Leaf-inhabiting genera of the Gnomoniaceae, Diaporthales - CBS

Studies in Mycology 62 (2008) 

Leaf-inhabiting genera of the Gnomoniaceae, 

Diaporthales 

M.V. Sogonov, L.A. Castlebury, A.Y. Rossman, L.C. Mejía and J.F. White 

CBS Fungal Biodiversity Centre, 

Utrecht, The Netherlands 

An institute of the Royal Netherlands Academy of Arts and Sciences

Leaf-inhabiting genera of the Gnomoniaceae, Diaporthales 

St u d i es in My c o l o g y 62, 2008

Studies in Mycology 

The Studies in Mycology is an international journal which publishes systematic monographs of filamentous fungi and yeasts, and in rare 

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systematics. For instructions for authors see www.cbs.knaw.nl. 

Ex e c u t i v e Ed i t o r 

Prof. dr Robert A. Samson, CBS Fungal Biodiversity Centre, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. 

E-mail: r.samson@cbs.knaw.nl 

Lay o u t Ed i t o r 

Marianne de Boeij, CBS Fungal Biodiversity Centre, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. 

E-mail: m.deboeij@cbs.knaw.nl 

Scientific Ed i t o rs 

Prof. dr Uwe Braun, Martin-Luther-Universität, Institut für Geobotanik und Botanischer Garten, Herbarium, Neuwerk 21, D-06099 Halle, Germany. 

E-mail: uwe.braun@botanik.uni-halle.de 

Prof. dr Pedro W. Crous, CBS Fungal Biodiversity Centre, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. 

E-mail: p.crous@cbs.knaw.nl 

Prof. dr David M. Geiser, Department of Plant Pathology, 121 Buckhout Laboratory, Pennsylvania State University, University Park, PA, U.S.A. 16802. 

E-mail: dgeiser@psu.edu 

Dr Lorelei L. Norvell, Pacific Northwest Mycology Service, 6720 NW Skyline Blvd, Portland, OR, U.S.A. 97229-1309. 

E-mail: llnorvell@pnw-ms.com 

Dr Erast Parmasto, Institute of Zoology & Botany, 181 Riia Street, Tartu, Estonia EE-51014. 

E-mail: e.parmasto@zbi.ee 

Prof. dr Alan J.L. Phillips, Faculdade de Ciências e Tecnologia, Universidade Nova de Lisboa, Quinta de Torre, 2829-516 Caparica, Portugal. 

E-mail: alp@mail.fct.unl.pt 

Dr Amy Y. Rossman, Rm 304, Bldg 011A, Systematic Mycology and Microbiology Laboratory, Beltsville, MD, U.S.A. 20705. 

E-mail: amy@nt.ars-grin.gov 

Dr Keith A. Seifert, Research Scientist / Biodiversity (Mycology and Botany), Agriculture & Agri-Food Canada, KW Neatby Bldg, 960 Carling Avenue, 

Ottawa, ON, Canada K1A OC6. 

E-mail: seifertk@agr.gc.ca 

Prof. dr Jeffrey K. Stone, Department of Botany & Plant Pathology, Cordley 2082, Oregon State University, Corvallis, OR, U.S.A. 97331-2902. 

E-mail: stonej@bcc.orst.edu 

Dr Richard C. Summerbell, 27 Hillcrest Park, Toronto, Ont. M4X 1E8, Canada. 

E-mail: summerbell@aol.com 

Copyright 2008 CBS Fungal Biodiversity Centre, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. 

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the author"s moral rights. 

Publication date: 29 December 2008 

Published and distributed by CBS Fungal Biodiversity Centre, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. Internet: www.cbs.knaw.nl. 

E-mail: info@cbs.knaw.nl. 

ISBN/EAN : 978-90-70351-74-8 

Online ISSN : 1872-9797 

Print ISSN : 0166-0616 

Cover: From left to right: Asci of Gnomonia neognomon BPI 877465C. Perithecia of Ambarignomonia petiolorum on decaying petioles of Liquidambar 

styraciflua BPI 844274. Asci of Gnomonia alnea BPI 877462A. Bottom from left to right: Perithecia of Gnomonia gnomon on overwintered fallen leaves 

on Corylus avellana BPI 844273. Asci of Apiognomonia acerina BPI 877677. Perithecium of Plagiostoma euphorbia-verrucosae on overwintered stems of 

Euphorbia verrucosa BPI 877685.

Leaf-inhabiting genera of the Gnomoniaceae, 

Diaporthales 

M.V. Sogonov 

Department of Plant Biology and Pathology, Rutgers University, New Brunswick, NJ 08901, U.S.A. and Systematic Mycology & Microbiology Laboratory, 

USDA Agricultural Research Service, Beltsville, Maryland 20705–2350, U.S.A. 

L.A. Castlebury 

Systematic Mycology & Microbiology Laboratory, USDA Agricultural Research Service, Beltsville, Maryland 20705–2350, U.S.A. 

A.Y. Rossman 

Systematic Mycology & Microbiology Laboratory, USDA Agricultural Research Service, Beltsville, Maryland 20705–2350, U.S.A. 

L.C. Mejía 

Department of Plant Biology and Pathology, Rutgers University, New Brunswick, NJ 08901, U.S.A. and Systematic Mycology & Microbiology Laboratory, 

USDA Agricultural Research Service, Beltsville, Maryland 20705–2350, U.S.A. 

J.F. White 

Department of Plant Biology and Pathology, Rutgers University, New Brunswick, NJ 08901, U.S.A. 




CONTENTS 

Abstract ............................................................................................................................................................................................................ 1 

Introduction ...................................................................................................................................................................................................... 1 

Materials and methods ..................................................................................................................................................................................... 3 

Collection and observation of herbarium specimens ................................................................................................................................. 3 

Culture preparation and morphology ......................................................................................................................................................... 3 

Measurements and data management ...................................................................................................................................................... 4 

DNA amplification and sequencing ............................................................................................................................................................ 4 

Phylogenetic analyses .................................................................................................................................................................................4 

Results and discussion .................................................................................................................................................................................... 8 

Phylogenetic analyses ............................................................................................................................................................................... 8 

Revised concepts of accepted genera ....................................................................................................................................................... 8 

Evaluation of morphological and host characteristics .............................................................................................................................. 12 

Taxonomy ....................................................................................................................................................................................................... 14 

Key to the species of Gnomoniaceae in this study .................................................................................................................................. 14 

Descriptions of genera and species of the Gnomoniaceae ........................................................................................................................... 18 

GNOMONIA ............................................................................................................................................................................................. 18 

Gnomonia gnomon ............................................................................................................................................................................ 18 

Gnomonia alnea ................................................................................................................................................................................ 21 

New and revised species of Gnomonia ................................................................................................................................................... 21 

Gnomonia incrassata ......................................................................................................................................................................... 21 

Gnomonia monodii ............................................................................................................................................................................ 21 

Gnomonia neognomon ...................................................................................................................................................................... 22 

Gnomonia orcispora .......................................................................................................................................................................... 25 

Gnomonia ostryae ............................................................................................................................................................................. 25 

Gnomonia pendulorum ...................................................................................................................................................................... 26 

Gnomonia rodmanii ........................................................................................................................................................................... 30 

Gnomonia skokomishica ................................................................................................................................................................... 30 

Gnomonia virginianae ........................................................................................................................................................................ 32 

Additional species accepted in Gnomonia ............................................................................................................................................... 32 

Gnomonia amoena ............................................................................................................................................................................ 32 

Gnomonia arnstadtiensis ................................................................................................................................................................... 33 

Gnomonia carpinicola ........................................................................................................................................................................ 34 

Gnomonia pseudoamoena ................................................................................................................................................................ 34 

AMBARIGNOMONIA ............................................................................................................................................................................... 35 

Ambarignomonia petiolorum .............................................................................................................................................................. 36 

APIOGNOMONIA ..................................................................................................................................................................................... 36 

Apiognomonia veneta ........................................................................................................................................................................ 36 

Additional species of Apiognomonia ........................................................................................................................................................ 38 

Apiognomonia acerina ....................................................................................................................................................................... 38 

Apiognomonia borealis ...................................................................................................................................................................... 38 

Apiognomonia errabunda .................................................................................................................................................................. 38 

Apiognomonia hystrix ........................................................................................................................................................................ 40 

GNOMONIOPSIS .................................................................................................................................................................................... 41 

Type species of Gnomoniopsis ................................................................................................................................................................ 41 

Gnomoniopsis chamaemori ............................................................................................................................................................... 41 

New and revised species of Gnomoniopsis ............................................................................................................................................. 44 

Gnomoniopsis clavulata .................................................................................................................................................................... 44 

Gnomoniopsis paraclavulata ............................................................................................................................................................. 44 

Additional species accepted in Gnomoniopsis ......................................................................................................................................... 47 

Gnomoniopsis comari ........................................................................................................................................................................ 47 

Gnomoniopsis fructicola .................................................................................................................................................................... 47 

Gnomoniopsis macounii .................................................................................................................................................................... 48 

Gnomoniopsis racemula .................................................................................................................................................................... 48 

Gnomoniopsis tormentillae ................................................................................................................................................................ 48 

OPHIOGNOMONIA .................................................................................................................................................................................. 48 

Type species of Ophiognomonia ............................................................................................................................................................. 48 

Ophiognomonia melanostyla ............................................................................................................................................................. 48 

New species of Ophiognomonia .............................................................................................................................................................. 51 

Ophiognomonia balsamiferae ............................................................................................................................................................ 51

Ophiognomonia pseudoclavulata ...................................................................................................................................................... 51 

Ophiognomonia vasiljevae ................................................................................................................................................................ 53 

Additional species accepted in Ophiognomonia ...................................................................................................................................... 55 

Ophiognomonia alni-viridis ................................................................................................................................................................ 55 

Ophiognomonia gei-montani ............................................................................................................................................................. 58 

Ophiognomonia intermedia ............................................................................................................................................................... 58 

Ophiognomonia ischnostyla .............................................................................................................................................................. 59 

Ophiognomonia leptostyla ................................................................................................................................................................. 62 

Ophiognomonia micromegala ............................................................................................................................................................ 63 

Ophiognomonia nana ........................................................................................................................................................................ 63 

Ophiognomonia padicola ................................................................................................................................................................... 63 

Ophiognomonia rosae ....................................................................................................................................................................... 64 

Ophiognomonia rubi-idaei ................................................................................................................................................................. 64 

Ophiognomonia sassafras ................................................................................................................................................................. 64 

Ophiognomonia setacea .................................................................................................................................................................... 64 

Ophiognomonia trientensis ................................................................................................................................................................ 64 

PLAGIOSTOMA ....................................................................................................................................................................................... 66 

Type species of Plagiostoma and synonymous genus, Cryptodiaporthe ................................................................................................. 67 

Plagiostoma euphorbiae .................................................................................................................................................................... 67 

Plagiostoma aesculi ........................................................................................................................................................................... 69 

New species of Plagiostoma .................................................................................................................................................................... 69 

Plagiostoma barriae ........................................................................................................................................................................... 69 

Additional species accepted in Plagiostoma ............................................................................................................................................ 70 

Plagiostoma amygdalinae ................................................................................................................................................................. 70 

Plagiostoma devexum ....................................................................................................................................................................... 70 

Plagiostoma euphorbiaceum ............................................................................................................................................................. 72 

Plagiostoma euphorbiae-verrucosae ................................................................................................................................................. 72 

Plagiostoma fraxini ............................................................................................................................................................................ 72 

Plagiostoma geranii ........................................................................................................................................................................... 72 

Plagiostoma petiolophilum ................................................................................................................................................................. 72 

Plagiostoma rhododendri ................................................................................................................................................................... 72 

Plagiostoma robergeanum ................................................................................................................................................................ 73 

Plagiostoma salicellum ...................................................................................................................................................................... 73 

Genera not included in this study or excluded from the Gnomoniaceae ................................................................................................. 74 

Acknowledgements ........................................................................................................................................................................................ 75 

References ..................................................................................................................................................................................................... 76 

Index .............................................................................................................................................................................................................. 78

available online at www.studiesinmycology.org 

doi:10.3114/sim.2008.62.01 

St u d i es in My c o l o g y 62: 1–79. 2008. 

Leaf-inhabiting genera of the Gnomoniaceae, Diaporthales 

M.V. Sogonov 1,2 , L.A. Castlebury 2 , A.Y. Rossman 2 , L.C. Mejía 1,2 and J.F. White 1 

1 

Department of Plant Biology and Pathology, Rutgers University, New Brunswick, NJ 08901, U.S.A.; 2 Systematic Mycology & Microbiology Laboratory, USDA Agricultural 

Research Service, Beltsville, Maryland 20705-2350, U.S.A. 

*Correspondence: A.Y. Rossman, Amy.Rossman@ars.usda.gov 

Abstract: The Gnomoniaceae are characterised by ascomata that are generally immersed, solitary, without a stroma, or aggregated with a rudimentary stroma, in herbaceous 

plant material especially in leaves, twigs or stems, but also in bark or wood. The ascomata are black, soft-textured, thin-walled, and pseudoparenchymatous with one or 

more central or eccentric necks. The asci usually have a distinct apical ring. The Gnomoniaceae includes species having ascospores that are small, mostly less than 25 

μm long, although some are longer, and range in septation from non-septate to one-septate, rarely multi-septate. Molecular studies of the Gnomoniaceae suggest that the 

traditional classification of genera based on characteristics of the ascomata such as position of the neck and ascospores such as septation have resulted in genera that are 

not monophyletic. In this paper the concepts of the leaf-inhabiting genera in the Gnomoniaceae are reevaluated using multiple genes, specifically nrLSU, translation elongation 

factor 1-alpha (tef1-α), and RNA polymerase II second largest subunit (rpb2) for 64 isolates. ITS sequences were generated for 322 isolates. Six genera of leaf-inhabiting 

Gnomoniaceae are defined based on placement of their type species within the multigene phylogeny. The new monotypic genus Ambarignomonia is established for an unusual 

species, A. petiolorum. A key to 59 species of leaf-inhabiting Gnomoniaceae is presented and 22 species of Gnomoniaceae are described and illustrated. 

Taxonomic novelties: New genus: Ambarignomonia. New species: Gnomonia incrassata, G. monodii, G. neognomon, G. orcispora, G. pendulorum, G. rodmanii, G. skokomishica, 

G. virginianae, Gnomoniopsis paraclavulata, Ophiognomonia balsamiferae, O. pseudoclavulata, O. vasiljevae, Plagiostoma barriae. New combinations: Ambarignomonia 

petiolorum; Apiognomonia hystrix; Gnomonia alnea, G. carpinicola, Gnomoniopsis clavulata, G. comari, G. fructicola, G. macounii, G. racemula, G. tormentillae; Ophiognomonia 

alni-viridis, O. gei-montani, O. intermedia, O. ischnostyla, O. leptostyla, O. micromegala, O. nana, O. rubi-idaei, O. setacea, O. trientensis; Plagiostoma aesculi, P. amygdalinae, 

P. robergeanum, and P. salicellum. 

Key words: Foliicolous fungi, multilocus phylogenetics, polyphasic taxonomy, species identification, species recognition. 

INTRODUCTION 

The ascomycete order Diaporthales includes a number of plant 

pathogenic fungi. The most notorious of these is the chestnut blight 

fungus [Cryphonectria parasitica (Murrill) M.E. Barr] that killed all 

of the American chestnut trees [Castanea dentata (Marsh.) Borkh.] 

in a few decades and thus altered the landscape of eastern North 

America (Anagnostakis 1987). Additional tree diseases are caused 

by members of the Diaporthales particularly in the Gnomoniaceae 

G. Winter. These include oak anthracnose [Apiognomonia 

errabunda (Roberge ex Desm.) Höhn.], cherry leaf scorch [A. 

erythrostoma (Pers.) Höhn.], sycamore canker [A. veneta (Sacc. 

& Speg.) Höhn.] (Sinclair & Lyon 2005) and ash anthracnose 

[Gnomoniella fraxini Redlin & Stack, now Plagiostoma fraxini 

(Redlin & Stack) Sogonov, anamorph Discula fraxinea Redlin & 

Stack]. Dogwood anthracnose, a disease that has killed dogwood 

trees (Cornus florida L., C. nuttallii Audubon ex Torr. & A. Gray) 

on both the east and west coasts of North America, is caused 

by Discula destructiva Redlin (1991), an asexually reproducing 

species in the Gnomoniaceae for which no sexual state is known 

(Zhang & Blackwell 2001, Castlebury et al. 2002). Recently it 

was discovered that the cause of butternut canker (Sirococcus 

clavigignenti-juglandacearum Nair et al.), a fungus that threatens 

to destroy another North American tree species (Juglans cinerea 

L.) belongs in the Gnomoniaceae (Ostry 1996, Mejia et al. 2008). 

The Diaporthales are a well-defined order of the 

Sordariomycetes, Sordariomycetidae, as demonstrated using 

a four-gene phylogeny (Zhang et al. 2006). Diaporthalean fungi 

are characterised morphologically by brown to black perithecial 

fruiting bodies immersed in a stroma or the substrate, lack of true 

paraphyses at maturity, and unitunicate asci that float free within 

the centrum at maturity and often have a conspicuous ring in the 

apex (Barr 1978, Samuels & Blackwell 2001). The ascospores 

vary from non-septate to multi-septate or muriform, ellipsoidal to 

elongate, and hyaline or pale yellow to dark brown, rarely black. 

The asexual states of Diaporthales are generally coelomycetous, 

producing phialidic, often annellidic conidiogenous cells, and 

mostly non- or one-septate conidia in acervuli or pycnidia with 

or without a well-developed stroma, although some anamorphic 

states produce dark brown, multi-septate conidia. 

Within the Diaporthales up to eight variously conceived 

families have been included over the past 30 years. These familial 

classifications of the Diaporthales were summarised by Zhang & 

Blackwell (2001) comparing Wehmeyer (1975), Barr (1978, 1990), 

and Kirk et al. (2001). In a molecular study Castlebury et al. (2002) 

analysed nuclear large subunit ribosomal DNA sequence data and 

outlined six major lineages, mostly recognised as families, within 

the Diaporthales. Since then three families have been added. 

A recent review discusses the definition of the nine currently 

accepted families included in the Diaporthales (Rossman et al. 

2007). 

The family Gnomoniaceae based on the genus Gnomonia 

has been variously conceived since it was established by Winter 

(1886). This name was proposed for conservation by Hawksworth 

Copyright 2008 CBS Fungal Biodiversity Centre, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. 

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1

So g o n o v e t a l. 

& Eriksson (1988) against Obryzaceae Körber and the proposal was 

accepted (McNeill et al. 2006). The concept of the Gnomoniaceae 

as monographed by Monod (1983) is in general agreement with 

results of molecular studies that include Gnomonia and its many 

segregate genera (Castlebury et al. 2002, DeSilva et al. 2008, Mejia 

et al. 2008). Other concepts of the family such as those proposed 

by Kobayashi (1970), Barr (1978, 1990), Vasilyeva (1998), and 

Eriksson et al. (2001) differ significantly from these results. The 

most commonly accepted concept of the Gnomoniaceae prior to 

the molecular studies cited above was that of Barr (1978). She 

recognised the suborder Gnomoniineae with the two families 

Gnomoniaceae and Valsaceae Tul. & C. Tul. distinguished by the 

placement of the neck. The Gnomoniaceae was defined as having 

“perithecia upright; necks central, rarely eccentric, erumpent 

separately” and included three subfamilies, one of which was the 

Gnomonioideae that included four genera now recognised within the 

Gnomoniaceae, i.e. Apiognomonia Höhn., Gnomonia, Gnomoniella 

Sacc., and Ophiognomonia (Sacc.) Sacc. The Valsaceae was 

defined as having “perithecia oblique or horizontal; necks oblique 

or lateral, erumpent separately or converging through stromatic 

disc” with the subfamily Plagiostomoideae that included four genera 

now recognised in the Gnomoniaceae, i.e. Apioplagiostoma M.E. 

Barr, Plagiosphaera Petr., Plagiostoma Fuckel, and Pleuroceras 

Riess. Kobayashi (1970) followed Höhnel (1917) in placing all 

genera of the Diaporthales in one family, Diaporthaceae Höhn. 

The family Cryptosporellaceae Arx & E. Müll. (Von Arx & Müller 

1954) was established for the genus Cryptosporella Sacc. but this 

family name is considered invalid because of the lack of a Latin 

description (ICBN Art. 36.1). Mejia et al. (2008) demonstrated 

that Cryptosporella belongs to the Gnomoniaceae as outlined by 

Castlebury et al. (2002), thus the name Cryptosporellaceae is a 

synonym of the much older Gnomoniaceae. 

Species in the Gnomoniaceae are characterised by ascomata 

that are immersed, rarely erumpent or superficial, solitary, without 

a stroma, or aggregated with a rudimentary stroma, in herbaceous 

plant material, especially in leaves, twigs or stems, but also in 

bark or wood. The ascomata are dark brown to black, generally 

soft-textured, thin-walled, and pseudoparenchymatous with either 

central or eccentric necks. Generally the asci have a distinct apical 

ring although this is not the case for species having long ascospores 

as in Crytosporella. The ascospores are generally small, mostly 

less than 25 μm long, although some are longer especially those 

of Cryptosporella, and range in septation from non-septate to oneseptate, 

either in median or eccentric position. The asexual states 

of members of the Gnomoniaceae are acervular or pycnidial with a 

broad opening; conidiogenous cells are phialidic, and conidia are 

pallid, non-septate (Monod, 1983). 

The Gnomoniaceae sensu Monod (1983) included 22 genera, 

some of which were excluded from this family by Castlebury et al. 

(2002). According to the latter authors, the family comprised the 

teleomorph genera Apiognomonia, Apioplagiostoma, Ditopella De 

Not., Gnomonia, Gnomoniella, Gnomoniopsis (Sacc.) Berl, Linospora 

Fuckel, Ophiognomonia, Phragmoporthe Petr., Plagiostoma, and 

Pleuroceras as well as species of the anamorph genera Discula 

Sacc. and Sirococcus Preus. Some genera previously placed 

in the Gnomoniaceae sensu Monod (1983) have been removed 

such as Mazzantia Mont., now placed within the Diaporthaceae, 

and Sydowiella Petr., type of the Sydowiellaceae Lar.N. Vassiljeva 

(Rossman et al. 2007). Two genera, namely Cryptodiaporthe and 

Cryptosporella with its synonym Ophiovalsa on woody substrates, 

were placed in the Valsaceae by Barr (1978) and not considered by 

Monod (1983); however, Castlebury et al. (2002) determined that 

these genera belong in the Gnomoniaceae. 

Considerable confusion exists about the generic concepts in the 

Diaporthales including the Gnomoniaceae such that one species 

may have been placed in several different genera. For example, 

Ophiognomonia melanostyla, originally described in Sphaeria, 

was transferred to Cryptoderis Auersw., Gnomonia, Gnomoniella, 

and Pleuroceras, all before 1899 when it was designated the type 

species of the genus Ophiognomonia. 

The genus Gnomonia includes nearly 280 specific and 

subspecific names. The type species, Gnomonia gnomon, and G. 

setacea (Pers. : Fr.) Ces. & De Not. were recently re-described 

by Sogonov et al. (2005). Species of Gnomonia typically have 

solitary, thin-walled, immersed perithecia with long necks and lack 

any stroma. In most species ascospores have one median septum. 

Species of Gnomonia generally occur on overwintered leaves and 

are relatively commonly collected in temperate regions. Recent data 

show that the genus Gnomonia is not monophyletic (Sogonov et al. 

2005); some species have been transferred to the Sydowiellaceae 

(Moročko & Fatehi 2007, Rossman et al. 2007). 

The genus Apiognomonia has been distinguished from 

Gnomonia by unequally septate ascospores (Barr 1978, Monod 

1983). Most of the 28 species and subspecific names placed in 

Apiognomonia were originally described in Gnomonia. Results of 

a molecular study demonstrated that the type species, A. veneta, 

is closely related but distinct from A. errabunda (Sogonov et al. 

2007). Both have a Discula asexual state. In molecular studies A. 

errabunda has previously grouped with Cryptodiaporthe aesculi 

and Plagiostoma (Mejia et al. 2008). 

Cryptodiaporthe Petr. is based on C. aesculi (Fuckel) Petr. that 

occurs on branches of Aesculus hippocastanum. Unlike typical 

members of the Gnomoniaceae, this genus occurs on woody plant 

parts as do species of Cryptosporella. Both genera were placed 

in the Valsaceae by Barr (1978) and Monod (1983) based on the 

presence of stromatic tissues. Castlebury et al. (2002) demonstrated 

that C. aesculi belongs in the Gnomoniaceae. At present 56 

species names have been placed in Cryptodiaporthe. Pathogenic 

species in Cryptodiaporthe include C. populi (Sacc.) Butin, cause 

of Cryptodiaporthe canker of poplar, and C. salicella (Fr.) Petr., 

cause of Cryptodiaporthe canker of willow (Sinclair & Lyon 2005). 

Cryptodiaporthe corni, cause of golden canker of alternate leaf 

dogwood, Cornus alternifolia L. f. (Redlin & Rossman 1991) has 

been excluded from the Gnomoniaceae and shown to belong in 

the Cryphonectriaceae (Castlebury et al. 2002, Gryzenhout et al. 

2006). 

The genus Plagiostoma was established for Gnomonia-like fungi 

having eccentric necks that result in horizontal or oblique ascomata 

and one-septate ascospores. Barr (1978) included this genus in the 

Valsaceae based on these characteristics of the ascomata, while 

Monod (1983) placed Plagiostoma in the Gnomoniaceae. The type 

species, P. euphorbiae (Fuckel) Fuckel, is known from dead stems 

of Euphorbia in Europe and has been included in molecular studies 

(Castlebury et al. 2002). At present about 32 additional species 

are included in Plagiostoma, most of which occur on overwintered 

herbaceous and woody plant parts of diverse dicotyledonous plants 

including hardwood trees. 

The genus Cryptosporella based on C. hypodermia (Fr.) Sacc., 

now includes the genus Ophiovalsa Petr., type species O. suffusa 

(Fr.) Petr., and occurs exclusively on woody substrates as recently 

monographed by Mejia et al. (2008). Species of Cryptodiaporthe 

have traditionally been defined as having one-septate ascospores. At 

present, Cryptosporella is a distinct genus within the Gnomoniaceae 

and includes nine species (Mejia et al. 2008). Unlike most other 

members of the Gnomoniaceae, Cryptosporella is characterised 

by a distinctly valsoid arrangement of ascomata. However, 

2

Le a f-i n h a b i t i n g g e n e r a o f t h e Gn o m o n i a c e a e, Di a p o r t h a l e s 

Cryptosporella is similar to other members of the Gnomoniaceae in 

having stromatal tissues that are prosenchymatous, forming small 

ectostromatic discs between the erumpent cluster of necks. This 

genus is not considered in detail here. 

The type species of Ditopella, D. ditopa (Fr.) J. Schröt., is common 

on woody branches of Alnus. In addition to being morphologically 

similar to the phragmosporous Phragmoporthe conformis (Berk. 

& Broome) Petr., Castlebury et al. (2002) showed their close 

phylogenetic relationship using LSU sequences. Species of Ditopella 

and Phragmoporthe are morphologically similar to Gnomonia except 

that their necks are individually surrounded by a rudimentary stroma 

and thus were placed in the tribe Ditopelleae of the Pseudovalsaceae 

M.E. Barr (Barr 1978). Thirteen species were described in Ditopella, 

of which two were excluded from the Diaporthales by Barr (1978). 

Ditopella is characterised by having one-septate, rarely non-septate 

ascospores in polysporous asci, while Phragmopothe differs from 

Ditopella by ascosporses having more than one septum in eightspored 

asci. In addition to the type, two other species are placed in 

Phragmoporthe, P. ploettneriana (Henn.) Petr. and P. pseudotsugae 

A. Funk. Two species placed in Phragmoporthe by Monod (1983) 

belong in Magnaporthe outside the Diaporthales (Kraus & Webster 

1972, Barr 1978). 

The genus Gnomoniella was established for Gnomonia-like 

species having non-septate ascospores. The type species, G. 

tubaeformis (Fr.) Sacc., occurs on overwintered leaves and petioles of 

Alnus in Europe and North America (Barr 1978). Gnomoniella fraxini 

was recognised as a member of the Gnomoniaceae by Castlebury et 

al. (2002). At present 85 species and subspecific names are included 

in Gnomoniella, most of which are poorly known. 

Gnomoniopsis was originally described as a subgenus within 

Gnomonia for species having ascospores that develop additional 

septa. The type species is Gnomoniopsis chamaemori (Fr.) Berl. Barr 

(1978) suggested that the development of additional septa was “of 

only occasional occurrence” and thus considered Gnomoniopsis to be 

a synonym of Gnomonia. The only other species in Gnomoniopsis, G. 

devexa (Desm.) Moesz & Smarods, was recognised as Plagiostoma 

devexum (Desm.) Fuckel by Barr 1978. 

The genus Ophiognomonia was based on Gnomoniella 

subgenus Ophiognomonia Sacc. for species having elongate, often 

septate ascospores. The type species, O. melanostyla (DC. : Fr.) 

Sacc., occurs on overwintered leaves and petioles of Tilia spp. in 

temperate regions. About 15 additional species are currently included 

in this genus but most of these are obscure. Two of these species 

are known as endophytes of woody plants, O. cryptica D. Wilson 

& M.E. Barr isolated from leaves of Quercus emoryi (Wilson et al. 

1997) and O. elasticae (Koord.) M. Monod on Ficus (Paulus et al. 

2007). Although O. cryptica is a dominant endophyte with interesting 

ecological implications, no living isolates of this species have been 

preserved (Wilson et al. 1997). 

With collection and culturing of fresh specimens it has become 

possible to re-evaluate the generic concepts in the Gnomoniaceae 

by analyzing the phylogenetic relationship of many species using 

multiple genes. Phylogenetic affinities of uncultureable species can 

be determined by sequencing multicopy genes and analyzing these 

sequences in relation to phylogenetically circumscribed genera. This 

study was undertaken to accurately define the leaf-inhabiting genera 

of the Gnomoniaceae including the type and additional species of 

as many genera as possible. In the course of this project many new 

species were collected and are described herein. 

MATERIALS AND METHODS 

Collection and observation of herbarium specimens 

Fresh specimens were collected by the first author in Canada (British 

Columbia, Ontario), Russia (Novgorod, Nizhniy Novgorod, Tver 

oblasts), Switzerland, and the United States (District of Columbia, 

Georgia, Hawaii, Louisiana, Maine, Maryland, Mississippi, New 

Jersey, New York, North Carolina, Pennsylvania, Tennessee, 

Virginia, Washington) in 2004–2007. Living and dead, attached 

or fallen, overwintered leaves, and overwintered dead parts of 

herbaceous plants were examined for the presence of ascomata 

or conidiomata. Those containing seemingly gnomoniaceous fungi 

were air dried and stored in paper bags or envelopes. Additional 

fresh material was collected by others and sent for use in this study 

from Austria, Bulgaria, Finland, Lithuania, Russia (Primorsky Kray), 

and the United Kingdom (Scotland). All specimens were deposited 

in the U.S. National Fungus Collections (BPI). 

Additional herbarium specimens were examined from the U.S. 

National Fungus Collections (BPI) as well as the Museum Botanicum 

Berolinense (B), Centraalbureau voor Schimmelcultures (CBS), 

Farlow Reference Library and Herbarium of Cryptogamic Botany 

in Harvard University (FH), Conservatoire et Jardin botaniques de 

la Ville de Genève (G), Royal Botanic Gardens at Kew (K), Leiden 

University branch of the Nationaal Herbarium Nederland (L), Musée 

et Jardins Botanique Cantonaux in Lausanne (LAU), Botanische 

Staatssammlung München (M), New York State Museum 

Mycological Collections Herbarium (NYS), Muséum National 

d’Histoire Naturelle (PC), Mycology Herbarium of Royal Ontario 

Museum (TRTC), Uppsala University (UPS), and Eidgenössische 

Technische Hochschule in Zürich (ZT). 

Fresh and herbarium specimens were first examined on natural 

substrates using a Wild M5A (Wild Heerbrugg Ltd., Heerbrugg, 

Switzerland) or Leica MZ APO (Leica Microsystems AG, Weitzlar, 

Germany) dissecting microscope and photographed with a DXM 

1200 digital camera (Nikon Instruments Inc., Melville, NY, U.S.A.). 

Perithecia and pycnidia-like conidiomata were extracted from 

leaf tissue using a sterile surgical scalpel under a dissecting 

microscope, placed into a drop of 3 % aqueous KOH, 7 % aqueous 

sodium acetate solution or water on a clean microscope slide. After 

rehydration, perithecia were examined and measured. Perithecia 

and pycnidia-like conidiomata were crushed to release their 

contents, which were transferred with an attenuated glass capillary, 

a scalpel or a micropipette to a clean area of the slide. For acervular 

conidiomata, a small part of the conidial mass with the underlying 

hyphal mat intermixed with leaf tissue was extracted to a slide. 

The material was covered with a cover slip and examined under 

Nomarski differential interference contrast (DIC) with an Axioplan2 

microscope (Carl Zeiss, New York, NY, U.S.A.) and photographed. 

Culture preparation and morphology 

For preparation of pure cultures, fresh material was rehydrated and 

crushed in sterile 7 % sodium acetate solution or water. Ascospores 

and asci or conidia were removed by means of an attenuated 

glass capillary or a micropipette and transferred to cornmeal agar 

(CMA, Sigma®, Sigma Chemical Co., St. Louis, MO, U.S.A.) plates 

containing 1 % (v/v) of an antibiotics solution (0.2 % streptomycin 

sulfate and 0.2 % neomycin sulfate in sterile distilled water). Plates 

were incubated at room temperature and periodically examined 

for germination of ascospores or conidia with a dissecting 

www.studiesinmycology.org 

3


microscope in transmitted light or the Axioplan2 microscope with 

low-magnification (×2.5–20) objectives. Germinated ascospores 

or conidia were transferred to fresh CMA or potato dextrose agar 

(PDA, Difco, Becton, Dickinson & Co., Sparks, MD, U.S.A.) and 

incubated at room temperature. Most cultures obtained in this study 

were deposited at the Centraalbureau voor Schimmelcultures 

(CBS, Utrecht, The Netherlands). For macroscopic descriptions 

of colonies, strains were grown on PDA, malt extract agar (MEA) 

containing 3 % malt extract (Bacto) and 1.5 % agar (Bacto), 

and malt yeast agar (MYA) containing MEA supplemented with 0.3 

% yeast extract (Bacto). Cultures were placed in an incubator 

with a 12 h light/dark cycle with blacklight (near UV) and cool white 

fluorescent light at 23 °C presented as (23 °C l/d) in the descriptions. 

In order to stimulate sporulation and/or perithecial formation by 

imitating natural conditions, some cultures were incubated on the 

same media as follows: 4 h blacklight/white fluorescent light at 2 °C, 

10 h same light at 10 °C, 1 h darkness at 10 °C, and 9 h darkness 

at 2 °C. This cycle is presented as 2/10 °C l/d in the descriptions. 

Cultures were observed for up to five mo. Colours were determined 

according to Kornerup & Wanscher (1978) with only colour names 

used herein. 

Measurements and data management 

Measurements in descriptions are given as minimum and maximum 

values in parentheses and ranges as intervals between the first and 

third quartile. Arithmetic means, standard deviations and number 

of measurements are given in parentheses. Thus, measurements 

are provided as length × width = (min–)Q 1 

–Q 3 

(–max) × (min–)Q 1 

– 

Q 3 

(–max) µm (mean1 × mean2, SD1, SD2, n). Measurement of 

microstructures are rounded to the nearest 0.5 μm. Images were 

processed with Adobe Photoshop 5.0 (Adobe Systems, Inc., San 

Jose, CA, U.S.A.). Original software (Sogonov 2005) built on MS 

Access 2000 (Microsoft Corporation, Bellevue, WA, U.S.A.) was 

used for collecting and storing data and images of the samples and 

for statistical evaluations. 

DNA amplification and sequencing 

Genomic DNA was extracted directly from actively growing surface 

mycelium scraped from PDA plates with the PUREGENE Cell and 

Tissue kit (Gentra Systems, Minneapolis, MN, U.S.A.) according to 

the manufacturer’s instructions using approximately 50 mg fresh 

mycelium. For some collections, ribosomal genes were amplified 

directly from perithecial or conidiomatal contents in one of two 

ways. A small amount of ascal or conidial masses was extracted 

from a perithecium or conidioma with a sterile scalpel under the 

dissecting microscope and placed on the inner sidewall of a 0.2 

mL PCR tube cap. Approximately 5 µL of PCR-grade water were 

added to the mass of spores with a micropipette. Alternatively, 

a perithecium or conidioma was placed in a drop of PCR-grade 

water on a fresh microscope slide and squeezed using a scalpel. 

Then approximately 5 µL of the water containing a cloud of asci 

or conidia was transferred either with a micropipette to the inner 

sidewall of a 0.2 mL PCR tube as above. PCR tubes containing 

spore suspensions were stored at -18 °C until amplification. The 

spore suspension was then spun to the bottom of the tube in a 

microcentrifuge (~30 s) after the PCR mix had been added to the 

tube. Before amplification, the spore suspensions were incubated 

for 5 min at 95 °C. 

The genes coding for the internal transcribed spacer regions 

1 and 2, including the 5.8S rDNA (ITS) and a region of the large 

ribosomal subunit (nrLSU), a fragment of the translation elongation 

factor 1-alpha (tef1-α) containing introns 4 and 5,and RNA polymerase 

II (rpb2) were amplified in 25 or 50 µL reactions on a GeneAmp 

9700 thermal cycler (Applied Biosystems, Foster City, CA, U.S.A.) 

under the following conditions: 0.2–0.3 ng/μL of genomic DNA, 4 

mM/μL each dNTP, 0.05 units/μL DNA polymerase (AmpliTaq®, 

Applied Biosystems, Foster City, CA, U.S.A. or GeneChoice®, Cat. 

No. T-12, GeneChoice, Inc., Frederick, MD, U.S.A.), 0.5 pmoles/ 

μL each primer and 10 % vol. of the manufacturer’s supplied 10× 

PCR buffer containing 15 mM MgCl 2 

. The thermal cycler program 

was as follows: 2 min at 95 °C followed by 35 cycles of 30 s at 94 

°C, 30 s at 55 °C, 1 min at 72 °C, with a final extension period of 

10 min at 72 °C. If no amplicon was obtained from a reaction under 

these conditions, the annealing temperature was decreased to 50 

or 52 °C and/or 4 % of DMSO (v/v) was added to the reaction 

mix. Following amplification, the PCR products were purified with 

ExoSAP-IT (USB Corporation, Cleveland, OH, U.S.A.) according to 

the manufacturer’s instructions. Internal transcribed spacer regions 

1 and 2, including the 5.8S rDNA, were amplified and sequenced 

using the primers ITS5 and ITS4 (White et al. 1990). A region of 

the tef1-α gene was amplified using primers EF1–728F designed 

by Carbone & Kohn (1999) and EF1–1567R designed by Rehner 

(2001). The tef1-α fragment was sequenced using primers EF1– 

983F and EF1–1567R (Rehner 2001). 

Phylogenetic analyses 

Sequences were edited using Sequencher v. 4.2 for Windows 

(Gene Codes Corporation, Ann Arbor, MI, U.S.A.). Alignments were 

manually adjusted using BioEdit v. 7.0.5.2 (Hall, http://www.mbio. 

ncsu.edu/BioEdit/) or JalView (Clamp et al. 2004). Sequences 

were deposited in GenBank and listed in Table 1 or as specimens 

sequenced for those not used in the phylogenetic analysis. 

Genes were aligned individually and concatenated in a text 

editor. The alignment consisted of nrLSU (791 bp), tef1-α (470 

bp), and rpb2 (1089 bp) sequences for a total of 2350 and 75 

taxa. Of these, 64 belonged to the Gnomoniaceae, three to the 

Melanconidaceae, and eight to other representatives of the 

Diaporthales. The alignment was partitioned by gene and by codon 

position for tef1-α and rpb2. Partitions were analysed for conflict 

using the 70 % reciprocal NJ bootstrap analysis as in Reeb et al. 

(2004) using distance settings identified by ModelTest (Posada 

& Crandall 1998) for the maximum likelihood analysis detailed 

below. Trees were inferred by maximum parsimony (MP) using 

the heuristic search option with random sequence addition (1000 

replications), MULTREES on and the branch swapping (tree 

bisection-reconnection) option of PAUP v. 4.0b10 (Swofford 2002). 

All characters were unordered and either given equal weight during 

the analysis or weighted according to a scheme of weight=3 for first 

and second codon positions, weight=1 for third codon positions and 

weight=2 for nrLSU. Gaps were treated as missing in the parsimony 

analysis. Relative support of branches was estimated with 1000 

bootstrap replications (Felsenstein 1985) with MULTREES and 

TBR on and 10 random sequence additions for the MP bootstraps. 

