau 1952, Hunt 1956).

Although Ceratostomella should include fungi withhyaline ascospores, nonstromatic perithecia andpersistent asci, von Arx and Muller (1954) treatedCeratostomella 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 aroundthe base of a projecting neck. For the next 20 years,Ceratostomella (under Endoxyla) was included in theDiaporthaceae (von Arx 1952, von Arx and Muller1954, Muller 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 thesystematics of the genus and it remained heteroge-neous. Munk (1965) doubted his former placementof Endoxyla in the Rhamphorioideae of the Dia-porthaceae and noted that some Endoxyla speciesmay even represent calosphaeriaceous elements. Barr(1978, 1990) excluded Endoxyla from the Dia-

porthales because of the carbonaceous perithecialwall, asci that do not float freely within the centrum,presence of free-ended paraphyses, occurrence on

wood as a secondary saprobe, and she classified thegenus in the family Clypeosphaeriaceae. Untereiner(1993) distinguished Ceratostomella from Endoxylaand suggested affinity of the former genus with theLasiosphaeriaceae while Endoxyla was accepted in theClypeosphaeriaceae. Eriksson et al (2003) followedBarrs concept and accepted Ceratostomellaas a mem-ber in the Clypeosphaeriaceae of the Xylariales.

Accepted for publication 8 Aug 2005.1 E-mail: [email protected]

Mycologia, 98(1), 2006, pp. 6893.# 2006 by The Mycological Society of America, Lawrence, KS 66044-8897

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Ceratostomella sensu Untereiner (1993) was re-tained for species with dark-colored, immersed,nonstromatic perithecia with opaque, pseudoparen-chymatous walls; central, elongate, separately pro-truding necks; true paraphyses; unitunicate, sessileasci arising from a crozier system with a distinct,

refractive, nonamyloid apical annulus and ellipsoidal,hyaline ascospores. The phylogenetic relationships ofCeratostomella have remained unknown and noanamorph connections have been reported.

Hohnel (1905) introduced a monotypic genus,LentomitellaHohn., for Ceratostomella vestitaSacc. Hebased the segregation of Lentomitella from Ceratosto-mella on the ornamented (longitudinally striate)ascospore walls of C. vestita and suggested otherspecies ofCeratostomellato belong to LentomitaNiesslor Ceratosphaeria Niessl. Lentomita brevicollis Niesslthe type of the generic name Lentomita(Niessl 1876),

was transferred by Muller and von Arx (1962) to

Chaetosphaeria, as C. brevicollis (Niessl) E. Mull., andLentomita was relegated to synonymy of Chaeto-sphaeria. It is difficult to find a difference inteleomorph morphology between Ceratosphaeria andCeratostomellaexcept that the ascospores are cylindri-cal in the type species, Ceratosphaeria lampadophora(Berk. & Broome) Niessl, but can range fromellipsoidal to oblong to subcylindrical in other speciescurrently attributed to the genus (Tsuda and Ueyama1977). The insufficiently delineated generic conceptand life histories continued to hamper the distinctionbetween Ceratostomella and Ceratosphaeria and con-siderable cross generic and species synonymy can befound in the literature.

When it was proposed originally (Saccardo 1878)two species were included in CeratostomellaSacc (viz.C. vestita and C. cirrhosa [Pers. : Fr.] Sacc). At thesame time Saccardo listed Ceratostoma rostratum(Tode : Fr.) Fuckel under Ceratostomella withoutcomment. Later (Saccardo 1882) transferred Ceratos-toma rostratum to Ceratostomella. The examination oftype and other herbarium material of the threespecies cited in the protologue of Ceratostomella(Saccardo 1878), revealed the genus to be heteroge-neous from the beginning and the three cited species

represent two distinct genera.Ceratostomella rostrata (Tode : Fr.) Sacc., the lecto-

type species of Ceratostomella (Clements and Shear1931), possesses dark, long-beaked perithecia; palebrown, suballantoid to oblong ascospores; branchingascogenous hyphae; broad-celled paraphyses, andclavate asci tapering toward the base from thesporiferous portion with a shallow, indistinct, non-amyloid apical annulus. Ceratostomella rostratamatches the generic concept of Wegelina Berl., asemended by Barr (1998) and it is identical to Wegelina

polyporinaM.E. Barr. Wegelinawas erected by Berlese(1900) to segregate taxa from CalosphaeriaTul. & C.Tul. in the Calosphaeriales. Both genera accommo-date species with tiny nonstromatic perithecia; hya-line, aseptate or delicately septate, allantoid asco-spores; clavate, long-stipitate asci with thickened ascal

apex and persistent paraphyses. Wegelina comprisesspecies with superficial, separately growing peritheciaand separately protruding necks, while Calosphaeriaencompasses species forming perithecia in ellipsoidalto circinate groups or valsoid configurations withshort or elongate, radially converging beaks piercingthe periderm. Seven species have been assigned toWegelina; W. discretaBerl. was designated as lectotypeby Clements and Shear (1931). Wegelinawas reducedto synonymy of Scoptria Nitschke (Barr 1985) orCalosphaeria (Barr et al 1993), but later it wasreinstated (Barr 1998) and one new species and twonew combinations were introduced to the genus. The

characteristics of W. polyporina and W. subdenudata(Peck) M.E. Barr (Barr 1998), including relativelylarge perithecia that form a crust on the substratum,thick perithecial wall (50100 mm thick vs the wall ofWegelina s. str. 2040 mm), sulcate necks and sub-allantoid to ellipsoidal, pale brown ascospores beforedischarge, led Barr (1998) to extend Berlesesconcept 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 inculture. These fungi match the concept of Ceratosto-mella sensu Untereiner (1993) and should be accom-modated in Lentomitella (Hohnel 1905). Lentomitellais compared here with phenotypically similar speciesof Ceratosphaeria, based on C. lampadophora, whichformed a harpophora-like anamorph in vitro.

During a survey of perithecial lignicolous ascomy-cetes in temperate regions of both Northern andSouthern Hemispheres, several collections of twofungi phenotypically similar to Ceratostomella sensustricto were encountered. In culture, colonies pro-duced a sterile mycelium. The unknown fungi re-semble Ceratostomella sensu stricto in having dark,long-beaked perithecia, densely branched ascogenous

hyphae, clavate asci floating freely within the centrumat maturity and pigmented ascospores with terminalpores before discharge. The two taxa differ from thecore species of Ceratostomella in thickness of theperithecial wall, morphology of paraphyses, shape ofasci and apical annulus, and shape and arrangementof ascospores. One of these fungi has been describedas Endoxyla avocetta (Cooke & Ellis) Romero &Samuels (Romero and Samuels 1991) or later asPhaeognomoniella avocetta (Cooke & Ellis) Romero(Romero 1999). The examination of the holotype

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material of Sphaeria avocetta Cooke & Ellis andherbarium material, on which the cited combinations

were based, revealed two distinct perithecial ascomy-cete genera. Sphaeria avocetta is a species ofEndoxylaand it is identical to E. operculata(Fr. : Fr.) Sacc, whileone of the unknown fungi is conspecific with E.avocetta sensu Romero and Samuels (1991); itsrelationship and evolutionary history are examinedhere. 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 relation-ships of Ceratostomella, Ceratosphaeria, Lentomitellaand the unknown fungi, sequences of nuclear small-

and large-subunit ribosomal DNA (SSU and LSUnrDNA) of their type species were analyzed in twoindependent sequence datasets using maximumparsimony and Bayesian analyses.

MATERIALS AND METHODS

Herbarium material and fungal strains.Dried herbarium

specimens were rehydrated in water and studied in

Melzers reagent or 90% lactic acid. All measurements

were made in Melzers reagent. Means 6 standard

errors (SE) for asci and ascospores are given, based on

2025 measurements. The length/width ratios (L/W)

for asci are given. Images were captured in Melzersreagent with differential interference microscopy (DIC)

and phase contrast (PC) and processed using Adobe

Photoshop 6.0 CE.

Single- and mass-ascospore isolates were obtained fromfresh material with the aid of a single-spore isolator(Meopta, Czech Republic). Cultures were grown onpotato-carrot agar (PCA, Gams et al 1998). Colonycharacters were taken from cultures grown on PCA for21 d at room temperature (24 C) under UV light. Culturesare maintained at the Institute of Botany, Academy ofSciences in Pruhonice; Centraalbureau voor Schimmelcul-

tures, Utrecht (CBS); and Landcare Research, Auckland(ICMP). Type material is preserved in Landcare Research,

Auckland (PDD), National Museum, Prague (PRM), otherherbarium specimens with M.R. numbers are preserved atthe Institute of Botany, Pruhonice (PRA). The isolates usedin this study and their sources are listed (TABLE I).

DNA extraction, amplification and sequencing.Methodsfor DNA extraction, amplification and sequencing of

the LSU and SSU nrDNA of Ceratostomella pyrenaica

Reblova & Fournier and Xylomelasma sordidaReblova are

identical to those described by Mostert et al (2004). The

LSU and SSU nrDNA sequences of other taxa newlysequenced in this study were generated by methods

identical to those described by Reblova and Seifert

(2004).

Sequence data analyses.Phylogenetic relationships were

examined using 52 LSU nrDNA and 43 SSU nrDNAsequences from 15 or 14 different orders or families of

Sordariomycetes, respectively. Members of the Dothi-

deomycetes 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 Reblova & Four-nier, Lentomitella cirrhosa(Pers. : Fr.) Reblova, L. crinigera

(Cooke) Reblova and Xylomelasma sordida Reblova.

Homologous LSU and SSU nrDNA sequences from

other 88 taxa were retrieved from GenBank; accession

numbers are given (FIGS. 1, 2).

All sequences were aligned manually in BioEdit 5.0.9 Hall1999). Predicted models of the secondary structure of theLSU and SSU rRNA molecules of Saccharomyces cerevisiaeMeyen ex E.C. Hansen (Gutell 1993, Gutell et al 1993) wereused to improve the alignment. The models of thesecondary structure of the LSU and SSU rRNA were highlyconsistent in all taxa. The insertion positions in the SSUnrDNA sequences are named for the 59 flanking nucleotideand 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.

TABLE I. List of substrates, localities, sources, relevant sexual and asexual states and accession numbers of taxa sequenced inthis study

Teleomorph Anamorph Locality and substrate Sourcea GenBank LSU SSU

Ceratosphaerialampadophora

Harpophora sp. France, Pyrenees Atlantiques, wood ofPopulus tremula

CBS 117555 AY761084AY761088

Ceratostomella pyrenaica Unknown Czech Republic, Southern Moravia,wood of Acer campestre

CBS 117116 DQ076323DQ076324

Lentomitella cirrhosa Unknown New Zealand, wood ofPodocarpus totara ICMP 15131 AY761085AY761090

Lentomitella crinigera Phaeoisariasp. New Zealand, wood ofNothofagus sp. CBS 113655 AY761086

Xylomelasma sordida Unknown Czech Republic, Southern Moravia,wood of Alnus glutinosa

CBS 116000 AY761087AY761089

aCBS 5 Centraalbureau voor Schimmelcultures, Utrecht, the Netherlands; ICMP 5 International Collection of Micro-organisms, Landcare Research, Auckland, New Zealand.

