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The Genera of Hyphomycetes

About the cover: Agar colonies of species of Fusarium, Epicoccum and Stilbella. Detail from an installation by Canadian artist Annie Thibault, in the exhibition Beyond Science, Metronòm – Fundació Rafael Tous d’Art Contemporani, with the collaboration of Institut de Biología Molecular de Barcelona, Spain, 03.12-22.01.2005 (photograph: José Nieva). The accompanying photograph shows a similar installation at a larger scale, from Fruitfulness and Multiplication, in the exhibition De-con-structions, curated by Josée Drouin Brisebois at the National Gallery of Canada, Ottawa, with the collaboration of Keith Seifert and Agriculture and Agri-Food Canada, 14.04-03.09.2007 (photograph: Christopher Varady-Szabo). Used with the permission of the artist; all rights reserved.

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Publication date: 1 April 2011

Published and distributed by CBS-KNAW Fungal Biodiversity Centre, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.

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THE GENERA of HYPHOMYCETES

CBS-KNAW Fungal Biodiversity CentreUtrecht, The Netherlands

An institute of the Royal Netherlands Academy of Arts and Sciences

Keith Seifert Ph.D. (Utrecht)

Biodiversity (Mycology and Botany), Eastern Cereal and Oilseed Research Centre,

Agriculture & Agri-Food Canada, Ottawa, Ontario, Canada.

Gareth Morgan-Jones Ph.D. (Nottingham), D.Sc. (Wales)

Department of Entomology and Plant Pathology, College of Agriculture, Auburn University, Alabama, USA.

Walter GamsPh.D. (Innsbruck)

CBS-KNAW Fungal Biodiversity Centre, Utrecht, The Netherlands (retired).

Bryce Kendrick Ph.D., D.Sc. (Liverpool), F.R.S.C.

Department of Biology,University of Waterloo, Ontario, Canada (retired).

CBS Biodiversity Series 9

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Keith Seifert has been with the Mycology Group of Agriculture and Agri-Food Canada, Ottawa, since 1990, after completing his Ph.D at CBS in 1985, and spending four years with Forintek Canada, Corp., in Ottawa. He is a specialist on synnematous hyphomycetes and the mycotoxigenic genera Fusarium and Penicillium, has published more than 150 papers on the tax-onomy of hyphomycetes, and is past chair of the International Commission on the Taxonomy of Fungi. He lives in a log house in a forest, where he throws Frisbees for dogs, and strums gui-tars. Dedicated with love to Charlene.

Walter Gams recently retired from the CBS after 40 years studying the little-differentiated step-children of fungal taxon-omy: the zygomycete genera Mortierella and Umbelopsis, and acremonium-like and verticillium-like hyphomycetes. He was secretary of the nomenclatural Committee for Fungi for sev-eral years and still defends dual nomenclature. He is a coauthor of the well-known identification manual Compendium of Soil Fungi, and is an author of several hundred scientific papers. He teaches mycology as Professor at the Technical University in Aachen (Germany). He collaborates with many international colleagues, and is a patron of art and music, dividing his time between Baarn, the Netherlands, and his own segment of the medieval castle of Bomarzo, Italy.

Bryce Kendrick retired as a professor in the Department of Biology at the University of Waterloo in 1994, after an active career as a teacher and researcher. Equally well known for his textbook The Fifth Kingdom, and his Kananaskis conference proceedings (Taxonomy of the Fungi Imperfecti and The Whole Fungus), Bryce was also an author of the Carmichael et al. pro-genitor of this book and about 300 other mycological publica-tions. He is now Principal of Mycologue Publications, based on Vancouver Island, and devotes much of his energy to environ-mental causes. Dedicated with love to Laurie.

Gareth Morgan-Jones has been a tenured faculty member at Auburn University, Alabama, USA, for 38 years, holding the rank and title of Distinguished University Professor since 1994. Educated at the Universities of Wales and Nottingham, Eng-land, he previously held positions at the former Biosystematics Research Institute, Ottawa, and the Commonwealth Mycologi-cal Institute, Kew. He has published widely on the taxonomy of both coelomycetes and hyphomycetes, authoring or co-authoring over 250 papers. His avocations include cultivating his two-acre woodland garden, tending his bonsai collection, and breeding, exhibiting and judging pedigreed dogs, for which he is approved by the American Kennel Club.

About the authors

Keith with Seamus

Walter with Chico

Bryce with Chelsea

Gareth with Tristan

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Acknowledgments ....................................................................................................................vi

Introduction ...............................................................................................................................1I. Overview ...........................................................................................................................3II. Classification and phylogeny .............................................................................................4III. Nomenclature ..................................................................................................................25IV. Using the dictionary ........................................................................................................26V. Notes on illustrations .......................................................................................................54VI. Internet resources .............................................................................................................54VII. Literature cited in the Introduction and additional references .........................................57

Dictionary of Hyphomycete Genera ..................................................................................... 61Synoptic Plates ......................................................................................................................481Keys .......................................................................................................................................867

Appendices1. Taxonomic classification of hyphomycete genera ............................................................9452. Alphabetical list of teleomorph-anamorph connections ...................................................9503. List of synanamorphies including hyphomycete genera ...................................................9574. Glossary ............................................................................................................................9615. Notes on and sources of illustrations ................................................................................9786. Index to plates ...................................................................................................................981

TABLE OF CONTENTS

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The preparation of this book occupied two decades, and we owe a special debt of gratitude to our families and many colleagues, who lived with its growing pains and suffered from our neglect as the work proceeded.

We are grateful to two authors of the 1980 book Genera of Hyphomycetes, Bill Carmichael and Lynne Sigler, who were unable to participate in this project, but provided the original computer files and information accumulated in the years following the original publication. The legacy of Ibra L. Conners, in the form of an extensive collection of original protocols of fungal genera housed with the DAOM herbarium, was an invaluable resource for our work. The University of Alberta Press graciously surrendered rights to the 1980 book, allowing our own project to proceed.

George Bills, Sarah Hambleton, Vadim Mel’nik, Bernhard Oertel, Robert Shoemaker and George White commented on drafts of earlier versions of the manuscript, and we are very appreciative of time they devoted to these reviews. We are grateful to many colleagues who reviewed parts of the text, provided insight into the taxonomy and nomenclature of genera in their areas of interest, sometimes allowing us access to unpublished results, including Sean Abbott, Andre Aptroot, Meredith Blackwell, Uwe Braun, Lori Carris, Pedro Crous, John David, Jens Frisvad, Kathie Hodge, Sybren de Hoog, Stan Hughes, Richard Humber, Karin Jacobs, Paul Kirk, Roland Kirschner, Linda Kohn, Matthias Lutz, Ludmila Marvanová, Franz Oberwinkler, Silvano Onofri, Kadri Põldmaa, Martina Réblová, Scott Redhead, Amy Rossman, Rob Samson, Gary Samuels, Markus Scholler, Hans-Josef Schroers, Emory Simmons, Lynne Sigler, Joost Stalpers, Jeff Stone, Richard Summerbell, Michael Vincent and Hermann Voglmayr. Bernhard Oertel assisted with some classical literature and brought several previously overlooked genera to our attention. We received assistance with some of the database and other computerized aspects of this project from John Brzustowski, and advice on layout from Helmuth Schmid.

The following colleagues provided excellent photographs for this book: Toni Atkinson, George Barron, Adolf Ceska, Pedro Crous, Dennis Desjardins, Diane Earl, Sarah Hambleton, Kathie Hodge, Karin Jacobs, Kent Loeffler, Gen Okada, Brian Perry, Martina Réblová, Scott Redhead, Gary Samuels, Rob Samson, Hans-Josef Schroers, James Scott and Adnan Uzunovic.

We are very grateful to Uwe Braun, Pedro Crous, Paul Diederich, Ludmila Marvanová, Gen Okada, Martina Réblová and Rasoul Zare for providing or contributing to identification keys for groups in their areas of expertise. Conrad Schoch provided invaluable assistance with the phylogenetic trees in the Introduction.

Annie Thibault, of Gatineau, Quebec, is warmly thanked for generously allowing the use of her unique artwork for the book cover and Richard D. Albee, at Auburn, for providing his graphics expertise in the process of refining the cover design. The latter also participated in the formatting of the line-drawing Introduction and Glossary figures for which we are most grateful.

Rob Samson expressed constant interest in this project, and we are grateful for his efforts in facilitating final pre-publication preparation.