Bootstrap values are indicated on Fig.1 under the respective 

branches. 

Trees were also inferred using maximum likelihood as 

implemented in PAUP v. 4.0b10. ModelTest v. 3.7 (Posada & 

Crandall 1998) was used to determine the model used for the 

analysis. Likelihood settings were as follows: base=(0.2419 0.2900 

4


Table 1. Specimens and cultures of Gnomoniaceae sequenced for this study.* 

GenBank Accession Numbers 

Taxon Specimen Culture Country Host Collector tef1-α ITS nrLSU rpb2 

Ambarignomonia petiolorum BPI 844274 CBS 121227 U.S.A. : VA Liquidambar styraciflua M.V. Sogonov EU221898 EU254748 EU255070 EU219307 

Amphiporthe hranicensis BPI 843515 CBS 119289 Austria Tilia platyphylla W. Jaklitsch EU221890 EU199178 EU199122 EU199137 

Apiognomonia borealis NA CBS 799.79 Norway Geranium sylvaticum M. Monod EU221971 EU255000 EU255169 EU219275 

Apiognomonia errabunda NA CBS 109747 Switzerland Fagus sylvatica M. Monod EU221914 DQ313525 AF408334 EU219261 

Apiognomonia hystrix CBSH 11343 CBS 911.79 Switzerland Acer pseudoplatanus M. Monod EU221986 DQ313549 EU255180 EU219260 

Apiognomonia veneta NA CBS 897.79 Switzerland Platanus orientalis M. Monod EU221910 DQ313532 EU255195 EU219259 

“Apioplagiostoma” aceriferum NA CBS 778.79 Switzerland Acer campestre M. Monod EU221970 EU254750 EU255072 EU219316 

Cryphonectria cubensis BPI 841768 CBS 101281 Cameroon Eucalyptus urophylla I. Gibson EU222012 NS AF408338 DQ862016 

Cryphonectria nitschkei BPI 747935 CBS 109758 Russia Quercus mongolica L. Vasilyeva DQ862031 NS AF408335 DQ862015 

Cryphonectria parasitica NA ATCC 38755 U.S.A.: CT Castanea dentata N. DePalma EU222014 NS EU199123 DQ862017 

Cryptosporella alnicola NA CBS 121074 U.S.A.: MN Corylus cornuta L. Vasilyeva EU221960 EU199204 EU255076 EU199160 

Cryptosporella betulae BPI 748448 CBS 109763 Austria Betula alba W. Jaklitsch EU221884 EU199180 AF408375 EU199139 

Cryptosporella confusa BPI 843580 CBS 121063 U.S.A.: TN Betula papyrifera W. Jaklitsch EU221958 EU199219 EU255079 EU199175 

Cryptosporella femoralis BPI 872326 CBS 121076 U.S.A.: NY Alnus rugosa L. Vasilyeva EU221951 EU199220 EU255080 EU199176 

Cryptosporella hypodermia NA CBS 171.69 The Netherlands Ulmus campestris H.A. van der Aa EU221881 EU199225 DQ862028 DQ862018 

Cryptosporella suffusa BPI 871231 CBS 121077 Austria Alnus incana W. Jaklitsch EU221891 EU199184 EU199124 EU199142 

Cryptosporella wehmeyeriana BPI 843485 CBS 121085 U.S.A.: NC Tilia sp. L. Vasilyeva EU221959 EU199205 EU255082 EU199161 

Diaporthe phaseolorum NA ATCC 64802 U.S.A.: MS Stokesia laevis F. Uecker EU222020 NS EU255083 EU219348 

Discula destructiva BPI 1107757 CBS 109771 U.S.A.: WA Cornus nuttallii J. Ammirati EU221897 EU199186 AF408359 EU199144 

Ditopella ditopa BPI 748439 CBS 109748 Austria Alnus glutinosa W. Jaklitsch EU221943 DQ323526 AF408360 EU219297 

Gnomonia amoena BPI 877469 CBS 121262 U.S.A.: TN Carpinus caroliniana M.V. Sogonov EU221983 EU254771 EU255091 EU219293 

Gnomonia gnomon NA CBS 199.53 Italy Corylus avellana M. Ribaldi? EU221885 AY818956 AF408361 EU219295 

Gnomonia neognomon BPI 877526C CBS 121265 Canada: BC Corylus californica M.V. Sogonov EU221982 EU254787 EU255098 EU219336 

Gnomonia orcispora BPI 877465C CBS 121247 U.S.A.: WA Corylus californica M.V. Sogonov EU221922 EU254788 EU255099 EU219314 

Gnomonia pseudoamoena BPI 877518 CBS 121261 Canada: BC Corylus californica M.V. Sogonov EU221984 EU254795 EU255102 EU219305 

Gnomonia rodmanii BPI 878211A CBS 121909 U.S.A.: GA Carpinus caroliniana M.V. Sogonov NS EU254796 NS EU219337 

Gnomonia skokomishica BPI 877465B CBS 121245 U.S.A.: WA Corylus californica M.V. Sogonov EU221929 EU254797 EU255103 EU219291 

Gnomonia virginianae BPI 844264 CBS 121913 U.S.A.: MD Ostrya virginiana M.V. Sogonov EU221900 EU254801 EU255105 EU219309 

Gnomoniopsis chamaemori NA CBS 803.79 Finland Rubus chamaemorus M. Monod NS EU254808 EU255107 NS 

Gnomoniopsis comari CBSH 12997 CBS 806.79 Finland Comarum palustre M. Monod NS EU254821 EU255114 EU219286 


5


Table 1. (Continued). 



Gnomoniopsis fructicola NA CBS 208.34 France Fragaria sp. G. Arnaud EU221968 EU254826 EU255116 EU219284 

Gnomoniopsis macounii BPI 871008 CBS 121468 U.S.A.: NY Spiraea sp. L. Vasilyeva EU221979 EU254762 EU255087 EU219243 

Gnomoniopsis paraclavulata BPI 877448 CBS 121263 U.S.A.: TN Quercus alba M.V. Sogonov EU221939 EU254839 EU255120 EU219248 

Gnomoniopsis racemula BPI 871003 CBS 121469 U.S.A.: MN Chamerion 

angustifolium 

L. Vasilyeva EU221889 EU254841 EU255122 EU219241 

Gnomoniopsis tormentillae NA CBS 904.79 Switzerland Potentilla erecta M. Monod NS EU254856 EU255133 NS 

Leucostoma niveum BPI 748232 CBS 109489 Russia Populus sp. L. Vasilyeva EU222015 NS AF362558 EU219343 

Mazzantia napelli BPI 748443 CBS 109769 Austria Aconitum vulparia W. Jaklitsch EU222017 NS AF408368 NS 

Melanconis alni BPI 748444 CBS 109773 Austria Alnus viridis W. Jaklitsch EU221896 DQ323523 AF408371 EU219300 

Melanconis marginalis BPI 748446 CBS 109744 Canada: BC Alnus rubra M.E. Barr EU221991 EU199197 AF408373 EU219301 

Melanconis stilbostoma BPI 748447 CBS 109778 Austria Betula alba W. Jaklitsch EU221886 DQ323524 AF408374 EU219299 

Ophiognomonia alni-viridis NA CBS 782.79 Switzerland Alnus viridis M. Monod EU221974 EU254864 EU255138 EU219333 

Ophiognomonia balsamiferae BPI 877606 CBS 121266 Canada:BC Populus balsamifera M.V. Sogonov EU221955 EU254870 EU255140 EU219322 

Ophiognomonia intermedia NA CBS 119194 United Kingdom Betula pubescens S. Green EU222008 EU254873 DQ323520 EU219321 

Ophiognomonia ischnostyla NA CBS 837.79 Switzerland Corylus avellana M. Monod EU221972 EU254890 EU255142 EU219334 

Ophiognomonia leptostyla NA CBS 844.79 Switzerland Juglans regia M. Monod EU221996 EU254910 EU255149 EU219338 

Ophiognomonia micromegala BPI 877615A CBS 121910 U.S.A.: DC Carya tomentosa M.V. Sogonov EU221944 EU254918 EU255150 EU219332 

Ophiognomonia nana NA CBS 883.79 Finland Betula nana M. Monod EU221949 DQ323534 DQ323522 EU219326 

Ophiognomonia nervisequa BPI 877467B CBS 121908 U.S.A.: NC Carpinus americana M.V. Sogonov EU221930 EU254902 EU255147 EU219330 

Ophiognomonia padicola NA CBS 845.79 Switzerland Prunus padus M. Monod EU221946 EU199192 EU255152 EU199150 

Ophiognomonia pseudoclavulata BPI 844280 CBS 121236 U.S.A.: PA Carya tomentosa M.V. Sogonov EU222004 EU254923 EU255153 EU219317 

Ophiognomonia rosae BPI 877636 CBS 121267 U.S.A.: ME Rosa sp. M.V. Sogonov EU221956 EU254936 EU255158 EU219319 

Ophiognomonia sassafras BPI 877639 CBS 121243 U.S.A.: PA Sassafras albidum M.V. Sogonov EU221941 EU254941 EU255159 EU219327 

Ophiognomonia setacea BPI 843499 CBS 116850 U.S.A.: TN Quercus sp. L. Vasilyeva EU222007 AY818953 AY818959 EU219339 

Ophiognomonia vasiljevae BPI 877671 CBS 121253 U.S.A.: TN Juglans nigra M.V. Sogonov EU221999 EU254977 EU255162 EU219331 

Phragmoporthe conformis BPI 748450 CBS 109783 Canada: BC Alnus rubra M.E. Barr EU221993 DQ323527 AF408377 NS 

Plagiostoma aesculi BPI 748430 CBS 109765 Austria Aesculus 

hippocastanum 

Plagiostoma amygdalinae NA CBS 791.79 Switzerland Euphorbia 

amygdaloides 

W. Jaklitsch EU221913 EU199179 AF408342 EU199138 

M. Monod NS EU254995 EU255165 NS 

Plagiostoma apiculatum BPI 843527 CBS 121466 Austria Salix alba W. Jaklitsch EU221957 EU254996 EU255166 EU219278 

Plagiostoma barriae BPI 877717B CBS 121249 U.S.A.: WA Acer macrophyllum M.V. Sogonov EU221947 EU254997 EU255167 EU219270 

Plagiostoma devexum BPI 843489 CBS 123201 U.S.A.: NY Polygonum sp. L. Vasilyeva EU221933 EU255001 EU255170 EU219258 

6


Table 1. (Continued). 



Plagiostoma euphorbiae NA CBS 340.78 The Netherlands Euphorbia palustris W. Gams EU219234 EU199198 AF408382 EU219292 

Plagiostoma fraxini BPI 746412 CBS 109498 U.S.A.: MD Fraxinus pennsylvanica S. Redlin EU221987 AY455810 AF362552 EU219263 

Plagiostoma geranii NA CBS 824.79 Switzerland Geranium sylvaticum M. Monod NS EU255009 NS EU219273 

“Plagiostoma” inclinatum NA CBS 772.79 Switzerland Acer campestre M. Monod NS EU255034 EU255183 EU219315 

Plagiostoma petiolophilum BPI 863769 AR 3821 U.S.A.: NY Acer sp. L. Vasilyeva EU221988 EU255039 EU255185 EU219257 

Plagiostoma rhododendri NA CBS 847.79 Switzerland Rhododendron 

hirsutum 

M. Monod NS EU255044 EU255187 EU219272 

Plagiostoma robergeana BPI 843593 CBS 121472 Austria Staphylea pinnata W. Jaklitsch EU221908 EU255046 EU255188 EU219262 

Plagiostoma salicellum BPI 747938 CBS 109775 Austria Salix sp. W. Jaklitsch EU221916 DQ323529 AF408345 EU199141 

Pleuroceras oregonense BPI 877719 CBS 121260 Canada: BC Salix sitchensis M.V. Sogonov EU221931 EU255060 EU255196 EU219313 

Pleuroceras pleurostylum NA CBS 906.79 Switzerland Salix helvetica M. Monod EU221962 EU255061 EU255197 EU219311 

Pleuroceras tenellum BPI 871059 CBS 121082 U.S.A.: NC Acer rubrum M.V. Sogonov EU221907 EU199199 EU255202 EU199155 

Sirococcus clavigignentijuglandacearum 

NA CBS 121081 U.S.A.: MN Juglans cinerea M. Ostry EU221998 EU199200 EU199133 EU199156 

Sirococcus conigenus BPI 871248 CBS 101225 Austria Picea abies R. Schneider EU221927 EU199201 EU199134 EU199157 

Valsa ceratosperma BPI 748459 CBS 109777 Austria Quercus robur W. Jaklitsch EU222016 NS EU255209 EU219344 

Valsella salicis BPI 748461 CBS 109754 Austria Salix fragilis W. Jaklitsch EU222018 NS EU255210 AF408389 

*ATCC = American Type Culture Collection, Manassas, VA U.S.A.; BPI = U.S. National Fungus Collections, USDA ARS, Beltsville, MD U.S.A.; CBS = Centraalbureau voor Schimmelcultures, Utrecht, The Netherlands; NA = None available; NS = Not 

Sequenced. 


7


0.2534), nst=6, rmat=(1.0000 3.8112 1.0000 1.0000 7.4314), 

rates=gamma, shape=0.7954 pinvar=0.5555. A heuristic search 

was performed with 10 random addition sequences using the MP 

tree as the starting tree. Maximum likelihood bootstrap analysis 

was not performed. 

MrModeltest (v. 2.2) was used to estimate the model that best 

fit the data for the alignment. Each gene was analysed individually 

and the entire alignment was analysed unpartitioned. All analyses 

resulted in the same model. A Bayesian analysis using the resulting 

GTR+I+G model was applied to the three partitions (genes) was 

conducted. Three hot and one cold chain with Markov Chain Monte 

Carlo 2 million generations in MrBayes v. 3.1.2 (Huelsenbeck & 

Ronquist 2001) was used for the analyses. Trees were sampled 

every 100 generations and the first 1 000 000 generations were 

eliminated (burn in period) after comparison in Excel when 

determining posterior probabilities (PP) for clades. Two separate 

runs were performed and posterior probabilities were pooled and 

indicated on Fig. 1 above the branches. Only probabilities greater 

than 95 % are shown. 

In this study, 322 ITS sequences were obtained from specimens 

or cultures and deposited in GenBank as EU254748—EU255069 

with host information, voucher specimen/culture and locality data. 

The ITS sequences could not be aligned across the whole family 

and were therefore not used in multigene phylogenetic analysis. 

However, ITS sequences were used to place taxa within genera 

through the use of a local BLAST server in BioEdit. The BLAST 

database contained 426 ITS sequences from taxa within the 

Gnomoniaceae and included previously sequenced isolates from 

GenBank. Taxa used in the multigene phylogenetic analysis were 

used as reference taxa for this database in determining generic 

placements. All genera are represented by their respective type 

species in the multigene analysis. 

RESULTS AND DISCUSSION 

Phylogenetic analyses 

Results of the 70 % reciprocal NJ bootstrap analyses showed 

no conflict among the genes (trees not shown). However, tef1-α 

and nrLSU individually did not resolve all of the genera identified 

in the multigene analysis. Combined analysis of the tef1-α and 

nrLSU partitions did resolve all genera with >70 % support. The 

rpb2 individual NJ bootstrap analysis resolved all genera with >70 

% support indicating that rpb2 is providing most of the signal for 

resolution of the genera. 

Maximum parsimony analyses resulted in 24 equally 

parsimonious trees (score= 5095) and six equally parsimonious 

trees (score= 7047) for the unweighted and weighted analyses, 

respectively. Strict consensus trees calculated for each parsimony 

analysis did not differ in the identification of the clades at the 

genus level, with minor backbone differences (trees not shown). 

Maximum likelihood analysis resulted in one tree with a –lnL score 

of 26702.18332 (Fig.1) with Bayesian PP and MP bootstraps shown 

above and below the branches Seven genera with multiple species 

were strongly supported by the multi-gene phylogenetic analysis at 

a Bayesian PP level of 95 % or greater and MP bootstrap support 

of 70 % or greater: Apiognomonia, Cryptosporella, Gnomoniopsis, 

Gnomonia, Ophiognomonia, Plagiostoma, and Pleuroceras. 

Ambarignomonia is newly recognised with a single species, A. 

petiolorum, and Amphiporthe is recognised with the type species, 

A. hranicensis, in the Gnomoniaceae. The type species of 

Ditopella and Phragmoporthe are morphologically and biologically 

similar differing primarily in the number of ascospores in the asci 

and appear to be congeneric in this analysis, but are not further 

considered due to the small number of isolates sampled. 

Bayesian analyis and MP boostrapping both supported a 

group containing Amphiporthe, Apiognomonia, Cryptosporella, 

Ditopella/Phragmoporthe, and Plagistoma, with Ambarignomonia, 

Ophiognomonia, and Pleuroceras forming a closely related 

second group of genera. The type species of Ophiognomonia, O. 

melanostyla, could not be sequenced for the multigene analysis. 

However, an ITS sequence was obtained directly from a specimen 

and was found by BLAST analysis to be very closely related to 

O. sassafras, which has been used as the reference taxon for 

Ophiognomonia in the multigene analysis. All other genera are 

represented by type species in the multigene analysis. In all 

analyses Gnomoniopsis is basal to other gnomoniaceaous taxa 

in this alignment with the Melanconidaceae forming a very closely 

related sister group to the Gmononiaceae. Sirococcus clavigignentijuglandacearum, 

the butternut canker pathogen with no known 

sexual state, is strongly supported as within Ophiognomonia. 

However, Sirococcus conigenus, the type species of the anamorph 

genus Sirococcus, is placed with Gnomoniopsis in these analyses. 

The dogwood anthracnose pathogen, Discula destructiva, is not 

strongly supported as belonging to any of the genera present in this 

tree, but it forms a consistent relationship with Ambarignomonia 

and Pleuroceras. ITS data suggest a close relationship with 

Pleuroceras (trees not shown). 

Revised concepts of accepted genera 

Based on the molecular data presented here the previously 

established concepts for many of the genera in the Gnomoniaceae 

must be rejected. These were based primarily on characteristics of 

stromal development, perithecial neck orientation, and ascospore 

septation (Barr 1978, Monod 1983). The new concepts of the 

genera in the Gnomoniaceae presented here cannot be defined 

based on a single morphological characteristic; however, some 

generalisations can be made about the characteristics of each 

genus as presented in Table 2. 

This study presents a revised concept of the genus Gnomonia. 

It includes relatively few species, some of which are newly 

described, that group with the type species G. gnomon in the 

multigene phylogeny (Fig. 1) or that ITS sequences show to be 

congeneric with G. gnomon. Gnomonina alnea, the type of the 

genus Gnomonina, is placed with Gnomonia based on ITS data 

and therefore Gnomonina is considered a synonym of Gnomonia. 

This revised concept of Gnomonia correlates with a number of 

morphological and host characteristics. All species occur on 

decaying leaves of woody trees and shrubs. The perithecia lack a 

stroma. Unlike most other species in the Gnomoniaceae except for 

a few species in Ophiognomonia that become partially erumpent, 

those species of Gnomonia without a neck become erumpent. 

If remaining immersed, the perithecia of species of Gnomonia 

have a short neck and lack a collar. A few species of Gnomonia 

have a collar, specifically G. amoena and G. pseudoamoena. The 

perithecia become concave or collapse from the top when dry, 

as illustrated in Figs 2, 5, 9, and 12, unlike other genera in the 

Gnomoniaceae that collapse from the base. The ascospores are 

ellipsoidal to fusiform, rarely acerose, bicellular with a median, 

occasionally supra-median, septum, or rarely non-septate, with 

8


100 

100 

100 

100 

100 

100 

100 

87 

100 

100 

100 

100 

100 

100 

99 

100 

100 

100 

100 

100 

98 

100 

100 

94 

100 

100 

100 

100 

100 

100 

100 

100 

100 

100 

100 

100 

100 

100 

Leucostoma niveum CBS109489 

Valsella salicis CBS109754 

Valsa ambiens CBS109777 

Diaporthe phaseolorum ATCC64802 

Mazzantia napelli CBS109769 

Plagiostoma amygdalinae CBS791.79 

Plagiostoma euphorbiae CBS340.78 

Plagiostoma fraxini CBS109498 

Plagiostoma devexum CBS123201 

Plagiostoma rhododendri CBS847.79 

Plagiostoma salicellum CBS109775 

Plagiostoma apiculatum CBS121466 

Plagiostoma barriae CBS121249 

Plagiostoma geranii CBS824.79 

Plagiostoma petiolophilum AR3821 

Plagiostoma aesculi CBS109765 

Plagiostoma robergeana CBS121472 

Apiognomonia errabunda CBS109747 

Apiognomonia veneta CBS897.79 

Apiognomonia hystrix CBS911.79 

Apiognomonia borealis CBS799.79 

Amphiporthe hranicensis CBS119289 

Gnomonia amoena CBS121262 

Gnomonia pseudoamoena CBS121261 

Gnomonia gnomon CBS199.53 

Gnomonia skokomishica CBS121245 

Gnomonia orcispora CBS121247 

Gnomonia virginianae CBS121913 

Gnomonia neognomon CBS121265 

Gnomonia rodmanii CBS121909 

Ditopella ditopa CBS109748 

Phragmoporthe conformis CBS109783 

Cryptosporella alnicola CBS121074 

Cryptosporella wehmeyeriana CBS121085 

Plagiostoma 

Apiognomonia 

Gnomonia 

Cryptosporella confusa CBS121063 

Cryptosporella betulae CBS109763 

Cryptosporella hypodermia CBS17169 

Cryptosporella femoralis CBS121076 

Cryptosporella suffusa CBS121007 

Ophiognomonia micromegala CBS121910 

Ophiognomonia pseudoclavulata CBS121236 

Ophiognomonia vasiljevae CBS121253 

Ophiognomonia padicola CBS845.79 

Ophiognomonia rosae CBS121267 

Ophiognomonia sassafras CBS121243 

Ophiognomonia alni-viridis CBS782.79 

Cryptosporella 

Ophiognomonia nana CBS883.79 

Ophiognomonia intermedia CBS119194 

Ophiognomonia balsamiferae CBS121266 

Sirococcus clavigignenti-juglandacearum AR3791 

Ophiognomonia ischnostyla CBS837.79 

Ophiognomonia nervisequa CBS121908 

Ophiognomonia setacea CBS116850 

Ophiognomonia leptostyla CBS844.79 

Pleuroceras oregonense CBS121260 

Pleuroceras pleurostylum CBS906.75 

Pleuroceras tenellum CBS121082 

Apioplagiostoma aceriferum CBS778.79 

Plagiostoma inclinatum CBS772.79 

Ambarignomonia petiolorum CBS121227 

Discula destructiva CBS109771 

Gnomoniopsis comari CBS806.79 

Gnomoniopsis tormentillae CBS904.79 

Gnomoniopsis fructicola CBS121226 

Gnomoniopsis chamaemori CBS803.79 

Gnomoniopsis racemula CBS121469 

Gnomoniopsis macounii CBS121468 

Gnomoniopsis paraclavulata CBS121263 

Sirococcus conigenus CBS101225 

Melanconis alni CBS109773 

Melanconis marginalis CBS109744 

Melanconis stilbostoma CBS109778 

Cryphonectria nitschkei AR3446 

Cryphonectria parasitica ATCC38755 

Cryphonectria cubensis CBS101281 

Amphiporthe 

Gnomoniopsis 

Ophiognomonia 

Pleuroceras 

Ambarignomonia 

Fig. 1. ML phylogenetic analysis (ML score = -lnL 26702.18832) of sequences for the tef1-α, nrLSU and rpb2 multigene analysis of genera in the Gnomoniaceae for 64 

gnomoniaceous taxa and 11 outgroup diaporthalean taxa. Bayesian posterior probabilities greater than 95 % are shown above each branch. MP bootstrap values greater than 

70 % are shown below each branch. 


9


Table 2. Characteristics of genera in the Gnomoniaceae. 

Habit of 

perithecia 

Stroma 

Perithecia 

Ascospores 

Colony 

growth rate 

Conidiomata 

formation in 

culture 

Host 

Gnomonia 

Single on leaves of trees 

and shrubs. 

Without stroma. Some 

species with collar 

around neck. 

Erumpent, concave 

when dry; or remaining 

immersed but then with 

very short necks or with 

collar around neck. 

One median or supramedian 

septum, rarely 

non-septate; ellipsoidal 

to fusiform or acerose, 

appendages short or 

long. 

Without stroma. 

With collar 

around neck. 

One median 

septum, fusiform, 

appendages 

medium. 

Without stroma 

or with weak 

stroma if on 

twigs. 

One septum, 

variable from 

submedian, 

median to 

supramedian, 

ellipsoidal, appendages 

absent 

or present. 

Cryptosporella 

In groups on twigs. 

Ambarignomonia 

Single on leaves 

of trees and 

shrubs. 

Apiognomonia 



shrubs and on 

herbaceous 

plants. In groups 

on twigs. 

Gnomoniopsis 

Single on 

leaves of trees 

and shrubs. 

Single or in 

groups on herbaceous 

plants 

or on twigs. 

Ophiognomonia 

Single on leaves of 

trees and shrubs and 

on herbaceous plants. 

Plagiostoma 



shrubs and on 

herbaceous 

plants. In groups 

on twigs. 

With weak stroma. Without stroma. Without stroma. Without stroma or 

with weak stroma 

if on twigs. 

Perithecia remaining immersed, convex when dry. 

Two species may 

have some irregularly 

shrunk or concave 

perithecia when dry, 

partly erumpent. 

Usually non-septate, 

rarely with one 

median septum, 

ellipsoidal, fusiform, 

femoroid to vermiculate. 

One submedian 

or median septum, 

ellipsoidal, 

slightly broader 

in their upper 

part with no 

apendages. 

One median septum, 

rarely submedian, 

supramedian in filiform 

ascospores, or absent, 

ellipsoidal or fusiform 

(acerose), rarely 

filiform, appendages 

short or long but not 

stout. 

Slow–moderate. Slow. Fast. Slow–moderate. Moderate–fast. Moderate–fast. Fast. 

Rarely. Never. Often, sometimes 

abundant. 

Strictly family Betulaceae, 

mostly subfamily 

Coryloideae. 

Known only from 

Liquidambar 

styraciflua 

(Hamamelidaceae). 

Diverse 

taxonomic 

groups (mostly 

Aceraceae, 

Fagaceae, 

Geraniaceae, 

Platanaceae, 

occasionally 

Anacardiaceae, 

Hippocastanaceae, 

Juglandaceae, 

Onagraceae, 

Rosaceae, 

Tiliaceae). 

In some species 

none, in some 

abundant. 

Betulaceae, Tiliaceae, 

Ulmaceae. 

Usually abundant. 

Diverse taxonomic 

groups 

(Ericaceae, 

Fagaceae, 

Rosaceae, 

Tiliaceae). 

Rarely. 

Mostly Fagales (Betulaceae, 

Fagaceae, 

Juglandaceae), a few 

species on Lauraceae, 

Rosaceae, Salicaceae, 

Tiliaceae. 

One median 

septum, rarely 

submedian or absent, 

ellipsoidal, 

appendages absent 

or present. 

Often, sometimes 

abundant. 

Diverse 

taxonomic groups 

(Aceraceae, 

Euphorbiaceae, 

Geraniaceae, 

Hippocastanaceae, 

Oleaceae, 

Polygonaceae, 

Salicaceae, 

Staphyleaceae). 

short to long appendages. The colonies in culture grow at a slow to 

moderate rate and rarely form conidiomata in culture. Similar to the 

stromatic Cryptosporella, the genus Gnomonia occurs primarily on 

members of the Betulaceae. 

The genus Ambarignomonia is established for the distinctive 

species, A. petiolorum, that is common on Liquidambar styraciflua 

(Hamamelidaceae), native to North America. Easy to recognise 

because of the white collar around the relatively long neck of the 

perithecia, Ambarignomonia is otherwise similar to members of 

the Gnomoniaceae in their occurrence on fallen leaves, lack of 

stromatic development, and perithecia that remain immersed in the 

substrate and collapse from the top when dry. The ascospores are 

fusiform, have one median septum, and bear appendages at both 

ends. The colonies are relatively slow-growing and do not produce 

conidiomata in cultures. In all analyses A. petiolorum appears to be 

unique among species in the Gnomoniaceae. 

The type species of Apiognomonia, A. veneta, and a second 

species, A. errabunda, were redescribed by Sogonov et al. (2007). 

In the present work three additional species have been determined 

to be congeneric with these species including A. hystrix, on woody 

substrates. Apiognomonia includes species producing solitary 

perithecia without a stroma or with a weakly developed stroma 

on decaying leaves and twigs. The perithecia remain immersed 

and become convex or collapse from the base when dry. The 

ascospores have one septum that is variable in placement ranging 

from median to supramedian. They are ellipsoidal with or without 

appendages. In culture species of Apiognomonia are relatively 

fast-growing and often produce abundant conidiomata. Species of 

Apiognomonia occur on a wide variety of woody plant hosts in the 

Aceraceae, Fagaceae, and Plantanaceae as well as herbaceous 

families such as the Anacardiaceae, Geraniaceae, Onagraceae, 

and Rosaceae. 

10


Fig. 2. Morphology on natural substrates, perithecia. A, B. Gnomonia gnomon. A. Epitype BPI 844273. B. BPI 596632. C–E. G. alnea. C, E. Epitype BPI 877462A. D. BPI 

799019. F, G. G. incrassata, holotype BPI 611818A. H, I. G. monodii, holotype BPI 877499A. A, C, D, F, H. Intact air-dry perithecia on leaves. B, E, G, I. Extracted and rehydrated 

perithecia. Scale 200 μm. 

The concept of Gnomoniopsis is herein expanded to include 

the type, G. chamaemori, and six additional species. Perithecia are 

generally single, rarely in groups, on decaying leaves or twigs of 

woody trees, shrubs or herbaceous plants. No stromatic tissues 

are associated with the perithecia. The perithecia remain immersed 

in the substrate and become convex collapsing from the base 

when dry. The ascospores are ellipsoidal, slightly broader in the 

upper portion, have one submedian or median septum, and lack 

appendages. In cultures these fungi are moderately fast growing 

and usually produce abundant conidiomata on PDA. Species of 

Gnomoniopsis occur on a vast range of plant families including 

the Ericaceae, Fagaceae, Rosaceae, and Tiliaceae. Ditopellopsis 

racemula is herein placed in Gnomoniopsis. 

Many species previously regarded as belonging to Gnomonia 

are now placed in Ophiognomonia. The perithecia occur singly on 

leaves of woody trees and shrubs as well as herbaceous plants. 

They lack a stroma and remain immersed becoming convex upon 

drying, although two species, O. balsamiferae and O. melanostyla, 

are partially erumpent and were found to collapse irregularly upon 

drying. The ascospores are ellipsoidal to fusiform with pointed ends, 

rarely filiform, have one median septum, with or without appendages 

of variable length. The cultures are moderately fast growing, rarely 

producing conidiomata. Most species of Ophiognomonia occur on 

members of the Fagales including the Betulaceae, Fagaceae, and 

Juglandaceae, but some also have been reported from other plant 

families. 

The genus Plagiostoma is herein recognised to include the type 

species P. euphorbiae, one new species, and eleven additional 


11


species transferred from other genera. The type species of the 

genus Cryptodiaporthe, C. aesculi, groups with P. euphorbiae and 

its relatives, thus Cryptodiaporthe is considered a synonym of 

Plagiostoma. Perithecia of Plagiostoma occur singly or in groups on 

leaves and twigs of woody trees and shrubs as well as herbaceous 

plants. Often the perithecia lack a stroma. Like all genera of the 

Gnomoniaceae dealt with in this study except Ambarignomonia 

and Gnomonia, the perithecia remain immersed in the substrate 

becoming convex with the base collapsing upward when dry. The 

ascospores are ellipsoidal, have one median septum that is rarely 

submedian or absent, and may or may not bear appendages. The 

species grow relatively fast in culture and often produce abundant 

conidiomata on PDA. Species of Plagiostoma occur on a diverse 

range of woody and herbaceous hosts. 

Evaluation of morphological and host characteristics 

The groupings of species based on the multigene phylogeny 

presented here suggest that the morphological characters 

previously used to define genera must be re-evaluated. The 

generic classification proposed by Barr (1978) and Monod (1983) 

are presented as rectangular tables, referred to as a “pigeon-hole” 

system in which columns and rows show genera corresponding 

to ascospore and perithecial/stromatal characteristics. The 

phylogenies resulting from the analysis of multiple genes do not 

agree with such a rigid definition of genera based on one or two 

characteristics. Little congruence can be found between the newly 

defined genera based on molecular data and the genera based on 

a single morphological characteristic. 

Host specificity is an important character in circumscription of 

genera and species of the Gnomoniaceae. The genus Gnomonia is 

almost strictly associated with plant hosts in the Betulaceae, mostly 

in the subfamily Coryloideae. Likewise, most of the species of 

Gnomonia are limited in their host range to a single genus and often 

to a single plant species. For example, the type species, G. gnomon, 

is restricted to species of Corylus except for one collection reported 

from a Populus seedling that may be an accidental colonisation 

of a non-specific host. The monotypic Ambarignomonia with A. 

petiolorum is likewise restricted to one plant host, Liquidambar 

styraciflua. The other genera of the Gnomoniaceae do not show 

such consistency in host associations. The genus Apiognomonia 

has the most diverse range of hosts that include hardwood trees 

as well as herbaceous plants. Species of Apiognomonia exhibit a 

diversity of host specificity with some species such as A. veneta 

occurring on plants in at least 10 different plant families while 

others such as A. acerina are restricted to one plant host. Species 

of Gnomoniopsis are mostly associated with either Fagaceae or 

Rosaceae also with the range of host specificity varying among 

species. While the type species G. chamaemori appears to be 

restricted to Rubus chamaemorus, other species are specific at the 

level of host genus such as G. clavulata on Quercus spp. Species 

of Ophiognomonia occur predominantly on members of the Fagales 

with some exceptions such as the type species O. melanostyla that 

infects overwintered leaves of Tilia spp. The genus Plagiostoma 

shows a broad host range with species occurring on a variety of 

hosts such as P. devexum on Persicaria, Polygonum, and Rumex 

(Polygonaceae) while others are species specific such as P. 

euphorbiae is known only from Euphorbia palustris. Few members 

of the Gnomoniaceae are known to infect hosts outside of the 

dicotyledonous plants; the asexual genus Sirococcus on conifers 

provides one exception (Rossman et al. 2007). 

Members of the Gnomoniaceae occur most commonly on 

fallen or still attached, overwintered leaves including petioles or 

herbaceous stems although some occur on woody substrates such 

as species of Cryptosporella but also, for example, Apiognomonia 

hystrix and Plagiostoma salicellum. When ascomata develop 

on woody substrates, these are often found on relatively small 

branches, one or two years old, that are dead but still attached 

to the host tree. A number of species of Gnomoniaceae have 

been reported as endophytes of woody plants (Viret & Petrini 

1994, Cohen 1999, 2004, Danti et al. 2002, Vujanovic & Britton 

2002, Green 2004, Moricca &Ragazzi 2008) but these are often 

not accurately identified. In addition, some species are pathogenic 

such as Apiognomonia veneta, cause of sycamore anthracnose, 

and Gnomoniopsis fructicola, cause of strawberry stem rot (Maas 

1998). 

One morphological character that was emphasised in earlier 

classification systems is the type and extent of stroma (Kobayashi 

1970, Barr 1978, Monod 1980, Vasilyeva, 1998). No members 

of the Gnomoniaceae have a well-developed stroma. Species of 

Cryptosporella as well as Amphiporthe hranicensis, Apiognomonia 

hystrix, and Plagiostoma salicellum, i.e. species that occur on 

woody substrates, produce limited stromatic tissues. 

These stromatic tissues may be associated with the rupture 

through the surface of the substrate. In addition, three species of 

Linospora, not considered in this study, produce a layer of tissue 

that covers the aggregated ascomata that are superficial or slightly 

immersed on leaves (Barr 1978, Monod 1983). In other families of 

the Diaporthales such as the Cryphonectriaceae, Diaporthaceae, 

Pseudovalsaceae, and Valsaceae, stromata are often welldeveloped 

(Castlebury et al., 2002, Gryzenhout et al. 2006, 

Voglmayer & Jaklitsch 2008). 

Most members of the Gnomoniaceae including all species 

of Ambarignomonia, Gnomonia, and Ophiognomonia, produce 

ascomata singly, immersed, although some species of Gnomonia 

become erumpent. Similar to the development of a rudimentary 

stroma, the formation of grouped perithecia appears to be more 

common in species that develop on woody substrates. This is 

exemplified by Apiognomonia hystrix. Apiognomonia hystrix as 

Cryptodiaporthe hystrix was traditionally placed in the Valsaceae 

(Barr 1978) because of its occurrence on woody substrates with 

ascomata developing a rudimentary stroma. One of its synonyms, 

Gnomonia cerastis, was traditionally placed in the Gnomoniaceae 

due to its occurrence on overwintered leaves with ascomata 

lacking any stroma. As suggested by Monod (1983), specimens of 

A. hystrix are known to occur both on woody substrates as well 

as on overwintered leaves. Some species of Gnomoniopsis and 

Plagiostoma, i.e. P. salicellum, produce grouped perithecia on 

woody substrates. All species of Cryptosporella occur on woody 

substrates and produce ascomata in groups (Mejia et al. 2008). 

Ascomata of species of Gnomoniaceae are perithecial, i.e. no 

cleistothecial members are known, dark brown to black, smooth, 

with or without an elongated neck. In a few species, the neck is 

surrounded by a distinct, powdery collar. The perithecial walls are 

thin-walled, less than 30 μm diam, composed of only one or two 

regions. The outer region is composed of textura angularis with cell 

walls dark brown, slightly thickened, 1–2 μm. The inner region is 

composed of hyaline, elongate cells. The structure of the ascomata 

is relatively constant within the family. However, one characteristic of 

the ascomata that has some taxonomic significance is the collapse 

upon drying. In species of Ambarignomonia and Gnomonia the 

ascomata collapse from the top becoming concave when dry while 

in other members of the Gnomoniaceae the ascomata collapse from 

12


the base becoming convex when dry. This difference in collapse 

appears to be associated with the structure of the ascomata in 

which the basal wall is relatively thin compared to the side walls 

(Klebahn 1918). 

One of the morphological characters formerly thought to be 

taxonomically significant at the generic level is the position of 

the neck categorised as either lateral or central. This character is 

sometimes difficult to assess in terms of discrete categories. Many 

species have necks that are centrally located. Necks may also 

be eccentric, e.g. not arising from the centre of the perithecium, 

in which case they can be either marginal and lateral. The term 

“lateral” is used herein only if the neck emerges from the margin 

of the perithecium and is oriented horizontal to the perithecium, 

at least at the base. Necks rising vertically from the margin of the 

perithecium are described as “marginal”. Necks often are eccentric 

i.e. not positioned in the exact centre of the perithecium, but neither 

are they truly central. We intentionally use the term “marginal” 

for the position of the neck in these genera to distinguish it from 

“lateral” in the narrow sense. This character may vary even within a 

single species e.g. Ophiognomonia setacea and does not correlate 

with the phylogenetically defined genera. Nevertheless, tendencies 

exist for some genera to have the neck in a certain position on the 

perithecium. For example, most species in Ophiognomonia have 

a central or slightly eccentric neck whereas eccentric to marginal 

necks are more common in other genera. Truly lateral necks do not 

occur in the genera treated in this paper although they are common 

in Pleuroceras (Barr 1978, Monod 1983). Most species have only 

one neck but one species, Gnomonia carpinicola, has perithecia 

each with 2–3 necks emerging from both sides of a leaf blade. 

In the Gnomoniaceae the asci are generally broadly clavate 

to broadly cylindric with a conspicuous ascal ring always present. 

The width of the ascal ring may vary and, for Gnomonia orcispora, 

the ascal ring that is over 4.5 μm diam is diagnostic. The shape of 

the ascus base varies from being rounded to narrowing to the base 

with a distinct stalk. Asci may accumulate at the top of the often 

elongated necks. All species of Gnomoniaceae have eight-spored 

asci except Ditopella ditopa which has 32 ascospores in each 

ascus. The arrangement of the ascospores in the asci varies with 

the shape of the ascospores from obliquely distichous for shorter 

ascospores to irregularly parallel for elongated ascospores. 

Within the Gnomoniaceae ascospores are hyaline but vary 

considerably in shape and septation including location of the 

septum, characters that traditionally been important for defining 

genera (Barr 1978, Monod 1983, Vasilyeva 1998). Ascospore 

shape and septation including the placement of the septum within 

the ascospore is no longer considered an important character for 

defining genera within the Gnomoniaceae. Ascospores range in 

shape from oval or short-fusiform to long cylindric and most species 

of Gnomoniaceae have non- or one-septate ascospores although 

Phragmoporthe conformis and three species of Pleuroceras have 

multi-septate ascospores. Species now known to be congeneric 

differ in ascospore shape and septation. Species with elongate, 

one-septate ascospores such as Ophiognomonia melanostyla and 

O. sassafras are congeneric with species having oval or fusiform 

ascospores such as O. intermedia and O. pseudoclavulata. 