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4.0b10 (Swofford 2002) using maximum parsimony; heu-ristic search with stepwise-addition option with 1000random taxon addition replicates and branch swapping

(tree bisection-recognition, TBR). All characters wereunordered and given equal weight during the analysis. Forall analyses ambiguously aligned positions were excluded.Gaps were treated as missing data. Branch support wasestimated by performing 1000 bootstrap replicates with a fullheuristic search consisting of 10 random-addition replicatesfor each bootstrap replicate.

The model of evolution that fits the data best wasdetermined by MODELTEST 3.5 (Posada and Crandall1998), showing that the best-fit model for the LSU sequencedataset was GTR+I+G model (Lanave et al 1984) withvariable base frequencies (freqA5 0.2221, freqC 5 0.2425,

freqG 5 0.3131, freqT 5 0.2223), different rates fortransitions and transversions R(a) [AC] 5 1.0263, R(b)[AG] 5 2.3442, R(c) [AT] 5 1.5809, R(d) [CG] 5

0.8755, R(e) [CT]5

6.7847, R(f) [GT]5

1.0000, anestimated proportion of invariable sites of 0.3137 anda gamma shape parameter for the rates of variable sites of0.4790. The best-fit model for the SSU sequence dataset wasdetermined as GTR+I+G model with variable base frequen-cies (freqA5 0.2420, freqC 5 0.2281, freqG 5 0.2840, freqT5 0.2459), different rates for transitions and transversionsR(a) [AC] 5 1.4452, R(b) [AG] 5 3.5075, R(c) [AT] 51.3198, R(d) [CG] 5 0.9220, R(e) [CT] 5 7.2559, R(f)[GT] 5 1.0000, an estimated proportion of invariable sitesof 0.5264 and a gamma shape parameter for the rates ofvariable sites of 0.6233.

FIG. 1. Phylogram of the most parsimonious tree from a heuristic analysis (MP1) of LSU rDNA sequences from 14ascomycetous orders. Bootstrap values 5 50% from 1000 replicates are included at the nodes. Thickened branches indicateposterior probabilities 5 95%. Branch lengths are drawn to scale.



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Bayesian analyses were performed with MrBayes v. 3.0b4

(Huelsenbeck and Ronquist 2001) to reconstruct phyloge-netic trees. The above models of evolution were implemen-ted and Bayesian analyses for both LSU and SSU sequencedatasets were conducted with the number of rate categoriesset to six, rates set to gamma and all remaining parametersestimated from the default prior probabilities. 5 000000generations were sampled every 100th generation resultingin 50 000 trees. The first 20 000 trees, which represented theburn-in phase of the analysis (number of trees to bediscarded after the likelihoods of the trees have convergedon a stable value), were discarded, and the remaining30 000 trees were used for calculating posterior probabilities

in the consensus tree. Posterior probabilities 5 95% are

indicated as thickened branches in the trees.

RESULTS

Insertions in the SSU rDNA.In the SSU sixinsertions from Xylomelasma sordidawere 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 59 flanking

nucleotide positions of X. sordida, corresponding

FIG. 2. Phylogram of one of the four equally parsimonious trees from a heuristic analysis (MP2) of SSU rDNA sequencesfrom 15 ascomycetous orders. Bootstrap values 5 50% from 1000 replicates are included at the nodes. Thickened branchesindicate posterior probabilities 5 95%. Branch lengths are drawn to scale.

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to the positions 330, 786, 904, 1072, 1184 and 1230of E. coli SSU nrDNA (Gargas et al 1995). Theinsertions of C. pyrenaica located at 393 and 142459 flanking nucleotide positions correspond to thepositions 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 thegroup I intron, the sixth insertion of X. sordidarepresents the spliceosomal intron with the 59- GTdonor and AG - 39 acceptor splice sites. Theseinsertions were excluded from all analyses.

Phylogenetic analyses of the LSU rDNA sequence data.A maximum parsimony analysis (MP1) was per-formed with 372 phylogenetically informativecharacters in an alignment including 1256 ntfrom 52 taxa. The first 75 positions in thealignment that are ambiguous were excluded.Two most parsimonious trees (MPT) were ob-

tained (tree length 2299, consistency index [CI] 50.375, retention index [RI] 5 0.553, homoplasyindex [HI] 5 0.625) (FIG. 1). The only source ofpolytomy was a clade containing the Annulatasca-ceae, the putative terrestrial Trichosphaeriaceaeand the fungi newly sequenced within this project.The two trees differed in position of C. pyrenaicaand X. sordida; C. pyrenaica was shown either asa sister to Cryptadelphia groenendalensis (Sacc. et al)Reblova & Seifert or to X. sordidawith no bootstrapsupport.

The consensus tree consisted of six major and wellsupported phylogenetic lineages of the Sordariomy-cetes (viz. a lineage [75% bootstrap support/100%posterior probability] consisting of four well sup-ported orders or groups), the Hypocreales (77/100)and the Microascales (98/100) clade, which is sister toa clade (87/100) of the Coronophorales (84/100)and Ascotaiwania/Conioscyphascus/Carpolignagroup(100/100); a lineage (64/100) consisting of sub-groupings 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 wellsupported Xylariales (89/100) and the Magnaportha-

ceae (83/100). Ceratosphaeria lampadophora nestedon a well supported branch with Ophioceras tenuis-porum Shearer et al (96/0) within the Magnaportha-ceae. Trichosphaeria pilosa (Pers. : Fr.) Fuckel, repre-senting the Trichosphaeriaceae, is shown basal to allother unitunicate ascomycetes in the tree.

In the sixth major lineage four separate clades werediscerned. In one of the two phylogenetic trees(FIG. 1), these clades were the Ophiostomatales(95/100), which were a sister to the larger un-supported clade consisting of X. sordida and C.

pyrenaica on one branch and the Annulatascaceae(75/100) subclade and the Cryptadelphia Reblova &Seifert, Lentomitella and Rhamphoria Niessl subcladeon the other branch. The two Lentomitella species(100%), Rhamphoria delicatula Niessl and C. groenen-dalensisrepresent a terrestrial sister to the freshwater

Annulatascaceae. The Annulatascaceae are represent-ed in the phylogeny by Annulusmagnus triseptatus J.Campbell & Shearer, Ascitendus austriacus (Reblovaet al) J. Campbell & Shearer, Annulatascus velatis-

porusK.D. Hyde and Aniptodera chesapeakensis Shear-er & M.A. Mill. Xylomelasma sordida and C. pyrenaicareside on a basal branch of this large clade with nobranch support.

In the consensus tree inferred from Bayesiananalysis the main topology of the six major lineages

was retained; the only lineage that did not receive anysupport was the one containing the Ophiostomatales,the Annulatascaceae, Cryptadelphia, Rhamphoria and

the newly sequenced taxa. Within this lineage threeclades were discerned (viz. a clade ofC. pyrenaica, X.sordida and R. delicatula [98 posterior probability],Cryptadelphiaand Lentomitellaclade [67] and a clade[97] containing the Annulatascaceae [100] as a sisterto the Ophiostomatales [100]).

Phylogenetic analysis of the SSU rDNA sequence data.A maximum parsimony analysis (MP2) was per-formed with 260 phylogenetically informativecharacters in an alignment including 1724 ntfrom 44 taxa. The first 27 positions in thealignment that are ambiguous were excluded.

Four MPT were obtained, one of which is shown(FIG. 2) (tree length 1092, CI 5 0.503, RI 5 0.632,HI 5 0.497). The trees differed in grouping of theSordariales, the Chaetosphaeriales and the Con-iochaetales within a single larger clade. Five majorlineages of the Sordariomycetes were discerned inthe MP2 analysis, (viz. a robust lineage [51/100]consisting of subgroupings of six orders orfamilies, the Coniochaetales [97/100], which area sister to the unsupported clade containing theCephalothecaceae on one branch and the Sordar-iales [97/100], the Boliniales, the Phyllachoralesand the Chaetosphaeriales [100/100] on the

other branch; a lineage of the Diaporthales[100/100] and the Calosphaeriales; a lineage ofthe Xylariales [80/100] and a larger lineage of theHypocreales and Microascales [89/100]; anda lineage consisting of members of the Magna-porthaceae [99/100], which are sister to a clade ofthe Ophiostomatales [100/100] with Ceratosto-mella, Lentomitella and Xylomelasma residing onseparate basal branches, respectively).

Anamorph-teleomorph connection.Single ascospores



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were isolated from L. cirrhosa, L. crinigera, C.lampadophora, C. pyrenaica and X. sordida. Thecultures derived from L. crinigera and C. lampado-

phora sporulated in 21 d on potato-carrot agar atroom temperature under UV light. Lentomitellacrinigera formed a phaeoisaria-like anamorph and

C. lampadophora formed a harpophora-like ana-morph in vitro. The cultures derived from C.

pyrenaica, L. cirrhosa and X. sordida never sporulat-ed, although the cultural characters of L. cirrhosaotherwise were identical with those of the culturederived from a morphologically similar L. crinigera.Because the cultures of L. crinigera and L. cirrhosahave identical appearance and the LSU sequencedata suggest their close relationship withina strongly supported monophyletic clade, it ismost likely explanation that a phaeoisaria-likeanamorph is also part of the life history of L.cirrhosa.

TAXONOMY

Taxonomic affinities of Ceratostomella rostrata.The lectotype species Ceratostomella rostrata wascited by Clements and Shear (1931). Although vonArx (1952) considered C. vestita as type species ofCeratostomella the lectotypification proposed byClements and Shear (1931) must be acceptedbecause even though it was not actually formallycombined in Ceratostomella at the time, it is clearthat Saccardo (1878) intended to include Ceratos-toma rostratum in Ceratostomella (P. Kirk, perscomm).

