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Multigene phylogeny of the Coronophorales: Morphology and new species in the order

Mycologia

January 2010
102(1):185-210

DOI:10.3852/09-043

Source
PubMed

Authors:

S.M. Huhndorf

Field Museum of Natural History

The phylogenetic relationships within Coronophorales have been debated because of uncertainty over the taxonomic usefulness of characteristics such as quellkörper, number of ascospores per ascus, presence of ascospore appendages, presence of subiculum and ascomatal vestiture. The phylogenetic relationships are examined with DNA sequence data from three nuclear genes targeting 69 taxa and 130 new sequences representing collections from Africa and the Americas. Analyses recovered monophyletic Bertiaceae, Chaetosphaerellaceae and Scortechiniaceae and a paraphyletic Nitschkiaceae. A single collection of Coronophora gregaria is included and Coronophoraceae is accepted. Bertiaceae is expanded to include Gaillardiella, and Thaxteria is synonymized with Bertia with a new combination, B. didyma. Three new species of Bertia are described: B. ngongensis from Kenya, B. orbis from Kenya and Costa Rica and B. triseptata from Ecuador and Puerto Rico. Bertia gigantospora is transferred from Nitschkia. Scortechiniaceae is confirmed for the quellkörper-bearing taxa including monotypic Biciliospora, Coronophorella, Neofracchiaea, Scortechiniella and Scortechiniellopsis. Tympanopsis is reinstated for T. confertula and T. uniseriata, while Scortechinia is more narrowly circumscribed to include S. acanthostroma and the new species, S. diminuspora from Ecuador. Cryptosphaerella is accepted in Scortechiniaceae including six new species from Kenya and Costa Rica, C. celata, C. costaricensis, C. cylindriformis, C. elliptica, C. globosa and C. malindensis. Spinulosphaeria is accepted in Coronophorales with uncertain family placement. The number of ascospores in the ascus is not phylogenetically useful in distinguishing genera within the order. The quellkörper continues to be an important character in defining the Scortechiniaceae, while taxa within the group show a mixture of morphological characteristics of varying phylogenetic importance. The presence of smooth versus spinulose subiculum aids in separating Tympanopsis and Scortechinia, and erumpent ascomata distinguish Cryptosphaerella species. Taxa within the Bertiaceae vary along the lines of robust, tuberculate, collapsing ascomata and large, hyaline to pigmented, septate ascospores.

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Short title: Coronophorales phylogeny

Multigene phylogeny of the Coronophorales: morphology and new species in the order

George K. Mugambi1

Botany Department, Field Museum of Natural History, Chicago, Illinois 60605–2496,

Department of Biological Sciences, University of Illinois at Chicago, 845 W. Taylor Street (MC

066), Chicago, IL 60607, and National Museums of Kenya, Botany Department, P.O. Box 45166,

00100, Nairobi, Kenya

Sabine M. Huhndorf

Botany Department, Field Museum of Natural History, Chicago, Illinois 60605–2496

Abstract: The phylogenetic relationships within Coronophorales have been debated because of

uncertainty over the taxonomic usefulness of characterisitics such as quellkörper, number of

ascospores per ascus, presence of ascospore appendages, presence of subiculum and ascomatal

vestiture. The phylogenetic relationships are examined with DNA sequence data from three

nuclear genes targeting 69 taxa and 130 new sequences representing collections from Africa and

the Americas. Analyses recovered monophyletic Bertiaceae, Chaetosphaerellaceae and

Scortechiniaceae and a paraphyletic Nitschkiaceae. A single collection of Coronophora gregaria

is included and Coronophoraceae is accepted. Bertiaceae is expanded to include Gaillardiella,

and Thaxteria is synonymized with Bertia with a new combination, B. didyma. Three new

species of Bertia are described: B. ngongensis from Kenya, B. orbis from Kenya and Costa Rica

and B. triseptata from Ecuador and Puerto Rico. Bertia gigantospora is transferred from

Nitschkia. Scortechiniaceae is confirmed for the quellkörper-bearing taxa including monotypic

Biciliospora, Coronophorella, Neofracchiaea, Scortechiniella and Scortechiniellopsis.

Tympanopsis is reinstated for T. confertula and T. uniseriate, while Scortechinia is more

In Press at Mycologia, published on July 17, 2009 as doi:10.3852/09-043

narrowly circumscribed to include S. acanthostroma and the new species, S. diminuspora from

Ecuador. Cryptosphaerella is accepted in Scortechiniaceae including six new species from

Kenya and Costa Rica, C. celata, C. costaricensis, C. cylindriformis, C. elliptica, C. globosa and

C. malindensis. Spinulosphaeria is accepted in Coronophorales with uncertain family placement.

The number of ascospores in the ascus is not phylogenetically useful in distinguishing genera

within the order. The quellkörper continues to be an important character in defining the

Scortechiniaceae, while taxa within the group show a mixture of morphological characteristics of

varying phylogenetic importance. The presence of smooth versus spinulose subiculum aids in

separating Tympanopsis and Scortechinia, and erumpent ascomata distinguish Cryptosphaerella

species. Taxa within the Bertiaceae vary along the lines of robust, tuberculate, collapsing

ascomata and large, hyaline to pigmented, septate ascospores.

Key words: Ascomycota, Coronophorales, LSU rDNA, phylogeny, quellkörper, rpb2,

tef1

INTRODUCTION

Members of the Coronophorales are common wood-inhabiting Ascomycete fungi with a

worldwide distribution. Coronophorales is characterized by taxa with mostly superficial

ascomata, sometimes with an extensive hyphal subiculum or well developed basal stroma that

often becomes cupulate or collapsed, and in some cases an ostiolar opening is either indistinct or

lacking. In many taxa a quellkörper is present in the centrum, and this structure is unique to this

group of ascomycetes. The quellkörper is a subcylindrical to inverted-conical structure attached

to the roof of the centrum, composed of thick-walled, concentrically arranged, hyaline cells that

functions by enlarging and forcibly breaking open the ascoma and allowing ascospore release.

Other important characteristics of the group include Munk pores that are found in the ascomatal

wall cells, thin-walled asci with a long stipe and in most cases without an apical ring and

ascospore numbers often greater than eight.

Coronophorales are found on woody substrates in many parts of the world. Collections from

temperate European localities form the majority of the early names in the group with some

additional collections made from North America (von Höhnel 1907; Fitzpatrick 1923; Nannfeldt

1975a, b). Over the years new species and collections have been added from tropical locations

such as French Guiana, India, Singapore and western Africa (Sivanesan 1974, Nannfeldt 1975b).

Until now Coronophorales have not been seriously studied in eastern Africa, although a few

species were described from Congo and Kenya (von Arx and Müller 1954, Nannfeldt 1975b).

Members of the Coronophorales have been treated in their own order (Nannfeldt 1932,

Müller and von Arx 1973, Subramanian and Sekar 1990) or in order Sordariales (Nannfeldt

1975a, b; Barr 1990; Hawksworth et al 1995) under either one or two families, Coronophoraceae

and Nitschkiaceae (Nannfeldt 1932; Nannfeldt 1975a, b) (TABLE I). Huhndorf et al (2004) used

molecular data to show that the group is not related to Sordariales and to demonstrate the

monophyly of the Coronophorales in subclass Hypocreomycetidae. They suggested the

separation of the taxa into four families (family Coronophoraceae was not sampled). However

their molecular study was based on limited taxon sampling and therefore did not adequately

address various conflicting taxonomic treatments previously proposed for taxa within the group

by different taxonomists.

Narrow generic circumscriptions for taxa in the Coronophorales were adopted by Müller and

von Arx (1973) (TABLE I). Nannfeldt (1975a, b) and Subramanian and Sekar (1990) presented

detailed historical perspectives of the Coronophorales and its components. The phylogenetic

relationships within group have long been debated because of uncertainty over the taxonomic

usefulness of distinguishing characterisitics, such as quellkörper, number of ascospores per

ascus, presence of ascospore appendages, presence of subiculum and ascomatal vestiture.

Nannfeldt (1975b) accepted five genera in the Nitschkiaceae and also published detailed

morphology, taxonomy and nomenclature of the many taxa. He considered most of the

morphological characters of value in the delimitation of the taxa at specific levels only and

therefore adopted broad generic concepts, which resulted in drastic reduction in the number of

genera recognized (TABLE I). Von Arx (1981) followed the generic circumscriptions of Müller

and von Arx (1973) (TABLE I), adding a few more taxa including Lasiobertia, which since has

been shown to belong outside the group (Huhndorf et al 2004). Subramanian and Sekar (1990)

also adopted narrow generic circumscriptions, differing from Nannfeldt (1975b) and chose to

recognize two families for the taxa (TABLE I).

Huhndorf et al (2004) in molecular studies of Coronophorales used large subunit ribosomal

nuclear DNA sequence data that resulted in their segregation of four families in the group,

including two new families, Chaetosphaerellaceae for Chaetosphaerella phaeostroma (Durieu &

Mont.) E. Müll. & C. Booth and Crassochaeta nigrita (Berk. & Broome) Réblová and

Scortechiniaceae for Scortechinia conferta (Schwein.) Subram. & Sekar, Neofracchiaea callista

(Berk. & M.A. Curtis) Teng and Euacanthe foveolata (Berk. & M.A. Curtis ex Berk.) Subram. &

Sekar. In their analyses Bertiaceae was represented by Bertia tropicalis Huhndorf, A.N. Mill. &

F.A. Fern. and B. moriformis (Tode) de Not. and a paraphyletic Nitschkiaceae by Nitschkia

meniscoidea Huhndorf, A.N. Mill. & F.A. Fern., N. grevillei (Rehm) Nannf., Acanthonitschkea

argentinensis Speg. and Fracchiaea broomeiana (Berk.) Petch.

Due to the limited number of taxa used by Huhndorf et al (2004), taxonomic relationships

and circumscriptions of some groups in the order remained uncertain. As a result in this follow-

up study we not only target broad taxon sampling but also take into account the great

morphological variability that is observed within currently recognized groups. Consequently taxa

in 19 morphologically recognized genera were targeted for phylogenetic analyses with DNA

partial sequence data of large subunit ribosomal nuclear DNA (LSU rDNA), ribosomal

polymerase II subunit 2 (rpb2) and translation elongation factor 1 alpha (tef1). The questions we

considered in this study include (i) what are the generic and familial circumscriptions within the

order, (ii) what are the taxonomic placements of previously unsequenced taxa and (iii) which are

the robust morphological characters that can be used in delineating taxa within Coronophorales?

MATERIALS AND METHODS

Taxon sampling and morphological analyses.—The taxa used in this study are listed (TABLE II) together with their

geographical locality, associated information and GenBank accession numbers. Ascomata were mounted in water

and replaced with lactophenol containing azure A. A minimum of 30 asci, ascospores were measured with Scion

Image (www.scioncorp.com) and measurements were made and images were captured of material in both mounting

fluids. Ascomata were sectioned at 5 μm for light microscopy following Huhndorf (1991) or were freehand

sectioned. Images were captured with photomacrography, bright field (BF), phase contrast (PH) and differential

interference microscopy (DIC), and photographic plates were produced following Huhndorf and Fernández (1998).

Representative species covering five families and 19 genera within the Coronophorales were included in the

analyses to determine the phylogenetic position of taxa currently accepted in the order. A total of 78 taxa were

included in the analyses; 69 were newly sequenced for this study (TABLE II). The full datasets contained respectively

72, 35 and 44 taxa for the LSU rDNA, rpb2, and tef1 genes, while a reduced dataset of 40 taxa from which at least

two genes were available was used in the combined analyses allowing for some missing data. All voucher specimens

are deposited in F and the Kenyan specimens also are deposited in EA.

DNA extraction, PCR amplification, sequencing and sequnce alignment.—Total DNA was extracted with the

Dneasy Plant Mini Kit (QIAGEN, Hilden, Germany) following the instructions of the manufacturer. Phylogenetic

analyses were conducted with partial sequences of three genes: translation elongation factor 1 alpha (tef1), nuclear

ribosomal large subunit (LSU rDNA) and RNA polymerase II subunit (rpb2). Nuclear LSU was amplified with

primers LROR, LR6 and LR3 (Vilgalys and Hester 1990); tef1 was amplified with primers EF1-526F, EF1-983F,

EF1-1567R, Ef-df and EF-gr obtained from the Assembling the Fungal Tree of Life Website

(http://ocid.nacse.org/research/aftol/primers.php), while rpb2 was amplified with fRPB2-5F and fRPB2-7cR (Liu et

al 1999).

Polymerase chain reaction (PCR) was carried out with this protocol: Final volume of the PCR reaction was 25

µL and contained 2.5 μL buffer, 2.5 μL dNTP mix, 1 μL each primer (10 μm), 5 μL BSA, 1.5 μL taq, 2 μL genomic

DNA extract and 9.5 μL deionized water. The reaction was allowed to run for 34 cycles. The annealing temperature

was set at 50 C for LSU rDNA and at 58 C for both tef1 and rpb2 reducing by 1 C each cycle for total of eight cycles

and then set at 50 C for the remaining cycles. The fragments were sequenced with the Big Dye Terminator reaction

kit (ABI PRISM, Applied Biosystems, Foster City, California). Sequencing was performed with the same set of

primers as PCR. The other sequences were obtained from GenBank. Sequences were aligned with multiple sequence

alignment program, MUSCLE® (Edger 2004) and further edited manually. Ambiguously aligned regions were

removed from the data matrices. Sequence data were deposited in GenBank and alignments in TreeBASE

(XXXXX).

Phylogenetic analyses.—Phylogenetic analyses were carried out with maximum parsimony (MP), maximum

likelihood (ML) and Bayesian analyses for single-gene data matrices and on combined gene dataset. Maximum

parsimony (MP) analyses were carried out with PAUP* 4.0b10 (Swofford 2002). Heuristic searches were performed

with random addition and TBR branch swapping procedures, characters unordered and MULTREES option in effect.