Previously all species with non-septate ascospores were placed 

in the genus Gnomoniella; however, based on this study, species 

having non-septate ascospores are placed Ophiognomonia, O. 

nana, and Plagiostoma, P. euphorbiae-verrucosae and P. fraxini. 

Considerable variation in ascospore morphology occurs in the 

wood-inhabiting genus Cryptosporella that range from ellipsoid or 

fusiform to femoroid to elongated cylindric although most of the 

ascospores are non-septate (Mejia et al. 2008). 

The location of the septum within the ascospore is variable in 

the Gnomoniaceae ranging from central i.e. in the middle of the 

ascospore, to below the septate, referred to as submedian or above 

the septum, referred to as supramedian. In this study the location of 

the septum was measured as the percent of the total length above 

the base of the ascospore, thus centrally located septa are generally 

45–55 %. Within genera, location of the ascospore septum may 

vary from usually median in Ophiognomonia and Plagiostoma to 

median and supramedian in Gnomonia or median and submedian 

in Gnomoniopsis, and submedian, median or supremedian in 

Apiognomonia. At the species level, the location of the septum is 

consistent. 

Many species of Gnomoniaceae bear appendages at both ends 

of the ascospore. These appendages vary in length from very short, 

stout to rather long, filiform but the appendages never envelop 

the entire ascospore. Within genera, the presence or absence of 

appendages is variable, although no species of Gnomoniopsis are 

known to have appendages. At the species level, the presence or 

absence of appendages is a useful diagnostic character although 

this may vary within species. 

Anamorphs of members of the Gnomoniaceae have been placed 

in a number of genera including Cylindrosporella, Discula, Disculina, 

Gloeosporium and Neomarssonina but in general the anamorphs 

are similar within genera. All anamorphs in Cryptosporella for which 

anamorphs are known have been placed in Disculina (Sutton 

1980). The conidia of species in the Gnomoniaceae are primarily 

non-septate, hyaline, and slimy. The asexual state appears in leaf 

spots often on the surface opposite the sexual state in late summer 

prior to leaf fall. Then, the sexual state develops on overwintered 

leaves in the spring as noted by Klebahn (1918). The nomenclature 

for the asexual states is complicated and is not dealt with here. 

Cultures of species of Gnomoniaceae usually produce pale to 

dark grey, brown or black pigments, often with other colors such as 

yellow and orange that diffuse into the media. This pigmentation 

may vary considerably within a single species. Perithecia develop 

in isolates of some species relatively quickly within 2 wk while 

others will do so after several mo at the warm/cold light/dark regime 

described in the Materials and Methods. Some species never 

produced perithecia or conidia in culture. 


13


TAXONOMY 

Following is a key to the 59 species of Gnomoniaceae included in this study. These represent the commonly encountered species in the five 

genera treated here. 

Key to the species of Gnomoniaceae in this study 

1. Perithecia immersed in woody substrates, developing in groups, with perithecial necks oriented toward the centre, 

often on dead, still attached, one–two year old branches .......................................................................................................................... 2 

1’. Perithecia immersed or erumpent on overwintered, fallen or attached leaves or on dead herbaceous stems; 

not grouped with necks oriented toward the centre ................................................................................................................................... 7 

2. Ascospores non-septate, ellipsoid to cylindric including femoroid, with broadly rounded ends ........ Cryptosporella (see Mejia et al. 2008) 

2’. Ascospores one-septate, ellipsoid to fusiform ........................................................................................................................................... 3 

3. On overwintered twigs and branches of Acer spp. .................................................................................................................................... 4 

3’. On overwintered twigs and branches of woody hosts other than Acer, specifically Aesculus, Salix and Spiraea ..................................... 5 

4. Ascospores 14–20 × 2–2.5 µm fide Barr (1978). On Acer pseudoplatanus and various other hardwoods .............. Apiognomonia hystrix 

4’. Ascospores 7–12 × 1–2.5 µm fide Barr (1978). On Acer saccharum and A. spicatum .................................... Plagiostroma petiolophilum 

5. Ascospores 7.5–10 × 1.5–2.5 µm fide Barr (1978). On Spiraea .......................................................................... Gnomoniopsis macounii 

5’. Ascospores greater than 10 µm long ........................................................................................................................................................ 6 

6. On Aesculus; ascospores 14–23 × 4.5–7 µm fide Barr (1978) ................................................................................... Plagiostoma aesculi 

6’. On Salix; ascospores 11–20 × 4.5–6 µm fide Barr (1978) ...................................................................................... Plagiostoma salicellum 

7. Individual necks surrounded with whitish powdery collars. Perithecia concave when dry, immersed in the 

substrate, on petioles or basal parts of major leaf veins ............................................................................................................................ 8 

7’. Lacking a collar around perithecial neck or neck lacking. Perithecia convex or concave when dry, in the 

latter case (partly) erumpent upon maturation ......................................................................................................................................... 10 

8. Ascospores (9–)11–12.5(–15) × 1.5–2 µm. On Liquidambar ......................................................................... Ambarignomonia petiolorum 

8’. Ascospores 2.5–3.2 μm wide. On Betulaceae ........................................................................................................................................... 9 

9. Ascospores 17–23 × 2.5–3 μm fide Monod (1983). On Carpinus betulus in Europe and U.S.A. (TN) ......................... Gnomonia amoena 

9’. Ascospores 11–14.5 × 2.7–3.2 μm fide Monod (1983). On Corylus avellana in Europe and Canada .............. Gnomonia pseudoamoena 

10. Perithecia small, to 200 μm diam, with 2–3 necks opening on both sides of a leaf blade. Ascospores 

septum submedian, 12–15 × 2.7–4 μm fide Monod (1983). On Carpinus betulus in Europe ................................... Gnomonia carpinicola 

10’. Perithecia with one ostiole ....................................................................................................................................................................... 11 

11. Perithecia concave when dry, (partly) erumpent upon maturation. Ascopores oval to fusiform with septum 

median or supramedian but ascospores always widest close to their middle. On Betulaceae ................................................................ 12 

11’. Perithecia convex when dry, remaining immersed in the substrate. If perithecia irregularly dented or concave 

when dry and erumpent with maturation, then ascospores filiform and upper cell wider than the lower, and septum supramedian ....... 23 

12. Lacking elongated perithecial necks. On Alnus, Carpinus or Ostrya ....................................................................................................... 13 

12’. Elongated perithecial necks present, at least on fully mature perithecia; elongated necks may be absent 

in immature perithecia on Corylus. On Carpinus, Corylus or Ostrya ....................................................................................................... 14 

13. On Alnus. Ascospores (13.5–)16–17.5(–20.5) × (3.5–)4–4.5(–5) µm .............................................................................. Gnomonia alnea 

13’. On Carpinus or Ostrya. Ascospores 17–23 × 3.5–4.5 μm fide Monod (1983) ..................................................... Gnomonia arnstadtiensis 

14. Ascospore septum median to slightly supramedian. Ascospores fusiform. Necks central, occasionally eccentric ................................. 15 

14’. Ascospore septum distinctly supramedian. Ascospores oval to fusiform. Necks eccentric to marginal except 

in G. incrassata and G. monodii ............................................................................................................................................................... 19 

15. Ascospores (13.5–)15–16.5(–18.5) × 2–2.5 µm. On Carpinus caroliniana in Georgia, U.S.A. ................................... Gnomonia rodmanii 

15’. On Corylus ............................................................................................................................................................................................... 16 

14


16. Ascospore septum slightly supramedian, located at 56 %. Ascospores (16.5–)17.5–19(–20.5) × 2–2.5(–3) µm. 

On Corylus californica in Washington, U.S.A. ....................................................................................................... Gnomonia skokomishica 

16’. Ascospore septum median, located at 48–50 %, mean ascospore length exceeding 19 μm .................................................................. 17 

17. Ascospores (20–)21–22(–23) × (2.5–)3(–3.5) µm. On Corylus californica in North America ................................. Gnomonia pendulorum 

17’. Ascospores narrower than 2.5 μm ........................................................................................................................................................... 18 

18. Ascospores (17–)19.5–21(–24.5) × 1.5–2(–2.5) µm. Perithecial necks central. On Corylus avellana in Europe ......... Gnomonia gnomon 

18’. Ascospores (18.5–)20–22.5(–24.5) × 2–2.5 µm. Perithecial necks eccentric. On Corylus californica in North America ........ Gnomonia neognomon 

19. Ascospores (14–)15.5–17.5(–19.5) × (4.5–)5–5.5(–6) µm, oval, ends blunt, distinct hila on bases of 

evanescent appendages; two large guttules per cell. Apical ring exceeding 4.5 μm diam. Necks marginal. 

On Corylus californica in North America ....................................................................................................................... Gnomonia orcispora 

19’. Ascospores oval to fusiform or lanceolate, tapering toward ends or ends blunt, no hila present. Apical 

ring less than 4 μm diam. Necks marginal, eccentric to central. On Corylus avellana in Europe or Ostrya 

in Europe or North America ...................................................................................................................................................................... 20 

20. Ascospores mostly lanceolate or oval, broadened in upper part and distal ends broadly rounded or 

nearly truncated. Ascospores (12.5–)15–17(–20.5) × (3–)3.5–4.5(–6) µm. On Ostrya carpinifolia in Europe ............. Gnomonia ostryae 

20’. Ascospore fusiform, tapering to both ends. Mean ascospore width smaller than 3.5 μm. 

On Corylus avellana in Europe or Ostrya virginiana in North America .................................................................................................... 21 

21. Necks marginal. Ascospores (12–)13–14(–14.5) × (2–)2.5–3 µm. On Ostrya virginiana in North America ............... Gnomonia virginianae 

21’. Necks eccentric or central. Mean ascospore length exceeding 14.5 μm. On Corylus avellana in Europe .............................................. 22 

22. Ascospores (13.5–)15.5–17(–18.5) × (2.5–)3–3.5(–4) µm, strongly constricted at septum ...................................... Gnomonia incrassata 

22’. Ascospores (14–)15–16(–18.5) × (2–)2.5(–3) µm, not or slightly constricted at septum ............................................... Gnomonia monodii 

23. Ascospores one-celled ............................................................................................................................................................................ 24 

23’. Ascospores two-celled ............................................................................................................................................................................. 26 

24. Ascospores 20–22.5 × 5.3–6 μm fide Monod (1983), with pointed ends. On Euphorbia in Europe.... Plagiostoma euphorbiae-verrucosae 

24’. Ascospores smaller, ends rounded .......................................................................................................................................................... 25 

25. Necks shorter than 200 μm. Ascospores (7.7–)8.6–12.7(–13.8) × (2.2–)2.8–5.9(–6.6) μm fide 

Redlin & Stack (1988). On Chionanthus and Fraxinus (Oleaceae) .............................................................................. Plagiostoma fraxini 

25’. Necks longer than 400 μm. Ascospores 8–10 × 2.5–4 μm fide Monod (1983). On Betula nana in Europe .............. Ophiognomonia nana 

26. Ascospores filiform, upper cell wider than lower cell. Necks long ............................................................................................................ 27 

26’. Ascospores oval, clavate or fusiform with variable septum position, or, if ascospores filiform, then 

septum median and cells of equal width. Necks short or long ................................................................................................................. 28 

27. Ascospores (30–)37–42.5(–44) × 1.5–2 µm. On Tilia spp., known from Europe and North America ................ Ophiognomonia melanostyla 

27’. Ascospores 38–65 × 1 μm fide Barr (1978). On Sassafras ............................................................................... Ophiognomonia sassafras 

28. Ascospore septum supramedian, 15–17 × 3.7–4.5 μm fide Monod (1983). On Geranium spp. in Europe ............ Apiognomonia borealis 

28’. Ascospore septum median or submedian ................................................................................................................................................ 29 

29. Ascospore septum submedian, if septum nearly median, than upper cell wider than lower, ascospores clavate ................................... 30 

29’. Ascospore septum median ...................................................................................................................................................................... 39 

30. Ascospores 13–16 × 4–5 μm fide Monod (1983), lower cell of ascospore conical, upper cell rounded. 

On overwintered but still attached pedicels and branches of Rhododendron spp. in Europe .............................. Plagiostoma rhododendri 

30’. Both cells of ascospores of similar shape ................................................................................................................................................ 31 

31. Majority of asci clavate with base tapering to long, narrow stalk. Apical ring exceeding 1.8 μm diam, 

bluntly hexagonal in side view. Ascospores equally wide in upper and lower parts or wider in lower part .............................................. 32 

31’. Majority of asci allantoid or obpyriform with bases rounded or shortly tapering. Apical ring less than 

2.5 μm diam, more or less circular in side view. Ascospores usually slightly wider in upper part ............................................................ 35 


15


32. Ascospores straight or slightly curved, 10–13 × 2–2.5 μm fide Monod (1983). 

On Geum montanum in the Alps (Europe) ...........................................................................................................................Ophiognomonia gei-montani 

32’. Ascospores curved, wider than 3.5. On other hosts ................................................................................................................................ 33 

33. Perithecia exceeding 250 μm diam on average. Ascospores 15–24 × 4.5–5.5 μm fide Monod (1983). On Acer spp. in Europe ..........Apiognomonia acerina 

33’. Perithecia less than 250 μm diam on average. Ascospores shorter than 17 μm and narrower than 

6 μm on average. Mostly on Fagus, Platanus, Quercus, and Tilia ........................................................................................................... 34 

34. Ascospores (13–)15.5–17.5(–23) × (3.5–)5–5.5(–7.5) μm fide Sogonov et al. (2007). 

On Platanus spp. in temperate regions ..................................................................................................................... Apiognomonia veneta 

34’. Ascospores (10.5–)15–16.5(–19.5) × (3.5–)4.5–5.5(–6.5) μm fide Sogonov et al. (2007). 

On various hosts in temperate regions ............................................................................................................... Apiognomonia errabunda 

35. Perithecia in groups of 3–9. Ascospores 7.5–11 × 2–3.5 μm fide Barr (1978). 

On Chamerion angustifolium in North America ..................................................................................................... Gnomoniopsis racemula 

35’. Perithecia scattered singly ....................................................................................................................................................................... 36 

36. Mean l:w of ascospores exceeding 3.5, upper ascospore cell slightly wider than lower. On Rosaceae .................................................. 37 

36’. Mean l:w of ascospores smaller than 3.5, upper ascospore cell distinctly wider than lower. On Fagaceae ............................................ 38 

37. Necks marginal. Ascospores 6.5–9 × 1.5–2 μm fide Monod (1983). On Potentilla spp. in Europe and North America .... Gnomoniopsis tormentillae 

37’. Necks central. Ascospores (10–)10.5–11.5(–13) × (2–)2.5(–3) µm ................... Gnomoniopsis chamaemori, G. comari, and G. fructicola 

(These three species are morphologically similar. Additional work is required to clarify their morphological limits.) 

38. Septum located above 37 % of ascospore length in the ascus. Ascospores (5–)8.5–9.5(–11) × (2–)3.5–4(–5.5) µm ....... Gnomoniopsis clavulata 

38’. Septum located below 37 % of ascospore length in the ascus. Ascospores (8–)9–10(–11) × (3–)3.5–4 µm . Gnomoniopsis paraclavulata 

39. Ascospores with l:w smaller than 3 .......................................................................................................................................................... 40 

39’. Ascospores with l:w exceeding 3.5 .......................................................................................................................................................... 41 

40. Necks 140–250 μm. Ascospores (6.5–)7.5–8(–9) × (2.5–)3–3.5 µm. On Carya in North America ............ Ophiognomonia pseudoclavulata 

40’. Necks 300–500 μm long. Ascospores 8–11 × 2.2–3 μm fide Monod (1983). 

On Alnus viridis in Europe and Canada (BC) .................................................................................................... Ophiognomonia trientensis 

41. Ascospores filiform, 39–51 × 1 μm fide Monod (1983). Necks central. On Prunus padus .................................. Ophiognomonia padicola 

41’. Ascospores shorter or ascospore width exceeding 4.2 μm ..................................................................................................................... 42 

42. Ascospores 26–36 × 5.5–10 μm fide Barr (1978). Necks eccentric or lateral, stout. On Carya ................... Ophiognomonia micromegala 

42’. Ascospores shorter .................................................................................................................................................................................. 43 

43. On Rosaceae. Necks typically tapering to pointed ends. Asci with long, narrow stipe at the base. 

Ascospores 13–22 × 1–1.5 μm fide Monod (1983), with long, filamentous appendages ............. Ophiognomonia rosae, Ophiognomonia 

rubi-idaei 

(These species cannot be reliably distinguished based on morphology. O. rubi-idaei occurs only on Rubus while O. rosae is found on various 

Rosaceae.) 

43’. Not on Rosaceae. Necks cylindrical or slightly tapering, their width exceeding 25 μm below apex ........................................................ 44 

44. Necks marginal. Ascospores 8–10 × 2–3 μm fide Monod (1983) On Persicaria and Polygonum, rarely on Rumex and 

Vitis .......................................................................................................................................................................... Plagiostoma devexum 

44’. Necks central or eccentric, few in a collection marginal. Ascospores generally longer than 10 μm. On other hosts ............................... 45 

45. Necks shorter than 170 μm. Ascospores shorter than 18 μm .................................................................................................................. 46 

45’. Necks mostly longer or, if necks shorter than 170 μm, then ascospores longer than 18 μm on average ................................................ 49 

46. Ascospores (11.5–)14–15.5(–17.5) × (2.5–)3.5–4(–4.5) µm. On Acer in Washington, U.S.A. .................................... Plagiostoma barriae 

46’. On Euphorbia in Europe .......................................................................................................................................................................... 47 

47. Necks shorter than 100 μm. Ascospores (12–)13–13.5(–15.5) × (3–)3.5(–4) µm ................................................ Plagiostoma euphorbiae 

47’. Necks longer than 100 μm ....................................................................................................................................................................... 48 

48. Ascospores 14–17.5 × 3.5–4.5 μm fide Monod (1983) .................................................................................. Plagiostoma euphorbiaceum 

48’. Ascospores 13–15.5 × 2.3–3 μm fide Monod (1983) .......................................................................................... Plagiostoma amygdalinae 

16


49. On dead stems of herbaceous plants, Geranium spp. in Europe. Ascospores 13–18 × 1.8–2.5 μm fide Monod (1983) .... Plagiostoma geranii 

49’. On overwintered leaves, one species also on twigs of trees and shrubs ................................................................................................ 50 

50. Necks shorter than 250 μm. Ascospores19–23 × 3.5 μm fide Monod (1983). On Juglans spp. in Europe and 

North America .................................................................................................................................................... Ophiognomonia leptostyla 

50’. Necks typically longer than 250 μm. If necks shorter than 250 μm, then ascospores shorter than 19 μm on average ........................... 51 

51. Necks 940–1150 μm. Ascospores (15–)18–19(–21) × 2.5–3(–3.5) µm. On Populus ....................................Ophiognomonia balsamiferae 

51’. Necks shorter; if longer than 900 μm, then ascospores shorter than 15 μm ........................................................................................... 52 

52. Ascospores (17.5–)18.5–19.5(–21) × (2.5–)3(–3.5) µm. Ascal apical ring 2.7–3.2 μm diam. On Juglans..........Ophiognomonia vasiljevae 

52’. Ascospores shorter than 17.5 μm on average ......................................................................................................................................... 53 

53. Ascal apical ring 3–3.5 μm diam. Ascospores 11–18 × 1.5–2.5 μm fide Monod (1983). On twigs of 

broad range of hosts, predominately on Acer ............................................................................................................ Apiognomonia hystrix 

53’. Ascal apical ring less than 2.7 μm diam. Ascospores mostly narrower than 2.6 μm. If ascospores wider 

than 2.6 μm, then ascospores shorter than 12 μm .................................................................................................................................... 54 

54. Ascospores (10–)11.5–13.5(–17) × (1.5–)2–2.5 µm with or without cuneiform appendages. 

On Castanea, Fagus and Quercus (Fagaceae) .................................................................................................... Ophiognomonia setacea 

54’. On Betulaceae, occasionally on other hosts ............................................................................................................................................ 55 

55. Ascospores (12.5–)13.5–15.5(–18.5) × (1.5–)2(–2.5) µm with cuneiform or long appendages. 

On Alnus, Betula, Corylus and Carpinus, possibly also Ostrya ....................................................................... Ophiognomonia ischnostyla 

55’. Ascospores generally smaller than 12 µm .............................................................................................................................................. 56 

56. Ascospores (9.5–)10–11(–13.5) × (2.2–)2.5–3(–3.6) µm without appendages. On Alnus and Betula ............. Ophiognomonia intermedia 

56’. Ascospores (10–)11–11.5(–12.5) × 2–2.5 µm with long appendages. On Alnus and Betula, 

occasionally Myricaria and Vaccinium............................................................................................................... Ophiognomonia alni-viridis 


17


DESCRIPTIONS OF GENERA AND SPECIES OF THE 

GNOMONIACEAE 

Following are the descriptions of five genera and their generic 

synonyms representing the the most common leaf-inhabiting 

species in the Gnomoniaceae. The genus Cryptosporella is not 

included as it has been dealt with elsewhere (Mejia et al. 2008). 

A description for each of the five genera is included along with 

a description of the type species and the type species of any 

synonymous genera. Thirteen new species are described in full. In 

addition, many new combinations are made for taxa that should be 

placed in the phylogenetically defined genera. 

GNOMONIA Ces. & De Not., Comment. Soc. Crittog. Ital. 1: 231. 

1863. Type: G. gnomon (Tode : Fr.) J. Schröt., designated by 

Höhnel (1917). 

= Gnomonina Höhn., Ber. Deutsch. Bot. Ges. 35: 635. 1917. Type: Gnomonina 

alnea (Fr.) Höhn., herein recognised as Gnomonia alnea (Fr.) Sogonov 

[≡ Laestadia Auersw., Hedwigia 8: 177. 1869 non Kunth ex Lessing, 

1832.] 

Perithecia solitary, without stroma, on overwintered, fallen or 

attached leaves of trees and shrubs, usually epiphyllous or on 

petioles, rarely hypophyllous. Perithecia black, immersed at 

first, later erumpent, rarely partly erumpent or with wide opening 

with white or pink powdery collar surrounding the neck, powdery 

substance not dissolving in water or 3 % KOH solution. Perithecia 

oblate-spheroidal to oblate when moist, concave when dry, circular 

in top view, rarely perithecia convex, nearly flat, with one neck, 

occasionaly with two or three necks emerging on both sides of a 

leaf blade. Necks central to marginal, never truly lateral, slightly 

curved to distinctly curved, their length mostly 1–2, sometimes 

to 5 times the perithecial diam, sometimes absent. Asci oval to 

fusiform, with an apical ring, with eight spores arranged unevenly 

parallel or irregularly multiseriate, occasionally obliquely uniseriate. 

Ascospores two-celled, fusiform to acerose, l:w 3–15, ends rounded; 

appendages cuneiform with diffuse ends or ovoid, subulate, or 

acicular, rarely absent. 

Cultures: Colonies reaching (0.5–)1–4 cm diam after 2 wk at 23 

°C dark/light on MEA/MYA and PDA. Colony surface velvety, pale 

grey, yellowish grey, greyish yellow, brownish grey, or brown. In 

some species fertile perithecia formed after 5–6 mo at 2/10 °C l/d 

on MEA, MYA, and PDA, rarely sterile perithecia noted within one 

month at 23 °C l/d. Conidiogenous structures not produced. 

Hosts: Limited to family Betulaceae, mostly in the subfamily 

Coryloideae. Individual fungal species are host specific at plant 

species level or, less commonly, at genus level. 

Gnomonia gnomon (Tode : Fr.) J. Schröt. in Cohn’s Krypt. Fl. 

Schles. 3(2): 390. 1897. Figs 2A,B; 3A–C; 4A–M. 

≡ Sphaeria gnomon Tode : Fr., Fungi Mecklenb. 2: 50. 1791: Syst. Mycol. 

2: 517. 1823. 

≡ Cryptosphaeria gnomon (Tode : Fr.) Grev., Fl. edin.: 360. 1824. 

≡ Gnomonia vulgaris Ces. & De Not., Comment. Soc. Crittog. Ital. 1: 232. 

1863. 

≡ Gnomoniella vulgaris (Ces. & De Not.) Sacc., Syll. Fung. 1: 416. 1882. 

≡ Gnomoniella gnomon (Tode : Fr.) Magnus, Pilze von Tirol: 490. 1906. 

Anamorph: Unknown. 

Perithecia hypophyllous, scattered randomly over leaf blade, 

immersed at first, erumpent at maturity, black, oblate when moist, 

(58–)163–206(–239) µm high × (144–)206–262(–318) µm diam 

(mean = 179 × 230, SD 35, 40, n1=51, n2=96), concave when dry. 

Necks central, occasionally eccentric, straight or slightly sinuous, 

(179–)250–335(–561) µm long (mean = 296, SD 70, n=71), (18–) 

27–37.5(–53) µm wide at base, (14.5–)23.5–33(–42.5) µm wide 

at apex. Asci fusiform with narrow tapering stipe, (28.5–)35.5– 

43(–56.5) × (5.5–)7–9.5(–11.5) µm (mean = 39.5 × 8, SD 6, 1.5, 

n=70), apical ring 1.5–2.5 µm diam, with eight ascospores more or 

less parallel. Ascospores fusiform, straight to slightly curved, (17–) 

19.5–21(–24.5) × 1.5–2(–2.5) µm (mean = 20.5 × 1.5, SD 1.5, 0.2, 

n=276), l:w (8.5–)11–13(–16.5), two-celled, slightly constricted at 

septum; septum located at (36–)47–51(–61) % (mean = 49, SD 3, 

n=266) of ascospore length; cells tapering to blunt, rounded ends, 

4–8 guttules per cell, usually one large guttule close to septum; 

appendages 2.5–4 µm long, cuneiform, sometimes whip-shaped, 

up to 27 µm long, sometimes absent. 

Cultures: Colony diam after 14 d at 23 °C 25–30 mm diam on PDA, 

5–25 mm on MEA, 10–20 mm on MYA. Colonies flat, radially or 

irregularly furrowed, velvety or with few loose felty tufts, pale grey 

to orange-grey, yellow-brown or dark brown; margin even or lobate; 

reverse yellow-brown to dark brown. On MEA colonies flat, orangebrown 

or greyish orange, aerial mycelium nearly lacking or whitish 

dendroid tufts formed; margin wavy; reverse greyish orange. On 

MYA colonies greyish orange to brownish orange, aerial mycelium 

scant; margin irregular; reverse brownish orange, pale brown or 

dark brown. Perithecia typically produced on PDA, MEA and MYA 

in cultures incubated at 2/10 °C dark/light regime, fertile or sterile 

depending on the strain, best sporulation observed on PDA. Asci 

and ascospores in culture not significantly different from those 

produced on natural substrates, although ascospores sometimes 

shorter, swollen, and with more guttules. 

Habitat: On overwintered leaves of Corylus spp., most common 

on C. avellana L. (Betulaceae), one collection on Populus nigra L. 

(Salicaeae). 

Distribution: Europe (Austria, Bulgaria, Czech Republic, Finland, 

Germany, Russia, Slovakia, Sweden, Switzerland, Ukraine, United 

Kingdom). 

Lectotype: Germany, Mecklenburg, Corylus avellana, 1791, H.I. 

Tode, illustration in ‘Fungi mecklenb.’, Tab. 16, Figs 125a–125f 

designated by Sogonov et al. (2005). 

Epitype: Finland, Helsinki, Helsinki University Botanical Garden, 

overwintered fallen leaves of C. avellana, 19 Apr 2004, D.S. 

Shchigel MS0036 (BPI 844273; ex-type culture AR 4062 = CBS 

116383) designated by Sogonov et al. (2005). 

Additional specimens examined: All on dead leaves of Corylus avellana except 

where noted. Austria, Lower Austria, Krems, Apr. 1871, Thümen (BPI 611815); 

Vienna, 1 May 2004, W. Jaklitsch WJ2501 (BPI 844279, culture CBS 116384); Czech 

Republic, Bohemia, Žlutice (Luditz), Krašov (Krasch), May 1913, R. Steppan (BPI 

611818); Bohemia, Turnov, 02 May 1907, J. M. Kabat (BPI 611820); Hranice, 1913, 

F. Petrak (BPI 611817); France, Haute-Savoie, Petit-Saleve, near Geneve, Feb. 

1852, collector unknown (BPI 596635). Germany, Brandenburg, Prignitz, Triglitz, 

14 Apr. 1906, O. Jaap, Fungi selecti exsiccati 220 (BPI 596638); Brandenburg, 

Prignitz, Triglitz, Willd., 06 Apr. 1911, O. Jaap (Jaap, Fungi selecti exsiccati 519, 

BPI 596287, BPI bound); Dillkreis, 02 Apr. 1934, collector unknown (BPI 596633); 

same details (BPI 596634); Dillkreis, Langenaubach, Apr. 1923, collector unknown 

(BPI 611435); Hessen, Oestrich, near Dillkreis, Langenaubach, 30 Apr. 1933, A. 

Ludwig (BPI 611808); Nossen, 24 Apr. 1889, W. Krieger (BPI 611809); Leipzig, May 

18


Fig. 3. Morphology on natural substrates, asci and ascospores. A–C. Gnomonia gnomon. A. Epitype BPI 844273. B. BPI 871054A. C. BPI 844279. D–F. G. alnea, epitype BPI 

877462A. G. G. incrassata, holotype BPI 611818A. H, I. G. monodii, holotype BPI 877499A. Scale 10 μm. 


19


Fig. 4. Gnomonia gnomon, cultures. A–F, M. Ex-type CBS 116383. G–L. CBS 121233. A–H. Colony habit, 40 d, 23 ºC. A, C, E, G. Surface. B, D, F, H. Reverse. I, J. Perithecia, 

4.5 mo, 2/10 ºC. K–M. Ascospores and asci, 4.5 mo, 2/10 ºC. A, B, G, H, J, M. PDA. C, D, K, L. MEA. E, F, I. MYA. Scale: A–H. 1 cm. I, J. 200 μm. K–M. 10 μm. 

1871, G. Winter (BPI 611810); same location, May 1874, G. Winter (BPI 611811); 

Schleussig, near Leipzig, May 1871, G. Winter (BPI 611812); Thüringen, Steiger 

near Erfurt, 13 May 1905, collector unknown (BPI 596632); locality unknown, date 

unknown, J.C. Schmidt & G. Kunze, Deutschlands Schwämme 57, BPI (611814); 

Russia, Nizhniy Novgorod oblast, Pil’na, birch park close to the river P’yana, 20 May 

2004, G.M. Sogonova MS0103 (BPI 863598); Novgorod oblast, Kholm, Dendropark, 

Jun. 2005, M.V. Sogonov MS0274b (BPI 877514C); Novgorod oblast, Kholm, ulitsa 

Naberezhnaya Reki Lovat’, 07 Jun. 2005, D.N. Borisov MS0275b (BPI 877517C, 

); Slovakia, Prenčov, 28 Mar. 1887, A. Kmet (BPI 611821); Sweden, E.M. Fries, 

Scleromyceti Sueciae 285, BPI (bound); Switzerland, Bischofszell, date unknown, 

H. Wegelin (ZT); Oberbuchsiten, 01 Mar. 1946, coll. J.A. von Arx (ZT); Changins, 

1 March 1976, M. Monod, No. 2 (LAU); Vaud, Bex, Le Bévieux, 13 May 1976, M. 

Monod, No. 47 (LAU); Valais, Populus nigra, 13 May 1977, A. Bolay, No. 267 (LAU, 

culture CBS 829.79); Misox, Grono, 17 May 1988 E. Müller (ZT); Domleschg, Rodel, 

04 May 1988, E. Müller (ZT); Albulatal, Filisur, Solis, 20 May 1988, E. Müller (ZT); 

Schanfigg, Lüen, 24 May 1989, E. Müller (ZT); Vorderrheintal, Panix, 13 June 1989, 

E. Müller (ZT); Valais, vicinity of Martigny, overwintered but still hanging leaves, 21 

May 2005, M. Monod MS0335a (BPI 877499B); Ticino, Monte San Salvatore, 28 

May 2005, M.V. Sogonov MS0205 (BPI 871054A, culture AR4189 = CBS 121233) 

GenBank EU254779; Vaud, St-Cergue, 20 May 2005, M.V. Sogonov MS0336 (BPI 

877492); same data, MS0392 (BPI 877490A) GenBank EU254780; Ukraine, Ivano- 

Frankivsk oblast, Kalkhingel, near Podluze, 12 May 1918, F. Petrak (BPI 611816); 

United Kingdom, England, 1873?, Plowright (BPI 611813). 

Notes: Gnomonia gnomon, the type species of the genus Gnomonia, 

was described and illustrated by Sogonov et al. (2005). This species 

20


is generally restricted to Corylus spp. in Europe although the one 

specimen on Populus exists for which the host identification has 

been verified. Species previously reported as G. gnomon from 

North America have been determined to be G. neognomon or G. 

pendulorum. 

Gnomonia alnea (Fr.) Sogonov, comb. nov. MycoBank MB 

512161. Figs 2C–E; 3D–F. 

Basionym: Sphaeria alnea Fr., Syst. Mycol. 2: 520. 1823. 

≡ Sphaeriella alnea (Fr.) Auersw. in Gonn. & Rabenh., Mycol. Europ. 5/6: 

Tab. 2, Fig. 15. 1869. 

≡ Laestadia alnea (Fr.) Auersw., Hedwigia 8: 177. 1869. 

≡ Guignardia alnea (Fr.) Schröt., Pilze Schlesiens 2: 330. 1894. 

≡ Gnomonina alnea (Fr.) Höhn., Ber. Deutsch. Bot. Ges. 35: 628. 1917. 

≡ Plagiostoma alneum (as alnea) (Fr.) Arx, Antonie van Leeuwenhoek 17: 

264. 1951. 

= Gnomonia perversa Rehm, Hedwigia 24: 70. 1885. 


Perithecia solitary, without stroma, hypophyllous, scattered 

randomly over leaf blade, immersed at first, erumpent or partly 

erumpent at maturity, black, oblate when moist, 100–140 µm high 

× 140–240 µm diam, collapsed concave or convex, occasionally 

irregularly wrinkled or flat when dry. Neck absent, ostiole marginal. 

Asci fusiform, (47.5–)58–67.5(–78) × (9.5–)12–13(–16) µm (mean 

= 63 × 12.5, SD 7, 1.5, n=19) , apical ring 4–4.5 µm diam, with eight 

ascospores arranged obliquely uniseriate or irregularly multiseriate. 

Ascospores fusiform (13.5–)16–17.5(–20.5) × (3.5–)4–4.5(–5) µm 

(mean = 17 × 4.5, SD 1.5, 0.5, n=89), l:w (3.1–)3.7–4.3(–5), twocelled 

with septum located at (44–)48–51(–55) % (mean = 49, SD 

3, n=60) of ascospore length, ends blunt, rounded, each cell with 

two large guttules; appendages ovoid to cuneiform, ca. 2 µm long, 

or absent. 

Habitat: On fallen overwintered leaves of Alnus glutinosa (L.) Gaertn., 

A. incana (L.) Moench and A. viridis (Chaix) DC. (Betulaceae). 

Distribution: Europe (Bulgaria, Czech Republic, Germany, Sweden, 

Ukraine). 

Lectotype (designated here): Sweden, date unknown, E.M. 

Fries, Scleromyceti Sueciae 59, specimen bound in Shear’s Types 

and Rarities (BPI 799019). 

Additional specimens examined: Bulgaria, Sredna Gora Mt (western), Lozenska 

Planina, along the track to Barbeka Lake locality, near river, 21 May 2005, D. 

Stoykov MS0310 (BPI 877462A) GenBank EU 254767. Czech Republic, Bohemia, 

1913, J.A. Stevenson (BPI 611541); Bohemia, Žlutice (Luditz), Krašov (Krasch), 

Jun. 1913, R. Steppan (BPI 611543); Germany, Bielatal near Königstein, Apr. 

1884, W. Krieger (BPI 611540, lectotype of Gnomonia perversa designated 

here); Ukraine, Ivano-Frankivsk oblast, Czarny Las, near Rybno, 08 Jun. 1918, F. 

Petrak (BPI 611542). 

Notes: Gnomonia alnea is restricted to species of Alnus in Europe. 

The synonymy of G. perversa is based on an examination of its type 

specimen as listed above. The taxonomic synonymy of Gnomonia 

vleugelii Kleb. listed by Monod (1983) is rejected because Monod’s 

description disagrees with the original one. Klebahn (1918) 

described the species as having necks up to 1 mm long while 

necks are absent in G. alnea. Gnomonia alnea is unlike the other 

species of Gnomoniaceae on Alnus in the lack of an elongated 

neck. Ophiognomonia alni-viridis, O. ischnostyla and O. trientensis 

all have elongated necks on the perithecium. 

New and revised species of Gnomonia 

Gnomonia incrassata Sogonov, sp. nov. MycoBank MB 512162, 

Figs 2F,G; 3G. 

Perithecia 130–310 µm alta × 190–380 µm diam. Rostrum 220– 

420 µm longum, basi 40–65 µm diam, apice 30–50 µm diam. 

Ascosporae fusiformes, leviter curvatae, (13.5–)15.5–17(–18.5) 

× (2.5–)3–3.5(–4) µm, L:l (4–)4.5–5(–6). Differt a speciebus aliis 

Gnomoniae ascosporis insigniter constrictis et septatis supra 

medio, et rostris crassis. Holotypus: BPI 611818A. 


Etymology: Refers to thickened ascospore cells separated by the 

septum and thick necks. 

Perithecia solitary, without stroma, hypophyllous, in irregular 

groups, immersed at first, erumpent at maturity, black, oblate to 

spheroidal when moist, 130–310 µm high × 190–380 µm wide, 

concave when dry. Necks eccentric or central, slightly sinuous, 

220–420 µm long, 40–65 µm wide at base, 30–50 µm wide at 

apex, compressed when dry. Asci fusiform, (46–)49–51(–61.5) × 

(9.5–)12.5–13.5(–15) µm (mean = 51.5 × 12.5, SD 5, 1.5, n=10), 

apical ring 2.5–3.5 µm diam, with eight ascospores arranged 

irregularly multiseriate. Ascospores fusiform, slightly curved (13.5–) 

15.5–17(–18.5) × (2.5–)3–3.5(–4) µm (mean = 16 × 3.5, SD 1, 0.5, 

n=72), l:w (3.9–)4.5–5.1(–6.2), two-celled, strongly constricted at 

septum, septum located at (55–)60–65(–75) % (mean = 63, SD 4, 

n=72) of ascospore length, ends blunt, rounded, each cell with 2 

(–3), large and sometimes several smaller guttules; appendages 

absent or ovoid to subulate to 7 µm long. 

Cultures: Not observed. 

Habitat: On overwintered fallen leaves of Corylus avellana 

(Betulaceae). 

Distribution: Europe (Czech Republic, France, Germany, 

Switzerland). 

Holotype: Czech Republic, Bohemia, Žlutice, Krašov, May 1913, 

R. Steppan (BPI 611818A). 

Additional specimen examined: France, Petit Saleve, Geneve, 05 Apr. 1851, J. 

Müller (BPI 596639); Germany, Thüringen, Steiger near Erfurt, 13 May 1905, H. 

Diedicke Mycotheca Germanica 482 (BPI bound); Switzerland, Calancatal, ob 

Buseno, 17 Jun. 1989, E. Müller (ZT). 

Notes: Some of the specimens here regarded as Gnomonia 

incrassata were identified as Apiognomonia ostryae by Monod 

(1983). 

Gnomonia monodii Sogonov, sp. nov. MycoBank MB 512163, 

Figs 2H, I; 3H, I. 

Perithecia 180–220 µm high × 270–310 µm diam. Rostrum 320– 


Ascosporae fusiformes, rectae vel inaequilaterae, (14–)15–16(– 

18.5) × (2–)2.5(–3) µm, L:l (5–)5.5–6.5(–7.5). Ad aliis cum rostro 


21


Fig. 5. Morphology on natural substrates, perithecia. A, B. Gnomonia neognomon. A. BPI 877466A. B. BPI 877526C. C–H. G. orcispora. C, D. BPI 877526A. E–H. Holotype BPI 

877465C. I–L. G. ostryae. I–K. BPI 611536. L. BPI 871051. A, C–F, I, J. Intact air-dry perithecia on leaves. B, H, K, L. Extracted and rehydrated perithecia. G. Semi-rehydrated 

perithecium on a small fragment of a leaf. Scale 200 μm. 

eccentrico Gnomoniae speciebus ascosporae magnitudine differt. 

Holotypus: BPI 877499A. 


Etymology: Named after Michel Monod, Lausanne, Switzerland, 

in recognition of his contribution to the taxonomy of the 

Gnomoniaceae. 