Tode (1791:14, Tab. IX, Fig. 79.) describedSphaeria rostrata with two varieties (viz. a. atra[perithecia black, tuberculate] and b. nigro-fusca[perithecia brown to black, glabrous]). No type orother authenticated material of S. rostrata is availablebecause none of Todes original collections arepreserved. The illustration accompanying the originaldescription is the only surviving original element(FIG. 22). The protologue does not include criticalfeatures and the illustrations of var. a and b show onlydark tuberculate or glabrous perithecia with long

central necks, respectively. Persoon (1801:58) citedthe name S. rostratavar. a. and referred to the Todesdescription. Fries (1823:473) based the description ofS. rostrata Tode : Fr. on material distributed in hisexsiccate collection Scleromyceti Sueciae 116 witha reference to Tode (1791) and Persoon (1801).Later Saccardo (1878) accepted S. rostrata inCeratostomella. The examination of the specimens ofScleromyceti Sueciae116 by the present author revealedthat S. rostrata is not congeneric with C. cirrhosa, C.vestita or other species currently attributed to

Ceratostomella sensu Untereiner (1993). Sphaeriarostrata differs from C. cirrhosa and C. vestita inhaving suballantoid to ellipsoid to reniform asco-spores that are pale brown before discharge. Asco-genous hyphae are branching, producing terminaland lateral, aseptate cells, from each of which one

ascus arises. The asci are clavate, short-stipitate,tapering toward the base from the sporiferousportion, floating freely within the centrum at matu-rity. The apical annulus is shallow, indistinct andnonamyloid. Paraphyses are broad-celled, constrictedat the septa and tapering.

The generic name Ceratostomella is reportedfrequently for species found on wood in late stagesof decay. The typification by Clements and Shear(1931) and the lectotype species C. rostrata areaccepted by the present author as valid. Because itnever will be possible to know what Tode actuallystudied, I accept Fries treatment of S. rostrata with

a reference to Tode (1791) and Persoon (1801) basedon material distributed in Scleromyceti Sueciae 116.Several exsiccate collections of S. rostrata underScleromyceti Sueciae 116 were studied (C, G, K, PRM,UPS) but only the G and PRM specimens containedmany intact mature perithecia in good condition. Toavoid confusion I herewith lectotypify the name S.rostratawith Todes original illustration and epitypify

with Fries material distributed in Scleromyceti Sueciae116 (PRM 666367!).

In this studyCeratostomellais redescribed based onthe lectotype species C. rostrata, and three otherspecies are accepted (viz. C. cuspidata, C. pyrenaicaand C. rhynchophora). Wegelina polyporinais relegatedto synonymy of C. rostrata. Other species currentlyattributed to Ceratostomella sensu lato that do notmatch the emended generic concept are underrevision by the author.

The generic concept of Wegelina based on thelectotype species W. discreta (Exsiccate. Mycotheca

Veneta 1450: ITALY. Belluno, on decorticated branchofAcer pseudoplatanus, autumn 1879, P.A. Saccardo(asCalosphaeria wahlenbergii; isotype of W. discreta, NY!)should follow Berleses treatment (Berlese 1900).

The distinction between the core species of

Ceratostomella and the phenotypically similar un-known fungi are based on these features: thickperithecial wall (ca. [60]80110 mm), clavate asci;broad-celled paraphyses; shallow, indistinct apicalannulus, and suballantoid to reniform to irregularlyellipsoidal ascospores 23-seriate or in a fascicle

within the ascus of Ceratostomella sensu stricto, vsthinner perithecial wall (ca. 5060[80] mm); cylin-drical asci; cylindrical, slightly constricted paraphyses,distinct apical annulus, and ellipsoidal ascospores,usually obliquely 1-seriate within the ascus of the

74 MYCOLOGIA


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unknown fungi. The LSU and SSU molecular data donot support closer relationship between Ceratosto-mella, represented by C. pyrenaica, and the unknownfungi. In the consensus tree of the two mostparsimonious trees C. pyrenaica resides either ona branch together with C. groenendalensis as sister to

Lentomitellaand Rhamphoriasubclade, which is sisterto the Annulatascaceae subclade, while the unknownfungus is shown on a basal branch of the whole clade,or both C. pyrenaica and the unknown fungus arenested on a basal branch of this large clade (FIG. 1).Based on the morphological characters and resultsfrom molecular analyses, both taxa are distinct fungiat the genus level. Because no ascomycete genuscould be found to match these characteristics, a newgenus, Xylomelasma, is introduced for the two un-known fungi and the new species X. novaezelandiaeand X. sordida are described. Xylomelasma sordidaincludes taxa formerly known as E. avocetta or P.avocetta (Romero and Samuels 1991, Romero 1999).

Taxonomic affinities of Ceratostomella cirrhosa andCeratostomella vestita.Sphaeria cirrhosa Pers. wasdescribed and illustrated by Persoon (1800: Tab.24, Fig. 3; 1801:59) for fungi with immersed tosemiimmersed, dark perithecia sparsely coveredwith short hairs and with sulcate necks. No typehas ever been designed for S. cirrhosa. Von Arx(1952) reported four collections of S. cirrhosafromthe Persoon herbarium (L) and said that only oneof them was a fungus that matched the descriptionofS. cirrhosa sensuPersoon (1801). That collection,according to von Arx (1952), contained fewascospores (810 3 3.5 mm). Unfortunately vonArx (1952) did not note which specimen con-tained ascospores; according to Untereiner (1993)and in my own experience, none of the fourcollections can be located in Persoons herbarium.

Von Arx (1952) considered C. cirrhosa and C.vestita conspecific and cited other 13 synonymousnames for C. cirrhosa. The examination of the typematerial of C. vestita by the present author revealeda fungus identical to C. cirrhosa sensuPersoon (1801)and von Arx (1952). Though the name S. cirrhosawas

sanctioned by Fries, no suitable herbarium materialmatching the concept of the species could be locatedin Fries herbarium. The only material ofS. cirrhosainFries herbarium was an exsiccate collection Scleromy-ceti Sueciae 346, which represents another fungus,Lentomitella crinigera(this study). Because no type orother authenticated material of S. cirrhosa could befound, I herewith lectotypify the name S. cirrhosawithPersoons original illustration of this species (Persoon1800: Tab 24, Fig. 3). The recent collection of thisfungus, from which the culture and LSU and SSU

sequences were derived, is designated here as anepitype (PDD 81434!).

Lentomitella (Hohnel 1905) was segregated fromCeratostomellafor the single species C. vestitaSacc. asL. vestita (Sacc.) Hohn. The name Lentomitella isavailable to accommodate taxa that do not match the

generic concept ofCeratostomellabased on C. rostrataand that possess hyaline, ellipsoidal ascospores;unitunicate, short-stipitate, sessile asci formed oncroziers and with a conspicuous, refractive, nonamy-loid apical ring, cylindrical paraphyses and thephaeoisaria-like anamorph. Therefore Lentomitella isreinstated and three species are accepted (viz. L.cirrhosa with L. vestita relegated to its synonymy, L.crinigera and L. tomentosa). Lentomitella cirrhosa andL. crinigera formed a strongly supported monophy-letic clade in the MP1 analysis. They were clearlyseparated from the phenotypically similar species ofCeratosphaeria, which resides on a well supportedbranch with Ophioceras Sacc. (96/100) within theMagnaporthaceae. The differences between the se-quence data, conidiogenesis (phaeoisaria-like ana-morph of Lentomitella vs harpophora-like anamorphof Ceratosphaeria) and ascospore shape warrant thedelimitation of the two holomorph genera, Lentomi-tella and Ceratosphaeria.

KEY TO THE GENERACERATOSPHAERIA, CERATOSTOMELLA,

ENDOXYLA, LENTOMITELLAAND XYLOMELASMA

1. Ascospores hyaline. . . . . . . . . . . . . . . . . . . . . . . . . 2

1. Ascospores dull brown. . . . . . . . . . . . . . . . . . . . . . 32. Ascospores ellipsoid; phaeoisaria-like ana-

morph . . . . . . . . . . . . . . . . . . . . . . . . . Lentomitella

2. Ascospores cylindrical to fusiform; harpo-phora-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, reniformto irregularly ellipsoidal; 23-seriate or in

a fascicle in the ascus; paraphyses broad-

celled; apical annulus indistinct . . . . . . Ceratostomella

4. Ascospores ellipsoid to oblong, obliquelyuniseriate in the ascus; paraphyses cylindri-

cal, slightly constricted at the distant septa;

apical annulus distinct. . . . . . . . . . . . . . Xylomelasma

Ceratostomella Sacc., Michelia 1:370. 1878, emend.ReblovaPerithecia solitary to densely crowded, nonstro-



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FIGS. 321. Ceratostomella spp. 36. C. cuspidata. 3, 4. Asci. 5. Longitudinal section of perithecial wall. 6. Ascogenoushyphae and paraphyses. 711. C. pyrenaica. 7, 8. Asci. 9. Young ascus on an ascogenous hypha. 10. Ascogenous hyphae andparaphyses. 11. Longitudinal section of perithecial wall. 1215. C. rhynchophora. 12, 13. Asci. 14. Ascogenous hyphae andparaphyses. 15. Longitudinal section of perithecial wall. 1618. C. rostrata. 16. Ascogenous hyphae and paraphyses. 17. Asci.18. Longitudinal section of perithecial wall. 1921. Ceratostomella sp. 19, 20. Asci. 21. Longitudinal section of perithecial wall.DIC: 4, 5, 7, 8, 1113, 15, 1719, 21; PC: 3, 6, 9, 10, 14, 16, 20. F IGS. 36 from M.R. 1813; 711 from J.F. 04071; 12, 15 fromPAD; 13, 14 from J.F. 02022; 16, 18 from DAOM 227786; 17 from M.R. 1887; 1921 from M.R. 2592. Bars: 3, 4, 610, 1214,16, 17, 1921 5 10 mm; 5, 11, 15, 18 5 50 mm.

76 MYCOLOGIA


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matic, 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 outerperithecial wall. Necks protruding separately, central,elongate, cylindrical, straight to slightly flexuous,

perpendicular, oblique to almost decumbent towardthe substratum, sulcate or glabrous, with periphysateostiole. Perithecial wallleathery to fragile, two-layered.Outer wall of brown, thick-walled cells, textura

prismaticato textura angularisto textura epidermoidea,and textura prismatica to porrecta in the neck; often

with a distinct, external crustose layer of heavilypigmented, dark brown cells with opaque walls. Innerlayer of thinner-walled, subhyaline to hyaline, elon-gated and compressed cells. Ascogenous hyphaebranching, discrete, with croziers, each producingseveral lateral and terminal, dehiscent cells sequen-tially and simultaneously, from each of which one

ascus arises as an outgrowth. Paraphyses abundant,unbranched, septate, hyaline, broad-celled and heavi-ly constricted at the septa, wider near the base,tapering toward the tip, apically free, longer thanthe asci, dissolving with age. Asci unitunicate, clavateto cylindrical-clavate, short-stipitate, truncate tobroadly rounded at the apex, tapering toward thebase 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 materialdeposited at the bottom part, visible after ascus

dehiscence from the ascogenous hypha. Ascosporesranging from suballantoid to irregularly ellipsoid toglobose to reniform, straight or curved, oftenflattened on one side, hyaline when young, palebrown before discharge, aseptate, smooth, sometimes

with terminal pores, arranged in a fascicle in theupper part of the ascus or 23-seriate in thesporiferous part.