Bootstraping (Felsenstein 1985) was performed with 1000 replicates to estimate branch support.

Modeltest 3.7 (Posada and Crandall 1998) was used to determine the best-fit model of evolution for Bayesian

and maximum likelihood analyses. Bayesian analyses employing Markov chain Monte Carlo (MCMC) were carried

out with MrBayes 3.1 (Huelsenbeck and Ronquist 2001). Four MCMC chains were run simultaneously for 1–5

million generations for the single gene and combined datasets with trees sampled every 100th generation. Maximum

likelihood (ML) analyses were carried out for each of the four datasets employing the best-fit model of evolution

obtained from Modeltest 3.7 (Posada and Crandall 1998) with Garli 0.951 (Zwickl 2006).

RESULTS

Sequence alignment and phylogenetic analyses.—Sequence alignment resulted in LSU rDNA

with 1139 characters. A total of 34 were ambiguously aligned and hence excluded from

analyses, 586 were constant, 91 parsimony uninformative ,while 428 were parsimony

informative. Tef1 had 824 characters. Three ambiguously aligned were excluded, 391 were

constant, 134 were parsimony uninformative and 296 were parsimony informative. In the case of

rpb2 1041 characters, 443 were constant, 72 were parsimony uninformative while 526 were

parsimony informative. The combined dataset had a total of 2954 characters, of which 1580 were

constant, 274 were parsimony uninformative, while 1100 were parsimony informative. The best-

fit model of evolution for LSU rDNA and combined datasets was GTR + I + G (Rodriguez et al

1990), while TRN + I + G (Tamura and Nei 1993) was the preferred model of evolution for tef1

and rpb2 datasets. Maximum parsimony analyses of LSU rDNA and combined datasets

generated respectively 12 and five most parsimonious trees, while tef1 and rpb2 generated two

trees each. Because there were no conflicts in the phylogenies produced by different analyses

(MP, ML and Bayesian), we present only maximum likelihood trees for LSU, tef1, rpb2 and

combined data analyses (FIGS. 1–4).

Taxon relationships.—The multiple gene genealogies all contain monophyletic clades

representing Bertiaceae, Chaetosphaerellaceae, Nitschkiaceae and Scortechiniaceae.

Coronophoraceae is represented in all trees except in the LSU rDNA by Coronophora gregaria

(Lib.) Fuckel. There were some differences among generic placements within individual families

resulting from different analyses, but most of these relationships were not supported. Bertiaceae

included species of Bertia, Thaxteria and Gaillardiella and formed a strongly supported

monophyletic group with high bootstrap (BS) and Bayesian posterior probabilities (PP) (FIGS. 1,

2). In our analyses Thaxteria didyma (Speg.) Sacc. is nested within Bertia (FIG. 1).

Chaetosphaerellaceae, composed of Chaetosphaerella and Crassochaeta, forms a strongly

supported clade (FIGS. 1–4). Genus Coronophora, represented by the type species C. gregaria,

does not group within Bertiaceae, Chaetosphaerellaceae, Nitschkiaceae or Scortechiniaceae

(FIGS. 2, 4). Spinulosphaeria whose placement in the Coronophorales has been debated is also

nested within the order but of unclear position (FIG. 1). Taxa in Nitschkia group together with

Acanthonitschkea in all analyses, but this relationship is weakly supported in tef1 and rpb2 gene

analyses (FIGS. 1–4). Genus Fracchiaea as currently circumscribed is polyphyletic (FIG. 2).

Scortechiniaceae form a strongly supported monophyletic group comprising Scortechinia,

Scortechiniella, Scortechiniellopsis, Biciliospora, Neofracchiaea, Euacanthe, Neotrotteria,

Cryptosphaerella, Tympanopsis and an unnamed Coronophorales (FIGS. 1–4). Scortechinia is

recognized for two species as is Tympanopsis (FIGS. 1–4). Cryptosphaerella species form a

monophyletic group within Scortechiniaceae in the tef1 and combined datasets with significant

PP (FIGS. 2, 4) but are polyphyletic in the LSU rDNA and rpb2 datasets (FIGS. 1, 3).

Coronophorella, Scortechiniella and Biciliospora form a supported clade with high PP in the

combined data analysis (FIG. 4).

Taxonomy.—Currently recognized families in the Coronophorales, Bertiaceae, Scortechiniaceae

and Chaetosphaerellaceae were found to be monophyletic. The concept of Bertiaceae is

expanded to include Gaillardiella (FIGS. 1, 2, 4), and Thaxteria is found to be nested within

Bertia. Scortechiniaceae contains the quellkörper-bearing taxa including six new species of

Cryptosphaerella (FIGS. 1–4). Family Coronophoraceae is recognized for taxa in Coronophora

(FIGS. 2–4). Nitschkiaceae is composed of Nitschkia, Fracchiaea and Acanthonitschkea, taxa

that group together in supported clade (FIGS. 1, 4). However we were not able to obtain type of

Nitschkia and therefore the circumscription of the genus and family remains equivocal until it is

included in analyses.

Bertia didyma (Speg.) Mugambi & Huhndorf, comb. nov. FIG. 5a–e

MycoBank MB 513306

≡ Bizzozeria didyma Speg., Boletín de la Academia Nacional de Ciencias de Córdoba 11(4):519. 1889. Basionym.

≡ Thaxteria didyma (Speg.) Sacc., Sylloge fung. 9:687. 1891.

Specimens examined: COSTA RICA. Guanacaste: Cantón Cañas, Sector Hacienda Montezuma, 715 m, 7 Mar

2000, on 10 cm diam branch, F.A. Fernández, G.M.Mueller, B. Strack, J.P. Schmit, L. Umaña, SMH4256 (F);

Puntarenas: Los Alturas Forest Biosphere Research Park, adjacent to Parque International la Amistad, 8°96′16″N;

82°82′00″7W, 21 Jan 2007, on decayed wood, J.L. Crane et al, CR-206 (ILLS). ECUADOR. Orellana Province:

Yasuni Biosphere Reserve, Tiputini Biological Station, Danta trail, 26 Mar 2002, on 25 cm log, F.A. Fernández, A.N.

Miller, SMH4719 (F).

Descriptions of this species are given by Fitzpatrick (1923) and Subramanian and Sekar

(1990). Both of these authors based their information on the type specimen, but they differ

somewhat in their interpretation of ascomatal and ascospore morphology. Fitzpatrick (1923)

describes the ascomata as not prominently tuberculate and the surface as brownish and fibrilose

with age, and Subramanian and Sekar (1990) describe ascomata as having a warty surface. Our

collections best match the description given by Fitzpatrick and also match the drawing given by

Müller and Booth (1972), which they also base on the type specimen.

Bertia gigantospora (Nannf.) Mugambi & Huhndorf, comb. nov. FIG. 5f–i

MycoBank MB 513307

≡ Nitschkia gigantospora Nannf., Svensk bot. Tidskr. 69(3):313. 1975. Basionym.

≡ Calyculosphaeria macrospora Fitzp., Mycologia 15:53. 1923. [non Nitschkia macrospora Teng, 1934; non Bertia

macrospora Sacc., 1878]

Specimens examined: KENYA. Rift Valley Province: Kajiando District, Ngong Hills Forest, near aviation

equipment, 1°24.682S, 36°38.182E, elev 2391 m, 16 Jun 2005, on branch on the ground, GKM1136 (EA).

Bertia gigantospora is represented in our trees by a single collection from Kenya that yielded

only rpb2 and tef1 sequences (FIGS. 2, 3). The collection matches the description of

Calyculosphaeria macrospora, which was transferred to Nitchkia by Nannfeldt (1972) and given

a new name because the macrospora epithet was already occupied in the genus. The same is true

within Bertia so the transfer must be made with the name given by Nannfeldt.

Bertia ngongensis Mugambi & Huhndorf, sp. nov. FIG. 5j–m

MYCOBANK MB 513308

Ascomata erumpentia, separata ad denso aggregata, tuberculata, turbinatascens in statu humectato, collabens in statu

sicco, 850–900 µm alta, 730–900 µm diam. Paries ascomatis cellulis pseudoparenchymatis. Munk pori preasens.

Asci clavati, curvi, octospori, 115–150 × 16–18 µm, partibus sporiferis 60–85 µm longitudine, stipitibus 45–75 µm

longitudine. Ascosporae cylindricae, geniculatae, hyalinae, 1-septatae, 26–33 × 7–8 µm.

Etymology: Refers to the collection locality.

Ascomata erumpent through cracks in the bark, solitary or in small to large groups, turbinate

when fresh, collabent when dry, tuberculate, with large sterile base, 730–900 µm wide, 850–900

µm high including the sterile base. Ascomatal wall composed of outer layer of brown thick-

walled pseudoparenchymatous cells and inner layer of flattened hyaline cells. Munk pores

present in ascomal wall cells, few per cell. Quellkörper absent, paraphyses few inflated,

branched and septate. Asci clavate, long stipitate, 115–150 × 16–18 µm, spore-bearing part 60–

85 µm, pedicel 45–75 µm, curved, thin-walled, no apical ring, 8-spored, biseriately arranged.

Ascospores cylindrical, hyaline, geniculate, 1-septate, without mucilagenous sheath or

appendages, 26–33 × 7–8 µm.

Habitat: On partialy decorticated woody branch on the ground in forested area.

Anamorph: Unknown.

Distribution: Presently known only from one tropical highland forest in Kenya.

Specimen examined: KENYA. Rift Valley Province: Kajiando District, Ngong Hills Forest near communication

towers, 1°23.934S, 36°38.287E, elev 2367 m, 7 Dec 2006, on woody branch, GKM1239 (EA HOLOTYPE, F

ISOTYPE).

Bertia orbis Mugambi & Huhndorf, sp. nov. FIG. 5n–r

MycoBank MB 513310

Ascomata superficialia, separata vel dense aggregata, tuberculata, turbinatascens in statu humectato, collabens in

statu sicco, 515–665 µm alta, 495–560 µm diam. Paries ascomatis cellulis pseudoparenchymatis, Munk pori

preasens. Sine quellkörper, paraphysoides ramosus, hyalinae, crassi. Asci cylindrici-clavati, annulus apicalis,

octospori, longe stipititae, 112–130 × 7–9 µm, partibus sporiferis 45–70 µm longitudine, stipitibus 45–65 µm

longitudine. Ascosporae cylindricae ad fusoidae, hyalinae, 1-septatae, saepe curvus, 17–20 × 3–4 µm.

Etymology: Orbis (lat.) = world, denotes the wide geographical occurrence of the species.

Ascomata superficial, solitary or in large groups, turbinate when fresh, collabent when dry,

with thick sterile base, tuberculate, 495–560 µm wide 515–665 µm high including the sterile

base. Ascomal wall composed of outer layers of brown isodiametric pseudoparenchymatous cells

and inner layer flattened hyaline cells, Munk pores present in ascomal wall cells. Quellkörper

absent, paraphyses few, inflated, branched and hyaline. Asci cylindrical-clavate, long stipitate,

112–130 × 7–9 µm, spore-bearing part 45–70 µm, pedicel 45–65 µm, ascus apex with ring, eight

spores partially biseriate. Ascospores cylindrical to broadly fusoid, hyaline, often slightly curved,

1-septate, median, without mucilagenous sheath or appendages, four guttulate, two per cell, 17–

20 × 3–4 µm.

Habitat: Found growing on decorticated woody substrates in tropical forests above 1500 m.

Anamorph: Unknown.

Distribution: Known from tropical forests in Kenya and Costa Rica.

Specimen examined: COSTA RICA. Alajuela: Parque Nacional Volcan Arenal, La Fortuna de San Carlos, Pilón

Trail, [10.4419, −84.7167], 15-VII-2001, on woody branch 1 cm diam, S.M. Huhndorf, F.A. Fernández, A.N. Miller,

M.P. DaRin, SMH4557 (USJ HOLOTYPE, F ISOTYPE); Punta Arenas: La Amistad Pacifico, Las Tablas,

8°56.47N 82°46.38W, elev 1680 m, 16-I-1999, on wood fragment, F. A. Fernández, E. Fletes SMH4035 (F).

KENYA. Western Province: Kakamega forest near south gate, 0°13N 34°51E, elev 1585 m, 17 Jan 2007, on woody

branch, GKM1271 (F, EA); Western Province: Kakamega Forest near north gate, 0°21N 34°51E, elev 1500 m, 17

Jan 2007, on woody branch, GKM1259 (F, EA).

Bertia triseptata Mugambi & Huhndorf, sp. nov. FIG. 6a–e

MycoBank MB 513311

Ascomata superficialia, dispersa vel dense aggregata, atrobrunnea, turbinatascens in statu humectato, collabens in

stactu sicco, basim grandis, tuberculata, 590–650 µm alta, 685–785 µm diam., paries ascomatis cellulis

pseudoparenchymatis. Munk pori preasens, sine quellkörper, paraphysoides crassi, septati. Asci clavati, longe

stipititae, octospori, 170–200 × 19–22 µm, partibus sporiferis 110–130 µm longitudine, stipitibus 75–100 µm

longitudine. Ascosporae cylindricae, hyalinae, geniculatae et 3-septatae, 29–43 × 6–9 µm.

Etymology: Refers to the septation of ascospores.

Ascomata superficial, dark brown, usually aggregated in small or large clusters, turbinate

when fresh, becoming collabent when dry, with large sterile base making the ascomata look as if

stalked, tuberculate, 685–785 µm wide and 590–650 µm high including the sterile base. Ascomal

wall ca. 95–136 µm wide, composed of outer layers of brown thick-walled

pseudoparenchymatous cells and inner layers of flattened hyaline cells. Munk pores present in

ascomatal wall cells. Quellkörper absent, paraphyses few, inflated, branched, hyaline and

septate. Asci clavate, long stipitate, 170–200 × 19–22 µm, spore-bearing part 110–130 µm,

pedicel 75–100 µm, thin-walled, no apical ring, eight spores partially biseriate to triseriate.