Perithecia solitary, without stroma, hypophyllous, on leaf blade or 

veins, in loose irregular groups, immersed at first, erumpent, black, 

oblate or suboblate when moist, concave when dry, 180–220 µm 

high × 270–310 µm diam. Necks eccentric, straight or slightly 

curved, 320–420 µm long, 40–45 µm wide at base, 35–40 µm 

wide at apex. Asci fusiform, (42.5–)45.5–46.5(–51) × 9–11(–13) 

µm (mean = 46 × 10.5, SD 3, 1.5, n=6), apical ring 2–2.5 µm diam, 

with eight ascospores arranged irregularly multiseriate. Ascospores 

fusiform, straight or inequilateral (14–)15–16(–18.5) × (2–)2.5(–3) 

µm (mean = 15.5 × 2.5, SD 1, 0.2, n=54), l:w (4.9–)5.7–6.4(–7.3) 

(mean = 6, SD 0.6, n=54), two-celled, septum located at (53–)60– 

64(–70) % (mean = 62, SD 4, n=54) of ascospore length, not or 

slightly constricted at septum, ends blunt, rounded, each cell with 

2(–3) large, some smaller, guttules; appendages absent or ovoid, 

cuneiform or subulate to whip-shaped, to 20 µm long. 

Habitat: On overwintered leaves of Corylus avellana (Betulaceae). 

Distribution: Europe (Denmark, Switzerland). 

Holotype: Switzerland, Valais, vicinity of Martigny, on overwintered 

but still attached leaves, 21 May 2005, M. Monod & M.V. Sogonov 

MS0335 (BPI 877499A). 

Additional specimen examined: Denmark, Jylland, Krabbesholm Skov, on leaves of 

Corylus avellana, 07 Apr. 1901, J. Lind (BPI 611819); Switzerland, Valais, La Tallaz, 

vallon de Gueuroz, 05 Jun. 1979, M. Monod, No. 741 (LAU). 

Notes: Two of the specimens here regarded as Gnomonia monodii 

were identified as Apiognomonia ostryae by Monod (1983). 

Gnomonia neognomon Sogonov, sp. nov. MycoBank MB 512254, 

Figs 5A,B; 6A–C; 7A–O. 

Perithecia 170–250 µm alta × 220–370 µm diam. Rostrum 

400–1075 µm longum, basi 35–52 µm diam, apice 31–40 µm 

diam. Ascosporae fusiformes, leviter curvatae, (18.5–)20–22.5(– 

24.5)×(2–)2(–2.5) µm, L:l (8–)9.5–11(–12.5). Ad alteris Gnomoniae 

speciebus ascosporae longitudine latitudineque differt. Ascosporae 

22


Fig. 6. Morphology on natural substrates, asci and ascospores. A–C. Gnomonia neognomon, holotype BPI 877465A. D–F. G. orcispora. D. BPI 877466B. E, F. Holotype BPI 

877465C. G, H. G. ostryae. G. BPI 611536. H. BPI 871051. Scale 10 μm. 


23


Fig. 7. Culture morphology. A–O. Gnomonia neognomon. A–F. CBS 121244. G–L. CBS 121265. M–O. Ex-type CBS 121246. P–X. G. orcispora. P–S. AR 4286. T–X. CBS 

121247. A–L, P–S, U–X. Colony habit, 40 d, 23 ºC. A, C, E, G, I, K, P, R, U, W. Surface. B, D, F, H, J, L, Q, S, U, X. Reverse. M–O, T. Perithecia, 2/10 ºC. M, N, T. 4.5 mo. O. 8 

mo. A, B, G, H, P, Q, U, V. PDA. C, D, I, J, N, R, S, W, X. MEA. E, F, K– M, O, T. MYA. Scale: A–L, P–S, U–X. 1 cm. M–O, T. 200 μm. 

longitudo latitudoque similares G. gnomon, sed perithecii rostris 

longioribus et semper eccentricis differt. Holotypus: BPI 877465A. 


Etymology: Refers to the morphological similarity with G. gnomon. 

Perithecia solitary, without stroma, hypophyllous, loosely scattered 

on blades and veins, immersed at first, erumpent at maturity, black, 

suboblate when moist, 170–250 µm high × 220–370 µm diam, 

concave when dry. Necks eccentric, straight or slightly sinuous, 

400–1075 µm long, 35–52 µm wide at base, 31–40 µm wide at 

apex. Asci fusiform, (39.5–)41–44.5(–48.5) × (8–)9–11(–12.5) µm 

(mean = 43.5 × 10, SD 3, 1.5, n=13), apical ring 1.5–3 µm diam, with 

eight parallel ascospores. Ascospores fusiform, slightly curved, (18.5– 

)20–22.5(–24.5) × 2–2.5 µm (mean = 21.5 × 2, SD 1.5, 0.2, n=29), l:w 

(7.9–)9.7–10.8(–12.7) (mean = 10.2, SD 1, n=29), two-celled, slightly 

constricted at septum, septum located at (45–)49–52(–57) % (mean 

= 50, SD 3, n=29) of ascospore length, cells tapering, at ends blunt, 

rounded, each cell with 4–7, guttules, usually one large guttule close 

to septum; appendages usually 2.5–4 µm long, cuneiform, sometimes 

absent, sometimes whip-shaped, to 16 µm long. 

Cultures: Colonies on PDA attaining 35 mm diam after 40 d at 23 °C, 

radially furrowed, velvety, greyish orange to brownish orange, with 

droplets of clear exudate; margin well-defined, wavy; reverse dark 

24


brown. Colonies on MEA attaining 60 mm diam after 40 d at 23 °C, 

flat, pale brown to dark brown, smooth with scant aerial mycelium; 

margin diffuse, wavy; reverse pale brown to dark brown. Colonies 

on MYA attaining 40 mm after 40 d at 23 °C, flat or radially furrowed, 

greyish brown or pale brown to dark brown, velvety, with scant 

droplets of clear exudate; margin well-defined, even or broadly 

wavy; reverse dark brown. Cultures incubated at 2/10 °C dark/light 

regime produce sterile perithecia after 4.5 mo on MEA and MYA but 

not on PDA. Perithecia remain sterile. 

Habitat: On overwintered fallen or hanging leaves of Corylus 

californica (A. DC.) Rose (Betulaceae). 

Distribution: Canada (British Columbia) and U.S.A. (WA). 

Holotype: U.S.A., Washington, Mason Co., Potlatch State Park, 

next to U.S. route 101, on overwintered fallen leaves, 16 May 2006, 

M.V. Sogonov MS0364 (BPI 877465A, ex-type culture AR 4287 = 

CBS 121246) GenBank EU254786. 

Additional specimens examined: Canada, British Columbia, Vancouver Island, 

Goldstream Provincial Park, on veins of overwintered but still hanging leaves, 11 

May 2006, M.V. Sogonov MS0408b (BPI 877526C, culture AR 4336 = CBS 121265); 

U.S.A., Washington, Mason Co., Potlatch State Park, on overwintered leaves, 16 

May 2006, M.V. Sogonov MS0363 (BPI 877466A, culture AR 4285 = CBS 121244). 

Notes: Specimens previously identified as Gnomonia gnomon on 

Corylus in North America may actually be G. neognomon or G. 

pendulorum. 

Gnomonia orcispora Sogonov, sp. nov. MycoBank MB 512164. 

Figs 5C–H; 6D–F; 7P–X. 

Perithecia 150–280 µm alta × 160–310 µm diam. Rostrum usque 

ad 350 µm longum, basi 32–42 µm diam, apice 24–29 µm diam. 

Ascosporae ovales, inaequilaterae, (14–)15.5–17.5(–19.5) × (4.5–) 

5–5.5(–6) µm. Ad plerumque Gnomoniae speciebus ascosporae 

longitudine latitudineque differt. Ascosporae longitudo latitudoque 

similares G. ostryae, sed ascosporis in extremitatibus hilo terminatis 

et guttulis majoribus in ascosporae cellulis differt. Holotypus: BPI 

877465C. 


Etymology: Refers to the appearance of spores from Latin orca 

(barrel-shaped) and spora (spore). 

Perithecia solitary, without stroma, hypophyllous, loosely scattered 

on blades and veins, immersed at first, erumpent at maturity, black, 

spherical when moist, 150–280 µm high × 160–310 µm diam, 

concave when dry. Necks marginal, straight or slightly sinuous, 

absent at ascospore formation, growing at maturity of ascospores, 

reaching 350 µm long, 32–42 µm wide at base, 24–29 µm wide 

at apex. Asci oval to fusiform, (53–)58.5–62.5(–69.5) × (19.5–) 

20–20.5(–22) µm (mean = 61 × 20.5, SD 5.9, 0.9, n=5), apical 

ring 5–5.5 µm diam, with eight ascospores with eight ascospores 

evenly or slightly unevenly parallel. Ascospores oval, inequilateral, 

(14–)15.5–17.5(–19.5) × (4.5–)5–5.5(–6) µm (mean = 16.5 × 5, SD 

1.1, 0.4, n=45), l:w (2.8–)3–3.3(–3.7) (mean = 3.2, SD 0.2, n=45), 

two-celled, constricted at septum, septum located at (54–)59–63(– 

66) % (mean = 61, SD 3, n=45) of ascospore length, ends blunt, 

rounded or somewhat truncated, each cell with two large guttules; 

appendages navicular, large to 15 μm long, 8 μm wide but very thin 

and often indistinct, with base surrounded by a pronounced hilum 

on ascospore wall. 

Cultures: Two cultures (CBS 121247 and AR 4286) differ significantly 

in their morphology. CBS 121247: Colonies on PDA attaining 18 

mm diam after 40 d at 23 °C, irregularly furrowed, shortly velvety, 

dark brown and dark grey to almost black, with whitish submerged 

mycelium at margin; margin diffuse, irregular; reverse dark brown, 

almost black with orange-brown to whitish margins. Colonies on 

MEA attaining 17 mm diam after 40 d at 23 °C, flat, glabrous with 

scant aerial mycelium, black with dark brown margins; margin clear, 

slightly wavy; black with dark brown margins. AR 4286: Colonies on 

PDA attaining 33 mm diam after 40 d at 23 °C, radially furrowed, 

velvety, whitish to greyish orange, with droplets of clear exudate; 

margin well-defined, slightly wavy; reverse reddish orange to dark 

brown; agar stained with orange pigment. Colonies on MEA attaining 

15 mm diam after 40 d at 23 °C, slightly radially furrowed, greyish 

orange overlaid by white short felty aerial mycelium; margin welldefined, 

slightly wavy; reverse reddish orange. Sterile perithecia 

observed in CBS 121247 incubated at 2/10 °C dark/light regime for 

4.5 mo on MYA. 

Habitat: On overwintered fallen or still hanging leaves of Corylus 

californica (Betulaceae). 

Distribution: Canada (British Columbia) and U.S.A. (WA). 


next to U.S. route 101, on overwintered fallen leaves, 16 May 

2006, M.V. Sogonov MS0364b (BPI 877465C, ex-type culture CBS 

121247). 

Additional specimen examined: Canada, British Columbia, Vancouver Island, 

Goldstream Provincial Park, on overwintered but still hanging leaves, 11 May 2006, 

M.V.Sogonov MS0408 (BPI 877526A) GenBank EU254789; U.S.A., Washington, 

Mason Co., Potlatch State Park, on overwintered fallen leaves, 16 May 2006, M.V. 

Sogonov MS0363a culture AR 4286 (BPI 877466B). 

Notes: Perithecia of Gnomonia orcispora have marginal necks and 

are thus distinct from other species of Gnomonia on Corylus in 

North America. 

Gnomonia ostryae De Not., Sfer. Ital. cent. 1, fasc. 1: 42. 1863. 

5. 8I–L; 6G–H; 8A–H. 

≡ Apiognomonia ostryae (De Not.) M. Monod, Beih. Sydowia 9: 50 1983. 

= Gnomonia veneta Speg., Michelia 1: 457. 1879. 

Perithecia solitary, without stroma, hypophyllous, on leaf blade or 

veins, in loose irregular groups, immersed at first, erumpent later, 

black, oblate or suboblate when moist, 130–200 µm high × 170–260 

µm diam, concave when dry. Necks eccentric to marginal, slightly 

curved, 175–290 µm long, 25–45 µm wide at base, 25–45 µm 

wide at apex. Asci fusiform, 49–57 × 12–16 µm, apical ring 2.9–3.8 

µm diam, with eight ascospores arranged irregularly multiseriate 

to obliquely uniseriate. Ascospores varying from fusiform with both 

ends gradually tapering although rounded at tips to oblanceolate 

or obovoid, broadened in upper part, with distal end extensively 

rounded or smoothly truncated, inequilateral, (12.5–)15–17(–20.5) 

× (3–)3.5–4.5(–6) µm (mean = 16 × 4, SD 1.5, 0.5, n=90), l:w 

(3–)3.6–4.1(–4.9) (mean = 3.9, SD 0.4), two-celled, constricted at 


25


Fig. 8. Culture morphology. A–H. Gnomonia ostryae CBS 121242. I, J. G. pendulorum ex-type CBS 121264. K–P. G. rodmanii ex-type CBS 121909. A–F, I–P. Colony habit, 40 

d, 23 ºC. A, C, E, I, K, M, O. Surface. B, D, F, J, L, N, P. Reverse. G. Conidiomata, 4 mo, 2/10 ºC. H. Conidia, 4.5 mo, 2/10 ºC. A, B, G, H, K, L. PDA. C, D, M, N. MEA. E, F, I, 

J, O, P. MYA. Scale: A–F, I–P. 1 cm. G. 200 μm. H. 10 μm. 

septum, septum located at (55–)62–65(–71) % (mean = 63, SD 3) 

of ascospore length, ascospore cells usually with two large or/and 

several small guttules; appendages subulate 2–5 µm long. 

Cultures: Colonies on PDA attaining 30 mm diam after 40 d at 

23 °C, radially furrowed, felty orange-white to greyish orange in 

central part, velvety, dark brownish grey at margin; droplets of clear 

exudate present all over the colony but more abundant at margin; 

margin well-defined, crenate; reverse dark brown with pale brown 

patterns. Colonies on MEA attaining 50 mm diam after 40 d at 23 

°C, flat, superficial, with almost no aerial mycelium, greyish orange, 

with tufts of orange-white aerial mycelium in central part; margins 

submerged; margin irregular; reverse greyish orange. Colonies on 

MYA attaining 30 mm diam after 40 d at 23 °C, radially furrowed, 

short felty orange-grey in central part, velvety, dark brownish grey 

at margin; droplets of clear exudate present all over the colony but 

more abundant at margin; margin well-defined, crenate; reverse 

dark brown with pale brown patterns. No perithecia observed in 

cultures at 2/10 °C dark/light regime. Cultures on PDA after 8 mo 

at 2/10°C regime produce amorphous slimy conidiomata. Conidia 

oval or allantoid, 4.9–7.1 × 1.9–2.7 µm. 

Habitat: On overwintered leaves of Ostrya carpinifolia Scop. 

(Betulaceae). 

Distribution: Europe (Bulgaria, Italy, Switzerland). 

Specimens examined: Italy, Varone near Riva (Valsesia), Garda-See, May 1900, H. 

Rehm (BPI 611536). Switzerland, Ticino, Monte San Salvatore, 28 May 2005, M.V. 

Sogonov MS0204, culture CBS 121242 (BPI 871051) GenBank EU254790. 

Notes: The asexual state of Gnomonia ostryae has been placed 

in Cylindrosporella (Monod, 1983). The varieties of G. ostryae 

as Apiognomonia ostryae defined by Monod (1983) are herein 

recognised as distinct species. 

Gnomonia pendulorum Sogonov, sp. nov. MycoBank MB 512165, 

Figs 8I,J; 9A,B; 10A,B. 

Perithecia 230–280 µm alta × 470–580 µm diam. Rostrum 

770–970 µm longum, basi 65–70 µm diam, apice 40–55 µm 

diam. Ascosporae fusiformes, leviter curvatae, (20–)21–22(– 

23)×(2.5–)3(–3.5) µm, L:l (6–)7–7.5(–8). Similares G. gnomon, 

sed ascosporae latioribus et perithecii rostris longioribus differt. 

Holotypus: BPI 877526B. 


Etymology: Refers to the fact that perithecia of this species were 

found on hanging, overwintered leaves. 

Perithecia solitary, without stroma, hypophyllous, mostly on major 

veins, immersed at first, later erumpent, black, suboblate when 

moist, 230–280 µm high × 470–580 µm diam, concave when dry. 

26


Fig. 9. Morphology on natural substrate, perithecia. A, B. Gnomonia pendulorum, holotype BPI 877526B. C, D. G. rodmanii, holotype BPI 878211A. E–G. G. skokomishica. E, 

G. Holotype BPI 877465B. F. BPI 877535. A, C, E, F. Intact air-dry perithecia on leaves. B, D, G. Extracted and rehydrated perithecia. Scale 200 μm. 


27


Fig. 10. Morphology on natural substrate, asci and ascospores. A, B. Gnomonia pendulorum, holotype BPI 877526B. C, D. G. rodmanii, holotype BPI 878211A. E, F. G. 

skokomishica, holotype BPI 877465B. Scale 10 μm. 

28


Fig. 11. Gnomonia skokomishica, culture morphology. A–L. Ex-type CBS 121245. M–Q. CBS 121398. A–F. Colony habit, 14 d, 23 ºC. G–L. Colony habit, 40 d, 23 ºC. A, C, E, 

G, I, K. Surface. B, D, F, J, L. Reverse. M–P. Perithecia, 2/10 ºC. M. 40 d. N. 40 d. O. 4.5 mo. P. 8 mo, ×1/2. Q. Asci and ascospores, 8 mo, 2/10 ºC. A, B, G, H, M. PDA. C, D, 

I, J. MEA. E, F, K, L, N–Q. MYA. Scale: A–L. 1 cm. M–O. 200 μm. P. 500 μm. Q. 10 μm. 


29


Necks central, straight or slightly sinuous, 770–970 µm long, 65–70 

µm wide at base, 40–55 µm wide at apex. Asci fusiform, 55–60 × 

10–11 µm, apical ring 2.5 µm diam, with eight ascospores arranged 

unevenly parallel or irregularly multiseriate. Ascospores fusiform, 

slightly curved (20–)21–22(–23) × (2.5–)3(–3.5) µm (mean = 21.5 

× 3, SD 0.5, 0.3, n=16), l:w (6–)6.8–7.4(–8.2) (mean = 7.1, SD 0.6), 

two-celled, constricted at septum, septum located at (47–)49–51 

% (mean = 49, SD 1.5) of ascospore length, ends blunt, rounded, 

each cell with 2–10 guttules, usually one large guttule close to 

septum; appendages whip-shaped, 7–32 µm long. 

Cultures: Not observed on PDA and MEA. Colonies on MYA 

attaining 30 mm diam after 40 d at 23 °C, thick, radially furrowed, 

short felty, orange-white; margin well-defined, even; reverse orange 

to brownish orange. Neither perithecia nor conidiomata observed in 

cultures at 2/10 °C after 4.5 mo. 

Habitat: On overwintered dead but still attached leaves of Corylus 

californica (Betulaceae). 

Distribution: Canada (British Columbia). 

Holotype: Canada, British Columbia, Vancouver Island, Goldstream 

Provincial Park, 11 May 2006, M.V. Sogonov MS0408a (BPI 

877526B; ex-type culture CBS 121264) GenBank EU254791. 

Notes: Among species of Gnomonia on Corylus, G. pendulorum in 

North America is similar to Gnomonia gnomon in Europe in having 

a central neck on the perithecia. 

Gnomonia rodmanii Sogonov, sp. nov. MycoBank MB 512166, 

Figs 8K–P; 9C,D; 10C,D. 

Perithecia 180–260 µm alta × 220–330 µm diam. Rostrum 300–530 

µm longum, basi 41–48 µm diam, apice 22–28 µm diam. Ascosporae 

fusiformes, rectae, interdum leviter curvatae, (13.5–)15–16.5(–18.5) 

× 2–2.5 µm, L:l (6–)6.5–7.5(–8.5). Abd aliis cum rostro eccentrico 

Gnomoniae speciebus ascosporae longitudine latitudineque differt. 

Ascosporae longitudo latitudoque similares G. monodii et G. 

virginianae, sed septi positione et hospitis genere differt. Holotypus: 

BPI 878211A. 


Etymology: Named after Dr. James Rodman in recognition of his 

promotion of taxonomic research. The holotype specimen of this 

species was collected during the 6th meeting of National Science 

Foundation Partnership for Enhancing Expertise in Taxonomy 

program that was initiated by Dr. Rodman. 

Perithecia solitary, without stroma, hypophyllous, mostly next to midrib 

but some scattered over leaf blade, immersed at first, erumpent at 

maturity, black, suboblate when moist, 180–260 µm high × 220–330 

µm diam, concave when dry. Necks eccentric, slightly sinuous, 

300–530 µm long, 41–48 µm wide at base, 22–28 µm wide at apex. 

Asci fusiform with narrow tapering stipe, (40.5–)43.5–46(–50.5) × 

(11–)13–14(–14.5) µm (mean = 45 × 13, SD 2.4, 1.1, n=11), apical 

ring 2–2.5 µm diam, with eight ascospores arranged irregularly 

multiseriate to obliquely uniseriate. Ascospores fusiform, straight, 

occasionally slightly curved, (13.5–)15–16.5(–18.5) × 2–2.5 µm 

(mean = 16 × 2.5, SD 1.1, 0.2, n=64), l:w (5.8–)6.6–7.3(–8.3) (mean 

= 6.9, SD 0.5), two-celled, not constricted at septum, septum located 

at (41–)48–50(–55) % (mean = 49, SD 2) of ascospore length, cells 

tapering, at ends blunt, rounded, each cell with 2–4 guttules, usually 

one large guttule close to septum; appendages whip-shaped, to 30 

µm long. 

Cultures: Colonies on PDA attaining 85 mm diam after 40 d at 23 

°C, flat, loosely velvety, granular from young perithecia in central 

part, dark brown; margin diffuse; reverse black. Colonies on MEA 

attaining 90 mm after 40 d at 23 °C, flat, dark brown to black, smooth 

with scant appressed aerial mycelium, with black dots of submerged 

young perithecia; margin diffuse; reverse black. Colonies on MYA 

attaining 85 mm diam after 40 d at 23 °C, shallowly and irregularly 

furrowed in centre, flat at marginal parts, velvety, greyish dark 

brown to black; margin diffuse; reverse dark brown to black. 

Habitat: On overwintered fallen leaves of Carpinus caroliniana 

Walter (Betulaceae). 

Distribution: U.S.A. (GA). 

Holotype: U.S.A., Georgia, Clarke Co., Athens, Botanical Garden, 

Orange Trail, 28 Mar. 2007, M.V. Sogonov MS0535 (BPI 878211A, 

ex-type culture CBS 121909). 

Notes: Among species of Gnomonia on Carpinus, G. rodmanii 

has an elongated, eccentric neck on each perithecium unlike G. 

arnstaeditensis in which an elongated neck is lacking. In addition, 

the necks of G. rodmanii lack a collar unlike G. amoena that has a 

central neck surrounded by a collar and is unlike G. carpinicola, a 

species having two or three necks on each perithecium. 

Gnomonia skokomishica Sogonov, sp. nov. MycoBank 

MB512167, Figs 9E–G; 10E,F; 11A–L. 


µm longum, basi (34.5–)36.5–42(–44) µm diam, apice (22.5–)29.5– 

34.5(–37) µm diam. Ascosporae fusiformes, rectae, (16.5–)17.5– 

19(–20.5) × 2–2.5(–3) µm, L:l (7–)7.5–8.5(–9.5). Similares G. 

gnomon, sed ascosporae latioribus et ascosporarum septis leviter 

sed semper supra medio differt. Holotypus: BPI 877465B. 


Etymology: Refers to the Native American tribe near whose 

reservation the holotype was collected. 

Perithecia solitary, without stroma, hypophyllous, scattered loosely on 

midribs or in dense groups on petioles, immersed at first, erumpent, 

partly erumpent or immersed at maturity, black, oblate when moist, 

190–340 µm high × 260–480 µm diam, concave when dry. Necks 

central, curved, 610–930 µm long (mean = 722, SD 134, n=6), 

(34.5–)36.5–42(–44) µm wide at base, (22.5–)29.5–34.5(–37) µm 

wide at apex. Asci fusiform with tapering stipe, (38.5–)40–46.5(–51) 

× (8.5–)10–10.5(–11.5) µm (mean = 43.5 × 10, SD 4, 1, n=12), 

apical ring 2–2.5 µm diam, with eight ascospores arranged evenly 

or unevenly parallel. Ascospores fusiform, straight (16.5–)17.5– 

19(–20.5) × 2–2.5(–3) µm (mean = 18.5 × 2.5, SD 1, 0.1, n=37), l:w 

(6.8–)7.5–8.5(–9.6) (mean = 8, SD 0.7, n=37), two-celled, constricted 

at septum, septum located at (51)54–58(–62) % (mean = 56, SD 3) 

of ascospore length, each cell with 2–3 large or one large and 3–5, 

30


Fig. 12. Morphology on natural substrate, perithecia. A–C. Gnomonia virginianae. A. BPI 878210. B. BPI 877565A. C. BPI 878209. D–F. G. amoena. D, E. BPI 877469. F. 

BPI 877468. G. G. arnstadiensis, BPI 877470. H. G. pseudoamoena, BPI 877516. A, B, D, F–H. Intact air-dry perithecia on leaves and petioles. C. Extracted and rehydrated 

perithecia. E. Air-dry perithecium on fragment of petiole with removed outer tissue. Scale 200 μm. 

smaller guttules, largest guttule close to septum; appendages whipshaped 

to 27 µm long. 

Cultures: Colonies on PDA and MYA at 23 °C initially forming velvety 

pale brown or brown surface, later overgrown by orange-grey to 

whitish (PDA) or snow-white (MYA) short felty mycelium expanding 

from the centre of colonies, attaining 80 mm diam after 40 d; margin 

well-defined, low, wavy or irregularly serrate; reverse greyish brown 

to dark brown. Colonies on MEA attaining 90 mm diam after 40 

d at 23 °C, flat, thin, greyish orange for the most part, with dark 

brown areas or almost entirely dark brown, thin, smooth with scant 

appressed aerial mycelium, with black dots of submerged young 

perithecia; margin diffuse; reverse of same colours as front side. 

Cultures at 2/10 °C produce sterile perithecia after 4.5 mo on PDA 

and MYA, asci and ascospores were observed only on MYA in CBS 

121245 after 8 mo at 2/10 C. Perithecia produced in culture often 

with 2–3, rarely 4 necks; necks often hairy. Ascospores similar to 

those on natural substrates. 

Habitat: On overwintered fallen leaves of Corylus californica 

(Betulaceae). 

Distribution: U.S.A. (WA). 


31



next to U.S. route 101, on overwintered fallen leaves, 16 May 

2006, M.V. Sogonov MS0364a (BPI 877465B, ex-type culture CBS 

121245). 

Additional specimen examined: U.S.A., Washington, King Co., Tiger Mountain State 

Forest, near U.S. route 18, 16 May 2006, M.V. Sogonov MS0393 (BPI 877535, 

culture CBS 121398) GenBank EU254798. 

Notes: This new taxon is similar to Gnomonia californica described 

by Monod (1983) as compared with the description and illustrations 

only. According to Monod the type specimen of G. californica is 

deposited in TRTC from where it was requested but was not found. 

Gnomonia skokomishica lacks a dark red collar as described for 

G. californica by Monod (1983) as “collum rubrum rostrum cingens 

ex substrato”, “le substrat forme une couronne rougeâtre, non 

pulvérulente à la sortie du bec”. In addition G. skokomishica has 

a submedian ascospore septum while G. californica is described 

as having a median septum (“ascosporae… septatae dimidio 

longitudinis”, “ascospores… cloisonnées à mi-longueur”) (Monod 

1983). 

Gnomonia virginianae Sogonov, sp. nov. MycoBank MB 512168. 

Figs 12A–C; 13A–C; 14A–F. 

Perithecia 115–160 µm high × 150–260 µm diam. Rostrum 200– 


Ascosporae fusiformes, leviter curvatae, (12–)13–14(–14.5) × (2–) 

2.5–3 µm, L:l (4–)4.5–5(–6). Ad plerumque cum rostro eccentrico 

Gnomoniae speciebus ascosporae longitudine latitudineque differt. 

Ascosporae longitudo latitudoque similares G. romanii, sed septi 

positione et hospitis genere differt. Holotypus: BPI 844264. 


Etymology: Refers to species epithet of the host plant. 

Perithecia hypophyllous, on leaf blade or veins, in loose irregular 

groups, immersed at first, erumpent later, black, suboblate to oblate 

spheroidal when moist, 115–160 µm high × 150–260 µm diam, 

concave when dry. Necks marginal, straight or slightly sinuous, 

200–430 µm long, 21–33 µm wide at base, 18–28 µm wide at apex. 

Asci fusiform with narrow tapering stipe, (33–)34.5–42(–47.5) × 

(8–)9.5–11(–14) µm (mean = 38.5 × 10.5, SD 5, 1.5, n=14) , apical 

ring 2–2.5 µm diam, with eight ascospores arranged unevenly 

parallel, less commonly in evenly parallel or irregularly multiseriate. 

Ascospores fusiform, slightly curved (12–)13–14(–14.5) × (2–)2.5– 

3 µm (mean = 13.5 × 2.5, SD 0.5, 0.2, n=47), l:w (4.2–)4.7–5.3(– 

5.8) (mean = 5, SD 0.4), two-celled, slightly constricted at septum, 

septum located at (55–)59–62(–66) % (mean = 61.5, SD 3) of 

ascospore length, cells tapering, at ends blunt, rounded, distal cell 

usually with two, basal cell with two or three large guttules, several 

smaller guttules may be present in each cell; appendages whipshaped, 

7–45 µm long. 

Cultures: Colonies on PDA attaining 30 mm diam after 40 d at 

23 °C, with knobbed surface, velvety, greyish orange, brownish 

orange, pale brown or brownish grey, with droplets of clear exudate; 

margin well-defined, irregular; reverse pale brown to dark brown. 

Colonies on MEA attaining 30 mm diam after 40 d at 23 °C, radially 

wrinkled and furrowed, velvety, orange-grey to brownish orange; 

margins with two brims differing from the rest of colony, 1–2 mm 

wide each, a dark brown velvety inner brim and greyish orange 

waxy outer brim; margin well-defined, even; reverse greyish orange 

and orange-grey to brownish orange and brown. Colonies on MYA 

attaining 22 mm diam after 40 d at 23 °C, densely radially wrinkled, 

velvety to felty, orange-white with pale brown 3 mm brim; margin 

well-defined, even; reverse greyish brown to pale brown. Neither 

perithecia nor conidiomata observed in cultures on PDA, MEA and 

MYA after 8 mo at 2/10 C. Scarce fertile perithecia were observed 

in cultures on CMA after 5 mo at 15 °C in darkness. 

Habitat: On overwintered fallen leaves of Ostrya virginiana 

(Betulaceae). 

Distribution: U.S.A. (AR, GA, MD, NC). 

Holotype: U.S.A., Maryland, Montgomery Co., Chesapeake & Ohio 

Canal National Historic Park, 10 April 2004, M.V. Sogonov MS0016 

(BPI 844264, ex type culture CBS 121913). 

Additional specimens examined: U.S.A., Arkansas, Ozark Natural Science Center, 21 

June 2006, L.N. Vasilyeva (BPI 877565A) GenBank EU254804; Georgia, Clarke Co., 

Athens, Oconee Forest Park, 30 March 2007, M.V. Sogonov MS0532 (BPI 878209) 

GenBank EU254805; Georgia, Clarke Co., Athens, Botanical Garden, Orange Trail, 

28 March 2007, M.V. Sogonov MS0534 (BPI 878210); Maryland, Mongomery Co., 

Chesapeake & Ohio Canal National Historic Park, 10 April 2004, M.V. Sogonov 

MS0023 (BPI 877564) GenBank EU254802; North Carolina, Wake Co., Raleigh, 

William B. Umstead State Park, hardwood forest, 03 April 2005, M.V. Sogonov MS0169 

(BPI 877566) GenBank EU254803; same collecting data MS0170 (BPI 877422). 

Notes: Gnomonia virginianae is the only species of Gnomonia 

on Ostrya in North America and is distinct from the two European 

species on Ostrya, G. arnestadtiensis and G. ostryae. Gnomonia 

arnstadtiensis has perithecia lacking an elongated neck while 

ascospores of G. ostryae are longer and wider than those of G. 

virginianae. Many specimens of Gnomonia virginianae were 

identified as Apiognomonia ostryae variété 2 by Monod (1983). 

Additional species accepted in Gnomonia 

Gnomonia amoena (Nees : Fr.) Ces. & De Not., Comment. Soc. 

Crittog. Ital. 1: 232. 1863. 

Figs 12D–F; 13D,E; 14G–L. 

≡ Sphaeria amoena Nees : Fr., Nova Acta Acad. Caes. Leop.-Carol. Nat. 

Cur. 9: 257. 1818 : Syst. Mycol. 2: 517. 1823. 

≡ Gnomoniella amoena (Nees : Fr.) Sacc., Syll. Fung. 1: 414. 1882. 

Habitat: On dead leaves and petioles of Carpinus betulus and C. 

caroliniana (Betulaceae). 

Distribution: Europe (Germany, Switzerland) and U.S.A. (TN). 

Specimens examined: Switzerland, Lausanne, on overwintered leaves of Carpinus 

betulus, 25 May 2005, coll. M.V. Sogonov (BPI 877468) GenBank EU254770; Vaud, 

near hospital St-Loup, on overwintered leaves of Carpinus betulus, 24 May 2005, 

EU254769. 

Notes: Gnomonia amoena is unique among species of Gnomonia 

on Carpinus in having a distinct collar around the central neck. 

Monod (1983) provides a detailed description of this species. 

Barr (1978) erred in reporting this species on Corylus based on 

specimens later identified by Monod (1983) as G. californica M. 

32


Fig. 13. Morphology on natural substrate, asci and ascospores. A–C. Gnomonia viriginianae. A. BPI 877565A. B. Holotype BPI 844264. C. BPI 878209. D, E. G. amoena, BPI 

877469. F, G. G. pseudoamoena. F. BPI 877518. G. Stirpes Cryptogamae Vogeso-Rhenanae 1251, BPI bound. Scale 10 μm. 

Monod. Specimens of G. amoena on Liquidambar styraciflua L. are 

reidentified as Ambarignomonia petiolorum. 

Gnomonia arnstadtiensis Auersw. in Gonn. & Rabenh., Mycol. 

Europ. 5/6: 22. 1869. Fig. 12G. 

≡ Plagiostoma arnstadtiense (Auersw.) M. Monod, Beih. Sydowia 9: 143. 

1983. 

= Hypospila rehmii Sacc., Syll. Fung. 2: 189. 1883 fide Monod 1983. 

≡ Gnomonina rehmii (Sacc.) Höhn., Ann. Mycol. 16: 52. 1918. 

≡ Plagiostoma rehmii (Sacc.) Arx, Antonie van Leeuwenhoek 17: 264. 

1951. 

Habitat: On fallen leaves of Carpinus betulus and Ostrya carpinifolia 

(Betulaceae). 

Distribution: Europe (Bulgaria, Germany, Switzerland, 

“Yugoslavia”) 

Specimens examined: Bulgaria, Mt Belasitsa, nearby Belasitsa challet, alt. ca. 750 

m, on overwintered leaves of Ostrya carpinifolia, 30 Apr 2005, Stoykov, D. (BPI 

877470) GenBank EU254772; Balkan foot-hill region, Golyama Zhelyazna village, 

Promkombinat locality, on overwintered leaves of Ostrya carpinifolia, 4 Apr 2005, 

Stoykov, D. (BPI 877472B) GenBank EU254773. 

Notes: Gnomonia arnstadtiensis is unique among species of 


33


Fig. 14. Culture morphology. A–F. Gnomonia virginianae ex-type CBS 121913. G–L. G. amoena CBS 121262. M–T. G. pseudoamoena CBS 121261. A–R. Colony habit. A–N, 

Q, R. 40 d, 23 ºC. O, P. 14 d, 23 ºC. S. A, C, E, G, I, K, M, O, Q. Surface. B, D, F, H, J, L, N, P, R. Reverse. Perithecium, 4.5 mo, 2/10 ºC. T. Asci, 4.5 mo, 2/10 ºC. A, B, G, H, 

M, N. PDA. C, D, I, J, O, P, S, T. MEA. E, F, K, L, Q, R. MYA. Scale: A–R. 1 cm. S. 200 μm. T. 10 μm. 

Gnomonia on Carpinus and Ostrya in lacking an elongated neck 

on the perithecium. 

having two or three necks emerging on both sides of the leaf blades. 

In addition the ascospores have a submedian septum. 

Gnomonia carpinicola (Höhn.) Sogonov, comb. nov. MycoBank 

MB 512169. 

Basionym: Plagiostomella carpinicola Höhn., Ann. Mycol. 16: 52. 1918. 

≡ Apioplagiostoma carpinicola (Höhn.) M.E. Barr, Mycol. Mem. 7: 103. 

1978. 

= Gnomonia stahlii Kleb., Haupt- und Nebenfruchtformen der Ascomyzeten: 

279. 1918 fide Monod 1983. 

= Apiospora carpinea Rehm, Ber. naturalist. Ver. Augburg 26: 119. 1881 non 

Gnomonia carpinea (Fr.) Kleb. 1918 fide Monod 1983. 

Habitat: On fallen leaves of Carpinus betulus L. (Betulaceae). 

Distribution: Europe (Bulgaria, Germany, Switzerland). 

Notes: The perithecia of Gnomonia carpinicola are unusual in often 

Gnomonia pseudoamoena M. Monod, Beih. Sydowia 9: 86. 1983. 

Figs 12H; 13F,G; 14M–T. 

Habitat: On fallen leaves of Corylus avellana and C. californica 

(Betulaceae). 

Distribution: Canada (British Columbia) and Europe (Bulgaria, 

Germany, Sweden, Switzerland) 

Specimens examined: Switzerland, Vaud, near hospital St-Loup, on overwintered 

leaves of Corylus avellana, 24 May 2005, coll. M.V. Sognov (BPI 877512) GenBank 

EU254794; Flühli, on overwintered leaves of Corylus avellana, 27 May 2005, coll. 

M.V. Sogonov (BPI 877513) GenBank EU254793; same as above (BPI 877516) 

GenBank EU254792. 

34


Fig. 15. Morphology on natural substrate, perithecia. Ambarignomonia petiolorum. A, B. Epitype BPI 844274. C, D. BPI 877511. A, C. Intact air-dry perithecia on petioles. B. 

Air-dry perithecium on fragment of petiole with removed outer tissue. D. Extracted and rehydrated perithecia. Scale 200 μm. 

Fig. 16. Morphology on natural substrate, asci and ascospores. Ambarignomonia petiolorum. A. Epitype BPI 844274. B, C. BPI 877509. Scale 10 μm. 

Fig. 17. Ambarignomonia petiolorum ex-type CBS 121227, culture morphology, colony habit. A, C, E. Surface. B, D, F. Reverse. A, B. PDA. C, D. MEA. E, F. MYA. Scale 1 cm. 

Notes: Gnomonia pseudoamoena is unique among species of 

Gnomonia on Corylus in having a distinct collar around the central 

neck. Stoykov & Denchev (2006) reported this species from 

Bulgaria as G. amoena. 

AMBARIGNOMONIA Sogonov gen. nov. MycoBank MB 512170. 

Perithecia solitaria, sine stromate, immerse, in foliis caducis, in sicco 

concava. Rostri centralia, recti, apice contracti, basi circumcincti 

collo albo pulveraceo. Ascosporae fusiformes, bicellulares. 

Holotypus: Ambarignomonia petiolorum (Schwein. : Fr.) Sogonov, 

comb. nov. 

Perithecia solitary, without stroma, on fallen leaves, on petioles and 

basal parts of major veins of fallen leaves, immersed, black, suboblate 

when moist, concave when dry, round in top view. Necks 

central, straight, tapering to their ends, at base surrounded by white 

powdery collar not soluble in water or 3 % KOH, length 3–4 times 

greater than perithecial diam. Asci fusiform, with apical ring, with 

eight spores arranged irregularly fasciculate. Ascospores fusiform, 

l:w ca. 7, two-celled; appendages ovoid to subulate. Colonies 

usually slowly to moderately growing, reaching 0.5 cm diam or less 

in 2 wk at 23 °C dark/light. Colony surface glabrous, sometimes 

velvety in central part, yellowish brown. Conidiogenous structures 

or perithecia never formed in culture. Known only from Liquidambar 

styraciflua (Hamamelidaceae). 


35


Ambarignomonia petiolorum (Schwein. : Fr.) Sogonov, comb. 

nov. MycoBank MB 512171. Figs 15A–D; 16A–C; 17A–F. 

Basionym: Sphaeria petiolorum Schwein. : Fr., Schr. Naturf. Ges. Leipzig 1: 41. 