Typus. Ceratostomella rostrata (Tode : Fr.) Sacc.,Syll. Fung. 1:409. 1882. (Lectotype designated byClements and Shear [1931].)

Anamorph. Hyphomycetous, dematiaceous; inculture only sterile mycelium seen.

KEY TO THE SPECIES OF CERATOSTOMELLA

1. Asci shorter than 30 mm . . . . . . . . . . . . . . . . . . . . 21. Asci longer than 30 mm . . . . . . . . . . . . . . . . . . . . . 32. Ascospores suballantoid to reniform, 45 3 2

3 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . C. cuspidata

2. Ascospores ellipsoidal to globose, 34 3 (2.5)34 mm . . . . . . . . . . . . . . . . . . . . . Ceratostomella sp.

3. Ascospores suballantoid, non-apiculate at the ends,4.56 3 1.52 mm . . . . . . . . . . . . . . . . . . C. rostrata

3. Ascospores ellipsoidal to oblong, slightly apiculateat the ends, usually flattened at one side . . . . . . . . 4

4. Ascospores ellipsoidal, 67 3 (3.5)45 mm . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . C. rhynchophora

4. Ascospores ellipsoidal to oblong, 79 3 34 mm . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. pyrenaica

Ceratostomella cuspidata (Fr. : Fr.) Reblova, comb.nov. FIGS. 36, 23A, B.

Basionym. Sphaeria cuspidata Fr. : Fr., Syst. Mycol.2:220. 1823.5 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 slightlyimmersed, subglobose to globose, 400500 mm high,380500 mm diam, glabrous, slightly tuberculate, dark

brown, surrounded by dense mycelium, especially atthe bottom; hyphae reddish brown, septate, 4.56 mm

wide; neck central, elongate, straight or slightlyflexuous, rounded at the top, sulcate with 45 ridges,(250)4001000 mm long, 110130 mm diam. Perithe-cial wall leathery to fragile, textura prismatica totextura angularis, 5580 mm thick, up to 90112 mmthick in places with tubercules, external crustoselayer 1017 mm thick. Some cells in the outer layer

with Munk pores. Paraphyses 611 mm wide nearthe base, tapering to 1.52.0 mm at the top. Ascicylindrical-clavate, 2130 3 (5)67 (mean 6 SE 524.7 6 0.4 3 6.3 6 0.1) mm, truncate at

the apex, tapering basally, short-stipitate, with anapical annulus, 33.5 mm diam, ca. 1 mm high.Ascosporessuballantoid to reniform, curved, flattenedon one side, 45 3 23 (mean6 SE 5 4.4 6 0.1 3 2.76 0.1) mm, pale brown, aseptate, smooth, in a fasciclein the upper part of the ascus or 23-seriate in theascus.

ISOTYPE. Fries Scleromyceti Sueciae 117, decayedwood (PRM 666203).

Additional specimens examined. CZECH REPUBLIC.SOUTHERN MORAVIA: Valtice, Randevous, decayed wood ofQuercussp., 4 Jun 1999, A. Vagner, M.R. 1813 (PRA). NEWZEALAND. WESTLAND: Mount Aspiring National Park,

Makarora Bush Walk, 500 m N of NP Headquaters inMakarora, decayed wood ofNothofagussp., 30 Mar 2005, M.Reblova (PRA).

Habitat. Saprobe on decayed wood of deciduoustrees.

Distribution. Czech Republic, New Zealand, Swe-den.

Commentary. Ceratostomella cuspidata is the mostsimilar to C. rostrata, which differs by largerperithecia and narrower suballantoid to somewhatoblong ascospores.



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Ceratostomella pyrenaica Reblova & Fournier, sp.nov. FIGS. 711, 23C, DPerithecia gregaria, immersa vel semiimmersa, globosa

vel subglobosa, brunnea, 400550 mm diam, 350550 mm

alta, ostiolo centrali elongato, cylindraceo, 6001130 mm

longo, 95110 mm diam. Paries perithecii 4565 mm crassus,

bistratosus. Paraphyses copiosae, hyalinae. Asci cylindraceo-

clavati, (30)3340 3 5.57 mm, breviter stipitati, 8-spori.

Ascosporae ellipsoideae vel oblongae, apiculatae, 79 3 3

4 mm, unicellulares, brunneae.


Perithecia gregarious, venter immersed to semiim-mersed, globose to subglobose, 350550 mm high,400550 mm diam, slightly roughened, dark brown,surrounded by sparse mycelium; hyphae dark brown,septate, flexuous, 45 mm wide; neck central, elon-gate, straight or slightly flexuous, rounded at theapex, sulcate with 45 ridges, 6001130 mm long, 95

110 mm diam. Perithecial wall leathery to fragile,textura prismatica, 4565 mm thick, external crustoselayer 1015 mm thick. Paraphyses1012 mm wide nearthe base, tapering to 22.5 mm at the top. Ascicylindrical-clavate, (30)33403 5.57 (mean6 SE 536.2 6 0.7 3 7.3 6 0.3) mm, truncate to broadlyrounded at the apex, short-stipitate, with an apicalannulus ca. 3 mm diam, 0.51 mm high. Ascosporesellipsoidal to oblong, slightly curved and apiculate atboth ends, flattened on one side, 79 3 34 (mean 6SE 5 8.2 6 0.1 3 3.16 0.1) mm, pale brown, aseptate,

smooth, often with two large drops, 23-seriate inthe ascus or in a fascicle in the upper part of theascus.

Characteristics in culture. Colonies reaching 1011 mm diam on PCA, grayish-brown. Aerial myceli-um well developed, denser in the center of colony;

hyphae subhyaline, 33.5 mm wide, septate,smooth. Substrate mycelium of brown, septatehyphae, 34 mm wide, smooth. Sporulation notseen. Margin gnawed. Reverse dark brown.

HOLOTYPE. FRANCE. PYRE NEES 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: Hodonn district, Skariny NatureReserve near Mikulcice, decayed wood of a trunk of Acercampestre, 24 Oct 2004, M. Reblova, M.R. 2912 (PRA,culture CBS 117116). FRANCE. PYRENEES ATLANTIQUES:

Rimont, Las Muros, Ruisseau de Peyran, decayed wood ofa trunk of A. glutinosa, 18 Jun 2001, J. Fournier J.F. 01114(PRA); ibid., Charente Maritime, La Villedieu, Foretd9Aulnay, decayed basidioma of Trametes gibbosaon a deadstump ofFagus sylvatica, 30 Apr 2004, J. Fournier J.F. 04071(PRA).

Etymology. Pyrenaica referring to the locality oforigin, Pyrenees Atlantiques.

Culture. CBS 117116 ex M.R. 2912.Habitat. Saprobe on decayed deciduous wood.

Distribution. Czech Republic, France.Commentary. Ceratostomella pyrenaica is character-

ized by pale brown, smooth, ellipsoidal to oblongascospores, slightly apiculate at the ends, a shapethat is well distinguishable within other Ceratosto-mella species.

Ceratostomella rhynchophora (De Not.) Reblova ,comb. nov. FIGS. 1215, 23EF

Basionym. Sordaria rhynchophora De Not., Comm.Crittog. Ital. 2:480. 1867.5 Ceratostoma rhynchophorum (De Not.) W. Kirschst.,

Krypt. Fl. Mark Brandenb. 7:249. 1911.5 Ceratostoma notarisii Sacc., Fung Ven. Ser. II, p. 308,

1874. (Illegit. name, Art. 52)

Anamorph. Unknown.

Perithecia solitary to gregarious, venter superficialto completely immersed, globose to subglobose, 480660 mm high, 500650 mm diam, slightly roughened,dark brown, surrounded by dense mycelium; hyphaedark brown, septate, branching, flexuous, 45 mm

wide; neck central, elongate, straight or slightlyflexuous, sulcate at the apex with 34 ridges, 4002000 mm long, 140190 mm diam. Perithecial wallleathery, textura prismatica to textura epidermoidea,75100 mm thick, external crustose layer 1217.5 mmthick. Paraphyses 811 mm wide near the base,

FIG. 22. Ceratostomella rostrata. A, B. Sphaeria rostrataa.atra. A. Habit sketch of perithecia on wood. B. Peritheciawith central elongate neck. C. Sphaeria rostrata b. nigro-fusca. Perithecia with central elongate neck. AC copiedfrom Tode (1791).

78 MYCOLOGIA


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tapering to 22.5 mm at the top. Asci cylindrical-clavate, (33)3544 3 78.5(10) (mean 6 SE 5 38.8

6 0.5 3 8.6 6 0.2) mm, truncate to broadly roundedat the apex, short-stipitate, with an apical annulus 2.53 mm diam, ca. 1 mm high. Ascospores ellipsoidal,sometimes flattened on one side, slightly apiculate atboth ends, 67 3 (3.5)45 (mean 6 SE 5 6.4 6 0.13 4.3 6 0.1) mm, midbrown, aseptate, smooth, witha minute pore at each end, 12-seriate in the ascus orin a fascicle in the ascus.

NEOTYPE. ITALY. Decayed wood of Prunusdomestica, P.A. Saccardo (PAD, as Ceratostoma notar-isii; neotype designated here).

Additional specimens examined. FRANCE. PYRE NE ESATLANTIQUES: Rimont, Las Muros, decayed wood of P.

domestica, 3 Feb 2002, J. Fournier J.F. 02022(PRA); ibid. 21Apr 2002, J. Fournier J.F. 02070 (PRA).

Habitat. Saprobe on decayed deciduous wood.

Distribution. France, Italy.

Commentary. Saccardo (1874:308) created anobligate synonym Ceratostoma notarisii based onSordaria rhynchophora with explanation that thename Ceratostoma rhynchophorum would have anidentical meaning as the already existing name

Ceratostoma rostratum.Two specimens of C. rhynchophora were found in

FIG. 23. Ceratostomella spp. A, B. C. cuspidata. Asci with ascospores, ascogenous hyphae, paraphyses and habit sketches ofperithecia. 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 withascospores, ascogenous hyphae, paraphyses and habit sketch of perithecia. A, B from M.R. 1813; C, D from PRM 902273; E, Ffrom J.F. 02022; G, H from DAOM 227780. Bars: A, C, E, G 5 10 mm; B, D, F, H 5 500 mm.



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PAD under the name Ceratostoma notarisii. Onecontained old and empty, minute, brown perithecia

with long, central filiform necks strongly similar tospecies of Melanospora. The second specimen wascollected by Saccardo and contained mature perithe-cia 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), thecollection from Saccardos herbarium labeled as C.notarisii is chosen as a neotype.