Ascospores cylindrical, hyaline, geniculate, 3-septate, without mucilagenous sheath or

appendages, 29–43 × 6–9 µm.

Habitat: On decorticated woody substrates in forested areas.

Anamorph: Unknown.

Distribution: Known from tropical forests in Ecuador and Puerto Rico, USA.

Specimen examined: USA. Puerto Rico: Luquillo Mountains, El Verde research area, 16 ha Grid 07.04.14,

18°19′28″N 65°48′59″W, elev 382 m, 18-VI-1995, on log 50 cm, 1 m above ground, S.M. Huhndorf SMH1525

(HOLOTYPE, F). ECUADOR. Orellana Province: Yasuni National Park, Bariso trail, 7-III-2001, on 10 cm log,

F.A. Fernández, A.N. Miller, R.Briones, SMH4379 (F).

Cryptosphaerella celata Mugambi & Huhndorf, sp. nov. FIG. 6f–j

MycoBank MB 513312

Ascomata immersa vel erumpentia, atrobrunnea, separata vel gregaria, in subiculo denso hyphis brunneis,

turbinatascens in statu humectato, collabens in stactu sicco, paries ascomatis cellulis pseudoparanchymatis, Munk

pori preasens, quellkörper conocus, 395 µm longitudine, basim 245 µm diam. Paraphyses absens. Asci clavati, longe

stipititae, multispori, 100–127 × 9–11 µm, partibus sporiferis 38–70 µm longitudine, stipitibus 45–75 µm

longitudine. Ascosporae oblongae ad cylindricae, curvae, hyalinae, aseptatae, 8–14 × 2–3 µm.

Etymology: Celatus (lat.) = hidden, describing the habit of the ascomata that are usually concealed under thick

subiculum.

Ascomata immersed becoming erumpent, dark brown, solitary or aggregated in small

groups, with thick, brown tomentum around ascomata and often completely covering them,

turbinate when fresh, collabent when dry. Ascomal wall ca. 80–115 µm thick, composed of outer

layer of brown thick-walled pseudoparenchymatous cells with the cells becoming hyaline and

flattened toward the locule, Munk pores present in ascomal wall cells. Quellkörper present, large,

conical, 395 µm long and 245 µm wide at the base, paraphyses absent. Asci clavate, long

stipitate, 100–127 × 9–11 µm, spore-bearing part 38–70 µm, pedicel 45–75 µm, thin-walled, no

apical ring, multispored, spores irregularly arranged. Ascospores oblong to cylindrical, curved,

hyaline, 1-celled, without mucilagenous sheath or appendages, two guttulate one at each end of

the spore, 8–14 × 2–3 µm.

Habitat: Found growing on partially decorticated woody branch on the ground in a disturbed

forest patch.

Anamorph: Unknown.

Distribution: Currently known only from a tropical highland forest in Kenya.

Specimen examined: KENYA. Rift Valley Province: Nanyuki District, behind Bantu Lodge, 0°6′91″S, 37°2′70″E,

1 Dec 2006, on woody branch, GKM1231 (HOLOTYPE EA).

Cryptosphaerella costaricensis Mugambi & Huhndorf, sp. nov. FIG. 6k–o

MycoBank MB 513313

Ascomata erumpentia, separata vel gregaria, turbinatascens in statu humectato, collabens in stactu sicco, brunnea

setosus, 490–555 µm alta, 785–825 µm diam, paries ascomatis cellulis pseudoparenchymatis. Munk pori preasens,

quellkörper conicus, 200 µm longitudine, basim 165 µm diam. Asci clavati, brevi stipititae, 95–115 × 11–12 µm,

apice rotundatus, multisporae. Ascosporae cylindricae, hyalinae, aseptatae, 7–10 × 2–3 µm.

Etymology: Refers to the country in which the species was collected, Costa Rica.

Ascomata erumpent appearing superficial when the surrounding tissue wears off, solitary or

in small groups, turbinate when fresh, collabent when dry, covered with short thick walled

brown, septate tomentum, 490–555 µm high and 785–825 µm wide. Ascomal wall composed of

outer layer of brown pseudoparenchymatous cells with the cells becoming hyaline and flattened

toward the locule, Munk pores present in ascomal wall cells. Quellkörper present, conical, 200

µm long and 165 µm wide at the base. Asci clavate, short stipitate, 95–115 × 11–12 µm, thin-

walled, rounded apex, no apical ring, multispored, spores irregularly arranged. Ascospores

cylindrical, slightly curved, hyaline, 1-celled, with two large guttules, without mucilagenous

sheath or appendages, 7–10 × 2–3 µm.

Habitat: Found growing on log in a forested area.

Anamorph: Unknown.

Distribution: Presently known only from one forest in Costa Rica.

Specimen examined: COSTA RICA. Guanacaste, Bagaces, Tempisque Conservation Area, Parque Nacional Palo

Verde, 10:21:26.5245N −85:19:10.0313W, 13 Nov 2001, on log, M. Oses MO2111 (HOLOTYPE F).

Cryptosphaerella cylindriformis Mugambi & Huhndorf, sp. nov. FIG. 6p–t

MycoBank MB 513314

Ascomata erumpentia, atrobrunnea, separata vel gregaria, in subiculo sparsi hyphis brunneis, turbinatascens in statu

humectato, collabens in stactu sicco, 380–485 µm alta, 735–835 µm diam, setae sparsi. Ostiolum nullum, spinalis

brevibus dentiformibus saepe ornata, paries ascomatis cellulis pseudoparenchymatis. Munk pori preasens,

quellkörper conicus, 285 µm longitudine, basim 255 µm diam, paraphyses absens. Asci cylindricae-clavati, longae

stipitatae, 90–220 × 9–12 µm, partibus sporiferis 38–55 µm longitudine, stipitibus 25–60 µm longitudine, apice

rotundatus, multispori. Ascosporae cylindrici vel oblonge, hyalinae, curvus, aseptatae, 5–7 × 1–2 µm.

Etymology: Refers to the cylindrical ascospores.

Ascomata erumpent through the bark sometimes appearing superficial when the surrounding

plant tissue weathers off, dark brown, sparse setae often on the surface, solitary or in small

groups, turbinate when fresh, collabent when dry, nonostiolate, 380–485 µm high and 735–835

µm wide, sparse brown subiculum at the bases of ascomata. Ascomal wall of thick brown-walled

pseudoparenchymatous cells, becoming flattened and hyaline in the inner cells, outer cells

toothed, Munk pores present in the cells. Quellkörper present, conical, 285 µm long and 255 µm

wide at the base, paraphyses absent. Asci cylindrical-clavate, long stipitate, 90–220 × 9–12 µm,

spore-bearing part 38–55 µm, pedicel 25–60 µm, thin-walled, apex rounded, no apical ring,

multispored, spores irregularly arranged. Ascospores cylindric to oblong, hyaline, curved, 1-

celled, without mucilaginous sheath or appendages, often with two guttules, 5–7 × 1–2 µm.

Habitat: Found growing on partially decorticated woody substrates on the ground in forested

areas.

Anamorph: Unknown.

Distribution: Known from highland forests in Kenya.

Specimen examined: KENYA. Coast Province: Taita Taveta District, Taita Hills, Ngangao forest, 3°22′30″S

38°20′45″E, 14 Nov 2006, on woody branch 5 cm diam, GKM434N (HOLOTYPE EA, ISOTYPE F); 10 Nov 2006,

on woody branch, GKM1187 (EA); 16 Apr 2005, on woody branch, GKM1042 (EA); Nairobi Province: Nairobi

Arboretum, 1°16′S, 36°48′E, 7 Jun 2005, on wood, GKM1075 (EA).

Cryptosphaerella elliptica Mugambi & Huhndorf, sp. nov. FIG. 7a–e

MycoBank MB 513315

Ascomata erumpentia, gregaria, turbinatascens in statu humectato, collabens in stactu sicco, 635–665 µm alta, 935–

1035 µm diam., spinalis brevibus dentiformibus saepe ornata, paries ascomatis cellulis pseudoparenchymatis. Munk

pori preasens, quellkörper conicus, 245–320 µm longitudine, basim 165–220 µm diam., paraphyses absens. Asci

clavati, longe stipiti, 295–325 × 25–30 µm, partibus sporiferis 140–155 µm longitudines, stipitibus 150–180 µm,

apice rotundatus, miltisporae. Ascosporae ellipsoidae, aseptatae, 5–8 × 3–4 µm.

Etymology: Refers to the elliptical ascospores.

Ascomata erumpent often appearing superficial when surrounding tissues fall off, in large

clusters, turbinate when fresh, collabent when dry, 935–1035 µm wide and 635–665 µm high.

Ascomal wall ca. 65–80 µm in thick, composed of external wall of brown

pseudoparenchymatous cells becoming flattened and hyaline in cells lining the locule, ascomal

wall outer layer with tooth-like projections, Munk pores present in the cells. Quellkörper present,

conical, 245–320 µm long and 165–220 µm wide at the base, paraphyses absent. Asci clavate,

long stipitate, 295–325 × 25–30 µm, spore-bearing part 140–155 µm, pedicel 150–180 µm, thin-

walled, apex rounded, multispored, spores irregularly arranged. Ascospores ellipsoid, hyaline,

single celled, without mucilagenous sheath or appendages, 1–2 large guttules present in the cell,

5–8 × 3–4 µm.

Habitat: On decorticated woody branch on the ground.

Anamorph: Unknown.

Distribution: Presently known only from a single forested locality in Ecuador.

Specimen examined: ECUADOR. Orellana Province: Yasuni Biosphere Reserve, Tiputini Biological Station,

Guacamayo Trail, beyond 500 m marker, [−.6361, −76.1528], 26-III-2002, on woody branch 4 cm diam, F.A.

Fernández, A.N. Miller SMH4722 (HOLOTYPE F).

Cryptosphaerella globosa Mugambi & Huhndorf, sp. nov. FIG. 7f–j

MycoBank MB 513316

Ascomata immersa vel erumpentia, separata vel gregaria, atrobrunnea, subiculum sparsum, lateralis collapsa in statu

sicco, paries ascomatis cellulis pseudoparenchymatis, Munk spori preasens, quellkörper subcylindricae, 450 µm

longitudine, basim 185 µm diam. Paraphyses absens. Asci clavati, partibus sporiferis rotundatus, 60–68 × 19-23 µm,

partibus sporiferis 28–35 µm longitudine, stipitibus 25–37 µm longitudine, multispori. Ascosporae cylindricae,

curvae, hyalinae, aseptatae, 8–12 × 2–3 µm.

Etymology: Refers to the globose asci.

Ascomata mostly immersed rarely becoming erumpent, dark brown, sparse brown

subiculum around the ascomata, solitary or aggregated in small groups, occasionally laterally

collabent. Ascomal wall ca. 65 µm thick, composed of brown pseudoparenchymatous cells,

becoming hyaline and flattened toward the locule, surface cells of ascomatal wall more darkly

pigmented and ridged. Munk pores present in ascomatal wall cells. Quellkörper present,

subcylindrical, large, 450 µm long and 185 µm wide at the base. Paraphyes absent. Asci clavate,

stipitate, 60–68 × 19–23 µm, spore-bearing part 28–35 µm, pedicel 25–37 µm, thin-walled, apex

rounded, no apical ring, multispored, spores irregularly arranged. Ascospores cylindrical,

allantoid, hyaline, aseptate, without mucilaginous sheath or appendages, wall often collapsing in

parts, 8–12 × 2–3 µm.

Habitat: Found growing on partially decorticated woody branches in forest at 1800 m.

Anamorph: Unknown.

Distribution: Known only from single highland forest in Kenya.

Specimen examined: KENYA. Coast Province: Taita Taveta District, Taita Hills, Ngangao Forest, 3°22′30″S

38°20′45″E, elev 1800 m, 14 Nov 2006, on woody branch, GKM471N (EA HOLOTYPE); on woody branch,

GKM396N, GKM414N (EA).

Cryptosphaerella malindensis Mugambi & Huhndorf, sp. nov. FIG. 7k–q

MycoBank MB 513317

Ascomata superficialia, atrobrunnea, in subiculo denso hyphis brunneis et ramosis, dispersa vel gregaria,

turbinatascens in statu humectato, collabens in stactu sicco, spinalis brevibus dentiformibus saepe furcatis ornata,

455–515 µm alta, 630–650 µm diam, paries ascomatis cellulis pseudoparenchymatis. Ostiolum nullum. Quellkörper

conicus, 385 µm longitudine, basim 153 µm diam, Munk pori preasens. Asci clavati, longe pedicellati, octospori,

75–130 × 7–11 µm, partibus sporiferis 25–45 µm longitudine, stipitibus 45–85 µm longitudine. Ascosporae fusoidae

ad ellipsoidae, hyalinae, aseptatae, 10–14 × 3–5 µm.

Etymology: Refers to locality where the collection was made in Kenya, Malindi.

Ascomata dark brown, superficial, embedded in thick brown, smooth, septate subiculum,

nonostiolate, turbinate when fresh, collabent when dry, 455–515 µm high, 630–650 µm wide.

Ascomal wall ca. 54–60 µm thick, composed of two layers, a fibrous outer layer composed of

loosely packed brown septate hyphae and an inner layer made of brown thick walled

pseudoparenchymatic cells with outer layers of darkly pigmented cells with short spines and an

inner layer hyaline flattened cells, Munk pores present, few per cell. Quellkörper present, large,

conical, 385 µm long and 185 µm wide at the base, paraphyses absent. Asci clavate, long

stipitate, 75–130 × 7–11 µm, spore-bearing part 25–45 µm, pedicel 45–85 µm, thin-walled, no

apical ring, 8-spored, partially biseriate,. Ascospores broadly fusoid to ellipsoid, hyaline,

aseptate, without mucilagenous sheath or appendages, containing 1–2 large guttules 10–14 × 3–5

µm.

Habitat: On decorticated woody branch under forested area.

Anamorph: Unknown.

Distribution: Known only from a coastal lowland forest in Kenya.