1822 : Syst. Mycol. 2: 517. 1823. 

≡ Gnomonia petiolorum (Schwein. : Fr.) Cooke, Grevillea 7: 54. 1878. 

≡ Gnomoniella amoena var. petiolorum (Schwein. : Fr.) Sacc., Syll. Fung. 

1: 414. 1882. 

Perithecia solitary, without stroma, evenly and densely distributed 

over petioles of fallen leaves, sometimes also on basal parts of 

major veins, immersed, black, suboblate when moist, 180–220 µm 

high × 300–420 µm diam, concave when dry, round in top view. 

Necks central, straight, 200–700 µm long, 65–80 µm wide at base, 

35–45 µm wide at apex. Asci fusiform with narrow tapering stipe, 

(24–)27.5–29.5(–33.5) × (6.5–)8–9.5(–11) µm (mean = 29 × 9, SD 

2.5, 1.2, n=25), apical ring 1.3–2 µm diam, with eight ascospores 

arranged irregularly multiseriate or unevenly parallel. Ascospores 

fusiform, slightly curved (9–)11–12.5(–15) × 1.5–2 µm (mean 

= 11.5 × 2, SD 1.5, 0.2, n=44), l:w (5–)6.1–6.9(–8) (mean = 6.5, 

SD 0.6), two-celled, not constricted at septum located at (36–)45– 

50(–54) % (mean = 47, SD 4) of ascospore length, each cell with 

several (usually 3–5) guttules, usually one large guttule close to 

septum; appendages subulate 1–4 µm long, rarely whip-like to 10 

µm long. 


°C, branched, at margin with patches of aerial mycelium connected 

with each other, in centre surface knobbed, aerial mycelium 

velvety to felty, orange-white, orange-grey or pale orange; margin 

well-defined, irregular, in some parts submerged; reverse dark 

brown; agar stained with orange-brown pigment. Colonies on MEA 

extremely slow growing, forming a brim of only 1–2 mm around 

original inoculum after 40 d at 23 °C, velvety, orange-grey; margin 

clear; reverse dark brown. Colonies on MYA attaining 10 mm diam 

after 40 d at 23 °C, mostly submerged, radially stringy, pale brown, 

with an entire spot of orange-grey felty aerial mycelium, orangewhite 

with pale brown 3 mm brim; margin diffuse; reverse pale 

brown to dark brown. Neither perithecia nor conidiomata observed 

in cultures on PDA, MEA and MYA after 4.5 mo at 2/10 C. 

Habitat: On petioles and basal vein parts of fallen leaves of 

Liquidambar styraciflua (Hamamelidaceae). 

Distribution: U.S.A. (AL, DE, GA, LA, MD, MS, NC, NJ, SC, TN, 

TX, VA). 

Lectotype designated here: U.S.A., North Carolina, L.D. Schweinitz 

(Shear Study Collection Types & Rarities Series I, BPI 800519). 

Epitype designated here: U.S.A., Virginia, Accomack Co., 

southern part of Assateague Island, 09 May 2004, M.V. Sogonov 

MS0037 (BPI 844274, ex-type culture CBS 121227). 

Additional specimens examined: U.S.A., Alabama, Septent, 2 June 1854, T.M. Peters, 

D.A. Watt Herb. 569, Missouri Bot. Gard. Herb. 60514 (BPI 611281); Delaware, 

Newark, 25 May 1908, H.S. Jackson 2159 (BPI 611282); Georgia, Darien, date 

unknown, H.W. Ravenel, Fungi Americani Exsiccati 374 (BPI 611547); Louisiana, 

04 March 1896, collector unknown, Herbarium of Rev.A.B. Langlois (BPI 596288); 

Louisiana, 22 Dec. 1888, and others, collector unknown, Herbarium of Rev. A.B. 

Langlois (BPI 596290); Louisiana, 23 March 1893, collector unknown, Herbarium 

of Rev. A.B. Langlois (BPI 596291); Louisiana, St. Martinsville, March 1890, Rev. 

A.B. Langlois, Ellis & Everhart 2543 (BPI 596292); same location, 11 Nov. 1890, 

Rev. A.B. Langlois, Herb. S.M. Tracy (BPI 596295); Louisiana, near St. Martinsville, 

05 Nov. 1899, Rev. A.B. Langlois, Flora Ludoviciana (BPI 596294); Maryland, St. 

Mary’s, 29 May 1921, C.L. Shear (BPI 611546); Maryland, Suitland, date unknown, 

H.H. Whetzel, R.W. Davidson et al. (BPI 611280); Maryland, Prince George Co., 

Greenbelt, Greenbelt Park, 21 Apr. 2004, M.V. Sogonov MS0028 (BPI 877511); 

Maryland, Prince George’s Co., Beltsville, BARC, forest near B011A, 06 Apr. 2005, 

M.V. Sogonov MS0178 (BPI 877510); Mississippi, Pike Co., Percy Quinn State 

Park, 27 Feb. 2006, M.V. Sogonov MS0331 (BPI 877509); New Jersey, Monmouth 

Co., Turkey Swamp Wildlife Management Area, 08 Jan. 1995, G. Bills (BPI 802807); 

South Carolina, date unknown, collector unknown, Michener Collection, Shear Study 

Collection Types & Rarities Series I (BPI 800520); ibid. (BPI 800521); Tennessee, 

Great Smoky Mountains National Park, Cosby Cabin, 13 May 2002, L.N. Vasilyeva 

(BPI 843530, culture CBS 116866); Tennessee, Great Smoky Mountains National 

Park, Tremont, 04 June 2002, L.N. Vasilyeva (BPI 863545); Texas, Houston, year 

1869, H.W. Ravenel (BPI 596289). 

Additional sequence from GenBank: U.S.A., North Carolina, 

Durham, Duke Forest, litter, date unknown, H.E. O’Brien, J.L. 

Parrent, J.A. Jackson, J.-M. Moncalvo, R. Vilgalys, nrDNA ITS1– 

5.8S–ITS2 (AY969703). 

Notes: Ambarignomonia petiolorum is an extremely common 

species on petioles of overwintered leaves of sweetgum 

(Liquidambar styraciflua) in eastern North America. This species 

is easily identified by the whitish powdery collar surrounding the 

central neck. 

APIOGNOMONIA Höhn., Ber. Deutsch. Bot. Ges. 35: 635. 1917. 

Type species: Apiognomonia veneta (Sacc. & Speg.) Höhn. 

Perithecia solitary, on fallen leaves, epiphyllous or on petioles, or 

on dead but still attached pedicels of trees and shrubs, or on dead 

parts of herbaceous plants, otherwise in groups of 5–15 perithecia 

with or without weakly developed stroma on twigs of trees and 

shrubs. Perithecia black, remaining immersed in substrate, oblate 

to spherical when moist, convex, sometimes with some irregular 

dents when dry, round in top view, with one neck. Necks central 

to marginal, never truly lateral, mostly 0.5–2 perithecial diam long 

but varying from almost lacking to length 3–4 times perithecial 

diam. Asci fusiform, with an apical ring, with eight spores arranged 

irregularly multiseriate or obliquely uniseriate. Ascospores mostly 

two-celled, rarely one-celled, oval to fusiform, l:w 2.5–6; ends 

mostly rounded, rarely pointed; appendages mostly absent or less 

commonly present, subulate, navicular or whip-shaped, to 30 µm 

long. 

Cultures: Colonies fast growing, often reaching edges of 90 mm Petri 

plates after 2 wk at 23 °C l/d or at least 60–70 mm diam. Colonies 

floccose or lanose all over surface or in lobes or concentric rings 

intermingled with glabrous or velvety areas. Colonies whitish, grey, 

orange-grey, brownish orange, dark brown, olive. Some species 

produce fertile perithecia in culture after 5–6 mo at 2/10 °C l/d. 

Conidiomata often produced after 2–4 wk at 23 °C l/d. 

Hosts: In diverse taxonomic groups (Aceraceae, Ericaceae, 

Euphorbiaceae, Fagaceae, Geraniaceae, Hippocastanaceae, 

Oleaceae, Platanaceae, Polygonaceae, Salicaceae). Most species 

are specific to one host species or genus, however, a few species 

are on a diverse range of plants. 

Apiognomonia veneta (Sacc. & Speg.) Höhn., Ann. Mycol. 16: 51. 

1918. 

≡ Laestadia veneta Sacc. & Speg., Michelia 1: 351. 1878. 

≡ Apiospora veneta (Sacc. & Speg.) Kleb., Z. Pflanzenkrankh. 7: 258. 

1902. 

36


Fig. 18. Morphology on natural substrate, perithecia. A–C. Apiognomonia acerina. A, C. BPI 877677. B. BPI 877678. D, E. A. hystrix. D. Monod 464, LAU bound. E. BPI 877692. 

A, B, D, E. Intact air-dry perithecia on stems, twigs, leaves and petioles. C. Extracted and rehydrated perithecia. Scale 200 μm. 

Fig. 19. Morphology on natural substrate, asci and ascospores. A, B. Apiognomonia acerina, BPI 877677. C–E. A. hystrix. C. BPI 877697. D. BPI 877698. E. BPI 877692. Scale 

10 μm. 


37


Fig. 20. Apiognomonia borealis CBS 799.79, culture morphology, colony habit, 40 d, 23 ºC. A, C, E. Surface. B, D, F. Reverse. A, B. PDA. C, D. MEA. E, F. MYA. Scale 1 cm. 

[≡ Gnomonia veneta (Sacc. & Speg.) Kleb., Jahrb. Wiss. Bot. 41: 533. 

1905 non Speg. 1879.] 

≡ Gnomonia platani Kleb., Vortr. Ges. Bot. 1: 28. 1914. 

Habitat: On overwintered leaves of Platanus occidentalis and P. 

orientalis (Platanaceae), rarely, Tilia sp. (Tiliaceae). 

Distribution: Widespread in temperate regions including Canada 

(British Columbia), Europe (Bulgaria, France, Germany, 

Switzerland), New Zealand, and U.S.A. (MD, TN). 

Notes: A detailed description of Apiognomonia veneta and its 

distinction from the closely related A. errabunda is provided by 

Sogonov et. al. (2007). 

Additional species of Apiognomonia 

The following taxa are accepted species of Apiognomonia based 

on their inclusion in multigene and ITS phylogeny. 

Apiognomonia acerina (Starbäck) M. Monod, Beih. Sydowia 9: 

63. 1983. Figs 18A–C; 19A,B. 

≡ Gnomonia acerina Starbäck, Bih. K. Svenska Vetensk Akad. Handl. 14, 

Afd. 3, n. 5: 17. 1889. 

Habitat: On fallen leaves of Acer opalus Mill., A. platanoides L., and 

A. pseudoplatanus L. (Aceraceae). 

Distribution: Europe (Bulgaria, Germany, Switzerland) 

Specimen examined: Switzerland, Valais, Salvan/Les Marécottes, Pont du Triège, 

1300 m a.s.l., on overwintered leaves of Acer pseudoplatanus, May 2005, coll. M. 

Monod (BPI 877677) GenBank EU254990. 

Notes: Among species of Gnomoniaceae on Acer, Apiognomonia 

acerina is unique in having ascospores that are wider than 3.5 µm 

and having a central neck. Barr (1978) considered the basionym 

Gnomonia acerina to be synonym of Apioplagiostoma aceriferum 

(Cooke) M.E. Barr but Monod (1983) recognised this name as a 

distinct species and provides detailed descriptions of both species. 

ITS sequences of Apiognomonia acerina and Apioplagiostoma 

aceriferum show these species to be distinct. Fig. 1 shows 

Apioplagiostoma acerifum in Pleuroceras, a genus not detailed in 

this study. 

Apiognomonia borealis (J. Schröt.) M. Monod, Beih. Sydowia 9: 

61. 1983. Figs 20A–F. 

≡ Gnomonia borealis J. Schröt., Jahresber. Schles. Ges. Vaterl. Cult. 65: 

275. 1888. 

= Gnomonia pratensis Svrček, Česká Mykol. 28: 219. 1974 fide Monod 1983. 

Habitat: On overwintered leaves and stems of Geranium pratense 

L., G. sanguineum L., and G. sylvaticum L. (Geraniaceae). 

Distribution: Europe (Czech Republic, Finland, Norway, Sweden, 

Switzerland). 

Specimen examined: Switzerland, Vaud, col du Mollendruz, on Geranium 

sylvaticum, Monod 274, CBS 796.79, GenBank EU254999. 

Notes: Apiognomonia borealis is distinguished from other species 

of Gnomoniaceae on Geranium by the ascospores having a 

supramedium septum. Monod (1983) provides a detailed description 

of this species. 

Apiognomonia errabunda (Roberge) Höhn., Ann. Mycol. 16: 51. 

1918. 

≡ Sphaeria errabunda Roberge in Desm., Ann. Sci. nat. Bot., ser. 3 10: 

355. 1848. 

≡ Gnomonia errabunda (Roberge) Auersw. in Gonn. & Rabenh., Mycol. 

Eur. 5/6, p. 25. 1869. 

More synonyms are listed in Sogonov et al. (2007). 

Habitat: On overwintered leaves primarily of hardwoods trees in the 

Fagaceae, Salicaceae, and Tiliaceae as well as other woody and 

herbaceous plants including Chamerion angustifolium (L.) Holub, Rhus 

glabra L. and Sorbus aria (L.) Crantz as listed in Sogonov et al. (2007). 

Distribution: Widespread in northern temperate regions as listed in 

Sogonov et al. (2007). 

38


Fig. 21. Morphology on natural substrate, perithecia. A–C. Gnomoniopsis chamaemori, BPI 877438. D–G. G. clavulata. D–F. Epitype BPI 877443. G. Lectotype BPI 611339. 

H–K. G. paraclavulata, holotype BPI 877448. L, M. G. fructicola. L. BPI 877446. M. BPI 877454. N, O. G. racemula, BPI 871003. P. Q. G. cf. chamaemori. P. BPI 877452A. Q. 

BPI 877456. A, B, D, H, I, L–N, P, Q. Intact air-dry perithecia on leaves and stems. C, E–G, J, K, O. Extracted and rehydrated perithecia. Scale 200 μm. 


39


Fig. 22. Morphology on natural substrate, asci and ascospores. A–C. Gnomoniopsis chamaemori, neotype BPI 877438. D–F. G. clavulata. D. BPI 877440. E. BPI 877442. F. 

BPI 877441. G, H. G. paraclavulata. G. Holotype BPI 877448. H. BPI 877449. I. G. fructicola, BPI 877446. J. G. racemula, BPI 871003. K, L. G. cf. chamaemori. K. BPI 877455. 

L. BPI 877456. Scale 10 μm. 

Notes: Apiognomonia errabunda is the cause of oak anthracnose 

(Sinclair & Lyon 2005 as A. veneta). A detailed description including 

the differences between A. errabunda and the closely related A. 

veneta are provided by Sogonov et. al. (2007). Like many species 

in the Gnomoniaceae, A. errabunda is frequently isolated as an 

endophyte in woody plants. 

Apiognomonia hystrix (Tode : Fr.) Sogonov, comb. nov. 

MycoBank MB512172. Figs 18D,E; 19C–E. 

Basionym: Sphaeria hystrix Tode, Fungi Meckl. 2: 53. 1791. 

≡ Diatrype hystrix Tode : Fr., Sum. Veg. Scand.: 383. 1846. 

≡ Mamiana hystrix (Tode : Fr.) De Not., Comment. Soc. Crittog. Ital. 1: 

43. 1863. 

≡ Cryptospora hystrix (Tode : Fr.) Fuckel, Jb. Nassau Ver. Naturk. 23–24: 

194. 1870. 

≡ Diaporthe hystrix (Tode : Fr.) Sacc., Fung. Ven. 4: 6. 1873. 

≡ Chorostate hystrix (Tode : Fr.) Traverso, Fl. Ital. Crypt. 2: 212. 1906. 

40


Fig. 23. Gnomoniopsis chamaemori CBS 803.79, culture morphology, colony habit. A, C, E. Surface. B, D, F. Reverse. A, B. PDA. C, D. MEA. E, F. MYA. Scale 1 cm. 

≡ Cryptodiaporthe hystrix (Tode : Fr.) Petr., Ann. Mycol. 19: 119. 1921. 

= Valsa longirostris Tul. & C. Tul., Sel. Fung. Carp. 2: 200. 1863 fide Wehmeyer 

1933. 

≡ Diaporthe longirostris (Tul. & C. Tul.) Sacc., Syll. Fung. 1: 609. 1882. 

= Diaporthe mamiania Sacc., Syll. Fung. 1: 609. 1882 fide Wehmeyer 1933. 

≡ Chorostate mamiania (Sacc.) Traverso, Fl. Ital. Crypt. 2: 201. 1906. 

= Sphaeria cerastis Riess, Hedwigia 1: 24. 1853. 

≡ Gnomonia cerastis (Riess) Ces. & De Not., Comment. Soc. Crittog. Ital. 

1: 233. 1863. 

= Sphaeria petioli Fuckel, Jahrb. Ver. Naturkunde Herzogthume Nassau 15: 68. 

1860 fide Monod 1983. 

≡ Gnomonia petioli (Fuckel) Cooke in Rabenh., Fungi Europaei exsiccati 

927. 1866. 

= Gnomoniella brunaudiana Pass. in Brunaud, Champ. Saint. 5, 1, 1891 fide 

Monod 1983. 

= Gnomoniella hippocastani Brunaud, Bull. Soc. Bot. Fr. 36: 336. 1889 fide 

Monod 1983. 

= Gnomonia aesculi Oudem., Beih. bot. Centralbl. 11: 527. 1902 fide Monod 

1983. 

= Gnomonia cerastis Riess f. nedundinis Karakulin, Morbi plant. Script. Lect. 

Phyto. Hort. Bot. U.S.S.R. 14: 81. 1925 fide Monod 1983. 

Habitat: On overwintered leaves, twigs and branches of Acer 

pseudoplatanus (Aceraceae) and various other hardwoods. 

Distribution: Europe (Austria, Bulgaria, Czech Republic, Germany, 

Switzerland). 

Specimens examined: Canada, Ontario, Etobicoke, Dean West Park, on 

overwintered leaves of Acer saccharum, 1 Apr 2005, coll. M.V. Sogonov (BPI 877696) 

GenBank EU255019. Russia, Novgorod province, Kholm, Arboretum (Dendropark), 

on overwintered leaves of Fraxinus excelsior, 7 Jun 2005, coll. M.V. Sogonov (BPI 

877698) GenBank255022. The Netherlands, Baarn, Garen Eemnersserweg 90, on 

leaf spot of seedling of Acer negundo, Oct 1997, coll. H.A. van der Aa 12406 (CBS 

100566) GenBank EU255032. 

Notes: Apiognomonia hystrix as Cryptodiaporthe hystrix with its 

synonym Gnomonia cerastis and its relationship to members of the 

Gnomoniaceae was recognised by Monod (1983) who provides a 

detailed description of this species as C. hystrix. 

GNOMONIOPSIS Berl., Icon. Fung. 1: 93. 1894. 

Type species: Gnomoniopsis chamaemori (Fr.) Berl. 

Perithecia solitary or groups up to 5, without stroma, on fallen, 

overwintered leaves and twigs of trees and shrubs, usually 

epiphyllous or on petioles, on dead parts of herbaceous plants. 

Perithecia black, remaining immersed, spheroidal to suboblate 

when moist, convex or irregularly dented when dry, round in top 

view, with one neck. Necks central to lateral, slightly curved to 

curved, shorter or slightly longer than perithecial diam, sometimes 

almost absent. Asci oval to fusiform, with an apical ring, with eight 

spores arranged mostly biseriate or obliquely uniseriate, less 

commonly irregularly multiseriate. Ascospores two-celled, oval 

to fusiform, l:w 1.5–5, usually somewhat ovoid or pyriform; ends 

rounded; appendages absent. 

Cultures: Colonies fast growing, often reaching egdes of 90 mm 

Petri plates after 2 wk at 23 °C l/d, or moderately growing 40–60 

mm diam. Colony surface usually glabrous, velvety or lanose. 

Colonies whitish, grey, dark brown, olive. Some species produce 

fertile perithecia in culture after 5–6 mo at 2/10 Cl/d; rarely fertile 

perithecia produced after 2–4 wk at 23 °C l/d. Conidiomata 

produced by most species after 2–4 wk at 23 °C l/d. 

Hosts: In diverse taxonomic groups (Ericaceae, Fagaceae, 

Pinaceae, Rosaceae). Most species are specific at the host species 

or genus level; however, a few species occur on a diverse range 

of plant hosts. 

Type species of Gnomoniopsis 

Gnomoniopsis chamaemori (Fr.) Berl., Icon. Fung. 1: 93. 1894. 

Figs 21A–C; 22A–C; 23A–F. 

≡ Sphaeria chamaemori Fr., Syst. Mycol. 2: 519. 1823. 

Perithecia hypophyllous or less commonly epiphyllous, immersed, 

subepidermal, mostly on veins, black, oblate spheroidal when moist, 

150–220 µm high × 210–320 µm diam, convex when dry. Necks 

central, straight, 60–85 µm long, diam 45–65 µm. Asci fusiform or 

obclavate, 28–40 × 7–9 µm, apical ring 1.5–2 µm diam, with eight 

ascospores arranged irregularly multiseriate or obliquely uniseriate. 

Ascospores fusiform, straight to slightly curved, (10–)10.5–11.5(–13) 

× (2–)2.5(–3) µm (mean = 11 × 2.5, SD 0.5, 0.3, n=28), l:w (3.3–)4– 

4.8(–5.5) (mean = 4.4, SD 0.6), two-celled, not constricted or slightly 

constricted at septum, septum located at (27–)33–38(–46) % (mean 

= 36, SD 5) of ascospore length, ends blunt, rounded, each cell with 

two large guttules, or with one big guttule and several small ones, or 

several indistinct guttules; appendages absent. 


41


Fig. 24. Gnomoniopsis clavulata, culture morphology. A–F. CBS 121231. G–L. CBS 121257. M–R. Ex-type CBS 121259. S–W. G. cf. clavulata, CBS 119028. A–T, V, W. Colony 

habit, 40 d, 23 ºC. A, C, E, G, I, K, M, O, Q, S, V. Surface. B, D, F, H, J, L, N, P, R, T, W. Reverse. U. Conidiomata, 17 d, 23 ºC. A, B, G, H, M, N, S, T. PDA. C, D, I, J, O, P, U. 

MEA. E, F, K, L, Q, R, V, W. MYA. Scale: A–T, V, W. 1 cm. U. 1 mm. 

42


Fig. 25. Gnomoniopsis clavulata, culture morphology, 40 d, 23 ºC. A–F. Ex-type CBS 121259. G–I. CBS 121257. A–C. Conidiomata. D–I. Conidia. A, D, G. PDA. B, E, H. MEA. 

C, F, I. MYA. Scale: A–C. 1 mm. D–I. 10 μm. 

Cultures: Colonies on PDA usually attaining 90 mm after 40 d at 23 

°C, flat, velutinous to shortly woolly, dark brown in centre, gradually 

lightening to pale reddish grey at margin; margin diffuse; reverse 

of almost same colours as surface. Colonies on MEA attaining 90 

mm after 40 d at 23 °C, flat, almost glabrous, overlaid by loose and 

short woolly-like mycelium, pale reddish grey with indistinct pale 

orange-brown patterns in centre; margin diffuse; reverse of almost 

same colours as surface. Colonies on MYA attaining 90 mm after 

40 d at 23 °C, flat, dark brown in centre, brown with some shades 

of red, becoming pale reddish grey at margin, overlaid by whitish 

woolly, aerial mycelium; margin diffuse; reverse of same colours 

as surface. 


43


Habitat: On overwintered leaves of Rubus chamaemorus L. 

(Rosaceae). 

Distribution: Europe (Finland, Russia). 

Specimen examined: Russia, Novgorod oblast, Kholm raion, Rdeyskiy Zapovednik, 

vicinity of Fryunino, on overwintered leaves of Rubus chamaemorus, 11 Jun. 2005, 

M.V. Sogonov & D.A. Maykov MS0273 (BPI 877438) GenBank EU254809. 

Additional culture examined: Finland, Oulanka, on overwintered leaves of Rubus 

chamaemorus, 10 Jul. 1977, M. Monod, No. 345 (culture CBS 803.79). 

Specimens examined of G. aff. chamaemori: Bulgaria, Sredna Gora Mt (western), 

Lozenska Planina, above Pancharevo lake, near the track from ‘Stenata’ locality 

to VEC Kokaljane, on overwintered stems of Agrimonia eupatoria, 21 May 2005, 

coll. D. Stoykov (BPI 877452A) GenBank EU254812. Russia, Novgorod province, 

Kholm, on dead petioles of Potentilla anserina, 7 Jun 2005, coll. M.V. Sogonov (BPI 

877455), GenBank EU254811; Tver’ province, Toropets district, v. Bubonitsy, on 

overwintered stems of Potentilla canescens, 14 Jun 2005, coll. M.V. Sogonov (BPI 

877456) GenBank254810. 

Notes: Monod (1983) provided a detailed description of this species 

as Gnomonia chamaemori. 

New and revised species of Gnomoniopsis 

Gnomoniopsis clavulata (Ellis) Sogonov, comb. nov. MycoBank 

MB 512173. Figs 21D–G; 22D–F; 24A–W; 25A–I. 

Basionym: Gnomonia clavulata Ellis, Amer. Nat. 17: 318. 1883. 

≡ Didymiella clavulata (Ellis) Sacc., Syll. Fung. 9: 666. 1891. 

≡ Cercidospora clavulata (Ellis) Kuntze, Rev. Gen. Pl. 3 (2): 453. 1898. 

Perithecia solitary, without stroma, hypophyllous, scarce, mostly in 

upper and marginal parts of leaf blades, spheroidal when moist, 

110–150 µm high × 120–140 µm wide, convex when dry. Necks 

central, slightly flexuous, (158–)160–166(–169) µm long (mean = 

163, SD 8, n=2), (37–)37.5–39(–39.5) µm wide at base, (34.5–)36– 

39(–40.5) µm wide at apex. Asci fusiform to cylindrical, (28–)33.5– 

41.5(–47) × (6.5–)7–10(–11) µm (mean = 38 × 8.5, SD 5.5, 1.5, 

n=34), apical ring 1.5–2.5 µm diam, with eight ascospores biseriate 

or obliquely uniseriate. Ascospores pyriform, inequilateral, (5–)8.5– 

9.5(–11) × (2–)3.5–4(–5.5) µm (mean = 9 × 4, SD 1, 0.5, n=149), 

l:w (1.8–)2.2–2.4(–3) (mean = 2.3, SD 0.2), two-celled, constricted 

at septum; septum located at (29–)37–43(–49) % (mean = 40, SD 

4) of ascospore length; ends broadly rounded, distal cell with 2–3 

and basal cell with 1–2 small guttules, sometimes both cells without 

guttules; appendages absent. 

Cultures: Colonies on PDA usually attaining 90 mm after 40 d 

at 23 °C, in CBS 121257 slower growing, attaining 50 mm, flat, 

velutinous to woolly, pale brown to brown, overlaid by scant or 

abundant orange-grey, greyish orange or brownish orange slimy 

conidial mass drops, ex-epitype culture CBS 121259 with pale grey 

woolly mycelium; margin clear, even to lobate; reverse pale brown 

or orange-brown to dark brown; agar stained by yellow soluble 

pigment in some strains; conidia oval to oblong, sometimes slightly 

obovoid, straight or curved, allantoid or sigmoid, (5–)6–6.5(–8) 

× (2–)2.5–3(–4) µm (mean = 6.5 × 3, SD 0.5, 0.3, n=285), l:w 

(1.4–)2.1–2.6(–3.7) (mean = 2.4, SD 0.4, n=285). Colonies on MEA 

attaining 90 mm after 40 d at 23 °C, flat, thin, semitransparent, 

loosely woolly, colourless or whitish, with areas of pale orange, 

brown or dark brown, with numerous orange-grey or greyish 

orange slimy conidial mass drops; margin diffuse; reverse of same 

colours as surface; conidia oval to oblong, sometimes slightly 

obovoid, straight or slightly curved (4.5–)6–7(–8) × (2–)2.5–3(–3.5) 

µm (mean = 6.5 × 2.5, SD 0.5, 0.3, n=315), l:w (1.4–)2.2–2.7(–3.8) 

(mean = 2.5, SD 0.4). Colonies on MYA attaining 90 mm after 40 

d at 23 °C, flat, with areas of whitish to greyish orange felty aerial 

mycelium and areas of brownish yellow, pale brown, brown or 

brownish grey woolly, partly fasciculate mycelium; orange-grey or 

greyish orange slimy conidial mass drops; margin clear, even or 

irregular; reverse greyish orange, orange-brown, greyish brown or 

dark brown; conidia oval to oblong, sometimes slightly obovoid, 

straight or slightly curved (4.5–)6–6.5(–8) × (2–)2.5–3(–3.5) µm 

(mean = 6 × 3, SD 0.6, 0.4, n=151), l:w (1.4–)2–2.5(–3.5) (mean 

= 2.2, SD 0.4). No perithecia observed in cultures at 2/10 °C after 

8 mo. 

Habitat: On overwintered leaves of Quercus spp. (Fagaceae). 

Distribution: U.S.A. (MD, NC, NJ, TN, VA). 

Lectotype designated here: U.S.A., New Jersey, Newfield, on 

Quercus nigra, May 1884, J.B. Ellis, North American Fungi 1685 

(lectotype BPI 611339; isotype BPI bound). 

Epitype designated here: U.S.A., Maryland, Prince George’s Co., 

Riverdale, Anacostia River Park, on Quercus marilandica, 12 Jun. 

2006, M.V. Sogonov MS0401 (BPI 877443, ex-epitype culture CBS 

121259) GenBank EU254820. 

Additional specimens examined: U.S.A., Maryland, Prince George’s Co., Beltsville, 

Beltsville Agricultural Research Center, near building 011A, on Q. falcata, 06 Apr. 

2005, M.V. Sogonov MS0181 (BPI 877441); same location, Q. rubra, 29 Jun. 2005, 

M.V. Sogonov MS0206 (BPI 877444); same location, Q. prinus, 19 May 2006, 

M.V. Sogonov MS0371 (BPI 877477); same location, Q. rubra, 19 May 2006, M.V. 

Sogonov MS0434 (BPI 877522); same location, Q. falcata, 08 Jun. 2006, M.V. 

Sogonov MS0397 (BPI 877439, culture CBS 121255) GenBank EU254818; North 

Carolina, Wake Co., Raleigh, Carl Alwin Schenk memorial forest, on Q. ilicifolia, 03 

Apr. 2005, M.V. Sogonov MS0161 (BPI 877442, culture CBS 121239) GenBank 

EU254816; Tennessee, Sevier Co., Greenbrier, University of Tennessee field 

station, Conley Huskey Way, on Q. falcata, 25 May 2004, M.V. Sogonov MS0399 

(BPI 877440, culture CBS 121257) GenBank EU254819; Virginia, Albermarle Co., 

Charlottesville, University of Virginia campus, between Edgement Road and U.S. 

route 29 BYP, on Q. prinus, 02 Mar. 2005, M.V. Sogonov MS0139 (BPI 871056, 


Additional cultures examined: U.S.A., Maryland, Prince George’s Co., Patuxent 

Wildlife Research Center, on Quercus rubra, S. Cohen (R153 = AR 4123 = CBS 

121911); same data, (R154 = AR 4124) GenBank EU254814. 

Additional culture G. cf. clavulata: Switzerland, isol. from Fagus 

sylvatica, AR 4183 = CBS 119028 (BPI 871052) GenBank 

EU254817. 

Notes: Gnomoniopsis clavulata is common on overwintered leaves 

of oak (Quercus spp.) in eastern North America and was frequently 

isolated as an endophyte from Quercus rubra (Cohen, 1999; 

2004) mistakenly identified as Discula umbrinella. Gnomoniopsis 

clavulata and G. paraclavulata are distinct from most species of 

Gnomoniaceae on Quercus in having ascospores with a submedian 

septum. The ascospores of G. clavulata are larger than those of G. 

paraclavulata. 

Gnomoniopsis paraclavulata Sogonov, sp. nov. MycoBank MB 

512174. Figs 21H–K; 22G,H; 26A–M. 

Perithecia (139–)149–170(–180) µm alta × (156–)188–231(–241) 

44


Fig. 26. Gnomoniopsis paraclavulata, culture morphology. A–M. Ex-type CBS 121263. N–S. G. cf. paraclavulata CBS 121269. A–F, N–S. Colony habit, 40 d, 23 ºC. A, C, E, 

N, P, R. Surface. B, D, F, O, Q, S. Reverse. G–J. Conidiomata, 40 d, 23 ºC. K–M. Conidia, 40 d, 23 ºC. A, B, G, K, N, O. PDA. C, D, H, I, L, P, Q. MEA. E, F, J, M, R, S. MYA. 

Scale: A–F, N–S. 1 cm. G–J. 1 mm. K–M. 10 μm. 


45


Fig. 27. Culture morphology. A–N. Gnomoniopsis fructicola CBS 121226. O–S. G. macounii CBS 121468. T–Y. G. racemula CBS 121469. A–F, O, P, R–Y. Colony habit, 40 d, 

23 ºC. A, C, E, O, R, T, V, X. Surface. B, D, F, P, Q, U, W, Y. Reverse. G. Perithecia and coniodiomata, 40 d, 23 ºC. I. Perithecia immersed in agar, reverse view, 40 d, 23 ºC. J. 

Coniodiomata, 40 d, 23 ºC. K, L. Asci and ascospores, 40 d, 23 ºC. M. Conidia, 40 d, 23 ºC. N. Conidia, 40 d, 2/10 ºC. Q. Conidia, 4.5 mo, 2/10 ºC. A, B, G, K, M–P, T, U. PDA. 

C, D, I, L,Q, V, W. MEA. E, F, J, R, S, X, Y. MYA. Scale: A–F, O, P, R–Y. 1 cm. G–J. 1 mm. K–N, Q. 10 μm. 

46


µm diam. Rostrum (157–)180–21)180–210(–216) µm longum, basi 

(37.3–)38.9–41(–41.5)–41.5) µm diam, apice (40.8–)41.1–42.9(– 

44.3) µm diam. Ascosporae pyriformes, inaequilaterae (8–)9–10(– 

11) × (3–)3.5–4 µm, L:l (2.1–)2.4–2.8(–3.6). Similis to G. clavulatae, 

sed ascosporarum Longitudo/latitudo ratione leviter majore et septi 

positione inferiore differt. Holotypus: BPI 877448. 

Etymology: Refers to similarity and affinity with G. clavulata. 

Perithecia solitary, without stroma, hypophyllous, scarce, mostly in 

upper and marginal parts of leaf blades, spheroidal when moist, 

(139–)149–170(–180) µm high × (156–)188–231(–241) µm diam 

(mean = 159 × 206, SD 20, 44, n=3), convex when dry. Necks 

central, slightly flexuous, (157–)180–210(–216) µm long (mean = 

192, SD 31, n=3), (37–)39–41(–42) µm wide at base, 41–43(–45) 

µm wide at apex. Asci fusiform to cylindrical, (38–)45–48(–51.5) × 

7–8.5(–10.5) µm (mean = 46 × 8, SD 4, 1.5, n=7), apical ring 2–2.5 

µm diam, with eight ascospores biseriate or obliquely uniseriate. 

Ascospores pyriform, inequilateral, (8)9–10(–11) × (3–)3.5–4 µm 

(mean = 9.5 × 3.5, SD 0.5, 0.3, n=24), l:w (2.1–)2.4–2.8(–3.6) 

(mean = 2.6, SD 0.3), two-celled, constricted at septum; septum 

located at (25–)31–37(–44) % (mean = 34, SD 4) of ascospore 

length; distal and basal cells usually with correspondingly 1–5 and 

0–2 small guttules; appendages absent. 

Cultures: Colonies on PDA usually attaining 90 mm after 40 d at 23 

°C, flat, with pale red to greyish orange, smooth to shortly woolly 

and whitish to reddish white woolly areas, overlaid by abundant pale 

orange to orange-grey slimy conidial masses; margin submerged, 

irregular; reverse with areas of yellow-grey, pale orange and 

brownish orange; conidia oval to oblong, sometimes slightly 

obovoid, straight or slightly curved (6–)7.5–8(–9.5) × (2–)3–3(–3.5) 

µm (mean = 7.5 × 3, SD 0.5, 0.3, n=108), l:w (1.6–)2.4–2.9(–4.2) 

(mean = 2.6, SD 0.4). Colonies on MEA attaining 90 mm after 40 d at 

23 °C, flat, thin, semitransparent, colourless with brown centre and 

scattered flocks of white aerial mycelium, with black conidiomata 

which, at maturity, produce orange-grey slimy conidial mass drops; 

margin diffuse; reverse greyish orange to orange-grey; conidia oval 

to oblong or obovoid, straight or slightly curved (6.5–)7.5–8.5(–9.5) 

× (3–)3–3.5 µm (mean = 8 × 3.5, SD 0.5, 0.2, n=75), l:w (2–)2.3– 

2.6(–3.2) (mean = 2.5, SD 0.2). Colonies on MYA attaining 90 mm 

after 40 d at 23 °C, flat, whitish to orange-white felty, with small 

areas of shorter brownish grey woolly, partly fasciculate mycelium; 

moderate number of orange-grey or greyish orange slimy conidial 

mass; margin irregular, partly submerged; reverse greyish orange, 

orange-brown, greyish brown or brown, conidia obovoid to oblong, 

straight or slightly curved (6–)7–7.5(–8.5) × (2.5–)3(–3.5) µm (mean 

= 7 × 3, SD 0.5, 0.2, n=60), l:w (1.6–)2.3–2.6(–3.4) (mean = 2.4, SD 

0.3). No perithecia observed in cultures at 2/10 °C after 8 mo. 

Habitat: On overwintered leaves of Quercus alba L. (Fagaceae). 

Distribution: U.S.A. (MD, TN). 

Holotype: U.S.A., Tennessee, Sevier Co., Greenbrier, University of 

Tennessee field station, Conley Huskey Way, 22 May 2004, M.V. 

Sogonov MS0406 (BPI 877448, ex-type culture CBS121263) 

Additional specimens examined: U.S.A., Maryland, Prince George’s 

Co., Beltsville, Beltsville Agricultural Research Center, near building 

011A, 14 Feb. 2005, M.V. Sogonov MS0127 (BPI 877450) GenBank 

EU 254837; same location, 20 Mar. 2005, M.V. Sogonov MS0152 

(BPI 877449) GenBank EU 254838. 

Additional cultures examined: U.S.A., Maryland, Prince George’s Co., Patuxent Wild 

Life Research Center, S. Cohen W623 = AR 4125; same data W633 = AR 4126, 

GenBank EU254835; same data (W645 = AR 4127 = CBS 123202). 

Notes: Gnomoniopsis clavulata and G. paraclavulata are distinct 

from most species of Gnomoniaceae on Quercus in having 

ascospores with submedian septum. The ascospores of G. 

clavulata are larger than those of G. paraclavulata. 

Additional species accepted in Gnomoniopsis 

Gnomoniopsis comari (Karst.) Sogonov, comb. nov. MycoBank 

MB 512175. 

Basionym: Gnomonia comari Karst., Mycol. Fenn. 2: 122. 1873. 

≡ Gnomoniella comari (Karst.) Sacc., Syll. Fung. 1: 415. 1882. 

Habitat: On overwintered leaves of Comarum palustre L. 

(Rosaceae). 

Distribution: Europe (Finland, Germany, Switzerland) 

Specimen examined: Finland, on Comarum palustre, Monod 

366, CBS 806.79, GenBank EU254821; Monod 353, CBS 807.79, 

GenBank EU 254822. 

Notes: The concept of Gnomoniopsis comari is conceived here in 

a much narrower sense than by Monod (1983), thus the numerous 

taxonomic synonyms listed by Monod (1983) are not included. The 

multigene phylogeny presented here suggests that G. comari is 

distinct from G. fructicola (Fig. 1). 

Gnomoniopsis fructicola (Arnaud) Sogonov, comb. nov. 

MycoBank MB 512176. Figs 21L,M; 22I; 27A–N. 

Basionym: Gnomonia fragariae f. fructicola Arnaud, Traité de Pathol. Veg. p. 1558. 

1931. 

≡ Gnomonia fructicola (Arnaud) Fall., Can. J. Bot. 29: 309. 1951. 

Habitat: On overwintered leaves and fruits of Fragaria spp. 

(Rosaceae), occasionally pathogenic on fruits causing strawberry 

stem-end rot. The causal organism has often been referred to as 

Gnomonia comari, now considered Gnomoniopsis comari. 

Distribution: Canada (British Columbia), Europe (Belgium, France) 

and U.S.A. (MD, NY). 

Specimens examined: Belgium, on Fragaria sp., CBS 255.61, GenBank EU254828. 