Ceratostomella rostrata (Tode : Fr.) Sacc., Syll. Fung.1:409. 1882. FIGS. 1618, 23G, H

Basionym. Sphaeria rostrata Tode, Fungi Mecklenb.Sel. 2:14. 1791 : Fries, Syst. Mycol. 2:473. 1823.5 Ceratostoma rostratum (Tode : Fr.) Fuckel, Jahrb.

Nassau. Ver. Naturk. 2324:127. 1870.

5 Ceratosphaeria rostrata (Fr.) Sacc., Syll. Fung. 2:227.1883. (as [Kickx] Sacc.)

5 Endoxyla rostrata (Tode : Fr.) Munk, Dansk Bot. Ark.17(1):196. 1957.

5 Wegelina polyporina M.E. Barr, Cryptogamie, Bryol.Lichenol. 19:170. 1998.

5 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, 700750 mm high, 650750 mm diam, tuberculate, dark brown to black,surrounded by sparse mycelium; hyphae dark brown,septate, flexuous, branching, 67 mm wide; neckcentral, elongate, straight or slightly flexuous, taper-ing toward the apex and narrowly rounded, sulcate

with 45 ridges, 8002500 mm long, 130170 mmdiam. Perithecial wall leathery to fragile, textura

prismatica to textura angularis, 6285 mm thick, 110125 mm thick in places with tubercules, externalcrustose layer 1225 mm thick. Some cells in theouter layer with Munk pores. Paraphyses 812 mm

wide near the base, tapering to 2.53 mm at the top.Asci cylindrical-clavate, (26)3039 3 56 (mean 6SE 5 33.4 6 0.4 3 5.9 6 0.1) mm, truncate to broadly

rounded at the apex, short-stipitate, with an apicalannulus 1.52 mm diam, ca. 1 mm high. Ascosporesallantoid to suballantoid, 4.56 3 1.52 (mean 6 SE5 5.3 6 0.1 3 1.9 6 0.1) mm, pale brown, aseptate,smooth-walled, usually in a fascicle in the upper partof the ascus or 23-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, decayedwood (PRM 666367, epitype designated here).

Additional specimens examined. Fries Scleromyceti Sueciae116, decayed wood (C, G, K 114933, UPS, as Sphaeriarostrata). CZECH REPUBLIC. SOUTHERN BOHEMIA: SumavaMountains National Park, Volary, Medvedice Nature Re-serve, decayed wood of Ulmus glabra, 26 Oct 2001, M.Reblova M.R. 1887, 1925 (PRA). FINLAND. Wehmersalmi,on hymenium of Fomes fomentarius, 12 Aug 1992, U.

Soderholm, L. Kasoner 2052 (DAOM 227785). GERMANY.Bredowen Forst, Orthavelland, in hymenium of the decayedbasidioma of F. fomentarius on Betula sp., 6 Oct 1916, W.Kirschstein (B700006552, holotype of Ceratostoma grumsi-nianum). POLAND. W. Siemaszko: Fungi Biaowiezensesexsiccati no. 158, Biaystok, Biaowieza, National ParkPuzcza Biaowieszka, in hymenium of the decayed basi-dioma of Fomes sp., Aug 1922, W. Siemaszko (PRM 655803,as Ceratostoma rhynchophorum). SWEDEN. Narke, Glan-shammar Parish, Skolv, on hymenium of F. fomentarius, 20May 1993, K.G. Nilsson(DAOM 227786, isotype ofWegelina

polyporina); Oland, Vickleby Parish, Nature Reserve Vick-leby angar 2 km NW of Vickleby, decayed bracket basidio-mata of F. fomentarius on Betula pendulosa, 4 Jun 2001, N.Lundqvist N.L. 21416.

Habitat. Saprobe on decayed deciduous woodand decayed aphyllophoraceous basidiomata.

Nomenclatural note. Saccardo (1883:227) pro-posed a new combination Ceratosphaeria rostrata(Kickx) Sacc. based on Sphaeria rostrata Kickx, Fl.Crypt. Flandres 1:338. 1867, where: Sphaeriarostrata Fr., Kx. Rech. Cent. III. p. 17; West. Herb.Crypt. fasc. 11, No. 552; Dryinosphaera rostrataDmtr.Desm. Pl. Crypt. fasc. 20, No. 973 is given.In Kickxs earlier work cited above (Kickx 1847: 17)Sphaeria rostrata Fr., Desm. Crypt., fasc. 20,

no. 973 is given. It is obvious that in bothpublications, Kickx referred to Fries concept ofS. rostrata. It is probable that Saccardo (1883)misinterpreted the name Sphaeria rostrata andincorrectly ascribed the name to J. Kickx.

Kirschstein (1911) proposed a new combination forCeratostoma rhynchophorum, however according to itsdescription and comparison to the material of C.rhynchophora and C. rostrata this species was mis-identified; it is C. rostrata. No herbarium material ofC. rhynchophorum could be found in Kirschsteinherbarium (B).

Commentary. Ceratostomella rostratais similar to C.

cuspidata in having tuberculate perithecia withsulcate neck. However size of perithecia, shapeand width of ascospores and length of asci clearlydistinguish the two taxa; in C. cuspidata asci areusually shorter than 30 mm, and ascospores aresuballantoid to reniform, 23 mm wide.

Untereiner (1993) cited two other taxa in thesynonymy of S. rostrata (viz. Endoxyla laevirostrisMunk [Denmark. Jylland, Staksrode near Juelsminde,distr. 24, rotten wood of a branch of Fagus sylvatica,12 Oct 1963, A. Munk, C!] and Ceratostomella

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ampullasca [Cooke] Sacc. [U.K. Shere, on rotten

wood ofQuercussp., Feb 1869, K 84424!]). Accordingto the study of their type material by the presentauthor these two taxa are neither conspecific nor arethey species of Ceratostomella. Additional culture andmolecular data are needed to confirm their systematicposition.

Ceratostomellasp. FIGS. 1921Anamorph. Unknown.Perithecia superficial to semiimmersed, sitting

densely on wood around the margin of a resupinatebasidiomycete, globose to subglobose 530570 mm

high, 500550 mm diam, tuberculate, dark brown,surrounded by dense mycelium; hyphae dark brown,septate, branching, flexuous, 2.53 mm diam; neckcentral, elongate, straight or slightly flexuous, sulcate

with four ridges, 6001000 mm long, 110130 mmdiam. Perithecial wall fragile, textura prismatica totextura angularis, 8792 mm thick, 97125 mm thick inplaces with tubercules, external crustose layer 1012 mm thick. Some cells in the outer layer withMunk pores. Paraphyses 68 mm wide near thebase, tapering to 2.53 mm at the top. Asciclavate, 25

32 3 56 (mean 6 SE 5 28.7 6 0.9 3 5.5 6 0.1) mm,truncate to broadly rounded at the apex, short-stipitate, with an apical annulus 22.5 mm diam, 0.51 mm tall. Ascospores ellipsoidal to globose, 34 3(2.5)34 (mean 6 SE 5 3.6 6 0.1 3 3.2 6 0.1) mm,pale brown to stramineous, aseptate, smooth-walled,

12-seriate in the ascus.Specimen examined. SWEDEN. SCANIA: Skurup, Svane-

holm, near the west shore of the lake, decayed wood ofFagus sylvatica in association with Steccherinum ochraceum,10 Nov 2000, S.-A. Hanson, M.R. 2592.

Habitat. Saprobe on decayed deciduous wood inassociation with old basidiomata.

Distribution. Sweden.Commentary. The shape of the ascospores is

distinctive and unique within the reported asco-spore variability of Ceratostomella. However, thematerial of this fungus is too poor to designatea holotype and additional material is needed for

a more detailed study to describe a new species.

Ceratosphaeria lampadophora (Berk. & Broome)Niessl, Verh. Naturf. Vereins Brunn 14:203. 1876.

FIG. 24Anamorph. Harpophora-like.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 thecenter. Aerial mycelium scarcely developed in thecenter of the colony and on the inoculating block;hyphae subhyaline to hyaline 23 mm wide, septate,smooth. Sporulation copious, widespread through-out the colony in 1 mo. Phialides or shortconidiophores arising on aerial hyphae, conidialheads slimy, inconspicuous, and transparent. Re-verse dark yellow. Conidiogenous cells phialidic,terminal or intercalary, hyaline to subhyaline,cylindrical, 1418 3 22.5 mm, tapering below thecollarette to 1.5 mm. Conidia cylindrical withcurvature, hyaline, narrowly rounded at both ends,911(12) 3 11.5 mm, aseptate, smooth.

Culture. CBS 117555 ex J.F. 01065.HOLOTYPE. UNITED KINGDOM. ENGLAND: Bath,

Combe Hay, decayed wood, Apr 1855, C.E. Broome(holotype ofSphaeria lampadophora, K).Additional specimens examined. CZECH REPUBLIC.

SOUTHERN BOHEMIA: Sumava Mountains National Park,Mount Spaleniste near Stozec, decayed wood ofF. sylvatica,12 Nov 2000, M. Reblova M.R. 1734 (PRA). FRANCE.PYRENEES ATLANTIQUES: Rimont, Ariege, Las Muros, ondecorticated wood of Populus tremula, 11 Apr 2001, J.

Fournier J.F. 01065(PRA; culture CBS 117555); ibid. 18 Jun2001, J.F. 01115 (PRA). SWITZERLAND. Grange canalnear Geneve, decayed wood of a trunk, 8 Feb 1859, J. Muller(Herb. Barbey-Boissier 605, G).

FIG. 24. Ceratosphaeria lampadophora. Harpophora-likeanamorph in vitro. A. Conidiophores. B. Conidia. FromCBS 117555 ex J.F. 01065 (PCA, 1 mo). Bar 5 10 mm.



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LentomitellaHohnel, Ann. Mycol. 3:552. 1905.Perithecia solitary to gregarious, nonstromatic,

venter superficial, semiimmersed or immersed, glo-bose, 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. Perithecialwall 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 tohyaline, elongated and compressed cells. Ascogenoushyphae not extensive, proliferating through croziers.Paraphysesabundant, cylindrical, tapering toward thetip, septate, slightly constricted at the septa ornonconstricted, hyaline, apically free, longer than

the asci, dissolving partly at maturity. Asciunitunicate,cylindrical-clavate, short-stipitate, sessile, truncate tobroadly rounded at the apex, with a distinct, re-fractive, nonamyloid apical annulus, 8-spored. Asco-spores ellipsoidal, hyaline, aseptate or several-septate,smooth or with few longitudinal ridges, obliquely 1-seriate or 12-seriate in the ascus.