Specimen examined: KENYA. Coast Province: Malindi District, Arabuko-Sokoke National Park, 3°19′30″S

39°57′10″E, elev 6 m, 30 Oct 2006, on wood fragment, GKM1150 (HOLOTYPE EA).

Scortechinia diminuspora Mugambi & Huhndorf, sp. nov. FIG. 7r–v

MycoBank MB 513318

Ascomata superficialia, in subiculo denso hyphis brunneis, spinalis et ramosis, ostiolum nullum, turbinatascens in

statu humectato, collabens in stactu sicco, coriacea, 210–245 µm alta, 215–235 µm diam, paries ascomatis cellulis

pseudoparenchymatis. Munk pori preasens, quellkörper subcylindrici, 175 µm longitudine, basim 75 µm diam,

paraphyses absens. Asci clavati, longe stipitatae, 25–27 × 6–7 µm, partibus sporiferis 15–18 µm longitudine,

stipitibus 7–11 µm longitudine, octospori. Ascosporae ellipsoidae, hyalinae, aseptatae, 5–6 × 2–3 µm.

Etymology: diminutus (lat.) = made small, refers to the size of the ascospores.

Ascomata superficial, embedded in thick subiculum whose hyphal terminations are

characteristically spiny, subglobose, dark brown, nonostiolate, collabent when dry, coriaceous,

210–245 µm high and 215–235 µm wide. Ascomal wall membranous, thin, composed of brown

pseudoparenchymatous cells, Munk pores present in ascomal wall cells. Quellkörper present,

subcylindrical, 175 µm long and 75 µm wide at the base, paraphyses absent. Asci clavate, long

stipitate, 25–27 × 6–7 µm, spore-bearing part 15–18 µm, pedicel 7–11 µm, apex rounded, no

apical ring, eight spores irregularly arranged. Ascospores ellipsoid, hyaline, single celled,

without mucilagenous sheath or appendages, 5–6 × 2–3 µm.

Habitat: On decorticated woody substrate on the ground.

Anamorph: Unknown.

Distribution: Known only from one forested locality in Ecuador.

Specimen examined: ECUADOR. Orellana Province: Yasuni Biosphere Reserve, Tiputini Biological Station,

Puma Trail, [−.6361, −76.1528], 29-III-2002, on log, F.A. Fernández, A.N. Miller SMH4763 (HOLOTYPE F).

Spinulosphaeria nuda Mugambi & Huhndorf, sp. nov. FIG. 8a–h

MycoBank MB 513319

Ascomata ovoidea, superficialia, dispersa, nigra, metallice nitentia, tuberculata, 455–530 µm alta, 475–525 µm

diam. Paries ascomatis cellulis pseudoparanchymatis, Munk pori preasens, quellkörper absens, paraphysiodes

hyalino, crassi et septata. Asci clavati, longe stipitatae, octospori, 125–145 × 14–19 µm, partibus sporiferis 48–60

µm longitudine, stipitibus 70–85 µm. Ascosporae ellipsoidea ad oblongae, brunnea, 1-septata, habens mucosae

vaginae, 12–15 × 4–6 µm.

Etymology: Refers to ascomatal surface devoid of spines.

Ascomata ovoid, superficial, scattered, black with metallic irridescence, glabrous,

noncollabent, tuberculate, 455–530 µm high and 475–525 µm wide. Subiculum is absent,

ascomatal wall is composed of brown pseudoparenchymatous cells, munk pores present in the

cell, few per cell. Quellkörper absent, paraphyses are broad, longer than asci, septate and

constricted at the septa. Asci clavate, long stipitate, 125–145 × 14–19 µm, spore-bearing part 48–

60 µm, pedicel 70–845 µm, thin-walled, lacking apical ring, eight spores partially biseriate,

obliquely arranged,. Ascospores are ellipsoid to oblong, hyaline when young becoming brown at

maturity, verruculose, 1-septate, with large persistent mucilaginous sheath, 12–15 × 4–6 µm.

Habitat: On decorticated woody substrate on the ground.

Anamorph: Unknown.

Distribution: Known only from a tropical rain forest in Puerto Rico, USA.

Specimen examined: USA: Puerto Rico: Luquillo Mountains, El Verde Research Area, 16 ha Grid, 18°19′26″N,

65°48′58″W, elev 395 m, 18-I-1996, on log 20 cm diam, S.M. Huhndorf SMH1952 (HOLOTYPE F).

KEY TO THE FAMILIES

1. Ascomata with quellkörper …………….. Scortechiniaceae

1. Ascomata without quellkörper …………….. 2

2. Ascomata mostly tuberculate (if smooth ascospores uniformly brown), ascospores longer than 15 µm

…………….. Bertiaceae

2. Ascomata smooth or with short spines …………….. 3

3. Ascomata erumpent, laterally collapsing …………….. Coronophoraceae

3. Ascomata superficial …………….. 4

4. Asci 8-spored, ascospores versicolorous …………….. Chaetosphaerellaceae

4. Asci 4- to multispored, ascospores concolorous …………….. Nitschkiaceae

KEY TO THE GENERA INCLUDED IN THE PHYLOGENETIC ANALYSES

1. Ascomata with quellkörper …………….. 2

1. Ascomata without quellkörper …………….. 11

2. Ascomata immersed, becoming erumpent (occasionally appearing superficial through wearing away of the

substrate) …………….. Cryptosphaerella

2. Ascomata superficial …………….. 3

3. Ascospores with appendage-like extensions on both ends …………….. 4

3. Ascospores without extensions …………….. 5

4. Asci 8-spored …………….. Biciliospora

4. Asci multispored …………….. Scortechiniella

5. Ascomata with spiny setae on the surface …………….. 6

5. Ascomata lacking spiny setae …………….. 7

6. Asci 8-spored …………….. Euacanthe

6. Asci multispored …………….. Neotrotteria

7. Ascomata with brown, flexuous tomentum …………….. Neofracchiaea

7. Ascomata without tomentum …………….. 8

8. Asci 8-spored …………….. 9

8. Asci multispored, ascospores reniform …………….. Scortechiniellopsis

9. Ascospores allantoid …………….. Coronophorella

9. Ascospores ellipsoid to globose …………….. 10

10. Ascomata on thick subiculum with hyphal terminations characteristically spiny …………….. Scortechinia

10. Ascomata on smooth subiculum …………….. Tympanopsis

11. Ascomata mostly tuberculate (if smooth, then ascospores becoming uniformly brown), collabent or collapsing

laterally, sometimes not collapsing …………….. Bertia

11. Ascomata not tuberculate …………….. 12

12. Ascomata immersed becoming erumpent, collapsing laterally …………….. Coronophora

12. Ascomata superficial, collapsing collabent or not …………….. 13

13. Ascomata spinulose …………….. 14

13. Ascomata not spinulose …………….. 17

14. Asci multispored …………….. Fracchiaea

14. Asci 8-spored …………….. 15

15. Ascospores versicolorous …………….. 16

15. Ascospores hyaline …………….. Acanthonitschkea

16. Ascomata collabent or not, spines restricted toward the base of fruit body …………….. Chaetosphaerella

16. Ascomata not collabent, with spiny setae on the surface and on the subiculum …………….. Crassochaeta

17. Ascospores hyaline, smaller than 15 µm long …………….. Nitschkia

17. Ascospores brown …………….. 18

18. Ascomata collabent, ascospores without mucilagenous sheath …………….. Gaillardiella

18. Ascomata not collabent ascospores with mucilagenous sheath …………….. Spinulosphaeria nuda (the type

species, S. thaxteri has spinulose ascomata and lacks ascospore sheath)

KEY TO THE BERTIA SPECIES INCLUDED IN THE PHYLOGENETIC ANALYSES

1. Ascospores hyaline, not turning brown …………….. 3

1. Ascospores becoming brown …………….. 2

2. Ascospores cylindrical to broadly allantoid, curved, three septate …………….. B. didyma

2. Ascospores fusiform, straight or slightly curved, 3- or more septate …………….. B. multiseptata

3. Ascospores 1-septate …………….. 4

3. Ascospores 3-septate …………….. B. triseptata

4. Ascospores fusiform, straight or slightly curved …………….. 5

4. Ascospores cylindrical curved or oblong straight …………….. 7

5. Ascospores greater than 20 µm long …………….. 6

5. Ascospores less than 20 µm long …………….. B. orbis

6. Ascospores 25–30 × 6.5–8 µm, slightly curved geniculate …………….. B. sinensis

6. Ascospores 35–50 × 4–6.5 µm, straight …………….. B. moriformis

7. Ascospores cylindrical, strongly curved geniculate …………….. 8

7. Ascospores wide oblong, straight …………….. B. gigantospora

8. Ascomata erumpent through cracks in bark, known only from Kenya …………….. B. ngongensis

8. Ascomata occurring widely on substrate, known mainly from the Caribbean and Central America

…………….. B. tropicalis

KEY TO THE CRYPTOSPHAERELLA SPECIES INCLUDED IN THE PHYLOGENETIC ANALYSES

1. Asci multispored …………….. 2

1. Asci 8-spored …………….. C. malindensis

2. Ascomata with outer surface vestiture …………….. 3

2. Ascomata without vestiture …………….. 5

3. Ascomal wall with tooth-like projections …………….. C. elliptica

3. Ascomal wall with tomentose covering …………….. 4

4. Ascomata with thick, brown tomentum …………….. C. celata

4. Ascomata with short tomentum …………….. C. costaricensis

5. Ascomata mostly remaining immersed in the substrate, ascospores 8–12 × 2–3 µm …………….. C. globosa

5. Ascomata erumpent to superficial, ascospores 5–7 × 1–2 µm …………….. C. cylindriformis

DISCUSSION

Based on LSU rDNA data, Huhndorf et al (2004) accepted a monophyletic Coronophorales

composed of four families, Chaetosphaerellaceae, Bertiaceae, Scortechiniaceae and a

paraphyletic Nitschkiaceae. Based on the larger taxon sampling and three nuclear DNA markers,

the analyses presented here corroborate these findings. However the circumscriptions of

Bertiaceae and Scortechiniaceae are expanded and we accept Coronophoraceae as distinct from

Nitschkiaceae (FIGS. 1–4). Subramanian and Sekar (1990) took a broad concept of the

Coronophoraceae and placed all Coronophorales genera except Bertia and Spinulosphaeria in

the family. Here we limit Coronophoraceae to the species of Coronophora and possibly some

taxa that currently are placed in Fracchiaea (FIG. 2). At the same time we propose some changes

in the generic circumscriptions within the order.

Bertia generally has been treated as an isolated genus within the Coronophorales and more

recently was placed in its own family, Bertiaceae (Subramanian and Sekar 1990, Huhndorf et al

2004). In our analyses Bertiaceae forms a strongly supported clade including taxa in Thaxteria

and Gaillardiella. Three collections of T. didyma (= B. didyma) group together in a clade that is

nested within Bertia (FIG. 1). Based on these results we propose that Thaxteria should be

synonymized with Bertia. The position of Thaxteria has been debated for long time with the

genus recieving varied taxonomic placement over the years. Nannfeldt (1975) was of the view

that Thaxteria showed affinities to taxa in Lasiosphaeriaceae (Sordariales) and therefore placed it

in that family. Subramanian and Sekar (1990) transferred the genus back into Coronophorales

but synonymized it with Nitschkia. The inclusion of the genus in Bertiaceae had been suggested

based on its morphology (e.g. by Huhndorf et al 2004), however our analyses do not support its

separation from Bertia. Its combination of morphological characters agrees with those observed

in species of Bertia. The three collections of B. didyma match the descriptions given by

Fitzpatrick (1923) who examined the type collection. Bertia didyma has vertically elongate,

smooth ascomata that collapse laterally, long stipitate asci and cylindrical curved to widely

allantoid ascospores that remain hyaline and 1-septate for prolonged time but the ascospores later

become 3-septate and turn brown. This ascospore character was used by Fitzpatrick (1923) to

distinguish it from species of Nitschkia. Bertia multiseptata (Sivan.) Huhndorf, A.N. Mill. &

F.A. Fern. also has ascospores that become brown and septate, but these are long and narrow

instead of short and wide as in B. didyma. Along with the ascospores B. didyma differs primarily

from the other species of Bertia by its smooth laterally collapsing ascomata.

Our two collections of Gaillardiella pezizoides Pat. forms a basal clade in strongly

supported sister relationship with the taxa in Bertia (FIGS. 1–4). Its close relationship to Bertia

had been suggested based on its morphology (Huhndorf et al 2004), and in this study we

corroborate these findings and support its inclusion in the Bertiaceae. The species in the genus

resembles those of Bertia in their collabent ascomata but differs by having a circular thickening

around the edge, forming a cup-like structure.

Bertia occurs in both tropical and temperate regions, and in this study the genus is

represented by 21 specimens comprising eight species (including B. didyma) from a wide

geographic range. Three new species are described in the genus. Bertia orbis is represented by

four geographically diverse collections that group together in a well supported clade (FIG. 1).

Bertia orbis shows some morphological similarity to B. moriformis (Tode) de Not. in the shape

of asci and ascospores but differs in the size of both. The ascomata of Bertia orbis are collabent

and much smaller than those of B. moriformis. The two species also differ in molecular sequence

data (FIG. 1). Bertia turbinata Petch was not included in our analyses but was described as

having superficial, clustered, collabent ascomata with hyaline, 1-septate, straight or slightly

curved ascospores (Petch 1922). Bertia orbis differs from this species in the smaller sizes of asci

and ascospores.

Bertia triseptata differs from the other species of Bertia by possessing 3-septate, geniculate

ascospores, a combination of characters not previously reported in the genus. Geniculate

ascospores also occur in B. tropicalis, B. convolutispora K.D. Hyde, B. ngongensis and to some

extent B. sinensis J.C. Krug & Corlett, but in all these cases the ascospores are 1-septate. In the

LSU rDNA analyses B. triseptata forms a supported sister relationship with B. multiseptata (FIG.