Canada, Ontario, on Fragaria sp., CBS 275.51, GenBank EU254829. France, on 

Fragaria sp. coll. G. Arnaud, CBS 208.34, GenBank EU254826. U.S.A., New York, 

Sullivan Co., Roscoe, area around Campbell Inn, on dead petioles of Fragaria sp., 

Jul 2005, coll. M.V. Sogonov (BPI 877446) GenBank EU254830. 

Specimens examined G. cf. fructicola: Russia, Novgorod province, Kholm, valley of 

Lovat’ river, on dead petioles of Geum rivale, 10 Jun 2005, coll. M.V. Sogonov (BPI 

877454) GenBank EU254832. 

Notes: Considerable confusion has existed among the species 

of Gnomoniopsis on Fragaria. Gnomonipsis fructicola is herein 

recognised to be distinct from G. comari. “Gnomonia” fragariae 

Kleb. causes another disease of strawberry in Europe called leaf 


47


blotch, root rot and petiole blight (Maas 1998, Moročko et al. 2006). 

Moročko & Fatehi (2007) determined that “Gnomonia” fragariae 

belongs outside of the Gnomoniaceae in the Sydowiellaceae. 

Gnomoniopsis macounii (Dearn.) Sogonov, comb. nov. 

MycoBank MB 512177. Figs 27O–S. 

Basionym: Diaporthe macounii Dearn., Mycologia 8:100. 1916. 

≡ Cryptodiaporthe macounii (Dearn.) Wehm., The Genus Diaporthe: 191. 

1933. 

Habitat: On overwintered branches of Spiraea douglasii Hook. var. 

menziesii (Hook.) C. Presl and Spiraea sp. (Rosaceae). 

Distribution: Canada (British Columbia) and U.S.A. (NH, NY). 

Notes: Barr (1978) and Wehmeyer (1933) provide a detailed 

description of G. macounii as C. macounii. 

Gnomoniopsis racemula (Cooke & Peck) Sogonov, comb. nov. 

MycoBank MB 512178. Figs 21N,O; 22J; 27T–Y. 

Basionym: Sphaeria racemula Cooke & Peck in Peck, Ann. Rep. New York State 

Museum 26: 87. 1874 

≡ Diaporthe racemula (Cooke & Peck) Sacc., Syll. Fung. 1: 691. 1882. 

≡ Ditopellopsis racemula (Cooke & Peck) M.E. Barr, Mycol. Mem. 7: 91. 1978. 

Habitat: On overwintered stalks of Chamerion angustifolium 

(Onagraceae). 

Distribution: Canada (British Columbia) and U.S.A. (ME, MN, NY, 

OR). 

Notes: Gnomoniopsis racemula is unusual in this genus in having 

perithecia in groups of 3–9 that occur on fibrous overwintered 

stalks. Barr (1978) provided a detailed description of G. racemula 

as Ditopellopsis racemula. 

Gnomoniopsis tormentillae (Lind) Sogonov, comb. nov. 

MycoBank MB 512179. 

Basionym: Gnomoniella tormentillae Lind, Bot. Tidsskr. 41: 217. 1931. 

≡ Plagiostoma tormentillae (Lind) Bolay, Ber. Schweiz. Bot. Ges. 81: 436. 

1971. 

Habitat: On overwintered petioles, veins and stalks of Potentilla 

canadensis L. and P. erecta (L.) Raeusch (Rosaceae). 

Distribution: Europe (Switzerland) and U.S.A. (MA). 

Notes: Both Barr (1978) and Monod (1983) provide a detailed 

description of Gnomoniopsis tormentillae as P. tormentillae. The 

perithecial neck of this species is marginal. 

OPHIOGNOMONIA (Sacc.) Sacc., Syll. Fung. 14: 613. 1899. 

Lectotype species designated by Höhnel (1919): Ophiognomonia 

melanostyla (DC. : Fr.) Berl. 

≡ Gnomoniella subgenus Ophiognomonia Sacc., Syll. Fung. 1: 419. 

1882. 

Perithecia solitary, without stroma, on underside of leaf blade, 

petioles or rachises, occasionally on upper side of blade of 

overwintered fallen leaves, rarely on dead but attached pedicels, 

and on dead stems of herbaceous plants. Perithecia dark brown to 

black, remaining immersed or becoming partly erumpent at maturity, 

oblate when moist, convex or irregularly shrunk when dry, in some 

species, part of perithecia may be concave, round in top view, with 

one neck. Neck central to eccentric, rarely marginal, never truly 

lateral, mostly length 2.5–5 perithecial diam, in some species 

shorter, down to length of one perithecial diam. Asci oval to almost 

filiform, with an apical ring, with eight spores per ascus arranged 

mostly unevenly parallel, also irregularly multiseriate or obliquely 

uniseriate, occasionally evenly parallel. Ascospores mostly twocelled, 

rarely one-celled, oval to filiform, l:w 2.5–25; ends rounded, 

with or without appendages, may vary within species. 

Cultures: Colonies growing at a moderate rate, reaching 1–6 cm 

diam in 2 wk at 23 °C l/d, in some strains reach edges of 90 mm 

Petri plates in 2 wk on PDA. Colony surface leathery to coarsely 

farinose or velvety, in some species with floccose areas. Colonies 

mostly whitish, yellow, greyish yellow, pale orange, olive-brown. 

Some species produce fertile perithecia in culture after 5–6 mo at 

2/10 °C l/d, rarely sterile perithecia formed within one month at 23 

°C l/d. Conidiomata in cultures formed by a few species but then 

not requiring long-term cultivation at low temperatures. 

Hosts: Mostly on Fagales (Betulaceae, Fagaceae, Juglandaceae), 

a few species on Lauraceae, Rosaceae, Salicaceae, and Tiliaceae. 

Individual fungal species are host-specific at genus or, less 

commonly, at family level. 

Type species of Ophiognomonia 

Ophiognomonia melanostyla (DC. : Fr.) Berl., Icon. Fung. 2: 146. 

1899. Figs 28A–C; 29A–C. 

≡ Sphaeria melanostyla DC. : Fr., Fl. Franç., 5/6: 129. 1815 : Syst. Mycol. 

2: 517. 1823. 

≡ Gnomonia melanostyla (DC. : Fr.) Auersw. in Gonn. & Rabenh., Mycol. 

Europ. 5/6: 28. 1869. 

≡ Gnomoniella melanostyla (DC. : Fr.) Sacc., Syll. Fung. 1: 419. 1882. 

≡ Cryptoderis melanostyla (DC.: Fr.) G. Winter, Rabenhorst’s Kryptogamen- 

Flora I, Abt. 2: 592. 1887. 

Perithecia solitary, without stroma, hypophyllous, evenly distributed 

over large areas of leaf blades, sometimes on upper part of petioles, 

immersed at first, partly erumpent at maturity, oblate to suboblate 

when moist, 180–220 µm high × 220 350 µm diam, convex, 

occasionally irregularly dented or concave when dry. Necks central 

or eccentric, usually sinuous, 550–1100 µm long, 30–45 µm wide 

at base, 25–33 µm wide at apex. Asci narrowly fusiform, 55–65 

× 4.5–5 µm, apical ring 1–1.5 µm diam, with eight ascospores 

evenly or slightly unevenly parallel. Ascospores clavately filiform, 

slightly sinuous (30–)37–42.5(–44) × 1.5–2 µm (mean = 39 × 1.5, 

SD 4, 1, n=16), l:w (20.3–)22.4–26.4(–29.3) (mean = 24.6, SD 

2.6), two-celled, slightly constricted at septum, septum located at 

(55–)65–68(–71) % (mean = 66, SD 4) of ascospore length, ends 

blunt, rounded, basal cell is narrower than distal cell, ca. 1.2 µm 

wide, each cell with a few small guttules; appendages subulate to 

whip-shaped, 5–25 µm long. 

Habitat: On fallen overwintered leaves of Tilia spp. (Tiliaceae). 

48


Fig. 28. Morphology on natural substrates, perithecia. A–C. Ophiognomonia melanostyla. A. Lectotype G 00053951. B. Epitype BPI 877610. C. BPI 877611. D–F. O. balsamiferae, 

holotype BPI 877606. G, H. O. pseudoclavulata. G. BPI 877615B. H. BPI 877631. A, D, F, G. Intact air-dry perithecia on leaves and petioles. B, C, E, H. Extracted and rehydrated 

perithecia. Scale 200 μm. 


49


Fig. 29. Morphology on natural substrates, asci and ascospores. A–C. Ophiognomonia melanostyla. A, B. Epitype BPI 877610. C. BPI 877607. D, E. O. balsamiferae, holotype 

BPI 877606. F–H. O. pseudoclavulata. F. Holotype BPI 844280. G. BPI 877632. H. BPI 877633A. Scale 10 μm. 

50


Distribution: Europe (Austria, Bulgaria, Czech Republic, Germany, 

Switzerland, Ukraine), Canada (Ontario) and U.S.A. (NY, PA) 

Lectotype: Switzerland, vicinity of Geneva, Tilia sp., March, year 

unknown, M. Chaillet, (G 00053951). 

Additional specimens examined: Austria, Sonntagberg, near Rosenau, on Tilia 

sp., Apr., year unknown, P.P. Strasser (BPI 596571); Czech Republic, Moravia, 

Hranice na Moravě, Teplice, on Tilia platyphyllos, Apr. 1914, F. Petrak (BPI 

596581); same location, on Tilia sp., May 1924, F. Petrak (BPI 596572); Germany, 

Frankensteinerkopf near Oestrich (Nassau), on Tilia parvifolia, Spr. 1894, L. Fuckel 

(BPI 596576, BPI 596577); Oestrich (Nassau), on Tilia parvifolia, 1894, L. Fuckel 

(BPI 596575); Switzerland, Vaud, Lausanne, Parc Bourge, on Tilia cordata, 28 May 

2005, M.V. Sogonov MS0333 (BPI 877611) GenBank EU254913; Vaud, St. Cergue, 

on Tilia cordata, 20 May 2005, M.V. Sogonov MS0197 (BPI 877610) GenBank 

EU254911; Ukraine, Lviv oblast, Stryi raion, Pidhirtsi, on Tilia platyphyllos, 27 

Mar. 1918, F. Petrak (BPI 596579); U.S.A., New York, Heldenburg Mts., on Tilia 

americana, May, year unknown, C.H. Peck (BPI 596574); New York, Ithaca vicinity, 

Arnot forest, on Tilia americana, 10 Jul. 2002, L.N. Vasilyeva MS0353 (BPI 877609); 

New York, Sullivan Co., Roscoe vicinity, area around Campbell Inn, on Tilia 

americana, Jul. 2005, M.V. Sogonov MS0299 (BPI 877608) GenBank EU254912; 

Pennsylvania, Franklin Co., Cove Gap, Buchanan Birthplace State Park, on Tilia 

americana, 05 May 2006, M.V. Sogonov MS0358 (BPI 877607). 

Notes: Ophiognomonia melanostyla is relatively common on 

overwintered leaves of Tilia spp. The very long ascospores over 

35 µm long distinguish this species from other species of the 

Gnomoniaceae on Tilia. 

New species of Ophiognomonia 

Ophiognomonia balsamiferae Sogonov, sp. nov. MycoBank MB 

512180. Figs 28D–F; 29D,E; 30A–N. 

Perithecia 320–390 µm alta × 370–425 µm diam. Rostrum 940– 


Ascosporae fusiformes, leviter curvatae, (15–)18–19(–21) × 

2.5–3(–3.5) µm, L:l (4.9–)6.2–7.2(–8.1). Ad aliis Ophiognomoniae 

speciebus morphologiae characteribus combinatis differt. 

Singularis Ophiognomoniae species lecta in Salicaceis. Holotypus: 

BPI 877606. 

Etymology: Named after the epithet of the plant host. 

Perithecia solitary, without stroma, evenly and densely distributed 

over petioles, immersed or partly emerging, dark brown, oblate 

spheroidal when moist, 320–390 µm high × 370–425 µm diam, 

convex when dry, round from top. Necks central, straight, curved 

or flexuous when dry, straight when moist, 940–1150 µm long, 

73–90 µm wide at base, 42–55 µm wide at apex. Asci fusiform 

with tapering stipe, 36–70 × 9–17 µm, apical ring 2.5–3 µm diam, 

with eight ascospores arranged obliquely uniseriate, irregularly 

multiseriate or unevenly parallel. Ascospores fusiform, slightly 

curved, (15–)18–19(–21) × 2.5–3(–3.5) µm (mean = 18.5 × 3, SD 

1.5, 0.3, n=29), l:w (4.9–)6.2–7.2(–8.1) (mean = 6.7, SD 0.7), twocelled, 

slightly constricted at septum; septum located at (44–)47– 

51(–52) % (mean = 48, SD 2) of ascospore length, cells tapering, 

at ends blunt, rounded or indistinctly truncated, each cell with 3–5, 

guttules, often one large guttule close to septum; appendages 

subulate to navicular, 10–15 µm long. 

Cultures: Colonies on PDA attaining 90 mm after 40 d at 23 °C, 

flat, short woolly in centre, velvety with loose tufts at margin, pale 

brownish grey to brown; margin very irregular; reverse dark brown. 

Colonies on MEA attaining 90 mm after 40 d at 23 °C, flat, woolly, 

whitish, with scarce dark brown amorphous conidiomata attaining 

500 µm diam; margin diffuse; reverse orange-white to orange and 

brownish orange; conidia oval, cylindrical, oblanceolate or allantoid, 

(6–)8.5–10(–13.5) × (1.5–)2–2.5(–3) µm (mean = 9.5 × 2.5, SD 1.5, 

0.5, n=61); l:w (1.8–)3.6–4.5(–7.2) (mean = 4.1, SD 0.9). Colonies 

on MYA almost attaining 90 mm after 40 d at 23 °C, flat, felty to 

woolly, pale brownish grey to dark brown, with droplets of clear 

exudate; margin irregular; reverse dark brown. Cultures at 2/10 °C 

after 4.5 mo produce dark thick-walled conidiomata with conidia on 

MYA, sterile conidioma-like structures on PDA with sparse conidia 

after 8 mo, on MEA, sterile after 8 mo. 

Distibution: Canada (British Columbia). 

Habitat: On overwintered petioles of Populus balsamifera L. 

(Salicaceae). 

Holotype: Canada, British Columbia, Manning Provincial Park, 

rest area at West Gate, beginning of Engineers Loop Trail, 13 May 

2006, M.V. Sogonov, MS0409 (BPI 877606, ex-type culture AR 

4320 = CBS 121266). 

Notes: Ophiognomonia balsamiferae has a central neck on the 

perithecium unlike other species of Gnomoniaceae on Populus, 

specifically Apioplagiostoma populi and Plagiostoma salicella 

in which the necks are lateral. Gnomonia gnomon is known to 

occur rarely on Populus but has ascospores that are considerably 

narrower, 1.5–2 µm wide, than those of O. balsamiferae. 

Ophiognomonia pseudoclavulata Sogonov, sp. nov. MycoBank 

MB 512181. Figs 28G–H; 29F–H; 30O–Z. 


µm longum, basi 37–54 µm diam, apice 34–44 µm diam. Ascosporae 

late ellipsoidal vel ellipsoidal, rectae vel leviter inaequilateralae, 

(6.5–)7.5–8(–9) × (2.5–)3–3.5(–3.6) µm (mean = 7.7 × 3.1, SD 

0.6, 0.3, n=112), L:l (2.12–)2.3–2.7(–3.4). Ad aliis Ophiognomoniae 

speciebus parvis peritheciis et brevibus ascosporis differt. Similis 

Gnomoniopsis clavulata et G. paraclavulata, sed ascosporis raro 

clavatis et septis ascosporarum fere semper in medio differt. 

Holotypus: BPI 844280. 

Etymology: Refers to the confusion with Gnomoniopsis clavulata. 

The oldest specimen of O. pseudoclavulata observed in this study 

was originally identified as G. clavulata. 

Perithecia solitary, without stroma, hypophyllous, mostly on and 

next to midrib, or scattered randomly over leaf blade, immersed, 

dark brown, oblate when moist, 170–190 µm high × 210–280 µm 

diam, convex when dry. Necks curved when dry, slightly curved 

when moist, 140–250 µm long, 37–54 µm wide at base, 34–44 

µm wide at apex. Asci ellipsoidal to fusiform, with tapering stipe, 

(30–)33.5–41.5(–46) × (6.5–)7–9.5(–11.5) µm (mean = 37.5 × 8.5, 

SD 4.5, 1.5, n=24), apical ring 2–3 µm diam, with eight ascospores 

arranged biseriate or irregularly multiseriate. Ascospores broadly 

ellipsoidal to ellipsoidal, often broader in upper part, straight or 

slightly inequilateral, (6.5–)7.5–8(–9) × (2.5–)3–3.5 µm (mean = 

7.5 × 3, SD 0.5, 0.3, n=112), l:w (2.1–)2.3–2.7(–3.4) (mean = 2.5, 

SD 0.3), two-celled, not constricted at septum; septum located at 


51


Fig. 30. Culture morphology. A–N. Ophiognomonia balsamiferae ex-type CBS 121266. O–Z. O. pseudoclavulata. O–T. Ex-type CBS 121236. U–Z. CBS 121232. A–L, O–Z. 

Colony habit. A, C, E, G, I, K, O, Q, S, U, W, Y. Surface. B, D, F, H, J, L, P, R, T, V, X, Z. Reverse. M. Conidiomata. N. Conidia. A–F. 14 d, 23 ºC. G–Z. 40 d, 23 ºC. A, B, G, H, 

O, P, U, V. PDA. C, D, I, J, M, N, Q, R, W, X. MEA. E, F, K, L, S, T, Y, Z. MYA. Scale: A–L, O–Z. 1 cm. M. 200 μm. N. 10 μm. 

52


(40–)47–51(–58) % (mean = 49, SD 3) of ascospore length; cells 

broadly rounded at ends, without guttules or with 2–5 small guttules; 

appendages absent or of different irregular shapes or filiform to 20 

µm long. 

Cultures: Two cultures (CBS 121232 and CBS 121236) differ 

significantly in their morphology, especially on MEA and MYA. 

CBS 121232: Colonies on PDA attaining 60 mm diam after 40 d at 

23 °C, with lobate convex concentric zones, felty to woolly, whitish; 

margin well-defined, lobate; reverse dark brown in central part, 

then greyish orange, at margin whitish. Colonies on MEA attaining 

22 mm diam after 40 d at 23 °C, slightly radially furrowed, velvety 

to woolly, orange-white; margin well-defined, slightly wavy; reverse 

dark brown in central part, then greyish orange, at margin orangewhite. 

Colonies on MYA attaining 30 mm diam after 40 d at 23 °C, 

slightly furrowed, in centre glabrous greyish orange with tufts of 

white aerial mycelium, then woolly, whitish, at margin felty, whitish; 

margin well-defined, lobate; reverse dark brown in central part, then 

greyish orange, at margin orange-white. CBS 121236: Colonies on 

PDA attaining 65 mm diam after 40 d at 23 °C, flat, felty to woolly, 

whitish; margin well-defined, serrately lobate; reverse dark brown 

in central part, then greyish orange, at margin whitish. Colonies on 

MEA attaining 70 mm diam after 40 d at 23 °C, flat, felty to woolly, 

white; margin diffuse, broadly indistinctly lobate; reverse orangewhite 

with small greyish orange area in centre. Colonies on MYA 

attaining 65 mm diam after 40 d at 23 °C, flat, felty, whitish; margin 

well-defined, even; reverse dark brown in central part, then orange. 

Neither perithecia nor conidiomata observed in cultures on PDA, 

MEA and MYA after 8 mo at 2/10 C. 

Habitat: On overwintered leaves of Carya spp., primarily C. 

tomentosa (Lam.) Nutt. (mockernut hickory) (Juglandaceae). 

Distribution: U.S.A. (DC, IL, IN, MD, NC, NJ, PA, TN, VA). 

Holotype: U.S.A., Pennsylvania, Kennett Square Co., vicinity of 

Philadelphia, near Phillips mushroom farm, Carya tomentosa, 17 

Apr. 2004, M.V. Sogonov MS0025 (BPI 844280, ex-holotype culture 

AR 4059 = CBS 121236). 

Additional specimens examined: U.S.A., District of Columbia, National Arboretum, 

Carya tomentosa, 03 May 2005, M.V. Sogonov MS0355a (BPI 877615B); Illinois, 

Hancock Co., St. Mary’s Township, Section 27, Apr. 2006, L.C. Castlebury MS0527 

(BPI 877520); Indiana, Clark State Forest, Carya sp., 16 Jan 2005, M.V. Sogonov 

MS113 (BPI 877630) GenBank EU254925; Maryland, Prince Georges Co., Beltsville, 

B.A.R.C., Entomology Rd., Carya tomentosa, 04 Apr 2004, M.V. Sogonov MS0012a 

(BPI 877613B) GenBank EU254922; Maryland, Prince George’s Co., Beltsville, 

B.A.R.C., forest near Building 011A, Carya sp., 13 Jan 2005, M.V. Sogonov MS0112 

(BPI 877631) GenBank EU254924; North Carolina, Wake Co., Raleigh, Carl Alwin 

Schenk memorial forest, Carya tomentosa, 03 Apr 2005, M.V. Sogonov MS0165 

(BPI 877633A) GenBank EU254927; New Jersey, Newfield, Carya sp., Apr 1891, 

J.B. Ellis, North American Fungi 3429 (BPI 611620); Tennessee, Blount Co., 

Great Smoky Mountains National Park, Cades Cove, Anthony Creek Trail, Carya 

tomentosa, 24 May 2006, M.V. Sogonov MS0488 (BPI 877521A); Tennessee, 

Blount Co., Great Smoky Mountains National Park, Cades Cove, Carya tomentosa, 

24 May 2006, A.Y. Rossman, MS0470 (BPI 877632); Tennessee, Sevier Co., Univ. 

of Tennessee field station, Conley Huskey Way, Carya tomentosa, 23 May 2006, 

M.V. Sogonov MS0469 (BPI 877519); same data MS0471a (BPI 877667B); Virginia, 

Albermarle Co., Charlottesville, University of Virginia Campus, between Edgement 

Road and highway US 29 BYP, Carya sp., 02 Mar. 2005, M.V. Sogonov MS0140 

(BPI 871057, culture CBS 121232) GenBank EU254926. 

Notes: Ophiognomonia pseudoclavulata is distinguished from 

similar species on Carya in the Gnomoniaceae by the relatively 

short ascospores compared to those of O. micromegala (Ellis 

& Everh.) Sogonov that are 26–36 × 5.5–10 μm and Gnomonia 

caryae Wolf that are (16–)22–30(–37) × (2–)3–5.5 µm fide Barr 

(1978). 

Ophiognomonia vasiljevae Sogonov, sp. nov. MycoBank MB 

512182. Figs 31A,B; 32A–B; 33A–I. 


µm logum, basi 75–85 µm diam, apice 32–42 µm diam. Ascosporae 

fusiformes, leviter curvatae (17.5–)18.5–19.5(–21) × (2.5–)3(–3.5) 

µm (mean = 19 × 3, SD 1, 0.2, n=31), L:l (5.4–)5.9–6.5(–7.4) 

(mean = 6.3, SD 0.5, n=31). Ad aliis Ophiognomoniae speciebus 

morphologiae characteribus combinatis differt. Holotypus: BPI 

877671. 

Etymology: Named after the Russian mycologist Larissa 

Vasilyeva in recognition of her contribution to the taxonomy of the 

Gnomoniaceae. 

Perithecia solitary, without stroma, in small loose groups on 

compound leaf rachises, immersed, from upper side not easily 

detachable from plant tissue, peroblate when moist, 310–390 µm 

high × 590–690 µm diam, convex when dry. Necks eccentric to 

lateral, slightly flexuous, 520–640 µm long, 75–85 µm wide at base, 

32–42 µm wide at apex. Asci fusiform with narrow stipe, (52.5–) 

58.5–64.5(–74) × (10–)11.5–13.5(–17) µm (mean = 62 × 12.5, SD 

5.5, 1.7, n=13) , apical ring 2.5–3 µm diam, with eight ascospores 

arranged obliquely uniseriate or irregularly multiseriate. Ascospores 

fusiform, slightly curved, (17.5–)18.5–19.5(–21) × (2.5–)3(–3.5) µm 

(mean = 19 × 3, SD 1, 0.2, n=31), l:w (5.4–)5.9–6.5(–7.4) (mean = 

6.3, SD 0.5), two-celled, not constricted at septum; septum located 

at (42–)48–50(–54) % (mean = 49, SD 3) of ascospore length; cells 

strongly tapering, at ends blunt, rounded, each cell with 2–3 large 

guttules with largest guttule close to septum or with numerous small 


Cultures: Colonies on PDA attaining 90 mm after 40 d at 23 °C, flat, 

short, loose, woolly, orange, in some areas overlaid with whitish 

aerial mycelium; margin diffuse; reverse orange to brownish 

orange. Colonies on MEA attaining 90 mm after 40 d at 23 °C, flat, 

short, loose, woolly, brownish orange, in some areas overlaid with 

whitish aerial mycelium; margin diffuse; reverse brownish orange. 

Colonies on MYA attaining 80 mm diam after 40 d at 23 °C, flat, 

woolly, whitish; margins submerged, orange; margin irregular; 

reverse orange. Cultures at 2/10 °C produce sterile perithecia and 

dark amorphous bodies on PDA after 4.5 mo, on MEA after 8 mo. 

No such structures were observed after 8 mo on MYA. 

Habitat: On overwintered leaf rachises of Juglans nigra L. 

(Juglandaceae). 

Holotype: U.S.A., Tennessee, Blount Co., Great Smoky Mountains 

National Park, along loop near the Methodist Church, 24 May 2006, 

M.V. Sogonov MS0388 (BPI 877671, ex-holotype culture CBS 

121253). 

Notes: In distinguishing species of Ophiognomonia on Juglans, O. 

vasiljevae is similar to O. ischnostyla in having a neck longer than 

250 µm but, unlike O. leptostyla, which has a neck less than 250 

µm long. Ascospores of O. vasiljevae are generally greater than 

17.5 µm long while those of O. ischnostyla are less than 17.5 µm. 


53


Fig. 31. Morphology on natural substrates, perithecia. A, B. Ophiognomonia vasiljevae, holotype BPI 877671. C. O. alni-viridis, BPI 877585A. D. O. gei-montani, BPI 877589. 

E–G. O. intermedia. E. BPI 877498. F. BPI 877598. G. BPI 877496. A, C–E, G. Intact air-dry perithecia on leaves and stems. B, F. Extracted and rehydrated perithecia. Scale 

200 μm. 

54


Fig. 32. Morphology on natural substrates, asci and ascospores. A, B. Ophiognomonia vasiljevae, holotype BPI 877671. C. O. alni-viridis, BPI 877600. D. O. gei-montani, BPI 

877589. E–I. O. intermedia. E. BPI 877498. F. BPI 877598. G. BPI 877602. H. BPI 877488B. I. BPI 877496. Scale 10 μm. 

Additional species accepted in Ophiognomonia 

Ophiognomonia alni-viridis (Podlahova & Svrček) Sogonov, 

comb. nov. MycoBank MB 512215. Figs 31C; 32C; 33J–S. 

Basionym: Gnomonia alni-viridis Podlahova & Svrček, Česká Mykol. 24: 129. 

1970. 

= Gnomonia intermedia var. alni M.E. Barr, Mycol. Mem. 7: 55. 1978 fide 

Monod 1983. 

Habitat: On overwintered leaves of Alnus viridis (Chaix) DC. (Betulaceae). 

Distribution: Canada (British Columbia), Europe (Bulgaria, Czech 

Republic, Switzerland) and U.S.A. (WA). 

Specimens examined: Switzerland, Valais, vicinity of Martigny, on overwintered leaves of 

Alnus viridis, 21 May 2005, coll. M. Monod (BPI 877585A) GenBank EU254866. Canada, 

British Columbia, 15 km S from Princeton, near Indian reserve #3, on overwintered leaves 

of ?Betula papyrifera, 13 May 2006, coll. M.V. Sogonov (BPI 877600) GenBank EU254869. 

U.S.A., Washington, King Co., Mount Baker-Snoqualmie National Forest, Snoqualmie 

Ranger District, near exit 42 on the highway US 90, road to mines, on overwintered leaves 

of Alnus viridis, 16 May 2006 (BPI 877595) GenBank EU254867. 

Notes: Ophiognomonia alni-viridis can be distinquished from the 

other species of Gnomoniaceae on Alnus. Gnomonia alnea lacks an 

elongated neck unlike O. alni-viridis, O. ischnostyla and O. trientensis. 

Ophiognomonia trientense has an ascospore l:w less than 3 while O. 

alni-viridis and O. ischnostyla both have an ascospore l:w greater than 3. 

The ascospores of O. alni-viridis are 10–12.5 × 2–2.5 µm fide Podlahova 

& Svrček (1970) while those of O. ischnostyla fide Monod (1983) are 

longer, 12.5–18.5 × 1.5–2.5 µm. For a more detailed description of O. 

alni-viridis, see Monod (1983) and Podlahova & Svrček (1970). 

Ophiognomonia gei-montani (Ranoj.) Sogonov, comb. nov. MycoBank 

MB 512183. Figs 31D; 32D. 

Basionym: Gnomonia gei-montani Ranoj., Ann. Mycol. 8: 362. 1910. 


55


Fig. 33. Culture morphology. A–I. Ophiognomonia vasiljevae ex-type CBS 121253. J–S. O. cf. alni-viridis CBS 121250. A–F, J–O. Colony habit, 40 d, 23 ºC. A, C, E, J, L, N. 

Surface. B, D, F, K, M, O. Reverse. G–I. Sterile perithecia or perithecium-like bodies, 2/10 ºC. P. Young perithecia, 2/10 ºC. Q, R. Perithecia, 2/10 ºC. G, H, Q, R. 4.5 mo. I. 8 

mo. P. 40 d. A, B, G, H, Q¬–S. PDA. C, D, I, P. MEA. E, F. MYA. G, H, Q–S. Scale: A–F, J–O. 1 cm. G–I, P–R. 200 μm. S. 10 μm. 

56


Fig. 34. Culture morphology, colony habit. A–F. Ophiognomonia cf. intermedia CBS 121229. G–L. O. cf. ischnostyla CBS 121234. M–R. O. cf. ischnostyla BPI 877467B. S–X. 

O. cf. ischnostyla CBS 121252. A, C, E, G, I, K, M, O, Q, S, U, W. Surface. B, D, F, H, J, L, N, P, R, T, V, X. Reverse. A, B, G, H, M, N, S, T. PDA. C, D, I, J, O, P, U, V. MEA. E, 

F, K, L, Q, R, W, X. MYA. Scale 1 cm. 


57


Fig. 35. Morphology on natural substrates, perithecia. A, B. Ophiognomonia ischnostyla. A. BPI 877514B. B. BPI 877620. C–E. O. cf. ischnostyla. C, D. BPI 877605A. E. BPI 

877616. A, C, E. Intact air-dry perithecia on leaves and petioles. B, D. Extracted and rehydrated perithecia. Scale 200 μm. 

Habitat: On overwintered leaves of Geum bulgaricum Panc., G. coccineum 

Sm., G. montanum L., and G. rhodopeum Stoj. & Stef. (Rosaceae). 

Distribution: Europe (Bulgaria, Switzerland) 

Specimen examined: Switzerland, Salvan, La Tendraz, 1600 m a.s.l., on dead leaves of 

Geum montanum, 28 May 2005, coll. M. Monod (BPI 877589) GenBank EU254872. 

Notes: See Monod (1983) for a detailed description. 

Ophiognomonia intermedia (Rehm) Sogonov, comb. nov. 

MycoBank MB 512185. Figs 31E–G; 32E–I; 33A–F. 

Basionym: Gnomonia intermedia Rehm, Ann. Mycol. 6: 489. 1908. 

Habitat: On overwintered leaves of Betula nana L., B. papyrifera 

Marshall, B. pendula Roth, and B. pubescens Ehrh. (Betulaceae). 

Distribution: Canada (British Columbia), Europe (Germany, Russia, 

Scotland, Switzerland, United Kingdom) and U.S.A (MD). 

Specimens examined: Canada, British Columbia, 15 km NE from Agassiz, route 

#7, on overwintered leaves of Betula papyrifera, 13 May 2005, coll. M.V. Sogonov 

(BPI 877599) GenBank EU 254884; Burnaby Lake Regional Park, on overwintered 

leaves of Betula papyrifera, 12 May 2006, coll. M.V. Sogonov (BPI 877602) GenBank 

EU254886. . Russia, Tver’ province, Toropets district, v. Kosilovo, on overwintered 

leaves of Betula pendula, 5 Jun 2005, coll. M.V. Sogonov (BPI 877488B) GenBank 

EU254887; Novgorod province, Kholm district, Rdeysky Natural Reserve, vicinity 

of the village Fryunino, on overwintered leaves of Betula nana, 11 Jun 2005, coll. 

M.V. Sogonov (BPI 877496) GenBank EU254881; Naberezhnaya reki Lovat’ str., 9, 

on overwintered leaves of Betula pendula, 23 Aug 2004, coll. M.V. Sogonov (BPI 

877498) GenBank EU254878. U.S.A., Maryland, Prince George’s Co., Beltsville, 

Little Paint Branch Park, on overwintered leaves of Betula nigra, 17 Mar 2005, coll. 

M.V. Sogonov (BPI 877597) GenBank EU254879; 11 Apr 2005, coll. M.V. Sogonov 

(BPI 877598) GenBank EU254880. 

Notes: Among other species of Ophiognomonia on Betula, 

Ophiognomonia intermedia with two-celled ascospores is distinct 

from O. nana with one-celled ascospores. Ophiognomonia 

intermedia is similar to O. alni-viridis and O. ischnostyla in ascospore 

size except that the ascospores of O. intermedia lack appendages 

and tend to have a length wide ratio of less than 5. Ophiognomonia 

intermedia causes a foliar disease of birch that also causes dieback 

of young shoots (Green 2004). Green & Castlebury (2007) proved 

the connection between O. intermedia as G. intermedia with the 

asexual state Discula betulae (Westend.) Pennycook (Pennycook, 

2007). 

58


Fig. 36. Morphology on natural substrates, asci and ascospores. A, B. Ophiognomonia ischnostyla. A. BPI 877514B. B. BPI 877619. C–H. O. cf. ischnostyla. C, D. BPI 877605A. 

E. BPI 877605B. F. BPI 877467B. G. BPI 877622. H. BPI 877616. Scale 10 μm. 

Ophiognomonia ischnostyla (Desm.) Sogonov, comb. nov. 

MycoBank MB 512186. Figs 34G–L; 35A–E; 36A–H. 

Basionym: Sphaeria ischnostyla Desm., Annals Sci. nat., Bot., sér. 3. 11: 357. 

1849. 

≡ Gnomonia ischnostyla (Desm.) Auersw. in Gonn. & Rabenh., Mycol. 

Europ. 5/6: 2. 1869. 

= Gnomonia setacea f. alni Kleb., Haupt- und Nebenfruchtformen der 

Ascomyzeten: 244. 1918 fide Monod 1983. 

= Sphaeronema amenticolum Ces., Bot. Z. 15: 173. 1857 fide Monod 1983. 

≡ Gnomonia amenticola (Ces.) Prihoda, Česká Mykol. 10:122. 1956 fide 

Monod 1983. 

Habitat: On overwintered leaves of Alnus, Betula, Carpinus 

(Betulaceae), Juglans (Juglandaceae), and other hardwood trees. 

Distribution: Europe (Bulgaria, France, Russia, Switzerland) and 

U.S.A. (TN). 

Specimens examined: Russia, Tver’ province, Toropets district, v. Kosilovo, on 

overwintered leaves of Alnus glutinosa, 5 Jun 2005, coll. M.V. Sogonov (BPI 877617) 

GenGank EU254907; Tver’ province, Toropets district, vicinity of v. Bubonitsy, 

biological research station “Chisty Les”, on fallen leaves of Betula ? pubescens, 

31 Aug 2004, coll. M.V. Sogonov (BPI 877616) EU254919; on overwintered leaves 


59


Fig. 37. Morphology on natural substrates, perithecia. A, B. Ophiognomonia micromegala. A. BPI 877613A. B. BPI 877634B. C. O. rosae, BPI 877586. D. O. rubi-idaei, BPI 

877637. A, C, D. Intact air-dry perithecia on leaves. B. Extracted and rehydrated perithecium. Scale 200 μm. 

of Alnus glutinosa, 14 Jun 2005, coll. M.V. Sogonov (BPI 877618) GenBank 

EU254908; Novgorod province, Kholm district, Rdeysky Natural Reserve, vicinity 

of the village Fryunino, on overwintered leaves of Alnus glutinosa, 11 Jun 2005, 

coll. M.V. Sogonov (BPI 877619) GenBank EU294900; Arboretum (Dendropark), 

near the tree #560, on overwintered leaves of Corylus avellana, Jun 2005, coll. 

M.V. Sogonov (BPI 877514B) GenBank EU254899. Switzerland, Wallis, Mörel, on 

overwintered leaves of Alnus incana, 28 May 2005, coll. M.V. Sogonov (BPI 877620) 


Specimen examined O. cf. ischnostyla: Russia, Tver’ province, Toropets district, 

vicinity, the beginning of the Ecotrail, on fallen leaves of Betula pubescens, 31 Aug 

2004, coll. M.V. Sogonov (BPI 877605A) GenBank EU254895. 

Notes: Ophiognomonia ischnostyla occurs on a variety of hardwood 

trees especially in the Betulaceae. Opiognomonia ischnostyla 

has an elongated neck unlike Gnomonia alnea on Alnus. The 

ascospores of O. alni-viridis and O. ischnostyla have a l:w greater 

than 3 while O. trientense has an ascospore l:w less than 3. The 

ascospores of O. alni-viridis are 10–12.5 × 2–2.5 µm while those of 

O. ischnostyla are (12.5–)13.5–15.5(–18.5) × (1.5–)2(–2.5) µm fide 

Monod (1983). For a more detailed description of O. ischnostyla, 

see Monod (1983). 

The name Gnomonia nervisequa (Wallr.) Fuckel based on 

Sphaeria nervisequa Wallr., was proposed by Monod (1983) as 

the correct name for O. ischnostyla but this is rejected. Wallroth’s 

(1833) original description of the basionym is poor, does not 

indicate ascospore size, and lacks reference to any type specimen. 

The host plant mentioned in the description, Salix caprea, has been 

never reported in any later study. Nevertheless, Fuckel (1870) used 

this epithet creating the new combination Gnomonia nervisequa 

(Wallr.) Fuckel in reference to his own collection on Carpinus 

betulus. None of the species of Ophiognomonia are known to be 

associated with hosts in the Salicales. The next available basionym 

for G. nervisequa sensu Monod (1983) is Sphaeria ischnostyla 

Desm. The type specimen of Sphaeria ischnostyla on Carpinus 

betulus in France was examined at BPI (Desmazieres, Pl. crypt. 

France 2084-bound). 

60


Fig. 38. Morphology on natural substrates, asci and ascospores. A, B. Ophiognomonia micromegala, BPI 877615A. C, D. O. rosae. C. BPI 877636. D. BPI 877588. E, F. O. 

rubi-idaei, BPI 877637. Scale 10 μm. 


61


Fig. 39. Culture morphology. A–H. Ophiognomonia rosae CBS 121263. I–Q. O. sassafras CBS 121243. A–F, I–N. Colony habit, 40 d, 23 ºC. A, C, E, I, K, M. Surface. B, D, F, 

J, L, N. Reverse. G, O, P. Perithecia. H, Q. Asci. G, H. 40 d, 23 ºC. O–Q. 4.5 mo, 2/10 ºC. A, B, I, J, P. PDA. C, D, G, H, K, L, Q. MEA. E, F, M, O. MYA. Scale: A–F, I–N. 1 cm. 

G, O. 500 μm. P. 200 μm. H, Q. 10 μm. 

Ophiognomonia leptostyla (Fr.) Sogonov, comb. nov. MycoBank 

MB 512187. 

Basionym: Sphaeria leptostyla Fr., Syst. Mycol. 2: 517. 1823. 

≡ Gnomonia leptostyla (Fr.) Ces. & De Not., Comment. Soc. Crittog. Ital. 

1(4): 232. 1863. 

Habitat: On overwintered leaves of Juglans spp. (Juglandaceae). 

Distribution: Canada (Ontario), Europe (Austria, Bulgaria, Germany, 

Poland, Russia, Switzerland) and U.S.A. (AL, DE, IA, IL, MA, MD, 

NY, PA, VA, WV). 

Specimen examined: Switzerland, on Juglans regia, coll. M. Monod 439, GenBank 

EU254910. 

Notes: Ophiognomonia leptostyla is the cause of walnut anthracnose 

or walnut leaf blotch, a disease that is particularly virulent in the 

midwestern and eastern United States (Neely & Black 1976, Berry 

1981, Juhasova et al. 2006). The anamorph of O. leptostyla has been 

called Marssoniella juglandis (Lib.) Hohn. but that anamorphic genus is 

a later homonoym of an alga, thus Braun (1991) established the genus 

Neomarssoniella U. Braun. For more detailed descriptions see Barr 

(1978) and Monod (1983). 