Typus. Lentomitella vestita (Sacc.) Hohn.Anamorph. Phaeoisaria-like.

KEY TO THE SPECIES OF LENTOMITELLA

1. Ascospores longitudinally striate. . . . . . . . . . . . . . . 21. Ascospores smooth . . . . . . . . . . . . . . . . L. tomentosa2. A s cos por es s h or ter th an 1 0 mm, (5.5)69

(10) 3 34.5(5) mm . . . . . . . . . . . . . . . L. cirrhosa

2. Ascospores longer than 10 mm, 1014 3 (4)56 mm . . . . . . . . . . . . . . . . . . . . . . . . . . L. crinigera

Lentomitella cirrhosa(Pers. : Fr.) Reblova, comb. nov.FIGS. 2528, 44AC

Basionym. Sphaeria cirrhosa Pers., Syn. Meth. Fung.p. 59. 1801 : Fries, Syst. Mycol. 2:475. 1823.5 Ceratostoma cirrhosum(Pers. : Fr.) Fuckel, Jahr. Nassau.

Ver. Naturk. 2324:127. 1870.5 Ceratostomella cirrhosa (Pers.: Fr.) Sacc., Michelia

1:370. 1878.5 Endoxyla cirrhosa (Pers.: Fr.) Arx & E. Mull., Beitr.

Krypt-Fl. Schweiz 11(1):355. 1954.5 Sphaeria investita Schw., Trans. Amer. Phil. Soc. 2,

Vol. 4:216. 1834.5 Ceratostoma investitum (Schw.) Ellis & Everh., North

Amer. Pyrenom. p 193. 1892.5 Ceratostomella investita (Schw.) Starback, Bih. Kongl.

Svenska Vet.-Akad. Handl. 19(2):26. 1894.5 Ceratostomella vestita Sacc., Michelia 1:370. 1878.

5 Lentomitella vestita (Sacc.) Hohn., Ann. Mycol. 3:548.1905.

5 Cerastomis vestita (Sacc.) Clem., Gen. Fungi p. 259.1931.

5 Endoxyla vestita (Sacc.) Munk, Bot. Tidsskr. 61:64.1965.

5 Ceratostomella vestitaSacc. var. varvicensisGrove, J. Bot.23:131. 1885.

5 Eriosphaeria conoidea Feltg., Pilz Flora Luxemb. 3:282.1903.


Peritheciasolitary or gregarious, venter immersed tosuperficial, globose to subglobose, 300410 mm high,300400 mm diam, slightly roughened, dark brown,surrounded by sparse mycelium; hyphae brown, 4.55 mm wide; neck dark brown, central, elongate,cylindrical, straight to slightly flexuous, sulcate withfour ridges at the apex, 300900 mm long, 100

120 mm diam. Perithecial wall leathery, 3050 mmthick, textura prismatica to textura angularis; exter-nally with a row of heavily melanized cells with opaque

walls. Paraphyses68 mm wide near the base, taperingto 2.53 mm. Asci cylindrical-clavate, truncate tobroadly rounded at the apex, (42)5074 3 67(8)(mean 6 SE 5 62.3 6 1.1 3 7.1 6 0.1) mm, sessile,

with an apical annulus 2.53 mm diam, 11.5 m high,8-spored. Ascospores ellipsoidal, (5.5)69(10) 3 34.5(5) (mean 6 SE 5 7.9 6 0.2 3 3.8 6 0.1) mm,hyaline, 1(3)-septate, not constricted at the septa,

with few longitudinal ridges, obliquely 1-seriate or 12-seriate.

Characteristics in culture. Colonies 812 mm diamon PCA, dark brown, cocoa brown in the center.Aerial mycelium well developed, densest in thecenter of the colony, hyphae subhyaline to palebrown, 34 mm wide, septate, smooth. Substratemycelium of brown, septate hyphae, 3.54 mm wide,septate, smooth. Sporulation not observed. Reversedark brown.

LECTOTYPE illustration: Persoon, Icones et De-scriptiones Fungorum minus Cognitorum. Fasc. 2,Tab. 24, Fig. 3. 1800 (lectotype illustration designatedhere).

EPITYPE. NEW ZEALAND. WESTLAND: Hokitika,

40 km S of Greymouth, Mananui Point, Lake Mahi-napua, Swimmers Beach walks, decayed wood ofPodocarpus totara, 5 Mar 2003, M. Reblova M.R.2705 (PDD 81434, culture ICMP 15131; epitypedesignated here).

Additional specimens examined. CZECH REPUBLIC.SOUTHERN BOHEMIA: Sumava Mountains. National Park,Volary, Spaleniste Mount near Stozec, decayed wood ofFraxinus excelsior, 16 Aug 1999, M. Reblova M.R. 1545; ibid.,Povydr National Nature Reserve, Cenkova pila, decayedwood of a trunk ofU. glabra, 27 Aug 2000, M. Reblova M.R.

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1677; ibid., Zelezna Ruda, glacial cirque of the Certovojezero lake, decayed wood of Picea abies, 22 Oct 1996, M.Reblova M.R. 858/96; ibid., Prasily, Mount Zdanidla, 22 Aug2000, M. Reblova M.R. 2234; Novohradske hory Mountains,Nove Hrady, Zofnsky prales National Nature Reserve,decayed wood of F. sylvatica, 20 May 1997, M. ReblovaM.R. 917 (PRA). Moravia: Flora Bohem. et Morav. exsicc.Lfg. 14, No. 675, Hranice na Morave, Podhor, decayedwood, 24 Mar 1913, F. Petrak (PRM 195089). BELGIUM.Decayed wood of Pyrus communis, 25 May 1902, Feltgen(LUX 043455!, holotype of Eriosphaeria conoidea). DEN-MARK. Snarup Mose SW of Kvaerndrup, inner side ofthe bark of Betula sp., 1 Apr 1999, J. Vesterholt J.V. 99-015 (C 45296). SJLLAND: Charlottenlund skov, rottenwood, 10 Nov 1964, A. Munk (C); ibid., Dyrehaven, rottenwood, 26 Nov 1964, A. Munk (C); ibid., Dyrehaven, rottenwood of a stump of F. sylvatica, 2 Dec 1963, A. Munk (C).FRANCE. PYRENEES ATLANTIQUES: Rimont, Ariege, LasMuros, 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 ofa branch of Betula sp., 21 Apr 2001, S.-A. Hanson. USA.PENNSYLVANIA: Northhampton County, Bethlehem, L.D.Schweinitz, S.F. 1621.476 (PH 01016198!, holotype ofSphaeria investita).

Culture. ICMP 15131 ex-epitype PDD 81434.Habitat. Saprobe on decayed wood.

Distribution. Austria, Czech Republic, Belgium,Denmark, France, Italy, New Zealand, Sweden,USA.

Commentary. Lentomitella cirrhosa is a commonsaprobe growing on decayed wood. It is character-ized by ellipsoidal, longitudinally striate ascosporeswith one, rarely three septa and long-neckedperithecia with the sulcate top of the neck.

Hohnel (1906) in his revision of Feltgens originalherbarium material distinguished between C. cirrhosaand L. debaryanaand considered them separate taxa.Hohnel (1906) considered E. conoidea conspecific

with Lentomita debaryana (Auersw.) Hohn. (as deBaryana) and listed four other species closely relatedto L. debaryana. Von Arx (1952) accepted L.debaryana and the four species cited by Hohnel

(1906) as synonyms of C. cirrhosa and listed othernine synonymous names but without studying theirtype material. Re-examination of the respective typesby the present author revealed: C. vestita (5 L.cirrhosa, PAD!), C. investita (5 L. cirrhosa, PH01016198!), E. conoidea (5 L. cirrhosa, LUX043455!), and Ceratostomella similis W. Kirschst. (5C. ampullasca, B 700009209!). The examination ofC.ampullasca (K 84424!) revealed that it is not a speciesof Lentomitellaand requires additional study. Ceratos-tomella rostrata is not congeneric with L. cirrhosa and

it is redescribed under Ceratostomella sensu stricto inthis study. The type material of Ceratostomellacapilliformis Bomm. et al, Ceratostomella albocoronata(Ellis) Sacc. and Ceratostomella multirostrata (Fuckel)Sacc. contained only empty perithecia. The typespecimen of Ceratostomella subpilosa (Fuckel) Sacc.

(Fungi rhenani2251, G!) contained several peritheciawith only few ascospores attached to the outerperithecial wall; ascospores ellipsoidal, slightly taper-ing toward the ends, 89 3 45 mm, finely verrucu-lose. The fungus is not conspecific with L. cirrhosa,but it is likely a member of Lentomitellabased on theascospore characters; however, fresh material isneeded to clarify its systematics. The type materialof Ceratostomella unedonis H. Fabre, Ceratostomelladebaryana (Auersw.) Sacc. and Ceratostomella lejocarpaSacc., which were cited by von Arx (1952) among thesynonyms of C. cirrhosa, could not be located.

Lentomitella crinigera (Cooke) Reblova, comb. nov.FIGS. 2933, 44DH

Basionym. Sphaeria crinigera Cooke, Grevillea 1:156.1873.; Ceratosphaeria crinigera (Cooke) Sacc. Syll. Fung.

2:227. 1883.

Anamorph. Phaeoisaria-like.Peritheciasolitary or gregarious, venter immersed to

superficial, globose to subglobose, 510700 mm high,500600 mm diam, slightly roughened, dark brown,surrounded by sparse mycelium; hyphae brown, 3.55 mm wide; neck dark brown, central, elongate,

cylindrical, straight to slightly flexuous, sulcate with4 ridges, 3001000 mm long, 120140 mm diam.Perithecial wall leathery, 3750 mm thick, textura

prismatica to textura angularis; externally with a rowof heavily melanized cells with opaque walls. Paraph-

yses68 mm wide near the base, tapering to 2.53 mm.Asci cylindrical-clavate, truncate to broadly roundedat the apex, (65)7898(103) 3 79(11) (mean 6SE 5 86.7 6 0.8 3 8.3 6 0.1) mm, sessile, with anapical annulus 2.53.5 mm diam, ca. 11.5 mm high.Ascosporesellipsoidal, 10143 (4)56 (mean6 SE 512 6 0.1 3 4.6 6 0.1) mm, hyaline, 13-septate, notconstricted at the septa, with few longitudinal ridges,

obliquely 1-seriate or 12-seriate.Characteristics in culture. Colonies reaching 8

10 mm diam on PCA, dark brown, cocoa-brownin the center of the colony. Aerial mycelium welldeveloped, denser in the center of colony; hyphaesubhyaline, 34 mm wide, septate, smooth. Substratemycelium of brown, septate hyphae, 3.54 mmwide, smooth. Sporulation copious, widespreadthroughout the colony. Short conidiophores aris-ing from aerial hyphae, conidial heads slimy,inconspicuous, transparent. Reverse dark brown.