Keith thanks the members of his lab, and the mycology group in Ottawa, for their industry and patience over the many years this project took to complete. In particular, Gerry Louis-Seize kept the laboratory running smoothly, and together with Ed Whitfield, helped in the final stages with the organization of files and checking of cross-references. The successive Directors at the Eastern Cereal and Oilseed Research Centre, and leaders of the Mycology group, especially Scott Redhead and André Lévesque, provided a research environment that has allowed biosystematic research to flourish. The IT-support team at AAFC, Robert Guilbault, Peter Miller and Neil O’Connor, was very patient as computing needs constantly exceeded the norm, and were very inventive at finding solutions to problems that never existed until Keith tried things that were not meant to be tried.

Gareth acknowledges with gratitude the support of successive Deans and Department Chairs in the College of Agriculture at Auburn University for allowing him the considerable freedom required to fulfill his part of the project. A much-valued colleague, Dr. Katheryn Lawrence, provided enthusiastic encouragement over the many years that it took to complete the work.

Walter is grateful to have had the facilities of CBS for 40 years, which allowed him to keep up with literature, culti-vate the fungi of his interest and maintain contact with numerous friends in the mycological world. Wieland Meyer, Martina Réblová and Rasoul Zare took the molecular lead in collaborative work that enabled him to stay abreast of recent developments in fungal taxonomy.

Bryce thanks his grandchildren (Celeste, Isabella, Mara, Skylar and Tray) for their constant stimulation, which has kept his attitude to life and the future positive; Laurie, his wife of 33 years, for her patience and for thousands of hikes that kept him healthy; his dog, Chelsea, who led those hikes for 16 years; friends, mycological and other, for ongo-ing support and encouragement; the main users of his consulting skills; Aerobiology Laboratory Associates, Dulles, Virginia (Suzanne Blevins and her analysts) and Pacific Environmental Consulting, Vancouver, B.C. (Peter Hansen and his analysts) for their patience when he was preoccupied by this book; and several gurus who kept his computers up and running over the years.

Acknowledgments

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INTRODUCTION

Seifert, Morgan-Jones, Gams & Kendrick

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Overview

IntroductionI. Overview

This book is a gateway to a hidden world invisible to most people. Its purpose is to assist scientists, students, and other naturalists to identify hyphomycetes (micro-scopic fungi that are often called moulds) to the level of genus. You can reach this goal by learning the ter-minology and concepts presented in this Introduction, scanning the synoptic plates, or by using the selection of dichotomous keys in the fourth part of the book. The Dictionary provides concise information on each genus and directs you to literature on species identification. This section also includes information on substrates, geographical distribution, teleomorphs, synanamorphs, DNA barcodes and the phylogenetic position of each genus. Our book is a complete revision and expansion of Carmichael, Kendrick, Conners & Sigler’s 1980 work Genera of Hyphomycetes, which was itself based on a book chapter by Kendrick & Carmichael (1973). We incorporated and expanded on successful elements of these works, added new features, and updated infor-mation to facilitate study of these fungi. The book contains four main sections, three appendi-ces, and a glossary. In the Introduction (pp. 1–60), we present basic information on the naming, biology and taxonomy of hyphomycetes. If you are unfamiliar with these organisms, careful reading of this section is essen-tial. Even if you are experienced with these fungi, this section will provide insight into the suites of traits used to characterize the accepted genera and how we use these to arrange them for identification purposes in the synoptic plates. The Introduction provides overviews of: (1) the history of the classification of hyphomycetes; (2) the rules that govern their names; and (3) the char-acters that are used to classify and identify them. We conclude with an explanation of the organization of the Dictionary, including definitions of the many abbrevia-tions, and recommendations to other helpful books and Web sites. The Dictionary (pp. 61–479) contains text records for every described genus of hyphomycetes (to the end of 2009, with some coverage for 2010 and 2011). Each genus record contains full bibliographic details, syn-onymies, key word descriptions that summarize genus concepts, details of substrates, geographical distribu-

tion, number of accepted species, anamorph–teleomorph connections (asexual reproductive structures linked to sexual reproductive structures), GenBank accession numbers for DNA barcodes, notes, and a bibliography. The targeted taxonomic level is explicitly the genus. The bibliographies for each genus include all relevant refer-ences to monographs, revisions or other publications that can be used to confirm genus identifications or identify individual species. Finally, because morphological vari-ation in some hyphomycete genera is considerable, and can be deceptive, we include a few additional drawings or photographs illustrating the variation among species of some genera. The Synoptic Plates (pp. 481–866) contain line-draw-ings of the type or other representative species of most accepted hyphomycete genera, arranged according to morphological similarities. Some genera that do not lend themselves well to illustration (such as sterile, yeast-like or sclerotial genera) are not illustrated. To be success-ful with this book, you should study the arrangement of plates to understand the underlying logic necessary to use them for identification purposes. Each plate has a thumb tab on the margin of the page noting its major unifying features. These synoptic plates should help you distin-guish similar genera, allowing you to compare genus concepts visually and intuitively without the need to consider the details and specialized terminology of the dichotomous keys. The Keys section (pp. 867–942) includes dichotomous or polychotomous keys to several ‘generic complexes’ or ecological groups, providing another way of arriving at a genus identification. Several keys were prepared for this book by taxonomic specialists, and we adapted oth-ers from the literature. The task of assembling a func-tional dichotomous key for all genera was not possible during this project, but we believe the included keys will be helpful. The Appendices (pp. 943–997) contain information extracted from the Dictionary, namely a putative classi-fication that distributes hyphomycete genera into asco-mycete or basidiomycete classes, orders and families suggested by their phylogenetic or teleomorphic rela-tionships, a list of anamorph–teleomorph connections

Fig. 1. Early illustrations of hyphomycetes. A. Alternaria tenuis from Nees (1816). B. Tubercularia vulgaris from Tode (1790). C. Acremonium alternatum from Ditmar in Sturm (1817). D. Stachybotrys atra from Corda (1837). E. A Hypocrea perithecium in section, with its Trichoderma anamorph growing from the top, from Tulasne & Tulasne (1863).


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Fig. 2. Triposporium elegans, as illustrated in Corda’s Prachtflora europäischer Schimmelbildungen (1839).

Wha

t are

fung

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arranged by teleomorph genus, and a list of synanamorph connections. The Glossary (pp. 961–977) is an alphabetical list of additional specialized terms used in this book, along with their definitions, to assist with the interpretation of genus descriptions and dichotomous keys. We have not written a section on methodology, but some information is provided in text boxes in the plates, as listed on p. 481.

II. Classification and phylogeny

A. What are fungi?

Mycologists define true fungi as heterotrophic, absorp-tive eukaryotes, generally with cell walls of β-glucan and chitin (although a few have lost chitin or even their entire wall during evolution). Fungi are generally aerobic, their organelles including mitochondria with flat cristae, and peroxisomes (although two small extant groups are obli-gately anaerobic and have lost these organelles). Fungi lack plastids. The earliest fungal cells had a single back-ward directed flagellum, but most extant fungi have lost this feature. The somatic expression of most fungi is hyphal, and the linear growth and branching of hyphae give rise to mycelial colonies that are extensive in some cases, but the fungal spectrum also includes many non-hyphal forms, and some extremely reduced unicells (see Cavalier-Smith 2001, Blackwell et al. 2006). By these criteria, estimates of the number of fungal species gener-ally vary from 1.5 to 15 million species, with fewer than 100,000 presently described (Hawksworth 2001). Although linked by their evolution, fungi are still diverse, particularly when you consider the giant colonies of Armillaria at one end of the spectrum (the ‘humun-gous fungus’), and Nosema, an intracellular insect patho-gen, or the unicellular Pneumocystis, which causes pneu-monia, at the other. Traditionally the group was even more heterogeneous, because fungi, slime moulds, and bacteria were sometimes lumped together as members of the Phylum Thallophyta of the plant kingdom. As related by Whittaker (1969), various authors suggested that fungi were not plants up to two centuries ago, and that slime moulds should be excluded from the fungi. It was not until the advent of the electron microscope in the 1950s that the majority of biologists accepted the fun-damental separation between what was first presented

as Lower Protists (bacteria and prokaryotic blue-green algae, which are now often called the Prokaryota) and Higher Protists (eukaryotic organisms: fungi, protozoa, and algae). Subsequently, the Lower Protists, the Fungi, and the Protozoa were recognized as separate Kingdoms (Whittaker 1969). In general, we now recognize that liv-ing things have evolved into three phylogenetic domains, a) Eukarya (eukaryotes), b) Bacteria and c) Archaea. The fungi are a phylogenetically coherent, kingdom-level subgroup of eukaryotes. The slime moulds have been accepted as protozoa by protozoologists (e.g., Kudo 1946) ever since de Bary (1859, Fig. 4A) first named them Mycetozoa. However, mycologists are reluctant to disown them, probably because it is they, and not protozoologists, who study this group. Here, we prefer to call the well-known ‘plas-modial slime moulds’ Phylum Mycetozoa (increasingly known as Myxostelida, or informally myxostelids) rather

Fig. 3. Early mycologists who had a profound influence on hyphomycete taxonomy. A– C.H. Persoon (1761–1836). B– E.M. Fries (1794–1878). C– J.H.F. Link (1767–1851). D– A.C.J. Corda (1809–1849). E– P.A. Saccardo (1845–1920) (public domain).