1). Bertia multiseptata was first described by Sivanesan (1978) as a variety of B moriformis and

raised to species level by Huhndorf et al (2004). It produces ascospores that are multiseptate (3–

7), mostly straight or slightly curved as opposed to smaller, 3-septate geniculate ones found in B.

triseptata. The single collection of Bertia ngongensis does not group with any species included

in the analyses and also differs from the rest of the taxa by having ascomata that are erumpent

through cracks in the bark and mature asci that are distinctly curved with geniculate ascospores.

Bertia latispora (Corlett & J.C. Krug) Lar. N. Vassiljeva was not included in our analyses but is

reported to occasionally have erumpent ascomata, but our collection differs from this species by

having much smaller ascospores and asci appearing curved. Bertia ngongensis shares geniculate

1-septate ascospore characters with B. sinensis and B tropicalis but differs from these species in

its erumpent ascomata, shape and size of the asci and in its phylogenetic placement (FIG. 1).

Bertia convolutispora, the other species with similar ascospore characters, differs from our

collection in its size of ascospores, superficial ascomata and habit. Bertia convolutispora was

described from wood submerged in water, with superficial ascomata that are solitary and rarely

clustered (Hyde 1995).

Subramanian and Sekar (1990) in their taxonomic revision of Coronophorales adopted a

broad concept of Coronophoraceae. They placed all the genera of Coronophorales except Bertia

and Spinulosphaeria within the family. In our analyses Coronophora is represented by a single

collection obtained from the USA whose morphology matches that of the type species

Coronophora gregaria (Lib.) Fuckel. The species shows no close affinities to the currently

recognized families, and therefore Coronophoraceae is retained for the taxa in the genus (FIGS.

2–4). However in the tef1 analyses C. gregaria groups in moderately supported clade with a

collection obtained from Kenya bearing morphology similar to that of Fracchiaea (FIG. 2). The

outcome was unexpected, but this grouping probably suggests that some of the taxa currently

accepted in Fracchiaea might belong in Coronophoraceae. More species of Coronophora need

to be included in analyses to better understand the generic and familial circumscriptions.

Huhndorf et al (2004) accepted Chaetosphaerellaceae for Chaetosphaerella and

Crassochaeta. Chaetosphaerella was described by Müller and Booth (1972) for versicolorous

species in Chaetosphaeria Tul. & C. Tul. Réblová (1999a, b) accepted two species, C.

phaeostroma (Durieu & Mont.) E. Müll. & C. Booth and C. fusca (Fuckel) E. Müll. & C. Booth,

in the genus. In our analyses C. fusca groups together with C. phaeostroma in a strongly

supported clade further suggesting close relationship of the two species (FIGS. 1–4). Collections

of Crassochaeta nigrita form a strongly supported sister relationship with species of

Chaetosphaerella further corroborating the finding of Huhndorf et al (2004) (FIG. 1).

We accept Nitschkiaceae for the taxa in Nitschkia, Fracchiaea and Acanthonitschkea. The

three genera group together with high PP in LSU rDNA and combined analyses (FIGS. 1, 4). This

relationship however is not supported for tef1 and rpb2 analyses (FIGS. 2, 3). In both cases

Fracchiaea groups separately but Nitschkia and Acanthonitschkea group together in all gene

trees (FIGS. 1–4). Despite this outcome we are inclined to retain Fracchiaea in Nitschkiaceae

until more taxa, especially in Nitschkia and Fracchiaea, are available for analyses to confirm the

relationship in the family. We were not able to obtain a collection of Nitschkia parasitans

(Schwein.) Nannf., the type species of Nitschkia, for the analyses, hence the circumscription of

the family remains unclear.

Fiztpatrick (1924) synonymized all the species that had been described in Fracchiaea with

F. broomeiana. Phylogenetic analyses presented here indicate that Fracchiaea might be

polyphyletic, with one collection of an unnamed species grouping with that of Coronophora

(FIG. 2). A collection identified as F. lunata Patw. & G.T. Joshi is represented by a single rpb2

sequence and it groups together with F. broomeiana. Another unnamed Coronophorales also

shares the ascomatal morphology of Fracchiaea but instead groups together with Neotrotteria

pulchella Sacc. (FIG. 1). More work is clearly needed in this group before the generic and the

species concepts become clear. In our analyses three collections of Nitschkia tetraspora group

together in a well supported clade with another collection of Nitschkia that consistently forms

eight ascospores per asci (FIG. 1). The morphology of this collection is in every way similar to

that of N. tetraspora except for the number of ascospores per ascus. In N. tetraspora four

ascospores abort leaving only four to attain maturity. Based on the close relationship of these

collections in the analyses and similarity in their morphology, we are inclined to think that they

represent a single species despite the differences in ascospores numbers. At this moment we treat

the ascospore number as a variable character within the species until more collections of both

kinds are included in analyses to acertain the validity of the character in the taxonomy.

Based on three collections Huhndorf et al (2004) established Scortechiniaceae for the taxa

that possess a quellkörper in the centrum. In our analyses composed of expanded taxon sampling,

the quellkörper-bearing taxa group together in a strongly supported clade (FIGS. 1–4),

corroborating Huhndorf et al (2004). Nannfeldt (1975a, b) did not consider presence or absence

of a quellkörper of taxonomic importance at the genus level and therefore grouped together taxa

with or without the character. He accorded prominence to other ascomal characters, such as

smooth versus tuberculate walls, and presence or absence of spines. In doing so he synonymized

quellkörper-bearing genera with Nitschkia and Acanthonitschkea. Subramanian and Sekar (1990)

reinstated the genera Nannfeldt (1975b) had placed in synonymy, and our molecular analyses

support the separation of these genera from Nitschkia and Acanthonitschkea. Phylogenetic

analyses presented here support the conclusion of Huhndorf et al (2004b) that presence or

absence of a quellkörper is a family-level taxonomic character. The analyses also demonstrate

that some of the currently used generic characters, such as presence or absence of subiculum,

presence or absence of spines on ascomata and the number of ascospores per ascus, are probably

homoplasious. Several monotypic genera have been erected historically based on single character

state changes (TABLE III), including Biciliospora velutina Petr., Scortechiniella similis (Bres.)

Arx & E. Müll. and Scortechiniellopsis leonensis Sivan. Biciliospora velutina and S. similis both

have a smooth subiculum and ascospores that bear elongate appendage-like wall extensions but

differ in their ascospore number, while S. leonensis has a spinulose subiculum and multispored

asci with ascospores that lack wall extensions. In our analyses these species group together in

fairly well supported clades (FIGS. 2–4) indicating the characters currently used in generic

delimitation in the family might include homoplasies and further evaluation of their usefulness is

needed.

Six new species of Cryptosphaerella Sacc. are proposed for morphologically distinct

collections that group within Scortechiniaceae (FIGS. 1–4). The collections were obtained from

Africa, Central and South America. The species do not group together in the LSU rDNA tree

(FIG. 1) but form a supported monophyletic clade in the tef1 and combined gene trees (FIGS. 2,

4). Cryptosphaerella was established for Coronophoralean taxa that are immersed becoming

erumpent, solitary or in small groups, cupulate, nonostiolate and bearing a quellkörper. Members

of the group have asci that are multispored, with ascospores irregularly arranged. Of the 13

names known in the genus Petrak (1962) accepted four species with the remainder belonging

elsewhere. However the genus is collected infrequently and aside from C. malabarica Subram. &

Sekar no modern illustrations are available. To the best of our knowledge none of the currently

described species match our new collections. Most of the species accepted by Petrak (1962) have

temperate distributions and differ from our collections by having smaller and/or allantoid

ascospores. Only C. marylandica Petr. is described as having oblong, subfusoid ascospores, a

shape similar to that found in several of our species. But again this species differs in smaller

spores and temperate distribution. Cryptosphaerella shearii Petr. has a tropical Hawaiian

distribution but has small allantoid ascospores. The tropical C. malabarica differs from all other

species in its 1-septate ascospores and its 16-spored asci. Cryptosphaerella cylindriformis differs

from C. elliptica by having smaller asci, and smaller, cylindrical, slightly curved ascospores.

Cryptosphaerella costaricensis differs primarily from the other species by having ascomata that

possess brown, thick-walled, septate tomentum and cylindrical ascospores that are much larger

than those observed in C. cylindriformis. Cryptosphaerella globosa differs from all the species

by possessing ascomata that are mostly immersed, rarely erumpent and collapse laterally as well

as having asci that are rounded, with a globose appearing spore-bearing part and a thin pedicel.

Ascospores in this species have walls that are collapsed in some places and therefore appear

uneven. Cryptosphaerella celata is unique in having ascomata that are erumpent but become

covered by thick, brown septate subiculum. The ascomata are cupulate and asci are clavate and

multispored. However the most unique species we recognize in this genus is C. malindensis,

collected from Kenya. This species consistently groups in a well supported clade with C.

cylindriformis, C. elliptica and C. costaricensis (FIGS. 1–4). The ascomata in this species are

superficial, seated on a thick subiculum, the asci are 8-spored and the ascospores are hyaline and

ellipsoid. However, apart from superficial ascomata and possession of eight ascospores per

ascus, other morphological characters are consistent with those of Cryptosphaerella, and

therefore based on morphological and molecular data we place it in the genus.

Scortechinia was described by Saccardo (Saccardo and Berlese 1885), and later Fitzpatrick

(1923) treated Scortechinia species under genus Tympanopsis Fitzp. Scortechinia was

recognized by Huhndorf et al (2004) based on S. conferta (Schwein.) Subram. & Sekar, a name

used by Subramanian and Sekar (1990). Fries' (1823) use of the name “Sphaeria conferta Fr.”

for another entity takes page precedence over Sphaeria conferta Schwein:Fr., and therefore the

combination into Scortechinia made by Subramanian and Sekar (1990) should have had the

“confertula” epithet (see Nannfeldt 1975a:59 for discussion). However we here propose that

genus Scortechinia should be limited to the clade with S. acanthostroma, the type species (Figs

1–4). This includes taxa with superficial collabent ascomata that are embedded in thick spinulose

subiculum, with thin ascomal walls as observed in S. acanthostroma. Members also have a long

subcylindrical quellkörper, while ascospores are fusoid, hyaline or brown and smooth or striate.

We reinstate genus Tympanopsis for the clade with T. confertula and T. uniseriata. They are

characterised by superficial ascomata, with firm ascomal walls, sparse to thick subiculum that is

smooth. However, although these two taxa constantly grouped together except in the LSU rDNA

tree, the relationship was mostly not supported and only in combined data analyses did they

recieve significant Bayesian support (FIGS. 1–4).

The taxonomic position of Spinulosphaeria has been debated since its description by

Sivanesan in 1974. The monotypic genus was described for S. thaxteri (Pat.) Sivan. and placed in

Coronophorales. However Nannfeldt (1975b) accepted it in Lasiosphaeriaceae (Sordariales) and

Subramanian and Sekar (1990) in Bertiaceae (Coronophorales). A specimen in our collection fits

the description as an additional species in the genus, and in our analyses it nested within the

Coronophorales (FIG. 1). The taxonomic placement is further supported by its possession of

morphological characters that are consistent with taxa in the order. We therefore accept

Spinulosphaeria in the Coronophorales, corroborating the placement of the genus in the order

(Sivanesan 1974, Subramanian and Sekar 1990, Huhndorf 2004). However the taxonomic

position of the genus remains unclear to us and based on the analyses it might represent a new

lineage within the order. We were not able to obtain DNA from S. thaxteri, and therefore the true

position of the genus will be clear only when this is included in analyses. Spinulosphaeria nuda

differs from S. thaxteri by having ascomata that lack spines, lacking a subiculum and possessing

paraphyses that are broad, septate and constricted at septa. The asci are much larger than those

reported for S. thaxteri and the ascospores possess a persistent mucilaginous sheath.

This study includes a mixture of specimens that represent temperate and tropical collecting

sites. By including different climatic types some biogeographic data can be inferred. Certain

species seem to be restricted to temperate areas (e.g. B. moriformis) or tropical regions (e.g. B.

tropicalis). In certain widespread taxa such as T. confertula there are few genetic differences

between collections from Kenya, USA or France. Fracchiaea broomeiana is another widespread

species that shows little genetic difference among the collections from Kenya, Venezuela and

USA. Scortechinia acanthostroma, a species thought to be restricted to a subtropical/tropical

distribution, was found to occur in central Illinois, albeit with some genetic differences from the

tropical collections. Specimens used in this study were collected in South and Central America

over a number of years and in Kenya in 2005 and 2006. Some species that were described from

or are known to occur in Africa were recollected in this study, including C. chaetomioides, E.

foveolata, F. broomeiana, N. pulchella, N. calyculus, N. tetraspora, S. acanthostroma and S.

leonensis. Other named species, such as B. gigantospora, B. velutina, C. fusca, G. pezizoides, F.

lunata, T. confertula, and T. uniseriate, previously were not found in eastern Africa. Seven of the

new species described are from Africa. Quellkörper-bearing taxa tended to predominate among

the collections from Africa, making up 12 out of the 22 species of Coronophorales collected. In

general quellkörper-bearing taxa show a tendency to be more diverse in tropical regions than

temperate areas but more studies are needed to confirm this hypothesis.