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FIGS. 2543. Lentomitellaspp., Xylomelasmaspp. 2528. L. cirrhosa. 25, 26. Asci. 27. Ascospores. 28. Longitudinal section ofperithecial wall. 2933. L. crinigera. 29. Ascospores. 30. Asci. 31, 32. Conidiophores and conidia of phaeoisaria-like anamorphofL. crinigera. Conidiophores and conidiain vitro. 33. Longitudinal section of perithecial wall. 34, 35. L. tomentosa. 34. Ascus.35. Longitudinal section of perithecial wall. 3638. X. novaezelandiae. 36. Asci. 37, 38. Ascogenous hyphae and paraphyses.

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Conidiophores macronematous, mononematous,arising terminally or laterally, hyaline, 2445 3 22.5 mm. Conidiogenous cells terminal or intercalary,hyaline, cylindrical, tapering toward the apex, 920(25) 3 22.5 mm, bearing 26 hyaline denticles0.51 mm wide, 0.51 mm long. Conidia ellipsoidalto globose, apiculate at the base, 46 3 22.5(mean 6 SE 5 5.26 0.23 2.26 0.3) mm, hyaline,aseptate, smooth.

HOLOTYPE. UNITED KINGDOM. ENGLAND: Nor-folk, Kings Lynn, decayed wood, C.B. Plowright(holotype ofSphaeria crinigera, K 84422!).

Additional specimens examined. CZECH REPUBLIC.

SOUTHERN BOHEMIA: Sumava Mountains National Park,Zelezna Ruda, glacial cirque of the Certovo jezero LakeNature Reserve, decayed wood of Abies alba, 28 Aug 1997,M. Reblova M.R. 997, M.R. 1034; ibid., 12 Aug 1999, M.Reblova M.R. 1544, M.R. 1585; ibid., 21 Aug 2000, M.Reblova M.R. 1672; ibid., glacial cirque of the Cerne jezeroLake Nature Reserve, decayed wood ofA. alba, 13 Aug 1999,M. Reblova M.R. 1526, M.R. 1546; ibid., Prasily, MountZdanidla, 24 Aug 2000, decayed wood of F. sylvatica, M.Reblova M.R. 1671 ; ibid., Boubnsky prales Nature Reserve,

decayed wood of P. abies, 17 Aug 1999, M. Reblova M.R.1611; ibid., Modrava, Ptac nadrz, decayed wood ofP. abies,14 Aug 1999, M. Reblova M.R. 1457; ibid., Modrava,Modravske slate, Pytlacky roh, decayed wood of a trunk of

FIG. 44. Lentomitella spp. A-C. L. cirrhosa. Asci, ascogenous hyphae, paraphyses, transverse section of ascospore and habitsketches 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. AC fromPDD 81434. DF from PDD 81435, G, H from CBS 113655 ex PDD 81435 (PCA, 14 d). Bars: A, B, D, E, G, H 5 10 mm; C, F 5500 mm.

r

39. Longitudinal section of perithecial wall. 4043. X. sordida. 4042. Asci. 43. Longitudinal section of perithecial wall. DIC:2531, 3336, 39, 4143; PC: 32, 37, 38, 40. FIGS. 25, 27 from PDD 81434; 26 from M.R. 2234; 29, 30, 33 from PDD 81435; 31, 32from CBS 113655 ex PDD 81435 (PCA, 14 d); 34, 35 from PRM 902274; 3639 from PDD 81433; 4042 from PRM 902275; 43from K.K. 236 (C). Bars: 2527, 2932, 34, 3638, 4042 5 10 mm 28, 33, 35, 39, 43 5 50 mm.



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P. abies, 14 Aug 1999, M. Reblova 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 ofGreymouth, Lake Stream track, 30 km SE of Reefton,decayed wood of a trunk of Nothofagus sp., 27 Feb 2003,M. Reblova M.R. 2659 (PDD 81435; cultures CBS 113655,ICMP 15124). SWEDEN. Fries Scleromyceti SueciaeNo. 346,decayed wood (UPS).

Cultures. CBS 113655, ICMP 15124, both isolatesex PDD 81435.

Habitat. Saprobe on decayed wood.Distribution. Czech Republic, Denmark, New

Zealand, Sweden, U.K.

Commentary. Lentomitella crinigerais characterizedby ellipsoidal, longitudinally striate, 13-septateascospores. The related L. cirrhosa is distinguishedfrom L. crinigera by shorter ascospores and asci andsmaller perithecia.

Lentomitella tomentosa Reblova & J. Fournier, sp.nov. FIGS. 34, 35, 45A, BPerithecia gregaria, immersa vel superficialia, globosa vel

subglobosa, brunnea, rubro tomentosa, 400500 mm alta,

350600 mm diam, ostiolo centrali elongato, cylindraceo,

200450 mm longo, 120180 mm diam. Paries perithecii 40

50 mm crassus, bistratosus. Paraphyses copiosae, hyalinae.

Asci cylindraceo-clavati, (75)8098 3 (9)1011 mm, bre-

viter stipitati, 8-spori. Ascosporae ellipsoideae, apiculatae,

1316(17) 3 (5)67 mm, continuae, hyalinae, laeves.

Anamorph. Unknown.Perithecia gregarious, venter immersed to entirely

superficial, globose to subglobose, 400500(600) mmhigh, 350600 mm diam, dark brown, collapsedlaterally on drying, surrounded by sparse hyphae 45 mm wide, growing out of the outer perithecial wall;neck dark brown, central, perpendicular to slightlydecumbent toward the substratum, elongate, cylin-drical, straight to slightly flexuous, sulcate with 34ridges, 200450 mm long, 120180 mm diam. Perithe-cial wall leathery, laterally 4050 mm thick, 3537 mmthick at the bottom, textura prismatica to texturaangularis; externally with a row of heavily melanizedcells with opaque walls. On the outer perithecial wall

a red waxy layer, 818 mm thick attached to thesurface of melanized cells; granules of red pigmentalso attached to the surface of the perithecial hairsand also staining the surrounding substratum. Outerperithecial wall of brown, thick-walled cells, innerlayer of thinner-walled, subhyaline to hyaline, elon-gated and compressed cells. Paraphyses 46 mm widenear the base, tapering to 33.5 mm, longer than theasci. Asci cylindrical-clavate, truncate to broadlyrounded at the apex, (75)8098 3 (9)1011 (mean6 SE 5 84.8 6 1.4 3 10.6 6 0.1) mm, sessile, with an

apical annulus 2.53.5 mm diam, ca. 1.5 mm high.Ascosporesellipsoidal, 1316(17)3 (5)67 (mean6SE 5 14.4 6 0.2 3 5.9 6 0.1) mm, hyaline, aseptate,smooth, obliquely 1-seriate or 12-seriate.

HOLOTYPE. FRANCE. Finistere, Plohars, Foret deCarnoet, Pont Douar, 40 m a.s.l., decorticated woodof F. sylvatica, 26 Oct 2002, J. Fournier J.F. 02196(PRM 902274).

FIG. 45. Lentomitella tomentosa. Asci, ascogenous hy-

phae, paraphyses and habit sketches of perithecia. FromPRM 902274. Bars: A5 10 mm; B 5 500 mm.

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Additional specimen examined. DENMARK. SJLAND :Dyrehaven, decayed wood of a stump, 3 Oct 1964, A. Munk(C); ibid., 20 Mar 1965, A. Munk (C); ibid., Silkeborg,Spring area near Alminds, decayed wood ofA. glutinosa, 4Sep 1953, A. Munk (C).

Etymology. Tomentosus (L), tomentose, refers tothe felted appearance of the surface of theperithecia.

Habitat. Saprobe on decayed wood.Distribution. Denmark, France.Commentary. Lentomitella tomentosa is character-

ized by aseptate, ellipsoidal, smooth ascosporesand perithecia with stout, sulcate, black neck incontrast to the finely tomentose venter with a redwaxy layer covering the surface of the wall andhairs; the red pigment dissolves in KOH and stainsthe nearby wood reddish.

Xylomelasma Reblova, 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 cylindra-

ceo-clavati, breviter stipitati, 8-spori. Ascosporae ellipsoideae

usque oblongae, apiculatae, unicellulares vel raro 1-septa-

tae, brunneae dum in asco.

Perithecia solitary to gregarious, nonstromatic,venter globose to subglobose, superficial, semiim-mersed or immersed, glabrous or roughened, darkbrown 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 towardthe substratum, sulcate or glabrous, ostiolum periph-

ysate. Perithecial wall leathery to fragile, two-layered.Outer wall of brown, thick-walled cells, textura

prismatica; externally with dark brown cells withopaque walls but without any distinct crustose layer.Inner layer of thinner-walled, subhyaline to hyaline,elongated and compressed cells. Ascogenous hyphaesparsely branching, discrete, each sequentially andsimultaneously producing several lateral and terminaldehiscent cells, each forming one ascus arises.

Paraphyses abundant, branching, septate, hyaline,cylindrical, slightly constricted, slightly tapering to-

ward the tip, apically free, longer than the asci. Asciunitunicate, cylindrical, short-stipitate, truncate tobroadly rounded at the apex, narrowly rounded at thebase, floating freely within the centrum at maturity,

with a distinct, refractive, nonamyloid apical annulus,8-spored. Ascus stipe usually containing a non-re-fractive material deposited at the bottom part, visibleafter ascal dehiscence from the ascogenous hyphae.Ascospores ellipsoidal to oblong, hyaline when young,

pale brown before discharge, aseptate or somedeveloping a median septum, smooth or finely

verruculose, sometimes with terminal pores, obliquely1-seriate in the ascus.

Typus. Xylomelasma sordida Reblova.Anamorph. Hyphomycetous, dematiaceous; in

culture only sterile mycelium seen.Etymology. Xylon (Gr) 5 wood; melas (Gr) black,

referring to the dark, opaque perithecia.

KEY TO THE SPECIES OF XYLOMELASMA

1. Ascospores 912 mm long. . . . . . . . . . . . . . X. sordida1. Ascospores 78 mm long . . . . . . . . X. novaezelandiae

Xylomelasma novaezelandiae Reblova, sp. nov.FIGS. 3638, 46A, B

Perithecia solitaria usque gregaria, immersa vel partim

erumpentia, globosa vel subglobosa, brunnea, 310340 mm

diam, 350400 mm alta, ostiolo centrali elongato, cylindra-

ceo, 95115 mm diam, 225375 mm longo. Paries perithecii

5262 mm crassus, bistratosus. Paraphyses copiosae, hyali-

nae. Asci cylindraceo-clavati, 5060(65) 3 78(9) mm,

breviter stipitati, 8-spori. Ascosporae ellipsoideae, 78 3

(3.5)45 mm, continuae, brunneae.