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than Myxomycetes or Myxomycota, because the endings ‘-mycetes’ or ‘-mycota’ imply a fungal nature, and we refer them to Kingdom Protozoa. The ‘cellular slime moulds’ (Phylum Dictyostelida), the ‘net slime moulds’ (Phylum Labyrinthulida) and the ‘endoparasitic slime moulds’ (Phylum Plasmodiophorida) are now also clas-sified in the Kingdom Protozoa. Even with slime moulds excluded, the organisms we generally call ‘fungi’ are not monophyletic. Organ-isms that look distinctly fungal under the light micro-scope, because in many cases their colonies are built up of branching hyphae, are now placed in two different Kingdoms. The ‘water moulds’ and ‘downy mildews’ of Phylum Oomycota (and the tiny Phylum Hyphochy-triomycota) are now understood to belong in Kingdom Chromista (sometimes called Stramenipila), the King-dom that also contains the diatoms and the brown algae. These Chromistan pseudofungi differ from true fungi in many important ways: their heterokont flagellation, a wall chemistry that includes cellulose, somatic ploidy, mitochondrial cristae, lysine biosynthesis, etc. After the slime moulds and the pseudofungi are excluded, the true Fungi comprising the Kingdom Eumycota remain, as defined in the first paragraph of this section. The hyphomycetes are all members of this Kingdom, but they are found in two different Phyla, the Ascomycota and the Basidiomycota. We currently recognize six Phyla (also known as Divi-sions) in Kingdom Eumycota or the Fungi (see Kendrick 2010): Phylum Chytridiomycota Phylum Zygomycota Phylum Glomeromycota Phylum Ascomycota Phylum Basidiomycota Phylum Microsporidia (recently recognized as redu-ced or secondarily simplified fungi, but still under zoo-logical nomenclatural rules). The phyla Ascomycota and Basidiomycota together comprise the subkingdom Dikarya. It is unnecessary for us to describe and differentiate the major fungal groups here. Motivated readers can pursue this matter in Cavalier-Smith (2001), Kirk et al. (2008), and Kendrick (2010), or other modern general mycology texts. The Nov.–Dec. 2006 issue of Mycologia includes the results of the ‘Assembling the Fungal Tree of Life’

project, and provides a comprehensive overview of all fungal groups from a phylogenetic perspective (Spata-fora 2005, Blackwell et al. 2007, Hibbett et al. 2007). The latter project provides the basis for most of the larger scale phylogenetic trees presented in this introduction, which show the relationships of selected hyphomycete genera in the series starting with Fig. 6.

B. What are hyphomycetes?

Hyphomycetes make up the majority of what are com-monly called moulds1, and some are regarded as the weeds of the fungal kingdom. In addition to growing on many natural substrates such as plant tissues (Fig. 5), wood and bark (Fig. 7), dung (Fig. 12), insects and other arthropods (Figs 14, 22), and other fungi (Fig. 16, 38B) including lichens (Fig. 18), and in a diversity of ecologi-cal habitats, moulds are involved in food spoilage, con-taminate many manufactured materials such as wood, paper and textiles, and are frequent visitors to the human indoor environment. Some hyphomycetes are asexually reproducing parts of the life cycle of sexually competent ascomycetous and basidiomycetous fungi. The asexually, or mitotically, reproducing structures are called anamorphs; the sexu-ally reproducing, or meiotic, counterparts of the same life cycle are called teleomorphs. Together, these forms of sporulation make up a whole fungus, or holomorph (Fig. 4, Hennebert & Weresub 1977, Weresub & Hennebert 1979). In some cases, anamorph and teleomorph develop side-by-side, but more commonly they mature at dif-ferent times, or on different substrates. Historically, the establishment of anamorph–teleomorph connections was difficult. Although thousands of connections are known, the majority of hyphomycetes remain orphaned. It seems almost certain that many anamorphs have permanently lost the potential to mate or to develop a teleomorph, and must be regarded as anamorphic holomorphs. Mycolo-gists must classify such anamorphic holomorphs prag-matically, using whatever characters are available. Mycologists generally recognize three major groups of anamorphic fungi. None of these three groups is a homo-geneous, phylogenetically based taxon: they are poly-

Fig. 4. The first anamorph-teleomorph connection. A. Discovered by Anton de Bary (1831–1888) (public domain). B. Aspergillus anamorph (green) and Eurotium teleomorph (yellow) in one agar colony. C. Conidiophores of Aspergillus anamorph of Eurotium. D. Eurotium teleomorph, optical section of ascoma on left, ascospores in asci on right. The teleomorph and anamorph together comprise the holomorph.

What are hyphom

ycetes?

1We consider ‘mould’ the correct spelling for a fungus, reflecting its etymological origin from the English ‘moul’, different from ‘mold’, i.e. a container for making a shape.


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phyletic assemblages, with lower-case initial letters and not italicized, as indicated here: •blastomycetes: asexually reproducing yeasts, includ-ing some ascomycetes, i.e., Saccharomycetales, and some basidiomycetes (i.e., Urediniomycetes such as the Sporidiales, Ustilaginomycetes, and in the Agarico-mycetes the Tremellomycetidae such as the Filobasidi-ales). •coelomycetes: pycnidial and acervular fungi, most ascomycetous, a few basidiomycetous. •hyphomycetes: moulds lacking pycnidial or acervular fruiting-bodies (conidiomata), most ascomycetous, some basidiomycetous. This book deals only with the third group. Hypho-mycetes are defined as those usually hyphal anamorphs in which the conidia (mitospores) or other reproduc-tive bodies (bulbils, sclerotia) are developed ‘out in the open’. This is in contrast to the conidia of the coelo-mycetes, which develop inside an initially closed struc-ture. Anamorph genera based on sterile mycelia and on mycelia producing only bulbils or sclerotia are also often dumped into the hyphomycetes (as the ‘Agonomycetes’), faute de mieux. Some of these genera are listed in the Dictionary, although generic names applied to sterile thalli or mycelium of lichenized fungi are not included. The name Deuteromycota (or Deuteromycetes) was long used for the anamorphic fungi and is still regrettably common. We consider this term archaic and misleading, because it implies that hyphomycetes and coelomycetes represent a phylum equivalent to the other six. As Ken-drick (1981, 2005), Taylor (1995) and others insisted, this completely artificial construct should be abandoned. The anamorphs of the Phylum Zygomycota, and of the classes Urediniomycetes and Ustilaginomycetes (Phylum Basidiomycota), are traditionally referred to the appro-priate holomorphic taxa in these Phyla, and are not usu-ally included in the hyphomycetes although they could be mistaken for such based on their appearance. Because of morphological similarities between anamorphs of some Zygomycetes and some hyphomycetes, there are several genera that were originally described as members of one group but later shown to belong to the other. Ana-morphic forms of the Zygomycota are mentioned in this book when they are morphologically similar to hypho-mycetes. The morphological distinctions between the hypho-mycetes and the other two groups, blastomycetes and coelomycetes, are not absolute, and intermediates occur.

Anamorph genera such as Geotrichum, Hormonema, and Aureobasidium, are usually placed in the hyphomycetes but could almost equally well be considered blastomy-cetes. Except for a few mycelial yeasts such as Geotri-chum, we do not attempt to deal with the asexual yeasts (anamorphs of the Saccharomycetales and Sporidiales), whose reduced morphology often does not lend itself to micromorphological identification. They are mono-graphed in other publications (e.g., Barnett et al. 2000, Kurtzman et al. 2010). Although there are groups of ascomycetes with pre-dominantly hyphomycetous anamorphs and other groups with predominantly coelomycetous anamorphs, other groups exhibit both types of anamorph. Some species produce both hyphomycetous and coelomycetous syn-anamorphs (e.g., many strains of the hyphomycete Epi-coccum nigrum produce a coelomycetous synanamorph classified in Phoma). Thus, the difference between these two morphological groups is phylogenetically ambigu-ous, but for morphological identification purposes the distinction is critical. Pycnidial and acervular anamorphs are usually considered to belong to the coelomycetes while sporodochial anamorphs are classified in the hypho-mycetes. However, intermediate forms occur between these three types of conidiomata (see Kendrick & Nag Raj 1979). The coelomycetous conidiomata known as acervuli are defined partly on their location in host tis-sue. Such conidiomata either cannot be formed in agar culture (cf. Barron 1968, pp. vii and 6), or they instead produce discrete conidiophores or sporodochium-like conidiomata that are similar to hyphomycetes. In our list, we include less complete documentation for some genera with acervular conidiomata when we thought this might be useful (e.g., Colletotrichum), but we neither intended nor attempted to cover coelomycetes. An identification guide to genera and species of coelomycetes, with many illustrations, was published by Sutton (1980), and illus-trations of many species appear in the multi-authored Icones Generum Coelomycetum (Morgan-Jones et al. 1972–1981) and in the superb book by Nag Raj (1993) on coelomycetes with appendaged conidia.