CONCLUSION

The Coronophorales is confirmed for species in the monophyletic families Bertiaceae,

Chaetosphaerellaceae, Scortechiniaceae and a paraphyletic Nitschkiaceae. A single collection of

Coronophora gregaria occurs separate from taxa in the other families so the Coronophoraceae is

applied for this genus. The Bertiaceae is expanded to include Gaillardiella and B. didyma, taxa

with ascospores that become brown. In G. pezizoides the ascomata are cup-shape with a circular

thickening at the top edge, and in B. didyma the ascomata are smooth and collapse laterally. With

these additions taxa within the family continue to vary along the morphological lines of large,

robust ascomata that mostly have warted or tuberculate walls and collapse either collabent or

laterally and ascospores that are commonly hyaline, can vary in septation and most often are

larger than in other taxa in the Coronophorales. The quellkörper continues to be an important

character in defining the Scortechiniaceae, but within the family the characters that define

relationships among taxa remain ambiguous. Cryptosphaerella species with mostly erumpent

ascomata separate from taxa with superficial ascomata, but one species of Cryptosphaerella is

superficial. Tympanopsis and Scortechinia are recognized as separate genera and differ in their

smooth versus spinulose subiculum. However other taxa with these character states also occur in

different places in the clade. Taxa with multispored asci also are dispersed within the clade. The

monotypic genera, Biciliospora, Scortechiniella and Scortechiniellopsis, show strong relatedness

indicating the generic-level morphological character-state changes that separate them probably

are untenable. Nitschkiaceae remains paraphyletic with Fracchiaea not consistently grouping

together or with Nitschkia and Acanthonitschkea. The correct placement of the generic type N.

parasitans remains elusive. Spinulosphaeria is expanded to include a species lacking spines and

subiculum, and the genus is included as a member within the order however with uncertain

family placement.

ACKNOWLEDGMENTS

This work was supported in part by NSF PEET Grant (Partnerships for Enhancing Expertise in Taxonomy) DEB–

0118695. The authors are grateful for the help of F.A. Fernández who provided access to specimens and of A.N.

Miller, INBio and D. J. Lodge for fieldwork assistance. The National Museums of Kenya provided logistical support

for the fieldwork carried out in Kenya. Sequences were generated in the Pritzker Laboratory for Molecular

Systematics and Evolution at the Field Museum of Natural History.

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LEGENDS

FIG. 1. Phylogram of the maximum likelihood analyses generated from nuclear LSU rDNA partial sequences.

Bootstrap values ≥ 70% are shown above or below branches. Thickened branches indicate Bayesian posterior

probabilities ≥ 95%. The Coronophorales families accepted in this study also are indicated. Roman numerals refer to

collections (TABLE II).

FIG. 2. Phylogram of the maximum likelihood analyses generated from translation elongation factor 1 alpha (tef1)

partial sequences. Bootstrap values ≥ 50% are shown above or below branches. Thickened branches indicate

Bayesian posterior probabilities ≥ 95%. The Coronophorales families accepted in this study also are indicated.

Roman numerals refer to collections (TABLE II).

FIG. 3. Phylogram of the maximum likelihood analyses generated from partial sequences of the ribosomal

polymerase II subunit 2 (rpb2). Bootstrap values ≥ 70% are shown above or below branches. Thickened branches

indicate Bayesian posterior probabilities ≥ 95%. The Coronophorales families accepted in this study also are

indicated. Roman numerals refer to collections (TABLE II).

FIG. 4. Phylogram of the maximum likelihood analyses generated from the combined genes (partial sequences of

ribosomal polymerase II subunit 2, Translation elongation factor 1 alpha and nuclear LSU rDNA). Bootstrap values

≥ 70% are shown above or below branches. Thickened branches indicate Bayesian posterior probabilities ≥9 5%.

The Coronophorales families accepted in this study also are indicated. Roman numerals refer to collections (TABLE

II).

FIG. 5a–e. Bertia didyma. a. Ascomata on substrate; b. Ascus; c. Inflated paraphyses; d. Mature hyaline and

brown ascospores; e. Asci showing thickened apical ring (arrow). f–i. Bertia gigantospora. f. Ascomata on

substrate; g. Munk pores on ascomatal wall; h. Ascospores and inflated paraphyses (arrow); i. Ascus. j–m. Bertia

ngongensis. j. Ascomata on substrate; k. Munk pores on ascomatal wall; l. Ascospores; m. Ascus and ascospores.

n–r. Bertia orbis. n. Ascomata on substrate; o. Ascus; p. Asci showing thickened apical ring (arrow); q. Inflated

paraphyses; r. Ascospores. Bars: a = 1 mm; f, j = 500 μm; n = 250 μm; b, c, i, o, q = 20 μm; d, e, g–i, k–m, p, r =

10 μm. a, f, j, n by photomacrography; d, e, g–i, k–m, o–r by DIC; b, c by PH. a from SMH4719; b–e from

SMH4256; f–i from GKM1136; j–m from GKM1239; n from GKM1259; o–r from GKM1271.

FIG. 6a–e. Bertia triseptata. a. Ascomata on substrate; b. Longitudinal section through ascoma; c. Section through

ascomatal wall; d. Ascus; e. Ascospores. f–j. Cryptosphaerella celata. f. Ascomata on substrate; g. Ascomatal

wall section with tomentum on outer surface (arrow); h. Ascus; i. Ascospores; j. Quellkörper. k–o.

Cryptosphaerella costaricensis. k. Ascomata on substrate; l. Longitudinal section through ascoma showing

quellkörper in place (arrow); m. Ascus; n. Ascospores; o. Quellkörper. p–t. Cryptosphaerella cylindriformis. p.

Ascomata on substrate; q. Section through ascomatal wall; r. Ascus; s. Ascospores; t. Quellkörper. Bars: a, f, k, p

= 500 μm; b = 200 μm; j, l, o, t = 100 μm; g, h, m, q, r = 20 μm; c–e, i, n, s = 10 μm. a, f, k, p by

photomacrography; g by BF; b–e, h–j, l–o, q–t by DIC. a–e from SMH1525; f–j from GKM1231; k–o from

MO2111; p–t from GKM434N.

FIG. 7a–e. Cryptosphaerella elliptica. a. Ascomata on substrate; b. Section through ascomatal wall showing outer

toothed cells (arrow); c. Quellkörper; d Ascus; e. Ascospores. f–j. Cryptosphaerella globosa. f. Ascoma on

substrate; g. Ascus; h. Section through ascomatal wall; i. Ascospores; j. Quellkörper. k–q. Cryptosphaerella

malindensis. k. Ascomata on substrate embedded in thick subiculum; l. Section through ascomatal wall; m. Ascus.

n. Ascospores; o. Smooth subicular hyphae; p. Munk pores (arrow) few and scattered; q. Quellkörper. r–v.

Scortechinia diminuspora. r. Ascomata on substrate embedded in thick subiculum; s. Spinulose subicular hyphae;

t. Quellkörper; u. Asci; v. Ascospores. Bars: a, f, k, r = 500 μm; c, j, q = 100 μm; t = 50 μm; d, h, o = 20 μm; b,

e, g, i, l–n, p, s, u, v = 10 μm. a, f, k, r by photomacrography; b–e, g–j, l–q, s–v by DIC. a–e from SMH4722; f–j

from GKM414N; k–q from GKM1150; r–v from SMH4763.

FIG. 8a–h. Spinulosphaeria nuda. a. Ascomata on substrate; b. Section through ascomatal wall; c. Asci; d.

Inflated paraphyses (arrow); e. Mature hyaline ascospores; f. Mature brown ascospores; g. Ascospores showing

surface roughening; h. Ascospores with mucilagenous sheath (arrow). Bars: a = 500 μm; b–d = 20 μm; e–h = 10

μm. a by photomacrography; b–h by DIC; a–h from SMH1952.

FOOTNOTES

Submitted 18 Mar 2009; accepted for publication 22 Jun 2009.

1Corresponding author. E-mail: gmugam1@uic.edu

100

100 100

100

Cryptosphaerella cylindriformis III

Cryptosphaerella cylindriformis I

Cryptosphaerella cylindriformis II

Cryptosphaerella elliptica

Cryptosphaerella costaricensis

Cryptosphaerella malindensis

Biciliospora velutina

Scortechiniella similis

Scortechiniellopsis leonensis

Scortechinia acanthostroma II

Scortechinia acanthostroma III

Scortechinia acanthostroma I

Scortechinia acanthostroma IV

Scortechinia diminuspora

Neofracchiaea callista

Coronophorella chaetomioides

Cryptosphaerella celata

Cryptosphaerella globosa IV

Cryptosphaerella globosa I

Euacanthe foveolata I

Euacanthe foveolata II

Tympanopsis uniseriata I

Tympanopsis uniseriata II

Tympanopsis confertula II

Tympanopsis confertula I

Tympanopsis confertula IV

Tympanopsis confertula III

Coronophorales

Neotrotteria pulchella

Bertia triseptata I

Bertia triseptata II

Bertia multiseptata I

Bertia multiseptata II

Bertia tropicalis III

Bertia tropicalis I

Bertia tropicalis IV

Bertia tropicalis II

Bertia didyma I

Bertia didyma II

Bertia didyma III

Bertia ngongensis

Bertia orbis III

Bertia orbis IV

Bertia orbis II

Bertia orbis I

Bertia moriformis II

Bertia moriformis III

Bertia moriformis I

Bertia sinensis

Gaillardiella pezizoides I

Gaillardiella pezizoides II

Acanthonitschkea argentinensis

Acanthonitschkea tristis

Nitschkia calyculus II

Nitschkia grevillei

Nitschkia tetraspora IV

Nitschkia tetraspora I

Nitschkia tetraspora II

Nitschkia tetraspora III

Nitschkia meniscoidea

Fracchiaea broomeiana III

Fracchiaea broomeiana II

Spinulosphaeria nuda

Chaetosphaerella fusca I

Chaetosphaerella fusca II

Chaetosphaerella fusca III

Chaetosphaerella phaeostroma I

Chaetosphaerella phaeostroma II

Crassochaeta nigrata I

Crassochaeta nigrata II

Niesslia exilis I

Nectria violacea

10 changes

100

93 100

100

Scortechiniaceae

Bertiaceae

Nitschkiaceae

Chaetosphaerellaceae

100

Chaetosphaerella phaeostroma II

Chaetosphaerella phaeostroma I

Chaetosphaerella fusca III

Fracchiaea sp

Coronophora gregaria

Fracchiaea broomeiana III

Fracchiaea broomeiana I

Fracchiaea broomeiana II

Nectria cinnabarina I

Niesslia exilis II

10 changes

Bertia orbis II

Bertia gigantospora

Gaillardiella pezizoides II

Cryptosphaerella cylindriformis I

Cryptosphaerella cylindriformis III

Cryptosphaerella cylindriformis II

Cryptosphaerella cylindriformis IV

Cryptosphaerella elliptica

Cryptosphaerella costaricensis

Cryptosphaerella malindensis

Cryptosphaerella globosa II

Cryptosphaerella globosa III

Cryptosphaerella globosa IV

Cryptosphaerella celata

Euacanthe foveolata I

Coronophorella chaetomioides

Biciliospora velutina

Scortechiniella similis

Scortechiniellopsis leonensis

Neofracchiaea callista

Tympanopsis confertula III

Tympanopsis confertula IV

Tympanopsis confertula I

Tympanopsis confertula II

Tympanopsis uniseriata I

Tympanopsis uniseriata II

Scortechinia acanthostroma II

Scortechinia acanthostroma III

Scortechinia acanthostroma I

Scortechinia acanthostroma IV

Nitschkia tetraspora IV

Nitschkia tetraspora I

Acanthonitschkea tristis

Acanthonitschkea argentinensis

Bertiaceae

Scortechiniaceae

Chaetosphaerellaceae

Coronophoraceae

Nitschkiaceae

100

Cryptosphaerella cylindriformis I

Cryptosphaerella cylindriformis II

Cryptosphaerella cylindriformis III

Cryptosphaerella cylindriformis IV

Cryptosphaerella elliptica

Cryptosphaerella costaricensis

Cryptosphaerella malindensis

Biciliospora velutina

Scortechiniella similis

Scortechiniellopsis leonensis

Coronophorella chaetomioides

Cryptosphaerella globosa IV

Cryptosphaerella celata

Tympanopsis uniseriata I

Tympanopsis uniseriata II

Scortechinia acanthostroma II

Scortechinia acanthostroma IV

Tympanopsis confertula II

Tympanopsis confertula IV

Tympanopsis confertula I

Neofracchiaea callista

Euacanthe foveolata I

Coronophora gregaria

Fracchiaea broomeiana I

Fracchiaea broomeiana III

Fracchiaea broomeiana II

Fracchiaea lunata

Bertia gigantospora

Nitschkia calyculus I

Acanthonitschkea argentinesis

Nitschkia tetraspora I

Chaetosphaerella phaeostroma I

Chaetosphaerella phaeostroma II

Hypocrea rufa

Nectria cinnabarina II

50 changes

Scortechiniaceae

Coronophoraceae

Bertiaceae

Chaetosphaerellaceae

Nitschkiaceae

100

Cryptosphaerella cylindriformis II

Cryptosphaerella cylindriformis III

Cryptosphaerella cylindriformis I

Cryptosphaerella cylindriformis IV

Cryptosphaerella elliptica

Cryptosphaerella costaricensis

Cryptosphaerella malindensis

Cryptosphaerella celata

Cryptosphaerella globosa IV

Biciliospora velutina

Scortechiniella similis

Scortechiniellopsis leonensis

Coronophorella chaetomioides

Tympanopsis confertula II

Tympanopsis confertula IV

Tympanopsis confertula I

Tympanopsis uniseriata I

Tympanopsis uniseriata II

Scortechinia acanthostroma II

Scortechinia acanthostroma III

Scortechinia acanthostroma I

Scortechinia acanthostroma IV

Neofracchiaea callista

Euacanthe foveolata I

Coronophora gregaria

Fracchiaea broomeiana III

Fracchiaea broomeiana I

Fracchiaea broomeiana II

Fracchiaea lunata

Nitschkia tetraspora IV

Nitschkia tetraspora I

Acanthonitschkea argentinensis

Acanthonitschkea tristis

Chaetosphaerella phaeostroma II

Chaetosphaerella phaeostroma I

Chaetosphaerella fusca III

Bertia orbis II

Bertia orbis I

Gaillardiella pezizoides II

Nectria cinnabarina

50 changes

100

Scortechiniaceae

Coronophoraceae

Nitschkiaceae

Chaetosphaerellaceae

Bertiaceae

TABLE I. Coronophoralean genera accepted in selected works since 1973

Müller and von Arx 1973, one

family Nannfeldt 1975a, b, one

family von Arx 1981, one

family Subramanian and Sekar 1990,

two families

Coronophoraceae,

Coronophorales

Acanthonitschkea Speg.