Anamorph. Unknown.Peritheciasolitary to gregarious, venter immersed or

partly erumpent, globose to subglobose, 350400 mmhigh, 310340 mm diam, smooth to slightly rough-ened, dark brown, surrounded by sparse mycelium;

hyphae brown, septate, sparsely branched, 34 mmwide; neck central, elongate, straight or slightlyflexuous, rounded at the top, not sulcate, 225375 mm long, 95115 mm diam. Perithecial wallleathery to fragile, textura prismatica, 5262 mmthick; external crustose layer 1114 mm thick. Paraph-

yses67 mm wide near the base, tapering to 2.53 mmat the top. Asci cylindrical-clavate, 5060(65) 3 78(9) (mean 6 SE 5 53.2 6 0.9 3 7.7 6 0.2) mm,truncate at the apex, short-stipitate, with an apicalannulus 2.53 mm diam, 11.5 mm high. Ascosporesellipsoidal, tapering toward both ends, narrowlyrounded and slightly apiculate, 78 3 (3.5)45

(mean 6 SE5

6.7 6 0.3 3 4.1 6 0.1) mm, palebrown, aseptate or rarely with one median septum atmaturity, smooth-walled, with two large drops; matureascospores become finely verruculose and darkbrown, with a minute pore at each end, 1-, rarely 12-seriate in the ascus.

HOLOTYPE. NEW ZEALAND. WESTLAND: Haast,300 km SW of Greymouth, Jackson River valley, trackto the Lake Ellery, S 43u52.95599, E 169u02.62119,rotten wood of a stump of Nothofagus sp., 10 Mar2003, M. Reblova M.R. 2787 (PDD 81433).



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Etymology. Novaezelandiae refers to country oforigin, New Zealand.

Habitat. Saprobe on decayed deciduous wood.Commentary. Attempts to germinate ascospores

of the cited specimen were unsuccessful.Although Xylomelasmaand Ceratostomellaare read-

ily distinguishable in the main characteristics of asci,apical annulus, ascospores and perithecial wall, X.novaezelandiae could be confused with C. rhyncho-

phora because of the similar ascospore shape; but

the ascospores are regularly ellipsoidal, not flattenedon a side and shorter in X. novaezelandiae. Ceratos-tomella rhynchophora differs from X. novaezelandiaealso in having shorter asci, ascospores 23-seriatelyarranged in the asci and larger perithecia witha thicker wall.

Xylomelasma sordida Reblova, sp. nov.FIGS. 4043, 46C, D

Perithecia solitaria usque gregaria, superficialia usque

immersa, globosa vel subglobosa, brunnea, 490550 mm

diam, 450600 mm alta, ostiolo centrali elongato, cylindra-

ceo, 120150 mm diam, 9501500 mm longo. Paries perithe-

cii 5562(80) mm crassus, bistratosus. Paraphyses copiosae,

hyalinae. Asci cylindraceo-clavati, (58)6076(81) 3 7

10(13) mm, breviter stipitati, 8-spori. Ascosporae ellipsoi-

deae, apiculatae, 912 3 46 mm, unicellulares, brunneae.


Perithecia solitary to gregarious, venter superficialto immersed, globose to subglobose, 490550 mm

diam, 450600 mm high, slightly roughened, darkbrown, surrounded by sparse mycelium; hyphae darkreddish brown, septate, slightly flexuous, 56 mm

wide; neck central, elongate, slightly roughened,sometimes unevenly thickened, rounded at the top,sulcate with four ridges, 9501500 mm long, 120150 mm diam. Perithecial wall leathery to fragile,textura prismatica to textura angularis, 5562(80)mm thick, external crustose layer 712 mm thick.Some cells in the outer layer with Munk pores.Paraphyses 610 mm wide near the base, tapering to 2

FIG. 46. Xylomelasmaspp. A, B. X. novaezelandiae. Asci, ascogenous hyphae, paraphyses and habit sketches of perithecia. C,

D. Xylomelasma sordida. Asci, ascogenous hyphae, paraphyses and habit sketch of perithecia. A, B from PDD 81433; C, D fromPRM 902275. Scale bars: A, C 5 10 mm ; B, D 5 500 mm.

88 MYCOLOGIA


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3 mm at the top. Asci cylindrical-clavate, (58)6076(81) 3 710(13) (mean6 SE 5 72.4 6 1.1 3 8.2 60.1) mm, truncate to broadly rounded at the apex,short-stipitate, with an apical annulus 22.5 mm diam,ca. 1 mm high. Ascospores ellipsoidal, sometimesslightly apiculate at both ends, 912 3 46 (mean 6

SE 5 10 6 0.1 3 5 6 0.1) mm, pale brown, aseptate,smooth, with a minute germ pore at each end, with1(2) oil drops, 12-seriate in the ascus.

Characteristics in culture. Colonieson PCA reaching1214 mm diam in 14 d, dark olivaceous brown,grayish in the center. Aerial mycelium scarcelydeveloped, densest in the center of colony; hyphaesubhyaline to pale brown, 33.5 mm wide, septate,smooth. Substrate mycelium of dark brown, septatehyphae, 34 mm. Sporulation absent. Margin dis-crete, somewhat gnawed. Reverse dark brown.

HOLOTYPE: FRANCE. PYRENEES ATLANTIQUES:Ariege, Lescure, Bois du Pas du Baup, 500 m a.s.l.,

rotten wood of A. glutinosa, 24 Feb 2004, J. Fournier04020 (PRM 902275, culture CBS 116000).

Additional specimens examined. ARGENTINA. BUENOSAIRES PROVINCE: Pdo. San Pedro, Gdor. Castro, Eucalyptusviminalis, May 1982, A. Romero 9, 19 (BAFC 32028, BAFC32030, BAFC 32031); ibid., Aug 1982, A. Romero 10 (BAFC51316); 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:Breclav district, obora Soutok near Lanzhot, on decayeddeciduous wood. 23 Oct 2004, M. Reblova M.R. 2904, 2911(PRA). DENMARK. SJLLA ND: 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.Bukk National Park in NE Hungary, decayed wood of F.sylvatica, Oct 2000, K. Kovacs K.K. 215, K.K. 236 (C).

Culture. CBS 116000 ex-type PRM 902275.

Etymology. Sordidus (L), dull, refers to the dullbrown ascospores.

Habitat. Saprobe on decayed wood.

Distribution. Argentina, Denmark, France, Hun-gary.

Commentary. Romero and Samuels (1991:43, Pl.3mp, Pl. 6gj) reported a fungus identical with X.sordidafrom Argentina, but they considered it to be

conspecific with Sphaeria avocettaCooke & Ellis andproposed a new combination Endoxyla avocetta.They based their identification on the comparisonwith isotype material of S. avocetta (BPI, asCeratostoma avocettum). Untereiner (1993) conclud-ed that E. avocetta could not be classified in

Endoxyla based on the description and illustrationby Romero and Samuels (1991) and suggested itsplacement in the Diaporthales. Romero (1999)introduced a new genus Phaeognomoniella Romerofor S. avocetta in this sense.

The revision of the holotype material ofS. avocetta(FIGS. 4750; USA. NEW JERSEY: Newfield, undeter-mined decayed wood, Jan 1876, J.B. Ellis, K 114931!,as Ceratostoma avocettum) by the present authorrevealed a fungus that possesses immersed, dark,conical, glabrous, monostichous perithecia withshort, stout, cohering necks protruding just abovethe substrate surface, with remnants of stromatictissue surrounding the projecting necks; paraphysesfiliform, hyaline, septate, apically free, ca. 2 mm wide,longer than the asci; asci unitunicate, cylindrical,long-stipitate, rounded apically, floating free withinthe centrum, 6688 mm long in pars sporifera, 67 mm

wide, stipe 3552 mm long, dissolving at maturity; witha nonamyloid indistinct apical annulus; ascosporesellipsoidal to subcylindrical, 1113 3 (3)3.54 mm,aseptate, dull brown before discharge, with a minutepore at one end and slightly tapering toward this end,smooth, 1-seriate in the ascus. These characteristics of

S. avocettaas seen on the holotype material are typicalof members of Endoxyla (Untereiner 1993). Sphaeriaavocetta is conspecific with Endoxyla operculata andshould be relegated to its synonymy. Phaeognomo-niella, although based on misidentified material,becomes a generic synonym of Endoxyla. For fullsynonymy of Endoxyla and E. operculata I refer toUntereiner (1993). The sizes of ascospores and asci in

pars sporifera of X. sordida and S. avocetta are similarand might be confusing but in all other respects bothfungi represent two distinct genera.

FIGS. 4750. Endoxyla parallela. 47, 48. Asci. 49. Asco-spores. 50. Longitudinal section of perithecial wall. DIC: 47,49, 50; PC: 48. From K 114931 (holotype of Sphaeriaavocetta). Scale bars: 4749 5 10 mm; 50 5 50 mm.



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DISCUSSION

Although morphological similarities among ascomy-cetes can lead to incorrect phylogenies, in this casethe DNA sequence data join phenotype characters toindicate thatCeratostomellaand Xylomelasmaare closerelatives. Xylomelasma resembles Ceratostomella inhaving pigmented ascospores before discharge, ascifloating within the centrum at maturity and similarbranching pattern of ascogenous hyphae, but itdiffers from it in the thickness of the perithecial wall;paraphyses that are cylindrical and only slightlyconstricted at the septa, and cylindrical and generallylarger asci with a distinct, refractive apical annulus,and ellipsoidal ascospores that are obliquely uniseri-ate within the ascus.

Among comparatively well known ascomycetes,species of Ceratostomella and Xylomelasma can becompared with some phenotypically similar taxa of

the Ceratostomataceae (e.g. Melanospora Corda orArxiomyces P. Cannon & D. Hawksw.) especiallybecause of the long-beaked perithecia and dullbrown, aseptate ascospores with terminal pores.However, based on morphology of the asci, ascogen-ous system and interthecial filaments, none of thespecies attributed to Ceratostomellaor Xylomelasmaaremelanosporaceous. According to molecular analysesof ribosomal DNA and RPB2, the phylogenetic affinityof Melanosporaand other members of the Ceratosto-mataceae lie with the Hypocreales (Zhang and Black-

well 2002). In all recent analyses Melanospora takesthis position. Ceratostomella bears also some resem-

blance to Cannonia Taylor & K.D. Hyde with thesingle species C. australis (Speg.) J. Taylor & K.D.Hyde in dark perithecia with immersed subglobose

venter, separately projecting necks and brown ellip-soidal

Documents

Ceratostomella Molecular Systematics M98 2006