C. Historical trends in hyphomycete taxonomy

As with most organisms, the first taxonomic research on hyphomycetes occurred in central and northern Europe, and was followed by explorations in North America. Although there were sporadic surveys in other parts of

Fig. 5. Hyphomycete colonies on plants. A. Monilinia fructigena brown rot of plums (B. Kendrick). B. Bud blast of Rhododendron caused by Seifertia azaleae (P. Crous). C. Ramularia diervillae leaf spots on Diervilla lonicera. D. Fusarium graminearum on maize (AAFC).

Wha

t are

hyp

hom

ycet

es?


7

Sordariomycetes

Leotiomycetes

Laboulbeniomycetes

Leotiomycetes

Geoglossomycetes

Lecanoromycetes

Eurotiomycetes

Lichinomycetes

Ascomycota

Pezizomycotina

Dothideomycetes

Arthoniomycetes

Pezizomycetes

Orbiliomycetes

Saccharomycetes

Taphrinomycotina

Saccharomycotina

Agaricomycetes Agaricomycotina

Basidiomycota

DacrymycetesTremellomycetesUstilaginomycetesExobasidiomycetes UstilaginomycotinaPucciniomycetesAgaricostilbomycetesMicrobotryomycetes

Pucciniomycotina

Fig. 6. Phylogenetic tree showing the relative arrangement of major classes and orders of the Dikarya, based on a multiple gene analysis. Taxa with a high concentration of hyphomycetous anamorphs are in green text. For all tree figures, methods and statistics are given in Appendix 5.

the globe, it was not until the mid-twentieth century that the moulds of the Indian subcontinent and then Japan and New Zealand were studied with any intensity. In the 1980s, many apparently endemic genera and species were described from the Caribbean (especially Cuba). In the 1990s, international interest in biodiversity stim-ulated studies of the hyphomycetes of Asia (especially Hong Kong and Thailand), South Africa, and in recent years China. As a consequence, we have the most com-prehensive knowledge of the hyphomycetes of the north-ern temperate regions and less developed understanding of many tropical and subtropical areas. Several regions are still essentially unexplored.

Most early mycologists, including Tode (1791, see Fig. 1B), Persoon (1801, Fig. 3A), Link (1809, Fig. 3C) and Fries (1832, Fig. 3B), included hyphomycetes in their classification systems. A remarkable degree of subtlety of description was achieved with a hand lens, and many of the genera and species proposed by these authors are still used today. The pioneer in the microscopic study of hyphomycetes was A.C.J. Corda (Fig. 3D), whose Icones Fungorum (Fig. 1D, 1837–1842), Prachtflora (Fig. 2, 1839), and contributions to Sturm’s Deutschlands Flora (1829–1837) were the first glimpses into the microfungal realm. Corda’s drawings and paintings of hyphomycetes and coelomycetes were sometimes fanciful but commu-

Fig. 7. Hyphomycete colonies on wood. A. Trichoderma sp. B. Bispora antennata (W. Gams). C. Cladosporium sp. D. Conoplea juniperi (K. Hodge, K. Loeffler).

Early hyphomycete taxonom

y

→Fig. 20


8

Fig. 8. Some conidiomatal types in hyphomycetes. A–F. Synnemata. A, B. Determinate with terminal, slimy conidial masses. C–E. Indeterminate with terminal and subterminal conidiogenous cells. F. Branched. G–J. Sporodochia in cross section. J. With cupulate hymenium, somewhat intermediate between typical sporodochia and acervular conidiomata.

nicated the complexity and beauty of these microfungi for the first time (Fig. 2). Preuss and Ditmar (in Sturm 1851, see Fig. 1C) improved the accuracy of the micro-scopic drawings. Then the Tulasne brothers (1861–1865) published magnificent microscopic illustrations prepared with a drawing prism (Fig. 1E), showing for the first time the physical connections between certain anamorphs and associated teleomorphs. Below, we focus on the two major historical devel-opments that continue to be relevant to modern hypho-mycete taxonomy, the sporological system developed by Saccardo and his followers, and the ontogenetic system developed by Hughes (1953) and subsequent workers.

1. The Saccardoan systemThe most successful early efforts in the taxonomy of hyphomycetes used characters that parallelled traits also

emphasized in teleomorph classifications. Saccardo (Fig. 3E) was an assiduous mycologist (1845–1920) who Lat-inized and compiled all published descriptions of fungi in a series called Sylloge fungorum hucusque cognitorum, which eventually spanned 26 volumes (some later vol-umes completed by his relatives or colleagues). Saccardo developed a system for classifying all fungi according to fruiting-body type, pigmentation, and spore morphology (the ‘sporological’ system). The taxonomic practice of establishing a rigid hierarchy of morphological charac-ters and then sorting organisms according to a strict inter-pretation of these phenotypes, regardless of whether the result is a phylogenetically natural classification, is often called ‘pigeon-holing’. Despite its artificiality, the practi-cality of Saccardo’s system made it very useful and it was applied by Clements & Shear (1931) in their influential book The Genera of Fungi. This system was also applied

Fig. 9. Sporodochial hyphomycetes. A. Bactridium flavum. B. Trichoderma spirale. C. Tubercularia vulgaris (yellow) and its teleomorph Nectria cinnabarina (red). D. Myrothecium inundatum.

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Dictionary of HypHomycete Genera


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63

The Genera of Hyphomycetes a-Ba

Abgliophragma R.Y. Roy & Gujarati 1966 –– Trans. Br. mycol. Soc. 49: 363 (363–365, Fig. 1, Pl. 13) / A. setosum R.Y. Roy & Gujarati 1966 = Wiesneriomyces Koord. 1907, fide Pirozynski 1972

Acalyptospora Desm. 1848 –– Annls Sci. nat. Bot., Sér. 3, 10: 342 (342–343, no illus.) / A. nervisequia Desm. 1848 = glandular leaf hairs, fide Mussat, Sacc. Syll. Fung. 15: 1, 1901

ACANTHELLORHIZA P. Roberts 1999 –– Rhizocto-nia-forming fungi, a taxonomic guide, p. 130 (no illus.) / A. globulifera P. Roberts 1999 no illus. Sterile mycelium with swollen, monilioid cells, and clamp connections. On wood: Europe, North America. One species. Teleo-morph: Heteroacanthella, fide Roberts 1999. notes: Anamorphic Basidiomycota (Heteroacanthel-laceae, Exidiales). Compare with Rhizoctonia and its segregates, in particular Opadorhiza, Oliveorhiza.

Acanthoderma Syd. & P. Syd. 1917 –– Annls mycol. 15: 234 (232–234, Fig. 2–4) / Acanthoderma memecyli Syd. & P. Syd. 1917 = Acanthoderma anamorph of Chaeto-placa memecyli Syd. & P. Syd. 1917 notes: The protologue described a phragmoconidial fungus with a thallus one cell thick and dark setae. The type has not been reexamined in modern times, and it is uncertain whether this name refers to a hyphomycete.

aCANTHODOCHIUM Samuels, J.D. Rogers & Nagas. 1987 –– Mycotaxon 28: 457 (453–459, Fig. 1 g–i, 3) / A. collodisculae Samuels, J.D. Rogers & Nagas. 1987 = Acanthodochium anamorph of Collodiscula japonica Hino & Katum. 1955 CDM: sporodochia or determinate synnemata, stroma-ta, dark. SET: none. CPH: branched, (sub)hyaline. CGC: sympodial, denticles, (sub)hyaline. CDA: amero, (sub)hyaline or brown, single, dry, schizo. pl. 150D. On bamboo litter (Phyllostachys, Pleioblas-tus, Sasa): Asia, North America. Three species. Teleo-morphs: Collodiscula, fide Samuels et al. 1987; Astro-cystis, Rosellinia, fide Ju & Rogers 1990. notes: Anamorphic Ascomycota (Xylariaceae, Xyla-riales). Compare with Amoenodochium, Dokmaia, Had-rotrichum, Hymenostilbe, Moelleroclavus. The unnamed anamorphs of Stilbohypoxylon spp. seem to be similar to Acanthodochium, cf. Rogers & Ju (1997), Petrini (2004). refs: Ju & Rogers, Mycologia 82: 342–349, 1990; Mycotax-on 73: 343–440, 1999 (teleomorphs). — Rogers & Ju, Mycol. Res. 101: 135–138, 1997 (as Stilbohypoxylon). — Hladki & Romero, Sydowia 55: 65–76, 2003 (as Stilbohypoxylon). — Petrini, Sydowia 56: 51–71, 2004 (revis. as Stilbohypoxylon).

aCAROCYBE Syd. 1937 –– Annls mycol. 35: 285 (285–286, no illus.) / A. hansfordii Syd. 1937 CDM: sporodochia or determinate synnemata, black, stromata. SET: none. CPH: branched, brown, monilioid.