62


Fig. 40. Morphology on natural substrates, perithecia. A–C. Ophiognomonia sassafras. A, B. BPI 877639. C. BPI 611592. D. O. cf. trientensis, BPI 877672. A, D. Intact air-dry 

perithecia on leaves. B, C. Extracted and rehydrated perithecia. Scale 200 μm. 

Ophiognomonia micromegala (Ellis & Everh.) Sogonov, comb. nov. 

MycoBank MB 512188. Figs 37A,B; 38A,B. 

Basionym: Diaporthe micromegala Ellis & Everh., Proc. Acad. nat. Sci. Philad. for 

1893: 449. 1894. 

≡ Plagiostoma micromegalum (Ellis & Everh.) M.E. Barr, Mycol. Memoir, 

7: 112. 1978. 

Habitat: On overwintered leaflets and rachises of Carya spp. 

(Juglandaceae). 

Distribution: U.S.A. (DC, DE, GA, MD) 

Specimens examined: U.S.A., Maryland, Montogomery Co., Chesapeake & Ohio 

Canal National Historic Park, on overwintered leaves of Carya tomentosa, 10 Apr 

2004, coll. M.V. Sogonov (BPI 877614) GenBank EU254916; Wheaton Regional 

Park, on overwintered leaves of Carya sp., 6 Mar 2005, coll. M.V. Sogonov (BPI 

877612) GenBank EU254917; Prince George’s Co., Beltsville, end of Entomology 

Road, on overwintered leaves of Carya tomentosa, 30 Mar 2004, coll. M.V. 

Sogonov (BPI 877634B) GenBank EU254914; 4 Apr 2004 (BPI 877613A) GenBank 

EU254915. 

Notes: Ophiognomonia micromegala with ascospores 32–45 × 

5.5–8 μm fide Barr (1978 as Plagiostoma micomegalum) is similar 

to Gnomonia caryae with ascospores (16–)22–30(–37) × (2–)3– 

5.5 μm fide Barr (1978), however, the latter species has thinner 

ascospores and central necks on the perithecia. Ophiognomonia 

pseudoclavulata has shorter ascospores than those of O. 

micromegala. Barr (1978 as Plagistoma micromegalum) provides 

a detailed description of this species as does Wehmeyer (1933 as 

Diaporthe micromegala). 

Ophiognomonia nana (Rehm) Sogonov, comb. nov. MycoBank 

MB 512189. 

Basionym: Gnomoniella nana Rehm, Hedwigia 42: 349. 1903. 

Habitat: On dead leaves of Betula nana L. (Betulaceae). 

Distribution: Europe (Finland, Germany, Switzerland). 

Notes: Ophiognomonia nana is distinguished from other species 

of Gnomoniaceae on Betula by the non-septate ascospores and 

beaks longer than 400 μm, and is unlike species of Gnomonia in 

having perithecia that do not become concave upon drying. For 

a more detailed description, see Monod (1983 as Gnomoniella 

nana). 

Ophiognomonia padicola (Lib.) M. Monod, Beih. Sydowia 9: 158. 

1983. 

≡ Sphaeria padicola Lib., Plant. Cryptog. Arduenn. Cent. 2, 149. 1832. 

≡ Gnomonia padicola (Lib.) Kleb., Z. Pflkrankh. 18:137. 1908. 

= Ophiognomonia padi Jaap, Verh. bot. Ver. Prov. Brandenburg 47: 87. 1905 

fide Monod 1983. 

Habitat: On overwintered leaves of Prunus padus L. (Rosaceae). 

Distribution: Europe (Germany, Switzerland) 

Notes: Ophiognomonia padicola is descried in detailed by Monod 

(1983) who placed the asexual state in Cylindrosporella. Gnomonia 

cerastis, previously reported on this host, is now considered a 

synonym of Apiognomonia hystrix. Ophiognomonia padicola 

has filiform ascospores more than 30 μm much longer than the 

ascospores of A. hystrix. 


63


Fig. 41. Morphology on natural substrates, asci and ascospores. A. Ophiognomonia sassafras, BPI 877639. B. O. cf. trientensis, BPI 877672. Scale 10 μm. 

Ophiognomonia rosae (Fuckel) Kirschst., Ann. Mycol. 37: 129. 

1939. Figs 37C; 38C,D; 39A–H. 

≡ Gnomonia rosae Fuckel, Jb. Nassau Ver. Naturk. 23–24: 122. 1870. 

≡ Gnomoniella rosae (Fuckel) Sacc., Syll. Fung. 1: 416. 1882. 

Habitat: On Rosa spp. and possibly other genera in the Rosaceae. 

Distribution: Europe (Germany, Russia) and U.S.A. (ME). 

Specimen examined: Russia, Tver’ province, Toropets district, v. Bubonitsy, near 

Bologovs’ house, on dead but attached twigs of Rosa sp., 14 Jun 2005, coll. M.V. 

Sogonov (BPI 877635) GenBank EU 254933. 

Notes: Ophiognomonia rosae having filiform ascospores has been 

reported on a number of rosaceous hosts; however, many of these 

specimens were not examined, thus a narrow concept of this 

species is retained. Monod (1983) provides a detailed description 

of this species as Gnomonia rosae. 

Ophiognomonia rubi-idaei (M. Monod) Sogonov, comb. nov. 

MycoBank 512190. Figs 37D; 38E,F. 

Basionym: Gnomonia rubi-idaei M. Monod, Beih. Sydowia 9: 106. 1983. 

Habitat: On overwintered leaves of Rubus idaeus L. (Rosaceae). 

Distribution: Canada (British Columbia) and Europe (Switzerland). 

Specimens examined: Canada, British Columbia, Manning Provincial Park, on 

overwintered leaves of Rubus sp., 13 May 2006, coll. M.V. Sogonov (BPI 877559B) 

GenBank EU254939; Victoria Island, Route 14, on overwintered leaves of Rubus 

spectabilis, 10 May 2006, coll. M.V. Sogonov (BPI 877638) GenBank EU 254938. 

Switzerland, on overwintered leaves of Rubus idaeus, 21 May 2005, coll. M.V. 

Sogonov (BPI 877637) GenBank EU254937). 

Notes: Ophiognomonia rubi-idaei is distinguished from other 

species in the Gnomoniaceae on Rubus by the filiform ascospores. 

Monod (1983) provides a detailed description as G. rubi-idaei. 

Ophiognomonia sassafras (Ellis & Everh.) M. Monod, Beih. 

Sydowia 9: 86. 1983. Figs 39I–Q; 40A–C; 41A. 

≡ Gnomonia sassafras Ellis & Everh., Bull. Torrey Bot. Club 10: 98. 1883. 

≡ Pleuroceras sassafras (Ellis & Everh.) M.E. Barr, Mycol. Mem. 7: 122. 1978. 

Habitat: On overwintered leaves of Sassafras officinale Nees. & 

Eberhm. (Lauraceae). 

Distribution: U.S.A. (MD, NJ, OH, PA). 

Specimen examined: U.S.A., Maryland, Howard Co., Columbia, Centennial Park, 

on overwintered leaves of Sassafras albidum, 9 Apr 2005, coll. M.V. Sogonov (BPI 

877642) GenBank EU254940. 

Notes: Ophiognomonia sassafras is the only species of 

Gnomoniaceae known on this plant host. For a detailed description, 

consult Barr (1978). 

Ophiognomonia setacea (Pers. : Fr.) Sogonov, comb. nov. 

MycoBank MB 512191. Figs 42A–X. 

Basionym: Sphaeria setacea Pers. : Fr., Syn. Method. Fung. p. 62. 1801 : Syst. 

Mycol. 2: 517. 1823. 

≡ Gnomonia setacea (Pers. : Fr.) Ces. & De Not., Comment. Soc. Crittog. 

Ital. 1: 232. 1863. 

Habitat: On overwintered leaves of Castanea dentata L., Castanea 

sp. and Quercus alba L., Q. bicolor Willd., Q. cerris L., Q. macrocarpa 

Michx., Q. montana Willd., Q. palustris Münchh., Q. phellos L., Q. 

pubescens Willd., Q. robur L., and Quercus sp. (Fagaceae). 

Distribution: Canada (Ontario), Europe (Austria, Bulgaria, Germany, 

Italy, Montenegro, Sweden, Switzerland) and U.S.A. (LA, MD, NJ, 

NY, OH, PA, TN, VA, WV). 

Notes: Ophiognomonia setacea was originally retained in 

Gnomonia by Sogonov et al. (2005) based on an analysis of the 

LSU of relatively few species in the Gnomoniaceae. However, 

the multigene phylogeny presented in this paper reveals that this 

species is allied with species of Ophiognomonia. Ophiognomonia 

setacea is conspicuous on overwintered leaves of chestnut and oak 

due to the very long, often over 500 μm, thin, black necks emerging 

from the leaf surface. Sogonov et al. (2005) provides an epitype 

and a detailed description of this species. 

Ophiognomonia trientensis (M. Monod) Sogonov, comb. nov. 

MycoBank MB 512192. Figs 40D; 41B. 

Basionym: Gnomonia trientensis M. Monod, Beih. Sydowia 9: 90. 1983. 

64


Fig. 42. Ophiognomonia setacea, culture morphology, colony habit, 40 d, 23 ºC. A–F. CBS 116406. G–L. CBS CBS116408. M–P. CBS 121235. Q, R. CBS 121237. S–X. O. cf. 

setacea CBS 121256. A, C, E, G, I, K, M, O, Q, S, U, W. Surface. B, D, F, H, J, L, N, P, R, T, V, X. Reverse. Scale 1 cm. 

Habitat: On overwintered leaves of Alnus viridis (Betulaceae). 

Distribution: Canada (British Columbia), Europe (Switzerland) and 

U.S.A. (WA). 

Specimens examined: Canada, British Columbia, Hope, on overwintered leaves of Alnus 

tenuifolia, 13 May 2006, coll. M.V. Sogonov (BPI 877672) GenBank EU254986; Manning 

Provincial Park, Engineers Trail, on overwintered leaves of Alnus viridis, 13 May 2006 (BPI 

877673) GenBank EU254987. U.S.A., Washington, King Co., Mount Baker-Snoqualmie 

National Forest, Snoqualmie Ranger District, near exit 42 on the highway US 90, road to 

mines, on overwintered but still hanging leaves of Alnus viridis, 16 May 2006 (BPI 877674) 


Notes: Ophiognomonia trientensis can be distinquished from the 

other species of Gnomoniaceae on Alnus. Gnomonia alnea lacks an 

elongated neck unlike O. alni-viridis, O. ischnostyla and O. trientensis. 

Ophiognomonia trientense has an ascospore l:w less than 3 while O. alniviridis 

and O. ischnostyla both have an ascospore l:w greater than 3. 

For a detailed description, see Monod (1983). 


65


Fig. 43. Morphology on natural substrates, perithecia. A–C. Plagiostoma euphorbiae, lectotype Fungi Rhenani 863, BPI bound. D–I. P. aesculi. D, E, I. Epitype BPI 748430. 

F–H. BPI 840942. J, K. P. barriae, holotype BPI 877717B. A, D, E, J. Intact air-dry perithecia on stems, twigs, leaves and petioles. B. Air-dry perithecium on fragment of bark, 

bottom view. C, H, I, K. Extracted and rehydrated perithecia. F. Air-dry perithecium on fragment of bark, side view. G. Air-dry perithecium on fragment of outer layers of stem, 

bottom view. Scale 200 μm. 

PLAGIOSTOMA Fuckel, Jb. Nassau Ver. Naturk. 23–24: 118. 

1870. 

Lectotype designated by Höhnel (1917): Plagiostoma euphorbiae 

Fuckel 

= Cryptodiaporthe Petr., Ann. Mycol. 19: 118. 1921. Lectotype designated 

by Clements & Shear (1931): Cryptodiaporthe aesculi (Fuckel) Petr. now 

recognised as Plagiostoma aesculi (Fuckel) Sogonov, comb. nov. 

= Rostrocoronophora Munk, Dansk Bot. Arkiv 15: 98. 1953. Type: R. geranii 

Munk, now recognised as Plagiostoma geranii (Hollos) Sogonov, comb. nov. 

Perithecia solitary, on fallen leaves, epiphyllous or on petioles, on 

dead but still attached pedicels of trees and shrubs, or on dead parts 

of herbaceous plants, in groups of 5–15 perithecia with or without a 

rudimentary stroma on twigs of trees and shrubs. Perithecia black, 

remaining immersed in substrate, oblate to globose when moist, 

convex, sometimes with irregular dents when dry, round in top 

view, with one neck. Necks central to marginal, never truly lateral, 

mostly length 0.5–2 times perithecial diam but varying from almost 

lacking to 3–4 perithecial diam long. Asci fusiform, with an apical 

ring, with eight spores arranged irregularly multiseriate or obliquely 

uniseriate. Ascospores mostly two-celled, rarely one-celled, 

oval to fusiform, l:w 2.5–6; ends mostly rounded, rarely pointed; 

appendages mostly absent or less commonly present, subulate, 

navicular or whip-shaped, to 30 µm long. 

Cultures: Colonies fast growing, often reaching edges of 90 mm Petri 

plates after 2 wk at 23 °C l/d or at least 60–70 mm diam. Colonies 

floccose or lanose all over surface or in lobes or concentric rings 

intermingled with glabrous or velvety areas. Colonies whitish, grey, 

orange-grey, brownish orange, dark brown, olive. Some species 

produce fertile perithecia in culture after 5–6 mo at 2/10 Cl/d. 

Conidiomata often produced after 2–4 wk at 23 °C l/d. 

Hosts: In diverse taxonomic groups (Aceraceae, Ericaceae, 

Euphorbiaceae, Fagaceae, Geraniaceae, Hippocastanaceae, 

Oleaceae, Platanaceae, Polygonaceae, Salicaceae). Most species 

are specific at the level of plant host species or genus, however, a 

few species occur on a wide diversity of plants. 

66


Fig. 44. Morphology on natural substrates, asci and ascospores. A. Plagiostoma euphorbiae, lectotype Fungi Rhenani 863, BPI bound. B, C. P. aesculi, epitype BPI 748430. 

D–F. P. barriae, holotype BPI 877717B. Scale 10 μm. 

Type species of Plagiostoma and synonymous 

genus, Cryptodiaporthe 

Plagiostoma euphorbiae (Fuckel) Fuckel, Jb. Nassau Ver. Naturk. 

23–24: 118. 1870. Figs 43A–C; 44A; 45A–F. 

≡ Sphaeria euphorbiae Fuckel, Enumeratio fungorum Nassoviae: 69. 

1860. 

≡ Gnomonia euphorbiae (Fuckel) Sacc., Michelia 2: 312. 1881. 

≡ Gnomoniella euphorbiae (Fuckel) Sacc., Syll. Fung. 1: 418. 1882. 

= Gnomonia tithymalina Sacc. & Briard, Revue mycol. 7: 209. 1885 fide Monod 

1983. 

Perithecia solitary, without stroma, randomly scattered on dead 

stems, black, suboblate to oblate-spheroidal when moist, 230–350 

µm high × 290–430 µm diam, convex when dry. Necks central or 

eccentric, short, only slightly projecting from plant tissue, 70–95 µm 

long, 80–90 µm diam. Asci oval, (33–)37.5–41(–52.5) × 10.5–12.5(– 

13.5) µm (mean = 41 × 12, SD 7.5, 1.5, n=5), apical ring 2.5–3 µm 

diam, with eight ascospores arranged obliquely uniseriate, obliquely 

biseriate ot irregularly multiseriate. Ascospores ellipsoidal, straight 

or inequilateral, (12–)13–13.5(–15.5) × (3–)3.5(–4) µm (mean 

= 13.5 × 3.5, SD 0.5, 0.2, n=33), l:w (3.1–)3.7–4(–4.5) (mean = 

3.9, SD 0.3), two-celled, not constricted at septum; septum located 

at (42–)45–49(–56) % (mean = 47, SD 3) of ascospore length; 

cells with parallel walls, rounded at ends, each cell with two large 



°C, flat, glabrous to velvety, dark brown to greyish brown; margins 

submerged, orange-grey; margin irregular; reverse dark brown to 

brownish orange. Colonies on MEA attaining 90 mm after 40 d at 23 

°C, flat, superficial and partly submerged, with no aerial mycelium, 

thin, consisting of dendroid branches, shades of orange-grey, 

brownish orange, brownish grey; margin irregular; reverse of same 

colours as surface. Colonies on MYA attaining 90 mm after 40 d 

at 23 °C, flat, short felty orange-grey in central part, glabrous to 

velvety, dark brown; margin submerged, greyish orange, irregular; 

reverse dark brown to brownish orange. 

Habitat: On dead stems of Euphorbia palustris L. and E. pannonica 

Host (Euphorbiaceae). 


67


Fig. 45. Culture morphology, colony habit. A–F. Plagiostoma euphorbiae CBS 817.79. G–R. P. barriae CBS 121249. S–X. P. devexum BPI 843489. A–F, M–X. 40 d, 23 ºC. G–L. 

14 d, 23 ºC. A, C, E, G, I, K, M, O, Q. Surface. B, D, F, H, J, L, N, P, R. Reverse. A, B, G, H, M, N. PDA. C, D, I, J, O, P. MEA. E, F, K, L, Q, R. MYA. Scale 1 cm. 

68


Distribution: Europe (Germany, Hungary, The Netherlands, Russia, 

Switzerland). 

Lectotype designated here: Germany, Freienweinheim, 1860 or 

before, K.W.G.L. Fuckel, Fungi Rhenani 863 (BPI bound). 

Additional cultures examined: The Netherlands, Baarn, 12 May 

1978, W. Gams, H.A. van der Aa 6449 (CBS 340.78); Switzerland, 

Vaud, lake shore between Yverdon and Yvonand, 14 Jun. 1978, M. 

Monod 466 (CBS 817.79). 

Notes: Plagiostoma euphorbiae is distinguished from the other 

species of Gnomoniaceae on Euphorbia by the short neck, less 

than 100 μm long, on each perithecium. 

Plagiostoma aesculi (Fuckel) Sogonov, comb. nov. MycoBank 

MB 512193. Figs 43D–I; 44B,C. 

Basionym: Cryptospora aesculi Fuckel, Jb. Nassau Ver. Naturk. 23–24: 193. 1870. 

≡ Cryptosporella aesculi (Fuckel) Sacc., Michelia 1: 30. 1877. 

≡ Diaporthe aesculi (Fuckel) Höhn., Ann. Mycol. 16: 116. 1918. 

≡ Cryptodiaporthe aesculi (Fuckel) Petr., Ann. Mycol. 19: 119. 1921. 

Perithecia in groups of 3–10, with loose stroma, on fresh dead 

twigs. Perithecia black, oblate spheroidal when moist, convex, 

usually with irregular dents on top when dry, 300–450 µm high × 

380–600 µm diam. Necks converged with others in group, eccentric 

to marginal, slightly curved, 420–700 µm long, 100–150 µm wide 

at base, 60–150 µm wide at apex. Asci fusiform, (45.5–)48.5–67(– 

78.5) × (10–)12.5–16(–21.5) µm (mean = 58.5 × 14.5, SD 11, 3, 

n=16), apical ring absent, with eight ascospores arranged obliquely 

biseriate to irregularly multiseriate. Ascospores variable in size and 

shape, ellipsoidal to fusiform, (13–)17.5–20(–23.5) × (3.5–)4–5(– 

6.5) µm (mean = 18.5 × 4.5, SD 2, 0.7, n=109), l:w (2.6–)3.5–4.4(– 

5.7) (mean = 4, SD 0.7), two-celled, constricted or not constricted 

at septum, ends rounded to tapering, distal cell often slightly wider 

than basal, septum located at (37–)46–50(–57) % (mean = 48, 

SD 4, n=98) of ascospore length; appendages usually absent, if 

present, subulate, length to 5 µm. 

Cultures: Not observed. 

Habitat: On overwintered twigs of Aesculus hippocastanum L. 

(Sapindaceae). 

Distribution: Europe (Austria, Czech Republic, Germany, United 

Kingdom). 

Lectotype designated here: Germany, Reichartshausen, on dry 

twigs of Aesculus hippocastanum, winter 1894 or before, K.W.G.L. 

Fuckel, Fungi Rhenani 2003 (BPI 601244). 

Epitype designated here: Austria, Vienna, 19 th district, 

Krapfenwaldgasse, Grinzing, Aesculus hippocastanum, 11 Nov. 

2000, W. Jaklitsch 1695 (BPI 748430, ex-type epiculture CBS 

109765). 

Additional specimens examined: Austria, Trieblach, St. Margareten im Rosental, 

Kaertnen, on dead twigs of A. hippocastanum, 14 Apr. 2001, W. Jaklitsch 1732 (BPI 

840942, culture CBS121905) GenBank EU254994; Czech Republic, Moravia, Hranice 

na Moravě, Aesculus sp., March 1913, F. Petrak (Flora Moravica, Missouri Bot. Gard. 

Herb. 43417 (BPI 617579); Moravia, Hranice na Moravě, Teplice, A. hippocastanum, 

May 1914, F. Petrak (BPI 617580); Germany, Saxony, near Köningstein, A. 

hippocastanum, 04 May 1907, W. Krieger Fungi Saxonici 2022 (BPI bound). 

New species of Plagiostoma 

Plagiostoma barriae Sogonov, sp. nov. MycoBank MB 512194. 

Figs 43J,K; 44D–F; 45G–R. 

Perithecia 140–170 µm alta × 180–240 µm diam, in sicco convexae. 

Rostrum 800–130 µm longum, basi 45–52 µm diam, apice 30–38 

µm diam. Ascosporae ellipsoideae vel ovales, rectae, (11.5–)14– 

15.5(–17.5) × (2.5–)3.5–4(–4.5) µm, L:l (3–)3.6–4.1(–5.1). Ad aliis 

Plagiostomae speciebus morphologiae characteribus combinatis 

differt. Holotypus: BPI 877717B. 

Etymology: Names for the Canadian mycologist Margaret E. Barr 

Bigelow in recognition of her contribution to the taxonomy of the 

Diaporthales. 

Perithecia solitary, without stroma, randomly scattered on overwintered 

petioles, black, suboblate when moist, 140–170 µm high × 180–240 

µm diam, convex when dry. Necks central, straight, 80–130 µm 

long, 45–52 µm wide at base, 30–38 µm wide at apex. Asci fusiform, 

(47.5–)48.5–53(–56.5) × (8.5–)9.5–10.5(–11) µm (mean = 51 × 10, 

SD 4, 1.2, n=4), apical ring 2.5–4 µm diam, with eight ascospores 

arranged biserate to irregularly multiseriate. Ascospores ellipsoidal 

to oval, straight to oval, (11.5–)14–15.5(–17.5) × (2.5–)3.5–4(–4.5) 

µm (mean = 15 × 4, SD 1.5, 0.5, n=58), l:w (3–)3.5–4(–5) (mean 

= 4, SD 0.5), two-celled, constricted at septum; septum located at 

(44–)47–49(–55) % (mean = 48, SD 3, n=20) of ascospore length; 

cells slightly tapering, at ends blunt, rounded, each cell with 2 large 

and sometimes 1–2 small guttules where the largest guttule close to 

septum; appendages absent or indistinct. 

Cultures: Colonies on PDA and MYA attaining 90 mm after 40 d 

at 23 °C, flat, woolly to floccose, with indistinct concentric zones, 

with areas of tint of orange in central part, with scarce-grey soft 

sclerotium-like bodies; reverse pale brown in centre to greyish 

orange at margin. Colonies on MEA attaining 90 mm after 40 d at 

23 °C, flat, thin, semitransparent, shortly felty, colourless or whitish 

with tint of brown in central part, with scattered flocks of white 

aerial mycelium, with dark brown matt or covered with pale grey 

felty mycelium soft sclerotium-like bodies; margin diffuse; reverse 

brownish orange to brown-grey. Neither perithecia nor conidiomata 

observed in cultures at 2/10 °C after 8 mo. 

Habitat: On overwintered petioles of Acer macrophyllum Pursh 

(Aceraceae). 

Holotype: U.S.A., Washington, Pierce Co., Gig Harbor, Narrows 

Park, 16 May 2006, M.V. Sogonov MS0367a (BPI 877717B, exholotype 

culture CBS 121249). 

Additional GenBank nucleotide sequence: U.S.A., Washington, 

Klickitat, young non-mildewed leaf of Acer macrophyllum, date 

unknown, C. Nischwitz, G. Newcombe, nrDNA ITS1–5.8S–ITS2 

(AY961407). 

Notes: Plagiostoma barriae having central necks on the perithecia 

differs from other species of Plagiostoma on Acer, specifically P. 

inclinatum (Desm.) Barr, P. petiolophilum, and P. pseudobavarica 

M. Monod, all species that have lateral necks. 


69


Fig. 46. Morphology on natural substrates, perithecia. A–E. Plagiostoma devexum. A, B, E. Plantes Cryptogames de France 367, BPI bound. C, D. BPI 843489. F, G. P. 

euphorbiae-verrucosae, BPI 877685. H. P. euphorbiaceum, BPI 871053. I. P. fraxini, BPI 877686. J. P. rhododendri, BPI 877701. A, C, D, F, H, J. Intact air-dry perithecia on 

stems, twigs, petioles and pedicels. B. Extracted air-dry perithecia. E, G, I. Extracted and rehydrated perithecia. Scale 200 μm. 

Additional species accepted in Plagiostoma 

Plagiostoma amygdalinae (Fuckel) Sogonov, comb. nov. 


Basionym: Gnomonia amygdalinae Fuckel, Jb. Nassau Ver. Naturk. 23–24: 121. 

1870. 

≡ Gnomoniella amygdalinae (Fuckel) Sacc., Syll. Fung. 1: 418. 1882. 

= Gnomoniella amygdalinae (Fuckel) Sacc. f. euphorbiae-stepposae Sandu- 

Ville, Studii Cerc. Biol., Bot. 18: 18. 1966 fide Monod 1983. 

Habitat: On overwintered leaves of Euphorbia amygdaloides L. and 

E. stepposa Zoz (Euphorbiaceae). 

Distribution: Europe (Bulgaria, France, Germany, Romania, 

Switzerland). 

Notes: Among the species of Plagiostoma on Euphorbia, Plagiostoma 

amygdalinae as well as P. euphorbiaceum and P. euphorbiae-verrucosae 

have perithecial necks longer than 100 µm and thus are distinct from 

P. euphorbiae having a shorter neck. Plagiostoma amygdalinae and 

P. euphorbiaceum have one-septate ascospores while those of P. 

euphorbiae-verrucosae are non-septate. Plagiostoma amygdalinae 

has ascospores that are 13–15.5 × 2.3–3 µm that are narrower than 

those of P. euphorbiae-verrucosae. See Monod (1983) for a detailed 

description of P. amygdalinae as Gnomonia amygdalinae. 

Plagiostoma devexum (Desm.) Fuckel, Jb. Nassau Ver. Naturk. 

23–24: 119. 1870. Figs 45S–X; 46A–E; 47A,B. 

≡ Sphaeria devexa Desm., Cryptog. de France, Edit. II, Sér. II, No 367. 

1856. 

≡ Gnomonia devexa (Desm.) Auersw. in Gonn. & Rabenh., Mycol. Europ. 

5/6: 23. 1869. 

≡ Gnomoniella devexa (Desm.) Sacc., Syll. Fung. 1: 417. 1881. 

≡ Gnomonopsis devexa (Desm.) Moesz & Smarods, Bot. Közl. 38: 68. 

1941. 

70


Fig. 47. Morphology on natural substrates, asci and ascospores. A, B. Plagiostoma devexum, BPI 843489. C. P. euphorbiae-verrucosae, BPI 877685. D. P. euphorbiaceum, 

BPI 871053. E. P. fraxini, BPI 877687. F. P. rhododendri, BPI 877701. Scale 10 μm. 

= Sphaeria euphorbiae f. polygoni Fuckel, Fungi Rhenani 864. 1864 fide 

Monod 1983. 

= Sphaeria excentrica Cooke & Peck, Ann. Rep. New York State Museum 25: 

105. 1873 fide Monod 1983. 

≡ Gnomoniella excentrica (Cooke & Peck) Sacc., Syll. Fung. 1: 418. 

1881. 

= Diaporthe sechalinensis Sacc., Atti del Congr. bot. di Palermo 1902: 52. 1902 

fide Monod 1983. 

= Ceriosporella polygoni A.L. Sm. & Ramsb., Trans. Br. mycol. Soc. 4: 325. 

1914 fide Monod 1983. 

Habitat: On overwintered stalks and leaves of Persicaria amphibium 

(L.) Delarbre, P. lapathifolia (L.) Gray, P. maculosa Gray., Polygonum 

sp. and Rumex longifolius DC. (Polygonaceae), and Vitis vitifera L. 

(Vitaceae). 

Distribution: Europe (Denmark, France, Germany, Sweden, 

Switzerland, United Kingdom) and U.S.A. (NY). 

Notes: Barr (1978) and Monod (1983) provide detailed descriptions 

of this species. 


71


Plagiostoma euphorbiaceum (Sacc. & Briard) Sogonov, comb. 

nov. MycoBank MB 512196. Figs 46H; 47D; 48A–F. 

Basionym: Gnomonia euphorbiacea Sacc. & Briard, Revue Mycol. 7: 208. 1885. 

Habitat: On dead branches of Euphorbia amygdaloides L. and E. 

palustris L. (Euphorbiaceae). 

Distribution: Europe (Germany, Switzerland). 

Specimen examined: Switzerland, Vaud, Arzier, on overwintered stems of Euphorbia 

amygdaloides, 25 May 2005, coll. M.V. Sogonov (BPI 871053) GenBank EU255004. 

Notes: Among the species of Plagiostoma on Euphorbia, P. 

euphorbiaceum as well as P. amydalinae and P. euphorbiaeverrucosae 

have perithecial necks longer than 100 µm and thus 

are distinct from P. euphorbiae having a shorter neck. Plagiostoma 

euphorbiaceum and P. amygdalinae have one-septate ascospores 

while those of P. euphorbiae-verrucosae are non-septate. 

Plagiostoma euphorbiaceum has ascospores that are 14–17.5 × 

3.5–4.5 µm and wider than those of P.amygdalinae. For a detailed 

description, see Monod (1983 as G. euphorbiacea). 

Plagiostoma euphorbiae-verrucosae (M. Monod) Sogonov, 

comb. nov. MycoBank MB 512197. Figs 15F,G; 29D. 

Basionym: Gnomoniella euphorbiae-verrucosae M. Monod, Beih. Sydowia 9: 42. 

1983. 

Habitat: On overwintered stalks of Euphorbia verrucosa L. 

(Euphorbiaceae). 

Distribution: Europe (Switzerland). 

Specimen examined: Switzerland, Les Plans sur Bex, Pont de 

Nant, on overwintered stems of Euphorbia verrucosa, 29 May 

2005, coll. M.V. Sogonov (BPI 877685) GenBank EU255006. 

Notes: Plagiostoma euphorbiae-verrucosae as well as P. 

amygdalinae and P. euphorbiaceum have perithecial necks longer 

than 100 µm and thus are distinct from P. euphorbiae having a 

shorter neck. The ascospores of P. euphorbiae-verrucosae 

are non-septate while those Plagiostoma amygdalinae and P. 

euphorbiaceum are one-septate. For a detailed description of P. 

euphorbiae-verrucosae, see Monod (1983 as G. euphorbiaeverrucosae). 

Plagiostoma fraxini (Redlin & Stack) Sogonov, comb. nov. 

MycoBank MB 512198. Figs 46I; 47E; 48G–L. 

Basionym: Gnomoniella fraxini Redlin & Stack, Mycotaxon 32: 185. 1988. 

Habitat: On living and overwintered leaves of Chionanthus retusus 

Lindl. & Paxton, Fraxinus americana L., and F. pennsylvanica 

Marshall (Oleaceae) 

Distribution: Canada (Manitoba, Saskatchewan) and U.S.A. (CA, 

DE, IA, IL, LA, KY, MI, MD, MS, NC, ND, NY, OK, OR, SD, VA, WI) 

Specimens examined: U.S.A., Maryland, Prince George’s 

Co., Riverdale, Anacostia, on overwintered leaves of Fraxinus 

americana, 12 Jun 2006, coll. M.V. Sogonov (BPI 877687) GenBank 

EU255008; Howard Co. Centennial Park, on overwintered leaves of 

Fraxinus americana, 9 Apr 2005, coll. M.V. Sogonov (BPI 877686) 


Notes: The anamorph of Plagiostoma fraxini is Discula fraxinea 

(Peck) Redlin & Stack under which this species has been reported 

to cause ash anthracnose (Holcomb 1998, Rossman et al. 2004) 

and anthracnose of fringetree (Gregory et al. 2004). For a detailed 

description, see Redlin & Stack (1988 as G. fraxini). 

Plagiostoma geranii (Hollós) Sogonov, comb. nov. MycoBank 

MB 512199. 

Basionym: Gnomonia geranii Hollós, Annls Mus. nat. hung. 7: 52. 1909. 

= Rostrocoronophora geranii Munk, Dansk Bot. Arkiv 15(2): 98. 1953. 

Habitat: On overwintered stalks of Geranium sanguineum L., G. 

sylvaticum L. (Geraniaceae). 

Distribution: Europe (Bulgaria, Denmark, Germany, Hungary, 

Sweden, Switzerland) 

Specimen examined: Bulgaria, Sredna Gory Mt (western), Lozenska Planina, along 

track to Vlakovete, on overwintered petioles and stems of Geranium sanguineum, 2 

May 2005, coll. D. Stoykov (BPI 877688) GenBank EU255010. 

Note: For a detailed description see Monod (1983) and Műller & Arx 

(1962) as G. geranii. 

Plagiostoma petiolophilum (Peck) Sogonov, comb. nov. 


Basionym: Sphaeria petiolophila Peck, Ann. Rep. New York State Museum 35: 144. 

1884. 

≡ Gnomonia petiolophila (Peck) Berl. & Voglino, Syll. Fung. Addit. 1–4: 

90. 1886. 

≡ Cryptodiaporthe petiolophila (Peck) Barr, Mycol. Mem. 7: 136. 1978. 

Habitat: On overwintered leaves, petioles, and twigs of Acer 

negundo, A. pensylvanicum, A. rubrum, A. saccharum, A. spicatum, 

and Acer sp. (Aceraceae) 

Distribution: Canada (Ontario) and U.S.A. (GA, MD, MI, NH, NY, 

TN). 

Specimens examined: U.S.A., Maryland, Prince George’s Co., Paint Branch Park, 

on overwintered petioles of Acer rubrum, 17 Mar 2006, coll. M.V. Sogonov (BPI 

877699) GenBank EU255040; Tennessee, Great Smoky Mountain National Park, 

on overwintered petioles of Acer sp., 10 May 2006, coll. L. Vasilyeva (BPI 878448, 


Notes: Plagiostoma petiolophilum has lateral necks on the 

perithecia unlike P. barriae with one central neck. Barr (1978 as C. 

petiolophila) provides a detailed description of this species. 

Plagiostoma rhododendri (Auersw.) Sogonov, comb. nov. 

MycoBank MB 512201. Figs 46J; 47F; 48M–R. 

Basionym: Gnomonia rhododendri Auersw. in Gonn. & Rabenh., Mycol. Europ. 5/6: 

26. 1869. 

≡ Apiognomonia rhododendri (Auersw.) Remler, Bibiotheca Mycologica 

68: 74. 1979. 

Habitat: On overwintered branches, flowers, and leaves of 

Rhododendron ferrugineum L., R. hirsutum L. (Ericaceae) 

Distribution: Europe (Austria, Germany, Italy, Switzerland) 

Specimens examined: Switzerland, Vaud, Pont de Nant, Botanical Garden, on dead 

inflorescenes of Rhododendron hirsutum, 29 May 2005, M. Monod (BPI 877701) 


72


Fig. 48. Culture morphology. A–F. P. euphorbiaceum CBS 121241. G–L. P. fraxini CBS 121258. M–R. P. rhododendri CBS 847.79. A, C, E, G, I, K, M, O, Q. Surface. B, D, F, 

H, J, L, N, P, R. Reverse. A, B, G, H, M, N. PDA. C, D, I, J, O, P. MEA. E, F, K, L, Q, R. MYA. Scale 1 cm. 

Note: Remler (1979 as A. rhododendri) and Monod (1983 as G. 

rhododendri) provide detailed descriptions of this species. 

Note: Wehmeyer (1933) provides a detailed description of this 

species as Cryptodiaporthe robergeana. 

Plagiostoma robergeanum (Desm.) Sogonov, comb. nov. 


Basionym: Sphaeria robergeana Desm., Ann. sci. nat., Ser. 3, 16: 306. 1851. 

≡ Diaporthe robergeana (Desm.) Niessl in Rabenh., Fungi Europ. 2222. 

1882. 

≡ Cryptodiaporthe robergeana (Desm.) Wehm., The Genus Diaporthe: 

200. 1933. 

Habitat: On overwintered, still attached branches of Staphylea 

colchica Steven and S. pinnata L. (Staphyleaceae) 

Distribution: Europe (Austria, Czech Republic, France, Germany, 

Poland, Russia, Switerland, United Kingdom) 

Plagiostoma salicellum (Fr.) Sogonov, comb. nov. MycoBank 

MB 512203. 

Basionym: Sphaeria salicella Fr., Syst. Mycol. 2: 377. 1823. 

[≡ Cryptodiaporthe salicella (Fr.) Wehm., The Genus Diaporthe, p. 193. 

1933 non Petrak 1921]. 

= Diaporthe spina Fuckel, Jb. Nassau Ver. Naturk. 23–24: 210. 1870 fide 

Wehmeyer 1933. 

≡ Gnomonia spina (Fuckel) Feltg., Vorst. Pilz. Lux., Nachtr. I: 214. 1899. 

= Valsa populina Fuckel, Jb. Nassau. Ver. Naturk. 25–26: 314. 1871 fide 


≡ Cryptodiaporthe populina (Fuckel) Petr., Ann. Mycol. 19: 119. 1921 fide 


= Cryptodiaporthe apiculata (Wallr.) Petr., Ann. Mycol. 19: 177. 1921 fide 


Habitat: On overwintered, still attached branches of Populus nigra 


73


L., P. tremula L. Salix appendiculata Vill., S. aurita L., S. caprea L., 

S. fragilis L., S. triandra L., and S. vitellina L., (Salicaceae). 

Distribution: Canada (Ontario, Quebec), Europe (Austria, Belgium, 

Bulgaria, Czech Republic, Germany, Poland, Sweden, Switzerland, 

United Kingdom) and U.S.A. (MA, NY). 

Notes: Wehmeyer (1933) suggests the relationship of this species 

to the Gnomoniaceae and provides a detailed description as 

Cryptodiaporthe salicella. 

Genera not included in this study or excluded from 

the Gnomoniaceae 

The genus Anisogramma Theiss. & Syd. includes two species 

pathogenic on woody plants, A. virgultorum (Fr. : Fr.) Theiss. & 

Syd., the type of the genus and cause of a disease of Betula in 

Europe (Froidevaux & Műller 1976), and A. anomala (Peck) E. 

Müll., cause of eastern filbert blight, a serious disease of Corylus 

avellana in North America (Gottwald & Cameron 1979, Johnson et 

al. 1996). Recent research on the phylogeny of these species has 

demonstrated that Anisogramma is sister to the Gnomoniaceae 

(DeSilva et al. 2008). 

The genus Apioplagiostoma M.E. Barr was established for 

species similar to Plagiostoma in having perithecia bearing an 

eccentric or lateral neck but with unequally septate ascospores 

(Barr 1978). Included in Apioplagiostoma is the type species, A. 

populi (E.K. Cash & Waterman) M.E. Barr, a species not included 

in this study. One species, Apioplagiostoma carpinicola, is herein 

transferred to Gnomonia carpinicola. One isolate of A. aceriferum 

was sequenced and determined to belong in Pleuroceras (Fig. 1). 

One additional species, A. hilberovae Schmid-Heckel (Schmid- 

Heckel 1988) is included in this genus. 

The genus Bagcheea E. Müll. & Menon includes only the type 

species, B. albomaculans (Fukui) Hino & Katumoto (= B. castaneae 

E. Müll. & Menon). It occurs on living leaves of Castaneopsis in 

India and Japan. Kobayashi (1970) collected this species but was 

unable to obtain it in culture. 

Clypeoporthella Petr. is based on C. brencklei Petr. occurring 

on Solidago in North America. However, based on the associated 

Phomopsis asexual state, it appears likely to be a synonym of 

Diaporthe. A specimen identified as this taxon (BPI 843482), grown 

in culture, and sequenced was determined to be a Diaporthe, thus 

Clypeoporthella is considered a synonym of Diaporthe. 

Dicarpella P. Syd. & Syd. based on D. bina (Harkn.) P. Syd. 