Bertia de Not.

Biciliospora Petr.

Calyculosphaeria

Coronophora Fuckel

Fracchiaea Sacc.

Gaillardiella Pat.

Nitschkia G. H. Otth

Scortechiniella Arx & E. Müll.

Thaxteria Sacc.

Tympanopsis Starbäck

Nitschkiaceae, Sordariales

Acanthonitschkea (syn.

Euacanthe Theiss.,

Neotrotteria Sacc.)

Bertia

Coronophora (syn.

Cryptosphaerella Sacc.)

Gaillardiella

Nitschkia (syn. Biciliospora,

Coronophorella Höhn.,

Fracchiaea,

Neofracchiaea Teng,

Scortechinia Sacc.,

Scortechiniella,

Scortechiniellopsis Sivan.,

Tympanopsis

Coronophoraceae,

Coronophorales

Acanthonitschkea

Bertia

Biciliospora

Coronophora

Fracchiaea

Gaillardiella

Lasiobertia Sivan.

Nitschkia

Spinulosphaeria Sivan.

Sydowinula Petr.

Tympanopsis

Coronophoraceae,

Coronophorales

Acanthonitschkea

Biciliospora

Biciliosporina Subram. & Sekar

Cryptosphaerella

Euacanthe

Fracchiaea

Gaillardiella

Janannfeldtia Subram. & Sekar

Neotrotteria

Nitschkia (syn. Thaxteria)

Schizocapnodium Fairman

Scortechinia

Bertiaceae, Coronophorales

Bertia

Spinulosphaeria

TABLE II. Taxa used in this study (new sequences in boldface)

Taxon Source Origin GenBank accession number

LSU rDNA tef1 rpb2

Acanthonitschkea

argentinensis

S. Huhndorf 1395 Puerto Rico,

Luquillo Mts.

AY695259 FJ969042 FJ968943

Acanthonitschkea

tristis

S. Huhndorf 4723 Ecuador,

Orellana Prov.

FJ968949 FJ969043 —

Bertia didyma I L. Crane CR-206 Costa Rica,

Puntarenas

FJ968996 — —

Bertia didyma II S. Huhndorf 4256 Costa Rica,

Guanacaste

FJ968950 — —

Bertia didyma III S. Huhndorf 4719 Ecuador,

Orellana Prov.

FJ968958 — —

Bertia gigantispora G. Mugambi 1136 Kenya, Ngong

hills

— FJ969008 FJ968937

Bertia moriformis I S. Huhndorf 5169 USA, WI,

Columbia Co.

FJ968951 — —

Bertia moriformis II S. Huhndorf 3344 USA, MI,

Marquette Co.

AY695261

Bertia moriformis

III

S. Huhndorf 4320 USA, MI,

Berrien Co.

AY695260

Bertia multiseptata I S. Huhndorf 1153 Puerto Rico,

Luquillo Mts.

FJ968953 — —

Bertia multiseptata

S. Huhndorf 3127 Puerto Rico,

Luquillo Mts.

FJ968952 — —

Bertia ngongensis G. Mugambi 1239 Kenya, Ngong

hills

FJ968954 — —

Bertia orbis I G. Mugambi 1259 Kenya,

Kakamega

forest

FJ968959 — —

Bertia orbis II G. Mugambi 1271 Kenya,

Kakamega

forest

FJ968955 FJ969009 —

Bertia orbis III S. Huhndorf 4035 Costa Rica,

Puntarenas

FJ968960 — —

Bertia orbis IV S. Huhndorf 4557 Costa Rica,

Alajuela

GQ184146

Bertia sinensis S. Huhndorf 4034 Costa Rica,

Puntarenas

FJ968961 — —

Bertia triseptata I S. Huhndorf 1525 Puerto Rico,

Luquillo Mts.

FJ968957 — —

Bertia triseptata II S. Huhndorf 4379 Ecuador,

Orellana Prov.

FJ968956 — —

Bertia tropicalis I S. Huhndorf 3132 Puerto Rico.

Luquillo Mts.

FJ968962 — —

Bertia tropicalis II S. Huhndorf 4046 Jamaica,

Manchester

Parish

FJ968963 — —

Bertia tropicalis III S. Huhndorf 1707 Puerto Rico.

Luquillo Mts.

AY695262

Bertia tropicalis IV S. Huhndorf 3513 Panama, Barro

Colorado Island

AY695263

Biciliospora

velutina

G. Mugambi 1268 Kenya,

Kakamega

forest

FJ968964 FJ969018 FJ968932

Chaetosphaerella

fusca I

A. Miller 605 USA, NC,

Blount Co.

FJ968965 — —

Chaetosphaerella

fusca II

A. Miller 852 USA, TN,

Sevier Co.

FJ968966 — —

Chaetosphaerella

fusca III

G. Mugambi

L124N

Kenya, Taita FJ968967 FJ969002 —

Chaetosphaerella

phaeostroma I

S. Huhndorf 4257 Costa Rica,

Guanacaste

AY695264 FJ969004 FJ968940

Chaetosphaerella

phaeostroma II

S. Huhndorf 4585 UK,

Northumberlan

AY346274 FJ969003 —

Coronophora

gregaria

A. Miller 1555 USA, TN,

Sevier Co.

— FJ969007 FJ968938

Coronophorales F. Fernández 1073 Costa Rica, San

José

FJ968968 — —

Coronophorella

chaetomioides

G. Mugambi 1099 Kenya, Ololua

forest, Nairobi

FJ968969 FJ969034 FJ968922

Crassochaeta

nigrita I

S. Huhndorf 1667 Puerto Rico.

Luquillo Mts.

AY695265

Crassochaeta

nigrita II

S. Huhndorf 2931 Puerto Rico.

Luquillo Mts.

AY695266

Cryptosphaerella

celata

G. Mugambi 1231 Kenya, Mt.

Kenya

FJ968975 FJ969035 FJ968929

Cryptosphaerella

cylindriformis I

G. Mugambi 434N Kenya, Taita FJ968972 FJ969031 FJ968934

Cryptosphaerella

cylindriformis II

G. Mugambi 1042 Kenya, Taita FJ968973 FJ969032 FJ968918

Cryptosphaerella G. Mugambi 1187 Kenya, Taita GQ217531 FJ969033 FJ968925

cylindriformis III

Cryptosphaerella

cylindriformis IV

G. Mugambi 1075 Kenya, Taita — FJ969030 FJ968920

Cryptosphaerella

costaricensis

M. Oses MO2111 Costa Rica,

Guanacaste

FJ968971 FJ969028 —

Cryptosphaerella

elliptica

S. Huhndorf 4722 Ecuador,

Orellana Prov.

FJ968974 FJ969029 FJ968944

Cryptosphaerella

globosa I

G. Mugambi 396N Kenya, Taita FJ968976 — —

Cryptosphaerella

globosa II

G. Mugambi 414N Kenya, Taita — FJ969037 —

Cryptosphaerella

globosa III

G. Mugambi 437N Kenya, Taita — FJ969038 —

Cryptosphaerella

globosa IV

G. Mugambi 471N Kenya, Taita FJ968977 FJ969036 FJ968935

Cryptosphaerella

malindensis

G. Mugambi 1150 Kenya, Taita FJ968970 FJ969027 FJ968923

Euacanthe foveolata

G. Mugambi 1221 Kenya, Taita FJ968978 FJ969026 FJ968927

Euacanthe foveolata

S. Huhndorf 4408 Ecuador,

Orellana Prov.

AY695267 — —

Fracchiaea

broomeiana I

G. Mugambi 1071 Kenya, Nairobi

arboretum

— FJ969040 FJ968919

Fracchiaea

broomeiana II

S. Huhndorf 347 Venezuela,

Guanare

FJ968979 FJ969041 FJ968947

Fracchiaea

broomeiana III

S. Huhndorf 2809 USA, IN, Lake

Co.

AY695268

(as

‘broomeana’)

FJ969039 FJ968942

Fracchiaea lunata G. Mugambi 1089 Kenya, Nairobi

arboretum

— —

FJ968921

Fracchiaea sp G. Mugambi 1250 Kenya, Ololua

forest, Nairobi

— FJ969005 —

Gaillardiella

pezizoides I

G. Mugambi 1144 Kenya, Ololua

forest, Nairobi

FJ968980 — —

Gaillardiella

pezizoides II

G. Mugambi 1245 Kenya, Ololua

forest, Nairobi

FJ968981 FJ969006 —

Hypocrea rufa GJS 90-97 EU341808

Nectria cinnabarina

GJS 89-107 AF543785

Nectria cinnabarina

GJS 91-111 AF545567

Nectria violacea MUCL40056 AF193242

Neofracchiaea

callista

S. Huhndorf 2689 USA, IL, Ogle

Co.

AY695269 FJ969020 FJ968941

Neotrotteria

pulchella

G. Mugambi 1255 Kenya, Ololua

forest, Nairobi

FJ968982 — —

Niesslia exilis I CBS357.70 AY489718

Niesslia exilis II CBS560.74 AY489614

Nitschkia calyculus

G. Mugambi 1243 Kenya, Ololua

forest, Nairobi

— —

FJ968931

Nitschkia calyculus

S. Huhndorf 918 French Guiana,

Saül

FJ968983 — —

Nitschkia grevillei S. Huhndorf 4663 USA, IL, La

Salle Co.

AY346294

Nitschkia

meniscoidea

S. Huhndorf 1523 Puerto Rico.

Luquillo Mts.

AY695270

Nitschkia tetraspora

G. Mugambi

L148N

Kenya, Taita FJ968987 FJ969011 FJ968936

Nitschkia tetraspora

G. Mugambi

L213N

Kenya, Taita FJ968985 — —

Nitschkia tetraspora

III

S. Huhndorf 4692 Ecuador,

Orellana Prov.

FJ968986 — —

Nitschkia tetraspora

S. Huhndorf 4787 Ecuador,

Orellana Prov.

FJ968984 FJ969010 —

Scortechinia

acanthostroma I

SMH5313 USA, IL, De

Witt Co.

FJ968990 FJ969013 —

Scortechinia

acanthostroma II

G. Mugambi 1164 Kenya,

Arabuko-

Sokoke

FJ968989 FJ969014 FJ968924

Scortechinia

acanthostroma III

G. Mugambi

L163N

Kenya, Taita FJ968991 FJ969015 —

Scortechinia

acanthostroma IV

S. Huhndorf 1143 Puerto Rico,

Luquillo Mts.

FJ968988 FJ969012 FJ968948

Scortechinia

diminuspora

S. Huhndorf 4763 Ecuador,

Orellana Prov.

FJ968992 — —

Scortechiniella

similis

S. Huhndorf 2006 Puerto Rico,

Luquillo Mts.

FJ968994 FJ969019 FJ968945

Scortechiniellopsis

leonensis

G. Mugambi 1269 Kenya,

Kakamega

forest

FJ968993 FJ969021 FJ968933

Spinulosphaeria

nuda

S. Huhndorf 1952 Puerto Rico,

Luquillo Mts.

FJ968995 — —

Tympanopsis

confertula I

A. Miller 1567 USA, TN,

Sevier Co.

FJ969001 FJ969025 FJ968939

Tympanopsis

confertula II

G. Mugambi 1242 Kenya, Ololua

forest, Nairobi

FJ968997 FJ969023 FJ968930

Tympanopsis

confertula III

S. Huhndorf 2648 USA, IL, Cook

Co.

AY695272

(as

‘Scortechinia

conferta’)

FJ969022 —

Tympanopsis

confertula IV

S. Huhndorf 4841 France, Midi-

Pyrénées

FJ968998 FJ969024 FJ968946

Tympanopsis

uniseriata I

G. Mugambi 1203 Kenya, Mt.

Kenya

FJ968999 FJ969016 FJ968926

Tympanopsis

uniseriata II

G. Mugambi 1228 Kenya, Mt.

Kenya

FJ969000 FJ969017 FJ968928

TABLE III. Morphological characteristics of included Coronophorales genera

Taxon (* indicates

monotypic)

Ascomatal position

on substrate

Ascomatal

surface

Ascomata

collapsing

when dry

Subiculum Paraphyses Quellkörper Ascospores

per ascus

Ascospore

appendage-

like wall

extensions

Acanthonitschkea Superficial Setose Collabent Sparse to dense,

spinulose

Absent Absent 8 No

Bertia Superficial Glabrous Collabent,

lateral or not

collapsing

Inconspicuous Wide,

inflated

Absent 8 No

Biciliospora* Superficial Glabrous Collabent Sparse to dense,

smooth

Absent Present 8 Yes

Chaetosphaerella Superficial Setose

around base

Collabent or

not collapsing

Dense,

spinulose

Wide,

inflated

Absent 8 No

Coronophora Immersed/erumpent Glabrous Lateral Inconspicuous Absent Absent Many No

Coronophorella* Superficial Glabrous Collabent Dense, smooth Absent Present 8 No

Crassochaeta Superficial Setose Not

collapsing

Dense,

spinulose

Wide,

inflated

Absent 8 No

Cryptosphaerella Immersed/erumpent Glabrous or

hyphal

Collabent or

lateral

Inconspicuous

to sparse

Absent Present Many No

Euacanthe Superficial Setose Collabent Dense,

spinulose

Absent Present 8 No

Fracchiaea Immersed/erumpent

/superficial

Setose,

barbed

Not

collapsing

Inconspicuous Absent Absent Many No

Gaillardiella Superficial Glabrous Collabent Inconspicuous Absent Absent 8 No

Neofracchiaea* Superficial Setose Collabent Sparse, smooth Absent Present Many No