CGC: monotretic, brown. CDA: (didymo) or phragmo, pale brown, rostrate, single, dry, schizo. pl. 225B. On leaves (Bridelia, Lychnodiscus): Africa, South America. Three species. notes: Compare with Acarocybiopsis. The synnema is comprised of a central, apically branched axis, with downward growing hyphae. refs: Ellis, Mycol. Pap. 76: 2–5 (1–36), 1960 (n. sp., key). — Ellis, Mycol. Pap. 82: 50–51 (1–55), 1961 (n. comb.). — Ellis, Demat. Hyphom., pp. 384–385, 1971 (illus.).

aCAROCYBELLA M.B. Ellis 1960 –– Mycol. Pap. 76: 5 (5–7, Fig. 3) / A. jasminicola (Hansf.) M.B. Ellis 1960 ≡ Cercospora jasminicola Hansf. 1944 CDM: none. SET: none. CPH: unbranched, brown. CGC: monotretic, brown. CDA: phragmo, scoleco, brown, rostrate, single, dry, schizo. pl. 241D. On leaves (Glyphaea, Jasminum, Phrygano-cydia): Africa, Caribbean. One species. notes: Compare with Alternaria. refs: Ellis, Demat. Hyphom., pp. 370–372, 1971 (illus.). — R.K. Verma et al., Forest Fungi of Central India, pp. 229–230, 2008 (illus.).

aCAROCYBELLINA Subram. 1992 –– Proc. Indian natn. Sci. Acad. B 58: 187 (179–190, no illus.) / A. aren-gae (Matsush.) Subram. 1992 ≡ Sporidesmium arengae Matsush. 1975 ≡ Brachysporiella arengae (Matsush.) Hol.-Jech. 1983 CDM: none. SET: none. CPH: unbranched or with 1–2 Y-shaped branches, brown. CGC: monoblastic, brown. CDA: phragmo, distoseptate, brown, single, dry, schizo. pl. 209a. On leaf litter (Arenga): Asia. One species. notes: A segregate from Sporidesmium characterized by both upward and downward, Y-shaped extensions of conidiophores emerging from spent conidiogenous loci, with hyphae growing downward from the branch point. Compare also with Veracruzomyces. refs: Matsushima, Ic. Microf. Mats. lect., p. 136, 1975 (Pl. 143, n. sp., as Sporidesmium). — Hughes, N. Z. Jl Bot. 16: 347–350 (311–370), 1978 (as Sporidesmium). — Holubová-Jechová, Česká Mykol. 37: 12–18, 1983 (n. comb., as Brachy-sporiella).

aCAROCYBIOPSIS J. Mena, A. Hern. & Mercado 1999 –– Mycol. Res. 103: 1032 (1032–1034, Fig. 1) / A. cubitaënsis J. Mena, A. Hern. & Mercado 1999 = Manoharachariomyces N.K. Rao, D.K. Agarwal & Kunwar, fide

Seifert, based on the protologue CDM: synnema-like, dark, with downward growing hyphae. SET: none. CPH: unbranched, nodose, brown. CGC: ?monoblastic, with a raised rim around the locus, brown. CDA: phragmo with a dark central band, brown, single, dry, schizo. pl. 206B. On dead plant material: Asia, Caribbean. Two species. notes: Compare with Acarocybe, Acarocybella. These unusual synnematous fungi produce one conidium at a time at the apex of the synnema, with downward grow-ing hyphae. The type of Manoharachariomyces, not yet transferred to Acarocybiopsis, is clearly congeneric, but appears to be a distinct species.


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64

a-B acaromyceS Boekhout, Scorzetti, Gerson & Sztejn-

berg 2003 –– In Boekhout, Theelen, Houbraken, Robert, Scorzetti, Gafni, Gerson & Sztejnberg, Int. J. syst. evol. Microbiol. 53: 1662 (1655–1664, Fig. 1 c, d) / A. ingoldii Boekhout, Scorzetti, Gerson & Sztejnberg 2003 CDM: none. SET: none. CPH: reduced, hyaline. CGC: sympodial, sometimes branched, conspicuous denticles, hyaline. CDA: scoleco or amero, hyaline, in short acro-petal chains, schizo. pl. 6c. On mites (Phyllocoptruta): Asia, Middle East. One species. ITS barcode: AY158671 (T). notes: Anamorphic Basidiomycota (Cryptobasi-diaceae, Ustilaginomycetes). Compare with Entylomella, Meira, Pseudozyma.

Acaulium Sopp 1912 –– Skr. VidenskSelsk. Christiania 11: 42 (42–53) / A. albonigrescens Sopp 1912 = Scopulariopsis Bainier 1907, fide Ellis 1971 notes: Considered an anamorph of Microascus by Emmons & Dodge (1931). Morton & Smith (1963) con-sidered A. albonigrescens unidentifiable; the type was not found. refs: Emmons & Dodge, Mycologia 23: 313–331, 1931 (revis., as Scopulariopsis). — Morton & Smith, Mycol. Pap. 86: 1–96, 1963 (revis.).

Acaulopage Drechsler 1935 –– Mycologia 27: 185 (177–205, Fig. 1) / Two original species Zygomycota, Zoopagales. notes: Endoparasites in amoebae and nematodes with conidia on short stalks, resembling a hyphomycete.

acepHaLa Grünig & Sieber 2005 –– Mycologia 97: 634 (628–640, Fig. 5 b, d) / A. applanata Grünig & Sie-ber 2005 Sterile mycelium. no illus. Associated with conifer roots: Europe. One named species. ITS barcode: AY078145 (T). notes: Anamorphic Ascomycota (Helotiales). Sterile pigmented mycelium, associated with conifer roots, phy-logenetically related to Phialocephala fortinii. See litera-ture under Phialocephala for more information.

Achitonium Kunze 1819 –– Flora, Jena 2: 49 (49–50, no illus.) / A. acicola Kunze 1819 = Chroostroma Corda 1837, fide Lindau 1910 = Leucosporium Corda 1837, fide Lindau 1910 ≡ Phymatostroma Corda 1837, nom. illegit. Art. 52 = Pactilia Fr. 1835, fide Lindau 1910 notes: The protologue described a gelatinous, orange, tuberculate fungus on needles of Pinus sylvestris. Its identity in modern terms is unknown. ref.: Lindau, Rabenhorst Krypt.-Fl., Ed. 2, Bd 1, Abt. 9: 456–457, 1910 (docum., as Pactilia).

Achorion Remak 1845 –– Diag. Pathol. Untersuch., p. 193 (193–215, no illus.) / A. schoenleinii Remak 1845 = Oidium schoenleinii Lebert 1845 ≡ Trichophyton schoen-leinii (Lebert) Langeron & Miloch. ex Nann. 1934 ≡ Grubyella M. Ota & Langeron 1923, nom. illegit. Art. 52 = Trichophyton Malmsten 1845, fide Emmons 1934, Ajello 1968

notes: Achorion was widely used for several species of dermatophytes (e.g., Dodge, Medical Mycology, 1935) that are now included in Trichophyton.

Aciculariella G. Arnaud 1954 –– Bull. trimest. Soc. mycol. Fr. 69: 298 (298–301, Fig. 16 o–q) / A. lasio-sphaeriae G. Arnaud 1954 nom. inval. Art. 36 notes: Perhaps similar to Mirandina or Microdochium. ref.: Mouchacca & Samson, Revue Mycol. 37: 267–275, 1973 (type, discussion).

aciDomyceS B.J. Baker, M.A. Lutz, S.C. Dawson, P.L. Bond & J.F. Banfield 2004 ex Selbmann, de Hoog & De Leo 2008 –– In Selbmann, de Hoog, Zucconi, Isola, Ruisi, Gerrits van den Ende, Ruibal, de Leo, Urzi & Ono-fri, Stud. Mycol. 61: 16 (16–20, Fig. 8) / A. acidophilus (Sigler & J.W. Carmich.) Selbmann, de Hoog & De Leo 2008 ≡ Scytalidium acidophilum Sigler & J.W. Carmich. 1974 = Acidomyces B.J. Baker, M.A. Lutz, S.C. Dawson, P.L. Bond & J.F.