& Syd. (≡ Physalospora bina Harkn.) is known only from the type 

collection on Quercus agrifolia Nee in California. The anamorphic 

state of Dicarpella, Tubakia B. Sutton is relatively common on oak 

leaves. 

Diplacella Syd., D. paullinae (Frag. & Cif.) Syd. is parasitic on 

leaves of Paullinia and other members of the Sapindaceae in Central 

and South America. Arx & Műller (1954) provide a description of 

this species. 

The genus Gaeumannomyces Arx & D.L. Olivier was included 

in the Gnomoniaceae by a number of authors such as Barr (1978) 

and Monod (1983), and considered as a synonym of Linocarpon by 

Kobayashi (1970). However, molecular data obtained from the type 

species and serious pathogen G. graminis (Sacc.) Arx & D.L. Olivier 

confirmed that this genus is not a member of the Diaporthales but 

that it belongs to the Magnaporthaceae (Castlebury et al. 2002, 

Zhang et al. 2008). 

The genus Gnomoniella Sacc., based on the type species, G. 

tubaeformis (Fr. : Fr. ) Sacc., was not included in this study for 

lack of living material. The type species occurs on overwintered 

leaves and petioles of Alnus spp. in Europe and North America. 

Several fresh collections were obtained of the non-type species G. 

alnobetulae Volkart, a species having four-spored asci but otherwise 

morphologically close to G. tubaeformis. Attempts to prepare a 

pure culture were not successful. Based on the ITS sequence of G. 

alnobetulae obtained by direct amplication from asci, this species 

may represent a distinct genus in the Gnomoniaceae. 

Hypospilina bifrons (DC. : Fr.) Traverso, type of the genus 

Hypospilina (Sacc.) Traverso, occurs on dead leaves of Quercus 

spp. in Europe and was not collected for this study. 

Lambro Racib. based on L. insignis Racib. produces necrotic 

spots in leaves of Sterculia subpeltata Blume in Indonesia (Műller 

& Arx 1962). Monod (1983) examined the type specimen and 

suggested that this species is related to Stegophora, thus it may 

belong in the Sydowiellaceae. 

The genus Linospora Fuckel based on L. capreae (DC. : Fr.) 

Fuckel) was distinguished by ascomata covered with a rudimentary 

stroma and elongate ascospores. Linospora capreae groups with 

Pleuroceras. Because Linospora was published after Pleuroceras, 

Linospora is considered a synonym of Pleuroceras. Fifty-seven 

names have been placed in Linospora but most of them have been 

little studied and are infrequently encountered. 

Mamiania Ces. & De Not. based on M. fimbriata (Fr.) Ces. & De 

Not. occurs on living leaves of Carpinus spp. in Europe, Asia, and 

North America. This species has a conspicuous stroma surrounding 

the perithecia and ascospores with a distinct, submedian septum. 

The genus Mamianiella Höhn., based on the species, M. coryli 

(Batsch : Fr.) Höhn., occurs on living leaves of Corylus. We have 

not been able to grow this species in culture. 

Mazzantia Mont. (syn. Clypeocarpus Kirschst., Paramazzantia 

Petr.) is based on M. galii (Fr.) Mont. Paramazzantia is based on 

Laestadia biennis Dearness on overwintered leaves of Solidago in 

North America. Clypeocarpus is based on C. alpinus Kirschst. occurs 

on Veratrum album and is considered a synonym of M. napelli by 

Von Arx & Műller (1954). Mazzantia napelli was sequenced and 

determined to belong in the Diaporthaceae by Castlebury et al. 

(2002). 

Phylloporthe Syd., based on P. vernoniae Syd., is parasitic on 

living leaves of Vernonia triflosculosa H.B.K. in Costa Rica. It is 

only known from the type specimen. A second species, P. orbiculata 

(Syd.) E. Műll. is now placed in Uleoporthe (Cannon 2001). 

Plagiosphaera Petr. based on P. immersa (Trail) Petr. (≡ 

Ophiobolus immersus Trail) occurs on overwintered stalks of 

Campanula and Urtica in Europe and was not collected during the 

course of this study. 

The genus Pleuroceras Riess includes Gnomonia-like fungi 

having ascomata with eccentric, lateral necks and elongated 

ascospores and was placed in the Valsaceae by Barr (1978). 

The type species is P. cryptoderis (Lév.) Höhn., which occurs on 

overwintered leaves of Populus alba. Many of the 23 species 

currently recognised in Pleuroceras were transferred from Gnomonia 

by Barr (1978) and Monod (1983). Most of the species included in 

Pleuroceras are found on overwintered leaves of hardwood trees in 

temperate regions. 

Although several species of Pleuroceras are included in the 

multigene phylogeny in this paper (Fig. 1), this genus is not treated 

74


in detail, because its type species, P. cryptoderis, was not available 

for sequencing. Several species of Pleuroceras were included in this 

study and form a well-supported monophyletic genus. One species 

of Pleuroceras, P. sassafras, is transferred to Ophiognomonia. 

Sphaerognomonia Potebnia based on S. carpinea (Fr.) 

Potebnia includes the synonym, Apiosporopsis (Traverso) Mariani 

based on the same type species. Sequences of this species (CBS 

617.72 and CBS 738.6) place this genus in the Diaporthales but 

outside of the Gnomoniaceae and Melanconidaceae (Castlebury, 

unpublished). 

Stegophora P. Syd. & Syd. based on S. ulmea (Schw. : Fr.) 

P. Syd. & Syd. is parasitic on living leaves of Ulmus spp. in North 

America. A second species, S. oharana (Y. Nisik. & H. Matsumoto) 

Petr. occurs in Japan. Molecular data suggests that S. ulmea 

belongs to the Sydowiellaceae (Castlebury, unpub. data) as 

circumscribed by Rossman et al. (2007). 

Uleoporthe Petr., typified by U. orbiculata (Syd.) Petr., was 

redescribed by Cannon (2001) based on a fresh specimen from 

Guyana. His redesciption, the occurrence of this species as a leaf 

parasite, and the presence of a distinct well-developed stroma 

suggest an affiliation with the Sydowiellaceae. 

ACKNOWLEDGEMENTS 

This work was funded by NSF PEET grant 0328634 for research on 

the systematics of the Gnomoniaceae, Diaporthales for which we 

are most grateful. The following individuals contributed significantly 

to this project: Michel Monod, Lausanne, Switzerland, was generous 

in sharing his knowledge and providing useful advice and comments 

as well as arranging the logistics for collecting in Switzerland. 

Jean-Louis Moret, Lausanne, Switzerland, allowed the first author 

access to specimens housed in the Botanical Garden Museum of 

University of Lausanne. Margaret Barr, recently deceased, Sidney, 

Canada, and Larissa Vassiljeva, Vladivostok, Russia, both experts 

in the Gnomoniaceae, willingly assisted in identifications and helped 

with collecting and providing fresh specimens. Walter Jaklitsch, 

Vienna, Austria, and Dimitar Stoykov, Bulgaria, sent numerous 

accurately identified fresh collections from which cultures were 

obtained. Dmitriy Maykov, Kholm, Novgorod oblast, Russia, also 

provided fresh collections and assistance with logistics while in 

Russia. Gary Samuels, Drew Minnis, John Wiersema, and Joseph 

Kirkbride, USDA-ARS, Beltsville, Maryland, provided advice on 

nomenclatural issues. Christian Feuillet kindly provided the Latin 

diagnoses. David Farr assisted with use of the microscope, digital 

photography, and computer aspects of this research. A number of 

technicians at the Systematic Mycology & Microbiology Laboratory 

were essential to this work, specifically Brandon Dyson, Franklin 

Hendrick, Aimee Hyten, Cindy Park, Suganda Patibanda, and 

Tunesha Phipps. Erin McCray, Collections Manager of the U.S. 

National Fungus Collections (BPI), was invaluable in locating, 

obtaining and accessioning the many specimens from other 

herbaria as well as those at BPI. The curators and directors of the 

numerous herbaria listed in the Material and Methods are thanked 

for providing reference collections essential for this research. The 

Centraalbureau voor Schimmelcultures especially Pedro Crous, 

Gerard Verkley and Trix Merkx were extremely cooperative in 

providing and accessioning the numerous cultures derived from 

specimens used in this study. Finally, the authors would like to 

acknowledge the thorough scrutiny given to this manuscript by two 

anonymous reviewers who provided many useful suggestions for 

improvements. 


75


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77


index 

A 

Acer 7, 14, 16-17, 38, 69, 72 

— saccharum 14, 41 

Aceraceae 10, 36, 38, 41, 66, 69, 72 

acerina, Gnomonia 38 

aesculi 2, 12, 14, 66-7 

Aesculus hippocastanum 2, 6, 69 

alnea 11, 19, 21 

alni-viridis 54-5, 58, 60, 65 

Alnus 3, 14, 17, 21, 55, 59, 65 

— glutinosa 5, 21, 59-60 

— rubra 6 

— viridis 6, 55, 64-5 

Ambarignomonia 1, 8-10, 12, 35 

— petiolorum 1, 5, 14, 33, 35-6 

Amphiporthe 8-9 

amygdalinae 1, 70, 72 

Anacardiaceae 10 

Apiognomonia 2, 8-10, 12-13, 38 

— acerina 16, 37-8 

— borealis 5, 15, 38 

— errabunda 1, 5, 16, 38, 40 

— hystrix 1, 5, 12, 14, 17, 40-1, 63 

— ostryae 21-2, 25-6 

— veneta 5, 12, 16, 36, 38 

Apioplagiostoma 2, 5, 74 

— aceriferum 38 

appendages 10-13, 17-18, 21-2, 24, 26, 30-2, 36, 41, 43-4, 47- 

8, 51, 53, 66-7, 69 

B 

balsamiferae 11, 49-51 

barriae 66-7, 72 

Betula 17, 58-9, 63, 74 

— nana 6, 58, 63 

— papyrifera 5, 55, 58 

— pendula 58 

Betulaceae 10-12, 14, 17-18, 21-2, 25-6, 30-4, 48, 55, 58-60, 

63-4 

C 

californica 32, 34 

Carpinus 14, 17, 30, 32, 34, 59 

— betulus 14, 32, 34, 60 

— caroliniana 5, 14 

Carya sp 53, 63 

— tomentosa 6, 53, 63 

chamaemori 11-12, 39-40, 44 

clavulata 12, 39-40, 42, 44, 47, 51 

Comarum palustre 5, 47 

Corylus 12, 14, 17, 30, 32, 35, 74 

— avellana 5-6, 14-5, 18, 21-2, 34, 60, 74 

— californica 5, 15, 25, 30-1 

Cryptoderis 74-5 

Cryptodiaporthe 2, 12, 67 

— hystrix 12, 41 

— robergeana 73 

v salicella 73-4 

Cryptosporella 2-3, 8-9, 12, 13-14 

D 

Diaporthalean fungi 1 

Diaporthales 1-5, 7, 9, 11-13, 15, 17, 19, 21, 23, 25, 27, 29, 31, 

33, 69, 73-5 

Diaporthe 48, 73-4 

Dicarpella 74 

Discula 2 

Disculina 13 

Ditopella 2-3, 8 

Ditopellopsis racemula 11, 48 

E 

epiphyllous 18, 36, 41, 66 

Ericaceae 10-11, 36, 41, 66, 72 

errabunda 2, 10, 38, 40 

Euphorbia 2, 15-16, 69-70, 72 

— amygdaloides 6, 70, 72 

— verrucosa 72 

Euphorbiaceae 10, 36, 66-7, 70, 72 

euphorbiaceum 70-2 

euphorbiae 2, 11-12, 70, 72 

euphorbiae-verrucosae 13, 70-2 

F 

Fagaceae 10-12, 16-17, 36, 38, 41, 44, 47-8, 64, 66 

Fagales 11-12, 48 

falcata 44 

femoroid 10, 13-14 

Fragaria sp 6, 47 

fraxini 13, 15, 70-2 

Fraxinus americana 72 

fructicola 1, 16, 39-40, 47 

G 

gei-montani 1, 54-5 

Geraniaceae 10, 36, 38, 66, 72 

Geranium sanguineum 72 

Gnomonia 1-3, 8-9, 10-13, 18, 20-1, 25, 30, 32, 34-5, 63-4, 74 

— alnea 1, 14, 18, 21, 55, 65 

— amoena 5, 14, 32 

— amygdalinae 70 

— arnstadtiensis 14, 32-3 

— carpinicola 13-14, 34, 74 

— euphorbiacea 72 

— fragariae 47-8 

— gnomon 2, 5, 11, 15, 18-20, 30, 51 

— incrassata 21 

— monodii 15, 21-2 

— neognomon 5, 15, 22-4 

— orcispora 25 

— ostryae 15, 25-6 

— pendulorum 15, 26-8 

— pseudoamoena 5, 14, 34-5 

— rodmanii 30 

— rosae 64 

— skokomishica 30 

— tithymalina 67 

— virginianae 5, 15, 31-2 

78


Gnomoniaceae 1-5, 7-15, 17-19, 21-3, 25, 27, 29, 31, 37-41, 44, 

47-9, 51, 53, 55, 63-5, 69, 73-5 

Gnomoniella 2-3, 74 

— amygdalinae 70 

Gnomoniella subgenus Ophiognomonia Sacc 3, 48 

Gnomoniopsis 2-3, 8-9, 11-13, 41, 44, 47 

— chamaemori 3, 5, 16, 39-41 

— clavulata 1, 16, 42-4, 47, 51 

— comari 5, 47 

— paraclavulata 44 

— racemula 6, 16, 48 

— tormentillae 6, 16, 48 

H 

Hamamelidaceae 10, 35-6 

Hippocastanaceae 10, 36, 66 

hippocastanum 69 

I 

ischnostyla 1, 5, 21, 53, 55, 58-60, 65 

J 

Juglandaceae 10-11, 48, 53, 59, 62-3 

L 

Leaf-inhabiting genera 1, 3, 5, 7, 9, 11, 13, 15, 17, 19, 21, 23, 

25, 27, 29, 31 

Linospora 12, 74 

Liquidambar styraciflua 5, 10, 12, 33, 35-6 

M 

melanostyla 3, 8, 11-12 

MycoBank MB 21-2, 25-6, 30, 32, 34-6, 40, 44, 47-8, 51, 53, 55, 

58-9, 62-4, 69-70, 72-3 

N 

nervisequa 60 

O 

Onagraceae 10, 48 

Ophiognomonia 2-3, 8-9, 11-13, 48, 51, 53, 55, 57-8, 60, 64, 75 

— alni-viridis 1, 6, 17, 21, 55 

— balsamiferae 51 

— intermedia 6, 17, 58 

— ischnostyla 6, 17, 58-60 

— leptostyla 6, 17, 62 

— melanostyla 2, 13, 15, 48-51 

— micromegala 6, 16, 60-1, 63 

— nana 6, 15, 63 

— padicola 6, 16, 63 

— pseudoclavulata 51 

— rosae 6, 16, 64 

— rubi-idaei 16, 64 

— sassafras 6, 15, 63-4 

— setacea 6, 13, 17, 64-5 

— trientensis 16, 64-5 

— vasiljevae 6, 17, 53-5 

Ostrya 14-15, 17 

— carpinifolia 33 

— virginiana 15 

ostryae 22-3, 25-6, 32 

P 

paraclavulata 39-40, 44, 47, 51 

Phragmoporthe 3 

Plagiostoma 2, 7-9, 11-13, 67, 69-70, 72, 74 

— amygdalinae 6, 16, 70, 72 

— barriae 1, 6, 16, 69 

— devexum 3, 6, 16, 70-1 

— euphorbiaceum 16, 72 

— euphorbiae 7, 16, 66-7, 69 

— euphorbiae-verrucosae 15, 72 

— fraxini 1, 7, 15, 72 

— petiolophilum 7, 14, 72 

Platanaceae 10, 36, 38, 66 

Pleuroceras 13, 74-5 

populi 2, 17, 21, 51, 74 

Populus nigra 18, 20, 74 

pseudoclavulata 1, 13, 49-52 

Q 

Quercus 16-17, 44, 47 

— rubra 44 

— spp 12, 44, 74 

R 

racemula 1, 39-40, 48 

rhododendri 70-1, 73 

rodmanii 1, 27-8, 30 

Rosaceae 10-12, 16, 41, 44, 47-8, 58, 63-4 

rosae 16, 60-1 

Rubus chamaemorus 5, 12, 44 

— idaeus 64 

S 

Salicaceae 10, 36, 38, 48, 51, 66, 74 

Salix 14 

Sydowiellaceae 2, 48, 74-5 

T 

Tilia americana 51 

— cordata 51 

— parvifolia 51 

— platyphyllos 51 

— sp 3, 5, 12, 15, 38, 48, 51 

Tiliaceae 10-11, 38, 48 

trientensis 1, 21, 55, 63-5 

U 

ulmea 75 

uncultureable species 3 

V 

Valsaceae 2, 12, 74 

vasiljevae 1, 53 

veneta 1-2, 10, 12, 40 


79

Studies in Mycology end of 2007 and 2008 


Black fungal extremes 

Edited by G.S. de Hoog and M. Grube 

Studies in Mycology 61: Black fungal extremes 

Edited by G.S. de Hoog and M. Grube 

Black fungi tend to be ecologically remarkable. Members of Dothideales / Capnodiales thrive under harsh environmental conditions, such as dryness, 

solar irradiation, high osmolarity or very low pH. Environments investigated range between Antarctic rock, Spitsbergen glaciers, Mediterranean saltpans, 

and even outer space. Mechanisms of stress tolerance are described, and new genera, species and varieties of extremotolerant fungi are reported. 

The human ‘infection’ tinea nigra by these species can be explained by halotolerant colonization. Members of the Chaetothyriales have ancestors with 

rock-dwelling life styles, but derived species are frequently found in human infections. Essential factors for ecology and evolution are described, such 

as oligotrophism, assimilation of aromatic hydrocarbons, and meristematic growth, and methods for selective isolation based on these features are 

proposed. Novel species are introduced, and tools for molecular identification are provided. 



An institute of the Royal Netherlands Academy of Arts and Sciences 

198 pp., fully illustrated with colour pictures (A4 format), paperback, 2008. €60 

Studies in Mycology 60: Neotropical Hypocrella (anamorph Aschersonia), Moelleriella, and 

Samuelsia 

Priscila Chaverri, Miao Liu and Kathie T. Hodge 

The present taxonomic revision deals with Neotropical species of three entomopathogenic genera that were once included in Hypocrella s. l.: Hypocrella 

s. str. (anamorph Aschersonia), Moelleriella (anamorph aschersonia-like), and Samuelsia gen. nov (anamorph aschersonia-like). Species of Hypocrella, 

Moelleriella, and Samuelsia are pathogens of scale insects (Coccidae and Lecaniidae, Homoptera) and whiteflies (Aleyrodidae, Homoptera) and are 

common in tropical regions. Phylogenetic analyses of DNA sequences from nuclear ribosomal large subunit (28S), translation elongation factor 1-α (TEF 

1-α), and RNA polymerase II subunit 1 (RPB1) and analyses of multiple morphological characters demonstrate that the three segregated genera can be 

distinguished by the disarticulation of the ascospores and shape and size of conidia. Moelleriella has filiform multi-septate ascospores that disarticulate 

at the septa within the ascus and aschersonia-like anamorphs with fusoid conidia. Hypocrella s. str. has filiform to long-fusiform ascospores that do not 

disarticulate and Aschersonia s. str. anamorphs with fusoid conidia. The new genus proposed here, Samuelsia, has filiform to long-fusiform ascospores 

that do not disarticulate and aschersonia-like anamorphs with small allantoid conidia. In addition, the present study presents and discusses the evolution 

of species, morphology, and ecology in Hypocrella, Moelleriella, and Samuelsia based on multigene phylogenetic analyses. 

66 pp., illustrated with colour pictures (A4 format), paperback, 2008. € 40 


Aspergillus systematics in the genomic era 

Studies in Mycology 59: Aspergillus systematics in the genomic era 

editors Robert A. Samson and János Varga 

Robert A. Samson and János Varga 

Since Pier Antonio Micheli described Aspergillus is his Nova Plantarum Genera in 1729 the genus attracted an immense interest. 

Many species were found as spoilage agents, or responsible for human and animal diseases. On the other hand Aspergilli were also 

found as beneficial micro organisms in the fermentation of Asian food and beverages. With the discovery of aflatoxins, the interest and 

research of the Aspergilli increased even more. In the present days Aspergillus research has grown to such a level, that it could be 

stated that Aspergilli might be the most studied fungi. 

This issue comprises 14 papers and is a summary of presentations and discussions of the international workshop entitled 

“Aspergillus systematics in the genomics era” with a multidisciplinary audience held in Utrecht, The Netherlands (12-14 April, 2007). 

The papers discuss topics such as the current species concept; what can comparative genomics tell us about species concepts 

in Aspergillus; sexual and vegetative compatibility genes in the aspergilli; secondary metabolite (including mycotoxins) profiling, 


growth profiles and other tools for species recognition; identification in the clinical setting; Aspergillus strain typing in the genomics 


An institute of the Royal Netherlands Academy of Arts and Sciences 

era and the biodiversity of Aspergillus species in some important agricultural products. Nomenclatural considerations in naming 

species of Aspergillus and its teleomorphs were discussed in a separate paper, while the recommendations of an international panel 

are included. Additionally, the polyphasic methods applied recently on aspergilli resulted in four monographs included in this issue: 

Aspergillus sections Candidi, Clavati, Fumigati and Usti. Diagnostic tools developed for the identification of the economically extremely important but taxonomically problematic 

black aspergilli (Aspergillus section Nigri) are also covered in a separate paper. This issue contains many colour illustrations, particularly in the four monographs. 

206 pp., fully illustrated with colour pictures (A4 format), paperback, 2007. € 65 

Studies in Mycology 58: The genus Cladosporium and similar dematiaceous hyphomycetes 

Pedro W. Crous, Uwe Braun, Konstanze Schubert and Johannes Z. Groenewald 

Species of Cladosporium are common and widespread, and interact with humans in every phase of life, from producing allergens in the 

indoor environment, to causing fruit decay and plant disease, or being associated with human mycoses. Although Cladosporium is one 

of the largest and most heterogeneous genera of hyphomycetes, only a mere fraction of these species are known from culture, and 

few have been characterised based on molecular data. The present volume consists of nine research papers, and introduces 71 new 

combinations, 53 new species, 15 genera, and one family. Specific conditions and media are recommended to study Cladosporium and 

allied genera, while the genus is circumscribed, and separated from morphologically similar taxa with which it has been confused in the 

past. Generic issues related to the anamorph genera and associated Dothideomycete teleomorph genera (Davidiella, Mycosphaerella, 

Venturia) are addressed, and multi-allelic data sets provided to facilitate species recognition. The volume is richly illustrated, with more 

than 100 colour plates, numerous line drawings, and scanning electron micrographs. 

253 pp., illustrated with colour pictures (A4 format), paperback, 2007. € 65 


The genus Cladosporium and similar 

dematiaceous hyphomycetes 

Pedro W. Crous, Uwe Braun, Konstanze Schubert 

and Johannes Z. Groenewald 

Centraalbureau voor Schimmelcultures, 



CBS Biodiversity Series 

No. 8: The genera of the Parmulariaceae 

Carlos Antonio Inácio and Paul F. Cannon 

The morphologically variable family Parmulariaceae (Fungi, Ascomycota, Dothideomycetes) is widespread in the tropics. The family now includes 34 accepted 

genera, with 24 further synonyms, and more than 100 species. The study was organized using a suite of computer databases, focusing on nomenclatural, 

geographical (floristic) and bibliographic information. More than 1000 scientific names were considered, and more than 1100 records of individual observations 

of these fungi have been gathered. All genera are fully redescribed and illustrated with drawings and microphotographs. A new key for identification of genera 

was constructed. A new formal taxonomic concept of the Parmulariaceae is introduced and two new genera Mintera and Viegasella have been re-described, 

Parmulariella is now included in the Parmulariaceae. Kentingia and Chaetaspis are now considered of uncertain family in the Dothideomycetes. The genus 

Apoa, previously placed as a synonym of Pachypatella, was found to be a distinct genus. 

196 pp., illustrated with 111 pictures including 15 colour plates (A4 format). Hardbound, 2008. € 55 

No. 7: Microfungi occurring on Proteaceae in the fynbos 

Seonju Marincowicz, Pedro W. Crous, Johannes Z. Groenewald and Michael J. Wingfield 

The fynbos is a shrubland characterised by a fire-prone ecosystem and Mediterranean climate. Although it is extremely rich in plant species, and has a high 

degree of floral endemism, very little is known regarding the fungi in this unique and fascinating environment. The present study investigated the saprobic 

microfungi that colonise and utilise leaf and twig litter and senescent flowerheads of Proteaceae. A total of 29 species and sub-species belonging to four 

genera of Proteaceae were sampled from 12 sites in the Western Cape Province of South Africa spanning a period of two years (2000–2001). An attempt 

was made to culture all fungi encountered, and where successful, the ITS and partial 28S nrDNA, and in some cases the translation elongation factor 1-α or 

the β-tubulin gene regions were sequenced. A total of 62 bags of litter yielded 316 individuals, consisting of 141 fungal species residing in 103 genera and 

43 families. Of these, 59 species, including eight species that had been previously published, represented novel taxa. Thirty-eight species reflected new 

records for South Africa, and 48 species were new reports on Proteaceae. Two new genera and one new combination were also introduced. Seventy-three 

species were represented by teleomorphs and 68 species by anamorphs, which were made up of 30 hyphomycetes and 38 coelomycetes. Eighty percent 

of the species occurred on only one type of substrate. The fungal community found on twig litter had the highest species richness, while flowerhead-styles 

yielded the highest percentage (100 %) of unique species. These results showed that the species richness for the fynbos Mycota was moderately high with 

every 2.2 collections representing a different fungal species. The percentage of new fungal taxa (43 % of the total species) was exceptionally high, and 

most of these probably host-specific. More than 80 % of the fungi collected in this study had hard and closed fruiting structures, indicating an adaptation to 

the constraints of the harsh fynbos environment. Other than providing a foundation for further studies, this investigation highlights a disturbing paucity of 

knowledge regarding the fynbos Mycota in one of the world’s most threatened and unusual floral Kingdoms. 

Microfungi occurring on Proteaceae 

in the fynbos 

Seonju Marincowitz 

Pedro W. Crous 

Johannes Z. Groenewald 

Michael J. Wingfield 

166 pp., illustrated with 93 colour pictures and 6 B&W pictures (A4 format). Hardbound, 2008. € 50 

No. 6: Alternaria An Identification Manual 

Emory G. Simmons 

This book will fill a very large void in the scientific literature and it is quite certain that the volume will become the standard 

reference for those needing to have critical access to Alternaria literature and taxonomic information. There are many scientists, 

both research and regulatory, who are in desperate need of resources like this book to facilitate routine identification. More than 1 

100 published names are associated with taxa that must be considered in the Alternaria context. Of these, 276 species with validly 

published names are maintained here as currently identifiable; these are keyed in the main text of the volume. An additional 16 

named taxa, although requiring expanded information and comparison, also are accepted. A few species that have been associated 

with the genus for years but which now are considered anomolous in the genus have been removed to other genera. Chapters of 

species and genus characterisations are followed by a comprehensive list of all the nearly 1 200 names involved historically with 

Alternaria taxonomy in the period 1796-2007. Each name is listed with its source, type, and an opinion on its validity and taxonomic 

disposition. A host index to all accepted species is followed by a comprehensive list of literature cited and a general index. Within 

the context of the manual, 88 names are assigned to newly described species and genera and to taxa whose epithets appear in 

new combinations. 

775 pp., with more than 288 line drawings (A4 format). Hardbound, 2007. € 170 

No. 5: Mycosphaerella and its anamorphs: 2. Conspectus of Mycosphaerella 

André Aptroot 

A revision of the species described in Mycosphaerella and Sphaerella is presented, together with observations on the types of most 

species or their disposition. The genus Stigmidium is expanded to encompass fungicolous species and internal parasites of algae, 

and includes the genus Mycophycias. 

Mycosphaerella and its anamorphs: 

2. Conspectus of Mycosphaerella 

André Aptroot 

173 pp., 115 plates (A4 format), paperback with spiral binding, 2005. € 50

CBS Biodiversity Series 

No. 4: Hypocreales of the Southeastern United States: An Identification Guide 

Hypocreales of the Southeastern 

United States: 

An Identification Guide 

Gary J. Samuels, Amy Y. Rossman, Priscila Chaverri, 

Barrie E. Overton and Kadri Põldmaa 

Gary J. Samuels, Amy Y. Rossman, Priscila Chaverri, Barrie E. Overton and Kadri Põldmaa 

An illustrated guide is presented to the members of the ascomycete order Hypocreales that are known to occur in the southeastern 

states of the United States, including North and South Carolina, Tennessee and Georgia. Species were selected mainly based 

on records in the United States National Fungus Collections (BPI). These states include or surround the Great Smoky Mountains 

National Park. Species of the Hypocreales are among the most numerous, and certainly most conspicuous, of the microfungi. The 

order also includes some of the most economically important fungi. This guide is intended for individuals who are participating in 

All Taxa Biological Diversity studies of the Great Smoky Mountains National Park as well as other interested professionals and 

amateurs. Short descriptions and colour illustrations of one-hundred and one species and two varieties in twenty genera are 

provided. Keys to genera and species are included. The new combination Neonectria ditissima is proposed. 

145 pp., over 120 colour pictures (A4 format), paperback with spiral binding, 2006. € 70 

No. 3: An illustrated guide to the coprophilous Ascomycetes of Australia 

Ann Bell 

Descriptions, keys and illustrations (many in colour). Ann Bell's observations of her own collections and some 2 000 microscope 

slides and assorted notebooks on Australian coprophilous fungi made by the late Major Harry Dade during his retirement years in 

Victoria. 

173 pp., 115 plates (A4 format), paperback with spiral binding, 2005. € 55 

No. 2: Cultivation and Diseases of Proteaceae: Leucadendron, Leucospermum and Protea 

Pedro W. Crous, Sandra Denman, Joanne E. Taylor, Lizeth Swart and Mary E. Palm 

The Proteaceae represent one of the Southern Hemisphere"s most prominent flowering plant families, the cultivation of which forms the 

basis of a thriving export industry. Diseases cause a loss in yield and also limit the export of these flowers due to strict phytosanitary 

regulations. In this publication the fungi that cause leaf, stem and root diseases on Leucadendron, Leucospermum and Protea are 

treated. Data are provided pertaining to the taxonomy, identification, host range, distribution, pathogenicity and control of these 

pathogens. Taxonomic descriptions and illustrations are provided and keys are included. Desease symptoms are illustrated with colour 

photographs. 

510 pp., (17 x 25 cm), paperback, 2004. € 55 

No. 1: Mycosphaerella and its anamorphs: 1. Names published in Cercospora and Passalora 

Pedro W. Crous and Uwe Braun 

This book contains a compilation of more than 3000 names that have been published or proposed in Cercospora, of which 659 are 

presently recognised in this genus, with a further 281 being referred to C. apii s.lat. Approximately 550 names of Passalora emend. (incl. 

Mycovellosiella, Phaeoramularia, Tandonella and Phaeoisariopsis p.p.) are treated in a second list. In total 5720 names are treated. 553 

taxonomic novelties are proposed. 

571 pp., 31 figures (17 x 25 cm), hard cover, 2003. € 80

Selection of other CBS publications 

Introduction to food- and airborne fungi 

Robert A. Samson, Ellen S. Hoekstra and Jens C. Frisvad 

Seventh edition with updated taxonomy and addition of some important species. The keys to the taxa were improved. The taxonomy 

and the nomenclature of Fusarium is revised and the number of Penicillium species has been increased because they are frequently 

encountered on food and indoor environments. The identification of Penicillium based on morphological characters remains difficult and 

therefore synoptic keys and tables are added to assist with the identification. 

389 pp., 120 plates (A4 format), paperback, 2004. € 55 

Identification of Common Aspergillus Species 

Maren A. Klich 

Descriptions and identification keys to 45 common Aspergillus species with their teleomorphs (Emericella, Eurotium, Neosartorya and 

Sclerocleista). Each species is illustrated with a one page plate and three plates showing the most common colony colours. 

116 pp., 45 black & white and 3 colour plates (Letter format), paperback, 2002. € 45 

A revision of the species described in Phyllosticta 

Huub A. van der Aa and Simon Vanev 

2936 taxa are enumerated, based on the original literature and on examination of numerous herbarium (mostly type) specimens 

and isolates. 203 names belong to the genus Phyllosticta s.str., and are classified in 143 accepted species. For seven of them new 

combinations are made and for six new names are proposed. The great majority, 2733 taxa, were redisposed to a number of other 

genera. A complete list of these novelties, as included in the book"s abstract, can also be consulted on the web-site of CBS. 

510 pp. (17 x 25 cm), paperback, 2002. € 55 

Atlas of Clinical Fungi 

G.S. de Hoog, J. Guarro, J. Gené and M.J. Figueras (eds) 

The second fully revised and greatly expanded edition of the Atlas of Clinical Fungi appeared in 2000. The modest and very competitive 

price of this standard work has certainly contributed to the popularity of the first edition. In recent years the application of molecular 

biology has become within reach for many routine laboratories. The new Atlas will provide ample molecular data for the majority of 

clinically relevant fungi. It will set a standard for innovative techniques in medical mycology. In addition, antifungal 

susceptibility data will be given for most species, which will provide essential knowledge for the clinician in view 

of adequate therapy. 

1126 pp., fully illustrated with line drawings and black & white photo plates (A4 format), hard cover, 2004. 

€ 140. 

Interactive CD-ROM v. of the Atlas € 65, Book plus CD-ROM € 180

Studies in Mycology (ISSN 0166-0616) 

The CBS taxonomy series “Studies in Mycology” is issued as individual booklets. Regular subscribers receive each issue automatically. Prices of backvolumes 

are specified below. 

For more information and ordering of other CBS books and publications see www.cbs.knaw.nl. and www.studiesinmycology.org 

62 Sogonov MV, Castlebury LA, Rossman AY, Mejía LC, White JF(2008). Leaf-inhabiting genera of the Gnomoniaceae, Diaporthales. 79 pp. € 40,00 

61 Hoog GS de, Grube M (eds) (2008). Black fungal extremes. 198 pp., € 60.00 

60 Chaverri P, Liu M, Hodge KT (2008). Neotropical Hypocrella (anamorph Aschersonia), Moelleriella, and Samuelsia. 68 pp., € 40.00 

59 Samson RA, Varga J (eds) (2007). Aspergillus systematics in the genomic era. 206 pp., € 65.00 

58 Crous PW, Braun U, Schubert K, Groenewald JZ (eds) (2007). The genus Cladosporium and similar dematiaceous hyphomycetes. 253 pp., € 65.00 

57 Sung G-H, Hywel-Jones NL, Sung J-M, Luangsa-ard JJ, Shrestha B, Spatafora JW (2006). Phylogenetic classification of Cordyceps and the 

clavicipitaceous fungi. 63 pp., € 40.00 

56 Gams W (ed.) (2006). Hypocrea and Trichoderma studies marking the 90 th birthday of Joan M. Dingley. 179 pp., € 60.00 

55 Crous PW, Wingfield MJ, Slippers B, Rong IH, Samson RA (2006). 100 Years of Fungal Biodiversity in southern Africa. 305 pp., € 65.00 

54 Mostert L, Groenewald JZ, Summerbell RC, Gams W, Crous PW (2006). Taxonomy and Pathology of Togninia (Diaporthales) and its Phaeoacremonium 

anamorphs. 115 pp., € 55.00 

53 Summerbell RC, Currah RS, Sigler L (2005). The Missing Lineages. Phylogeny and ecology of endophytic and other enigmatic root-associated fungi. 

252 pp., € 65.00 

52 Adams GC, Wingfield MJ, Common R, Roux J (2005). Phylogenetic relationships and morphology of Cytospora species and related teleomorphs 

(Ascomycota, Diaporthales, Valsaceae) from Eucalyptus. 147 pp., € 55.00 

51 Hoog GS de (ed.) (2005). Fungi of the Antarctic, Evolution under extreme conditions. 82 pp., € 40.00 

50 Crous PW, Samson RA, Gams W, Summerbell RC, Boekhout T, Hoog GS de, Stalpers JA (eds) (2004). CBS Centenary: 100 Years of Fungal Biodiversity 

and Ecology (Two parts). 580 pp., € 105.00 

49 Samson RA, Frisvad JC (2004). Penicillium subgenus Penicillium: new taxonomic schemes, mycotoxins and other extrolites. 253 pp., € 55.00 

48 Chaverri P, Samuels GJ (2003). Hypocrea/Trichoderma (Ascomycota, Hypocreales, Hypocreaceae): species with green ascospores. 113 pp., € 55.00 

47 Guarro J, Summerbell RC, Samson RA (2002). Onygenales: the dermatophytes, dimorphics and keratin degraders in their revolutionary context. 220 

pp., € 55.00 

46 Schroers HJ (2001). A monograh of Bionectria (Ascomycota, Hypocreales, Bionectriaceae) and its Clonostachys anamorphs. 214 pp., € 55.00 

45 Seifert KA, Gams W, Crous PW, Samuels GJ (eds) (2000). Molecules, morphology and classification: Towards monophyletic genera in the Ascomycetes. 

200 pp., € 55.00 

44 Verkley GJM (1999). A monograph of the genus Pezicula and its anamorphs. 180 pp., € 55.00 

43 Hoog GS de (ed.) (1999). Ecology and evolution of black yeasts and their relatives. 208 pp., € 55.00 

42 Rossman AY, Samuels GJ, Rogerson CT, Lowen R (1999). Genera of Bionectriaceae, Hypocreaceae and Nectriaceae (Hypocreales, Ascomycetes). 248 

pp., € 55.00 

41 Samuels GJ, Petrini O, Kuhls K, Lieckfeldt E, Kubicek CP (1998).The Hypocrea schweinitzii complex and Trichoderma sect. Longibrachiatum. 54 pp., € 

35.00 

40 Stalpers JA (1996). The aphyllophoraceous fungi II. Keys to the species of the Hericiales. 185 pp., € 25.00 

39 Rubner A (1996). Revision of predacious hyphomycetes in the Dactylella-Monacrosporium complex. 134 pp., € 25.00 

38 Boekhout T, Samson RA (eds) (1995). Heterobasidiomycetes: Systematics and applied aspects. 190 pp., € 35.00 

37 Aptroot A (1995). A monograph of Didymosphaeria. 160 pp., € 30.00 

36 Swertz CA (1994). Morphology of germlings of urediniospores and its value for the identification and classification of grass rust fungi. 152 pp., € 30.00 

35 Stalpers JA (1993). The aphyllophoraceous fungi I. Keys to the species of the Thelephorales. 168 pp. (out of stock). 

34 Reijnders AFM, Stalpers JA (1992). The development of the hymenophoral trama in the Aphyllophorales and the Agaricales. 109 pp., € 25.00 

33 Boekhout T (1991). A revision of ballistoconidia-forming yeasts and fungi. 194 pp. (out of stock). 

32 Gams W, Seifert KA, Aa van der HA, Samson RA (1990). Developments in the taxonomy of anamorphic fungi. Invited papers presented at the Symposium: 

Taxonomy of Fungi imperfecti during the Fourth International Mycological Congress, Regensburg. 104 pp. (out of stock). 

31 Aa HA van der, Gams W, Hoog GS de, Samson RA (1988). Memorial issue dedicated to J.A. von Arx. 212 pp., € 50.00 

30 Hoog GS de, Smith M Th, Weijman ACM (eds) (1987). The expanding realm of yeast-like fungi. (only available from Elsevier). 

29 Hoog GS de, Smith M Th, Guého E (1986). A revision of Geotrichum. 131 pp., € 25.00 

28 Rao V, Hoog GS de (1986). New or critical Hyphomycetes from India. 84 pp., € 20.00 

27 Seifert KA (1985). A monograph of Stilbella and some allied Hyphomycetes. 235 pp., € 35.00 

26 Hoog GS de (ed.) (1985). Taxonomy of the Dactylaria complex, IV–VI. 124 pp., € 20.00 

25 Schipper MAA, Stalpers JA (1984). A revision of the genus Rhizopus. 34 pp., € 10.00 

24 Stalpers JA (1984). A revision of the genus Sporotrichum. 105 pp., € 20.00 

23 Stolk AC, Samson RA (1983). The ascomycete genus Eupenicillium and related Penicillium anamorphs. 149 pp., € 25.00 

22 Hoog GS de (ed.) (1982). Character analyses of selected red yeasts. 74 pp. (out of stock). 

21 Plaats-Niterink AJ van der (1981). Monograph of the genus Pythium. 244 pp. (out of stock). 

20 Oorschot CAN van (1980). A revision of Chrysosporium and allied genera. 89 pp., € 20.00 

For a complete list of the Studies in Mycology see www.cbs.knaw.nl.

Leaf-inhabiting genera of the Gnomoniaceae, Diaporthales - CBS

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