Neotrotteria Superficial Setose Collabent Dense,

spinulose

Absent Present Many No

Nitschkia Superficial Glabrous Collabent Sparse to dense,

smooth

Absent Absent 8 or 4 No

Scortechinia Superficial Glabrous Collabent Dense,

spinulose

Absent Present 8 No

Scortechiniella* Superficial Glabrous to

hyphal

Collabent Dense, smooth Absent Present Many Yes

Scortechiniellopsis* Superficial Glabrous Collabent Dense,

spinulose

Absent Present Many No

Spinulosphaeria Superficial Broad

tooth-like

spines or

glabrous

Not

collapsing

Dense,

spinulose

Wide,

inflated

Absent 8 No

Tympanopsis Superficial Glabrous to

hyphal

Collabent Sparse to dense,

smooth

Absent Present 8 No

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Full-text available

May 2021

Antecedentes y Objetivos: Los ascomicetos son el grupo más diverso del reino Fungi; han sido descritos como saprobios, necrótrofos, biótrofos y endófitos. En la Isla de Cozumel, Quintana Roo, México, aunque recientemente se han citado 29 especies, no existe un estudio exclusivo de este grupo de hongos. El objetivo del presente estudio es hacer una lista de las especies de ascomicetos de tres Áreas Naturales Protegidas (ANPs) de la Reserva de la Biosfera Isla Cozumel, México. Métodos: Se realizaron cuatro exploraciones en las Áreas Naturales Protegidas de la Reserva de la Biosfera Isla Cozumel: Centro de Conservación y Educación Ambiental, Parque Ecológico Punta Sur y Zona Arqueológica de San Gervasio Chen-tuk, todas administradas por la Fundación Parques y Museos de Cozumel. Los ejemplares recolectados fueron estudiados, descritos y determinados de acuerdo con las técnicas tradicionales en micología. El material está depositado en el Herbario ENCB. Resultados clave: Se determinaron 40 especies de ascomicetos; de éstos, 15 se citan por primera vez para Quintana Roo y ocho se registran para la Isla de Cozumel. Bertia tropicalis se cita por primera vez para México y Platystomum mexicanum se describe como especie nueva para la ciencia. Conclusiones: La clase Sordariomycetes es la mejor estudiada con 28 especies, mientras que el orden Xylariales, con 25, tiene 62.5% del total de las 40 registradas. La familia Hypoxylaceae, con 40% de los taxa determinados, es la familia más representada. Hypoxylon lenormandii, Rhytidhysteron neorufulum y R. rufulum fueron encontradas en todas las ANPs estudiadas.

Especies conocidas y nuevos registros de Coronophorales (Ascomycota) en México

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Full-text available

Jun 2022

Antecedentes y Objetivos: El orden Coronophorales ha sido poco estudiado en México. Incluye seis familias y solo cuatro de ellas están representadas en el país. Se caracteriza por ascomas periteciales turbinados o globosos, con presencia o ausencia de estructuras quellkörper y/o poros de Munk. El objetivo de este trabajo es dar a conocer las especies de Coronophorales previamente conocidas del país y describir los nuevos registros. Métodos: Se realizaron exploraciones al azar para la recolección de ejemplares, entre 2011 y 2015, en áreas verdes urbanas y periurbanas de la ciudad de Xalapa, Veracruz, México. Se revisó material recolectado y previamente herborizado. El estudio de las muestras se hizo de acuerdo con las técnicas tradicionales en micología y todo el material estudiado está depositado en el herbario XAL. Resultados clave: Se describen 11 especies de Coronophorales. Nueve de ellas: Bertia sinensis, B. triseptata, Chaetosphaerella fusca, Fracchiaea callista, Gaillardiella pezizoides, Nitschkia collapsa, N. floridana, Tympanopsis confertula y T. uniseriata son nuevos registros para México; Fracchiaea broomeana se cita por primera vez para Veracruz y Nitschkia grevillii se registra de nuevas localidades. Se presentan dibujos o fotografías del material estudiado, además de una clave para determinar las especies hasta ahora conocidas del orden en el país. Conclusiones: El orden Coronophorales está poco representado en México. Previo a esta contribución se conocían diez especies del grupo. Sin embargo, con los nuevos registros que aquí se presentan, ahora suman 19 las especies conocidas de este grupo de hongos en el país.

Ascomycetes on Para rubber (Hevea brasiliensis)

Article

Jan 2021

Chanokned Senwanna

This study focuses on ascomycetes associated with Para rubber trees, collected from plantations in Thailand. We provide descriptions and phylogenies for one new genus, seven new species, two asexual-sexual morph connections, 20 new host records, and one reference specimen. Dothideomycetes are dominant among ascomycetes on Para rubber. Only three species from our collection have previously been reported from Para rubber in the Amazon Forest, where Para rubber originates. Most taxa found on rubber trees in this study have previously been recorded in Thailand, either on Para rubber or different hosts. It is apparent that the taxa jumped from unrelated hosts to colonize rubber. A checklist of fungi and fungus-like organisms associated with Para rubber is also provided. The checklist includes references for each taxon, life mode information, and distribution. The checklist comprises 785 species and 180 taxa identified only to genus from 59 countries. The taxa in the checklist belong in 67 orders, 168 families, and 513 genera

Ascomycetes from the relic forest of Oreomunnea mexicana, Oaxaca, Mexico

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Full-text available

Dec 2021

La Esperanza is an Oaxacan relic area of the Tertiary, dominated by the big tree Oreomunnea mexicana (Juglandaceae). The forest is part of the priority region for the conservation of La Chinantla and constitutes one of the most conserved Mexican tropical montane cloud forests. We studied the Ascomycetes fungi and found 63 species, of which 32 are new records for Oaxaca. Ascocoryne inflata, Calyculosphaeria macrospora, Cercophora costaricensis, Chaetosphaeria ellisii, Coccomyces limitatus, Lasiosphaeria ovina, Leptogidium dendriscum, Marthamyces quadrifidus, Stereocaulon didymicum and Thelonectria lucida are new records for Mexico. Xylaria was the most diverse genus with 12 species. The most abundant species were Xylaria arbuscula and Lachnum apalum. The main growth habit was lignicolous. The tropical montane cloud forest of La Esperanza has unique characteristics allowing great taxonomic diversity of Ascomycetes.

First occurrence of Nyssa endocarps and associated fungi in the Oligocene of South China: palaeogeographical and palaeoecological significance

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Full-text available

Nov 2021

Co-evolutionary relationships of plants and fungi are of great importance for the phylogeny of both groups. Nyssa was widely distributed in the northern hemisphere during the Cenozoic. Extant species of Nyssa exhibit a disjunct distribution between eastern North America, Central America and East Asia. Here, a new species, Nyssa nanningensis Xu & Jin, is described based on fruit endocarps from the upper Oligocene Yongning Formation of the Nanning Basin, South China. This new fossil record of Nyssa expands the known palaeogeographical distribution of the genus to the low latitudes of East Asia. Associated fossil fungal fruiting bodies on Nyssa endocarps are assigned to the new fossil genus and species, Yongnicta nyssae Tobias & Maslova. This new genus is similar to some members of extant wood destructor taxa Coronophorales and Amphisphaeriales (Sordariomycetes, Ascomycota). About 3% of the Nyssa endocarps studied were affected by Yongnicta nyssae. Low frequency of fruit damage indicates that endocarps could be released from mesocarps by animals eating fleshy parts of fruits, making them potentially available to wood-destroying fungi.

Taxonomic studies of some often over-looked Diaporthomycetidae and Sordariomycetidae

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Full-text available

Nov 2021
FUNGAL DIVERS

Sordariomycetes is an earlier-introduced and one of the widely distributed class of Ascomycota. The class was initially classified based on morphology in having inoperculate and unitunicate asci. With the development of DNA based phylogenetic analysis, several undetermined or polyphyletic members of Sordariomycetes were reclassified. However, not all species belonging to this class have been sequenced and analyzed. There are a number of species, especially those old and poorly studied ones which have never been sequenced before and not even recollected again for further taxonomic verification. One of the main objective in this study is to revise and update the taxonomy of several well-known early and poorly studied species whose classification are still obscure. Herein, we re-examined the type materials and/or authentic specimens together to explore 74 relatively poorly-studied genera, which mainly belong to Boliniales, Calosphaeriales, Chaetosphaeriales, Jobellisiales, and Sordariales classified under Diaporthomycetidae and Sordariomycetidae. We provide descriptions, notes, figures and/or drawings and discussed their phylogenetic relationships. As a result, the monotypic Jobellisiales is transferred from Hypocreomycetidae to Diaporthomycetidae. Based on phylogenetic analysis, the polyphyletic Lasiosphaeriaceae is divided into five families, Bombardiaceae (Apodospora, Bombardia, Bombardioidea, Fimetariella and Ramophialophora), Lasiosphaeriaceae (Anopodium, Bellojisia, Corylomyces, Lasiosphaeria, Mammaria and Zopfiella), Lasiosphaeridaceae (Lasiosphaeris), Strattoniaceae (Strattonia) and Zygospermellaceae (Episternus and Zygospermella). In addition, a new family Neoschizotheciaceae is established based on Neoschizothecium. Analysis of the type species of Boothiella, Stellatospora, Sulcatistroma and Tengiomyces placed them in Sordariaceae, Chaetomiaceae, Hypocreales and Coronophorales, respectively. We classify the genera lacking molecular data based on their morphology and expect them to be recollected; that is, Kacosphaeria in Calosphaeriales; Arnium, Biconiosporella, Camptosphaeria, Diffractella, Emblemospora, Eosphaeria, Periamphispora, Synaptospora and Tripterosporella in Sordariales; Conidiotheca in Sordariomycetes; Copromyces, Effetia, Endophragmiella and Tulipispora are accommodated in Ascomycota. Besides, we establish a new genus Neoschizothecium based on phylogenetic analysis. New combinations proposed: Camaropella amorpha, Cam. microspora, Cam. plana, Cladorrhinum grandiusculum, Cla. leucotrichum, Cla. terricola, Cla. olerum, Helminthosphaeria plumbea, Immersiella hirta, Jugulospora minor, Lasiosphaeris arenicola, Neoschizothecium aloides, Neo. carpinicola, Neo. conicum, Neo. curvisporum, Neo. fimbriatum, Neo. glutinans, Neo. inaequale, Neo. minicaudum, Neo. selenosporum, Neo. tetrasporum, Neurospora autosteira, Podospora brunnescens, P. flexuosa, P. jamaicensis, P. hamata, P. macrospora, P. spinosa, Strattonia petrogale and Triangularia microsclerotigena, T. nannopodalis, T. praecox, T. samala, T. tarvisina, T. unicaudata, T. yaeyamensis. New epithets are proposed for Apiorhynchostoma apiosporum and Podospora dacryoidea.

Taxonomic studies of Coronophorales and Niessliaceae (Hypocreomycetidae)

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Full-text available

Sep 2021

The number of species in Sordariomycetes has increased dramatically. However, some genera discovered earlier are poorly known, documented and thus need to be redescribed. This paper continues the work on 'Taxonomic studies of some often overlooked Diaporthomycetidae and Sordariomycetidae'. We provide descriptions, notes and figures or drawings for 71 overlooked genera belonging to Bertiaceae, Ceratostomataceae, Chaetosphaerellaceae, Coronophoraceae, Niessliaceae, Nitschkiaceae and Scortechiniaceae in Hypocreomycetidae based on observation of type materials, authentic specimens or studying the literature. The taxonomic placements of the genera are redefined based on morphology and/or phylogenetic results. As a result, Gonatobotrys and Neotrotteria, which are often overlooked and have unstable taxonomic placements in previous studies, belong to Ceratostomataceae based on morphology and phylogeny. The lichenicolous genus, Nitschkiopsis, which lacks DNA sequence data, is placed in Niessliaceae based on its setose ascomata and hyaline ellipsoidal ascospores similar to Niesslia species. The type species of Cryptosphaerella lacks molecular data and is morphologically similar to Cryptosphaeria or Coronophora, both of which have ostiolate ascomata. Therefore, it is placed in Sordariomycetes, not in Scortechiniaceae, whose species are characterized by ascomata that lack ostioles. In addition, the taxonomy of species with ascomata lacking ostioles in Cryptosphaerella have been revised and introduced as two new genera; Neocryptosphaerella and Pseudocryptosphaerella based on the phylogenetic result. Seven new combinations proposed are Fracchiaea myricoides (≡ Coronophora myricoides), Neocryptosphaerella celata (≡ Cryptosphaerella celata), N. globosa (≡ Cry. globosa), Pseudocryptosphaerella costaricensis (≡ Cry. costaricensis), P. cylindriformis (≡ Cry. cylindriformis), P. elliptica (≡ Cry. elliptica) and P. malindensis (≡ Cry. malindensis).

Ainsworth & Bisby's Dictionary of the Fungi

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Full-text available

Jan 1996

A Method for Sectioning Ascomycete Herbarium Specimens for Light Microscopy

Article

Jul 1991

S.M. Huhndorf

CONFIDENCE LIMITS ON PHYLOGENIES: AN APPROACH USING THE BOOTSTRAP

Article

Jul 1985
EVOLUTION

Joseph Felsenstein

The recently-developed statistical method known as the "bootstrap" can be used to place confidence intervals on phylogenies. It involves resampling points from one's own data, with replacement, to create a series of bootstrap samples of the same size as the original data. Each of these is analyzed, and the variation among the resulting estimates taken to indicate the size of the error involved in making estimates from the original data. In the case of phylogenies, it is argued that the proper method of resampling is to keep all of the original species while sampling characters with replacement, under the assumption that the characters have been independently drawn by the systematist and have evolved independently. Majority-rule consensus trees can be used to construct a phylogeny showing all of the inferred monophyletic groups that occurred in a majority of the bootstrap samples. If a group shows up 95% of the time or more, the evidence for it is taken to be statistically significant. Existing computer programs can be used to analyze different bootstrap samples by using weights on the characters, the weight of a character being how many times it was drawn in bootstrap sampling. When all characters are perfectly compatible, as envisioned by Hennig, bootstrap sampling becomes unnecessary; the bootstrap method would show significant evidence for a group if it is defined by three or more characters.

Fragmente zur Mykologie: IV. Mitteilung (Nr. 156 bis 168)

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