Banfield 2004, nom. inval., Arts 36, 37 CDM: none. SET: none. A-anamorph: CPH = CGC: thallic-arthric, brown. CDA: amero or didymo, brown, in chains, dry, schizo. no illus. One species: Europe, North America. ITS barcode: AJ244237. notes: Anamorphic Ascomycota (Teratosphaeriaceae, Capnodiales). Compare with Elasticomyces, Friedman-niomyces, Scytalidium. refs: Sigler & Carmichael, Can. J. Microbiol. 20: 267–268, 1974 (n. sp., as Scytalidium). — Sigler & Carmichael, Mycotax-on 4: 399–403 (349–488), 1976 (as Scytalidium). — Baker et al., Appl. envir. Microbiol. 70: 6264–6271, 2004 (nom. inval.). — Crous et al., Stud. Mycol. 58: 1–32, 2007 (LSU).

Acinula Fr. 1822 –– Syst. mycol. 2: 267 (267–268) / A. candicans Fr. 1822 notes: The name refers to sclerotia of uncertain iden-tity in modern terms.

Acladium Link 1809 –– Mag. Ges. naturf. Freunde, Ber-lin 3: 11 (Tab. I, Fig. 13) / A. conspersum Link 1809, lectotype fide Clements & Shear 1931 = Haplotrichum Link 1824 : Fr., fide Holubová-Je-chová 1976 notes: Name not sanctioned by Fries. ref.: Donk, Taxon 11: 76 (75–104), 1962 (nomenclature).

Acmosporium Corda 1839 –– Ic. Fung. 3: 12 (12–13, Tab. ii, Fig. 32) / A. botryoideum Corda 1839 notes: Hughes (1958) suggested a synonymy with Aspergillus but the illustration is more suggestive of Bo-trytis, fide Seifert.

Acontiopsis Negru 1961 –– Comun. Acad. Repub. Pop. Rom. 11: 839 (839–842, Figs 1–3) / A. crataegi Negru 1961 nom. inval. Art. 37 ?= Cylindrocladiella Boesew. 1982, fide Crous 2002


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The Genera of Hyphomycetes a-BAcontium Morgan 1902 –– J. Mycol. 8: 4 (4–5, no illus.) /

A. album Morgan 1902, lectotype fide Clements & Shear 1931 notes: Application uncertain. The protologue described a corticolous fungus with scolecosporous, aseptate, slimy, hyaline conidia 18–25 × 1 µm.

Acremoniella (Sacc.) Sacc. 1886 –– Syll. Fung. 4: 302 (illus. Fungi Italici del., no. 713, 1881) / A. atra (Corda) Sacc. 1886, lectotype fide Clements & Shear 1931 ≡ Acremonium atrum Corda 1837 = Harzia acremonioides (Harz) Costantin 1888 nom. illegit. Art. 52 = Harzia Costantin 1888 notes: Saccardo included the type species of Monoto-spora Corda 1837, fide Holubová-Jechová (1974). No type specimen of Acremonium atrum is available.

aCREMONIULA G. Arnaud ex Cif. 1962 –– Atti Ist. bot. Univ. Lab. Crittog. Pavia, Ser. 5, 19: 85 (85–86, no illus.) / A. suprameliola Cif. 1962 = A. sarcinellae (Pat. & Har.) G. Arnaud ex Deighton 1969 CDM: none. SET: none. CPH = CGC: monoblastic (sometimes sympodial), hyaline. CDA: amero, brown, single, dry, schizo. pl. 23B. On fungi (Asterina, Clypeolella, Echidnodes, Schiffnerula), leaves (Eugenia, Ocotea, Quercus) or branches (Fagus): Cosmopolitan. Four species. Synana-morph (hypho): Trichoconis, fide Hughes 1979. notes: Compare with Humicola. Ciferri’s type is lost, fide Hughes (1979). For A. fagi see Rhinocladium. refs: Arnaud, Bull. trimest. Soc. mycol. Fr. 69: 268–269, 1954 (Fig. 1v, nom. inval. Art. 36). — Deighton, Mycol. Pap. 118: 2–5 (1–41), 1969 (illus.). — Ellis, Demat. Hyphom., pp. 57–58, 1971 (illus.). — Ellis, More demat. Hyphom., pp. 36–37, 1976 (n. sp). — Hughes, N. Z. Jl Bot. 17: 165–166 (139–188), 1979 (n. comb., synanamorph). — Mercado et al., Mycotaxon 55: 491–499, 1995 (n. sp.).

aCREMONIUM Link 1809 : Fr. –– Mag. Ges. naturf. Freunde, Berlin 3: 15 (no illus.) : Fr., Syst. mycol. 1: xliv. 1821 / A. alternatum Link 1809 : Fr. 1832, lectotype fide Clements & Shear 1931 = Cephalosporium auct., non Corda, fide Gams 1971 ?= Cylindrophora Bonord. 1851, fide de Hoog 1978 = Gliomastix Guég. 1905, fide Gams 1971, but considered distinct

by Lechat et al. 2010 = Torulina Sacc. & D. Sacc. 1906, fide Dickinson 1968 (as Gliomas-

tix), Gams 1971 = Mastigocladium Matr. 1911, fide Gams 1971 = Andreaeana Palm & Jochems 1924, fide Gams ≡ Andreaea Palm & Jochems 1923, nom. illegit. Art. 53 ≡ Palmomyces Maire 1926, nom. illegit. Art. 52 = Basitorula G. Arnaud 1954, fide Dickinson 1968 (as Gliomastix),

Gams 1971 = Haplotrichella G. Arnaud 1954, fide Gams = Nematomyces Faurel & Schotter 1965, fide Gams 1971 = Pseudofusidium Deighton 1969, fide Gams 1971 = Sagrahamala Subram. 1972, fide Gams CDM: none. SET: none. CPH: unbranched or spar-ingly branched, hyaline or = cgc. CGC: phialides, hya-line (or pigment encrusted). CDA: amero (or didymo), hyaline, slimy or in basipetal chains, schizo. figs 1c, 18c, D, 27c, 100, pl. 39a. On many substrata: Cosmopolitan. More than one hundred species

(many undescribed or lacking anamorph epithets). Teleomorphs: Bulbithecium, fide Udagawa & Muroi 1990; Emericellopsis, fide Gams 1971; Cosmospora, Hydropisphaera, Ijuhya, Lasionectria, Nectriopsis, fide Rossman et al. 1999; Mycoarachis fide Malloch & Cain 1970; Nectriella, fide Lowen 1989, Rossman et al. 1999; acremonium-like anamorphs are also reported in Nectria, Mycocitrus, Peloronectriella, Hypocrea, Trichosphaerella, fide Gams 1971 (pp. 19–21); Hapsidospora, Nigrosabulum, fide Malloch & Cain 1970; Neocosmospora, fide von Arx 1974; Protocrea, fide Mercuri & Ranalli 1976, Rossman et al. 1999; Scopinella, fide Tsuneda & Hiratsuka 1981; Thielavia, fide Samson et al. 1977, Morgan-Jones & Gams 1982; Heleococcum, fide Tubaki 1967, Udagawa et al. 1995, Rossman et al. 1999; Pronectria, fide Lowen & Hawksworth 1986, Lowen 1995; Chaetom-ium, fide Morgan-Jones & Gams 1982; Hypomyces, fide Rogerson & Samuels 1993; Phytocordyceps, fide Shu & Wang 1986; Valetoniellopsis, fide Samuels & Barr 1997; Dimerosporiella, Ochronectria, Protocreopsis, Stilbocrea, Trichonectria, Valsonectria, Pseudohypocrea, fide Rossman et al. 1999; Sporophagomyces, fide Põldmaa et al. 1999; Neobarya, fide Candoussau et al. 2007; Verrucostoma, fide Hirooka et al. 2010; Rodentimyces, fide Doveri et al. 2010. Synanamorphs of acremonium-like fungi (hypho): Botryotrichum, Humicola, Trichocladium, fide Hennebert In Kendrick 1971; Diademospora, fide Söderstrom & Bååth 1979; Geminoarcus, fide Ando 1993; Stilbella, fide Seifert 1985; Ventrographium, fide Upadhyay et al. 1986; Volutella, fide Matsushima 1989; Mahabalella, Phialophaeoisaria, fide Matsushima 1995; Isthmolongispora, fide Descals 1997; Koorchaloma, fide Matsushima 2001; chlamydospores, fide de Hoog & Guarro 1995. ITS barcode: AY566992 (CBS 223.70). notes: Anamorphic Ascomycota (Bionectriaceae, Hypocreales), but acremonium-like anamorphs occur in several orders of Ascomycota. Compare with Apha-nocladium, Gabarnaudia, Haptocillium, Lecanicillium, Monilochaetes, Monocillium, Phaeoacremonium, Phia-losimplex, Pseudogliomastix, Sagenomella, Simplicil-lium, Stanjemonium, Taifanglania, Verticillium. Grass endophytes formerly classified in sect. Albo-lanata (teleomorphs: Epichloë) are now in Neotyphodium. Fur-ther synonyms are listed under Gliomastix. For A. falci-forme see Fusarium, for A. lactucae see Plectosporium. major refs: Gams, Nova Hedwigia 16: 141–145, 1968 (lectotype). — Gams, Cephalosporium-artige Schimmelpilze, 262 pp., 1971 (monogr., key). molecular systematics: Glenn et al., Mycologia 88: 369–383, 1996 (SSU). — Vicente et al., Mycol. Res. 103: 1173–1178, 1999 (RAPD A. cucurbitacearum). — Rossman et al., Mycologia 93: 100–110, 2001 (LSU). — Stchigel et al., Mycol. Res. 106: 975–983, 2002 (ITS, as Thielavia). — Summerbell et al., Stud. Mycol. 68: 141–164, 2011 (revis., LSU). — Kiyuna et al., Mycoscience 52: in press, 2011 (ITS, n. sp., n. comb.). new species and new combinations: Brown & Kendrick Trans. Br. mycol. Soc. 41: 499–500, 1958 (as Gliomastix). — Hughes & Dickinson, N. Z. Jl Bot. 6: 106–114, 1968 (n. sp., as Gliomastix). — Matsushima, Microf. Solomon Isl., pp. 11, 27–28, 1971 (Figs 115–117, 118:1, 121:2, Pl. 22: 2, as Cepha-


66


66

a-B

losporium, Gliomastix). — Tichelaar, Acta bot. neerl. 21: 197–199, 1972. — Gams & Lacey, Trans. Br. mycol. Soc. 59: 519–522, 1972. — Tubaki, Mycologia 65: 938–941, 1973 (see A. tubakii). — Morgan-Jones, Can. J. Bot. 52: 429–431, 1974. — Gams, Trans. Br. mycol. Soc. 64: 389–404, 1975. — Subra-manian, Kavaka 5: 93–98, 1977 (n. combs to Sagrahamala). — Purkayastha & Mallik, Nova Hedwigia 30: 869–872, 1978. — Smith & Davidson, Can. J. Bot. 57: 2122–2139, 1979. — Matsushima, Mats. Mycol. Mem. 4: 2, 9, 1985 (Figs 272, 273, 275, partly as Gliomastix). — Brady, Trans. Br. mycol. Soc. 87: 486–487, 1986 (cf. Monilochaetes). — Castañeda, Fungi Cub. 2: 2–3, 1987 (Fig. 1). — Iwatsu et al., Mycotaxon 32: 439–445, 1988 (as Phialophora = A. atrogriseum, fide Gams). — Matsu-shima, Mats. Mycol. Mem. 6: 2–4, 1989 (Figs 490–496, 498, P-16–26). — Okada et al., Trans. mycol. Soc. Japan 34: 171–185, 1993. — Matsushima, Mats. Mycol. Mem. 7: 3, 42, 1993 (Figs 583–585, P-126). — Samuels & Rogerson, Mycolo-gia 85: 248, 1993 (n. comb.). — Möller & Gams, Mycotaxon 48: 441–450, 1993. — Matsushima, Mats. Mycol. Mem. 8: 13–14, 51, 1995 (partly as Gliomastix, synanamorphs, many illustra-tions). — Ragazzi et al., Eur. J. For. Path. 26: 235–243, 1996. — Alfaro-García et al., Mycologia 88: 804–808, 1996. — Ito et al., Mycol. Res. 104: 77–80, 2000. — Roquebert & Dupont, Mycotaxon 75: 349–356, 2000. — Watanabe, Mycoscience 42: 591–595, 2001. — Matsushima, Mats. Mycol. Mem. 10: 3–5, 22–25, 2003 (synanamorph: Koorchaloma). — Sigler et al., Stud. Mycol. 50: 409–413, 2004 (ITS, BenA). — Zuccaro et al., Stud. Mycol. 50: 283–297, 2004 (ITS, BenA). — Weisenborn et al., Nova Hedwigia 90: 457–468, 2010 (rDNA, Plectosphaerel-laceae). teleomorphs: Tubaki, Trans. mycol. Soc. Japan 8: 5–10, 1967. — Malloch & Cain, Can. J. Bot. 48: 1815–1825, 1970. — Samuels, Mycologia 65: 401–420, 1973. — Mercuri & Ranalli, Physis B. Aires 35: 303–317, 1976. — Samuels, N. Z. Jl Bot. 14: 231–260, 1976 (key). — Samson et al., Trans. Br. mycol. Soc. 69: 417–423, 1977. — Tsuneda & Hiratsuka, Can. J. Bot. 59: 1192–1195, 1981 (teleomorph). — Morgan-Jones & Gams, Mycotaxon 15: 311–318, 1982 (n. spp., n. sect. for anamorphs of Chaetomium, Thielavia). — Ueda & Udagawa, Mycotaxon 16: 387–395, 1983 (n. sp.). — Shu & Wang, Mycotaxon 26: 337–344, 1986. — Lowen & Hawksworth, Lichenologist 18: 321–328, 1986. — Udagawa et al., Mycotaxon 33: 291–301, 1988. — Lowen, Mem. N. Y. Bot. Gdn 49: 243–252, 1989. — Udagawa & Muroi, Bull. natn. Sci. Mus. Tokyo, 16: 13–19, 1990. — Rogerson & Samuels, Mycologia 85: 248 (231–272), 1993 (key, n. comb.). — Samuels & Brayford, Sydowia 46: 75–161, 1994 (n. sp., synonymy, key to species with Nectria teleomorphs). — Lowen, Mycotaxon 53: 81–95, 1995 (n. sect., key to lichenicolous species). — Udagawa et al., Mycoscience

36: 37–43, 1995. — Pfenning, Sydowia 47: 65–69, 1995. — Samuels & Barr, Can. J. Bot. 75: 2165–2176, 1997. — Rossman et al., Stud. Mycol. 42: 1–248, 1999. — Põldmaa et al., Can. J. Bot. 77: 1756–1768, 1999. — Stchigel et al., Mycologia 95: 1218–1226, 2003 (as Thielavia). — Nong & W.Y. Zhuang, Fungal Diversity 19: 95–107, 2005 (Cosmospora). — Samuels et al., Hypocreales of the SE United States, pp. 20–27, 61, 80, 108, 130, 2006 (Cosmospora, Hydropisphaera, Nectriopsis, Neocosmospora). ― Candoussau et al., Sydowia 59: 179–215, 2007 (Neobarya). — Jaklitsch et al., Mycologia 100: 962–984, 2008 (Protocrea, rDNA). — Hirooka et al., Mycoscience 49: 281–290, 2008 (Cosmospora). ― Lechat, Bull. trimest. Soc. mycol. Fr. 124: 1–5, 2008 (Lasionectria). ― Lechat & Baral, Oest. Z. Pilzk. 17: 15–24, 2008 (Ijuhya). ― Hirooka et al., Mycologia 102: 418–429, 2010 (Verrucostoma). — Lechat et al., Mycotaxon 111: 95–102, 2010 (Hydropisphaera). ― Doveri et al., Fungal Diversity 42: 57–69, 2010 (Rodentimy-ces). — Lechat& Courtecuisse, Mycotaxon 113: 443–447, 2010 (Ijuhya). — Lechat et al., Persoonia 25: 120–121, 2010 (Fungal Planet 50, Nectriella). compilations: Barron, Gen. Hyphom. Soil, pp. 38, 114–116, 179–181, 1968 (as Cephalosporium and Gliomastix). — Subra-manian, Hyphomycetes, pp. 628–633, 1971 (key, as Cepha-losporium). — Hawksworth, Bull. Br. Mus. nat. Hist. (Bot.) 6: 192–195, 1979 (key to lichenicolous spp., nom. nov.). — Domsch et al., Compendium of Soil Fungi, pp. 16–29, 272–274, 1980; 2nd ed. pp. 30–38, 2007 (key to soil-borne spp., docum., partly as Emericellopsis). — Ellis & Ellis, Microf. Land Plants, pp. 293, 311, 484, 512, 1987 (repr. 1997) (Figs 1295, 1358, 1921, as Gliomastix). — Wang & Zabel, Fungi from Utility Poles, pp. 232–233, 1990 (as Gliomastix). — Moubasher, Soil fungi in Qatar and other Arab countries, pp. 27–29, 1993 (illus.). — de Hoog & Guarro, Atlas clin. Fungi, pp. 130–132, 370–384, 1995; 2nd ed., pp. 395–419, 2001 (medically import-ant spp.). — Tzean et al., Atlas of Entomopathogenic Fungi from Taiwan, pp. 90–93, 1997 (two misidentified spp., fide Gams). — Guarro et al., Clin. infect. Dis. 25: 1222–1229, 1997 (medically relevant species). ― Ellis & Ellis, Microf. on Misc. Substr., 2nd ed., pp. 48, 168, 169, 1998 (Figs 540, 541, as Gliomastix). — Samson et al., Food- & Airborne Fungi, 6th ed., pp. 58–61, 2000 (key). — Scholz, Schriftenr. Veg.Kunde 31, 298 pp., 2000 (lichenicolous spp.). — Y.Z. Wang et al., Mycosystema 21: 192–195, 2002 (key to spp. from China). — R.K. Verma et al., Forest Fungi of Central India, pp. 52–53, 2008 (illus.). other refs: Hennebert, In Kendrick (ed.), Tax. Fungi Imperfecti, pp. 202–223, 1971 (synanamorphs). — Raitviir & Tarasov, Mikol. Fitopatol. 9: 203–208, 1975 (analysis of section classification). — de Hoog, Persoonia 10: 33–81, 1978

fig. 100. Acremonium, variation in conidiophore branching and pigmentation. A. Conidia in chains. B. Conidia in slimy heads. C. Dark conidia in chains. D. Cymose branching. E. Didymoconidia.


67

The Genera of Hyphomycetes a-B(synonymy). — Söderstrom & Bååth, Trans. Br. mycol. Soc.

72: 340–342, 1979 (synanamorph). — von Arx, Gen. Fungi Sporul. Pure Cult. 3rd ed., pp. 259–261, 1981 (Fig. 68a). — Matsushima, Mats. Mycol. Mem. 3: 10, 1983 (Figs 152, 153, as Gliomastix). — Hawksworth, CMI Descr., no. 502, 1976; Booth, ibid. no. 624, 1979; Williams, ibid. nos 741–742, 1983; 931–935, Mycopathologia 100: 169–178, 1987 (illus.). — von Arx, Beih. Nova Hedwigia 87: 232, 1987 (no illus.). — Ando, Trans. mycol. Soc. Japan 34: 109–121, 1993 (synanamorph). — Descals, Mycotaxon 63: 431–466, 1997 (synanamorph). — Zuccaro et al., Mycol. Res. 107: 1451–1466, 2003 (18S, 28S probes). — Fernando et al., Mycotaxon 92: 323–338, 2005 (illus.). — Labuda, Czech Mycol. 57: 239–248, 2005 (illus.). — Onofri et al., Continental Antarctic Fungi, pp. 104–106, 2007 (Antarctic spp.).

Acrocladium Petr. 1949 –– Sydowia 3: 263 (263–264, no illus.) / A. andinum Petr. 1949 ≡ Periconiella andina (Petr.) Arx 1981 = Periconiella Sacc. 1885, fide von Arx 1981 ref.: Petrak, Sydowia 4: 568–569, 1950 (n. sp., no illus.).

aCROCONIDIELLA J.C. Lindq. & Alippi 1964 –– Darwiniana 13: 612 (610–613, Figs 1–6) / A. tropaeoli (T.E.T. Bond) J.C. Lindq. & Alippi 1964 ≡ Heterospo-rium tropaeoli T.E.T. Bond 1947 = Heterosporium Klotzsch ex Cooke 1877, fide von Arx 1983, but

considered distinct by David 1997, Crous & Braun 2003 (see Cla-dosporium)

CDM: none. SET: none. CPH: unbranched or sparing-ly branched, sympodial, brown. CGC: mono- or polytre-tic, scars, terminal and intercalary in cph, brown. CDA: phragmo, brown, single, dry, schizo. pl. 216a. On leaves (Eschscholtzia, Tropaeolum): Cosmopolitan. Three species. notes: Compare with Curvularia. For A. gracilis see Cladosporium. refs: Ellis, CMI Descr., no. 161, 1968 (illus.). — Ellis, Demat. Hyphom., pp. 461–462, 1971 (illus.). — Ellis, More demat. Hyphom., pp. 406–408, 1976 (n. comb.). — Mucho-vej, Mycologia 72: 1045–1047, 1980 (n. sp.). — Ellis & Ellis, Microf. Land Plants, p. 439, 1987 (repr. 1997) (Fig. 1696).

aCROCONIDIELLINA M.B. Ellis 1971 –– Mycol. Pap. 125: 22 (22–27, Fig. 20) / A. loudetiae M.B. Ellis 1971 CDM: none. SET: unbranched, brown. CPH: unbranched, brown. CGC: mono- or polytretic, scars, terminal and intercalary in cph, brown. CDA: didymo, brown, single, dry, schizo. pl. 165c. On leaves (Areca, Chloris, Cynodon, Loude-tia, Trichachne): Pantropical. Four species. Teleomorph: Zeuctomorpha, fide Sivanesan 1984. Synanamorph: Sclerotia. notes: Anamorphic Ascomycota (Pleosporaceae, Pleosporales). Compare with Acroconidiella. refs: Ellis, More demat. Hyphom., pp. 408–411, 1976 (key). — Sivanesan, Bitun. Ascomycetes, pp. 572–574, 1984 (teleo-morph). — Mercado et al., Hifom. demac. Cuba enterobl., pp. 22–25, 1997 (key).

Acrocylindrium Bonord. 1851 –– Handbuch allg. Mykol., p. 97 (97–98, Taf. vi, Fig. 147, Taf. viii, Fig. 172) / Sporo-trichum cylindrosporum Link 1824, lectotype fide Pfeiffer

1873 = Acrocylindrium roseum (Ehrenb.). Bonord. 1851 nomen dubium, fide Gams & Hawksworth, Kavaka 3: 57–60, 1975 notes: No type material of Bonorden’s material is preserved. For A. oryzae see Sarocladium. The later lec-totypification with A. elegans Bonord. by Clements & Shear (1931) is superfluous. ref.: Pfeiffer, Nomencl. bot. 1: 35, 1873 (lectotypification).

Acrodesmis Syd. 1926 –– Annls mycol. 24: 424 (424–426, no illus.) / A. cestri Syd. 1926 = Periconiella Sacc. 1885, fide Ellis 1967

Acrodictiella R.K. Verma, N. Sharma & Soni 2008 –– In R.K. Verma, N. Sharma, K.K. Soni & Jamaluddin, Forest Fungi of Central India p. 230 (230–233, Figs 327–331) / A. indica R.K. Verma, N. Sharma & Soni 2008 nom. illegit. Art. 53 [non Acrodictyella W. A. Baker & Partr. 2001 (hyphomycetes)]

aCRODICTYELLA W. A. Baker & Partr. 2001 –– In Baker, Partridge & Morgan-Jones, Mycotaxon 78: 30 (29–35, Fig. 1) / A. obovata W. A. Baker & Morgan-Jones 2001 CDM: none. SET: none. CPH: unbranched, brown. CGC: percurrent, brown. CDA: dictyo, hyaline when released, brown when mature, single, dry, schizo. pl. 276D. On wood: North America. One species. notes: Compare with Acrodictys, Ityorhoptrum, June-wangia, Pseudoacrodictys, Rhexoacrodictys.

aCRODICTYOPSIS P.M. Kirk 1983 –– Mycotaxon 18: 260 (260–264, Fig. 2) / A. lauri P.M. Kirk 1983 CDM: none. SET: none. CPH: unbranched, brown. CGC: monoblastic or percurrent, brown. CDA: dictyo or helico, brown with a darker basal cell, single, dry, schizo. pl. 290a. On leaves (Laurus): Europe. One species. notes: Compare with Digitoramispora, Zalerion. ref.: Ellis & Ellis, Microf. Land Plants, p. 582, 1987 (repr. 1997) (Fig. 2153).

aCRODICTYS M.B. Ellis 1961 –– Mycol. Pap. 79: 5 (5–7, Fig. 3) / A. bambusicola M.B. Ellis 1961 CDM: none. SET: none. CPH: unbranched, percurrent, brown. CGC: monoblastic, brown. CDA: dictyo, brown, single, dry, schizo. pl. 276a. On stems, bark and wood: Mostly pantro-pical, some temperate. About 38 species in the broad sense, or about 10 species in the restricted sense. Synana-morphs (hypho): idriella- or selenosporella-like fide Mat-sushima 1975; chlamydospores, fide Matsushima 1981; Dactylaria fide Matsushima 1985. notes: Compare with Acrodictyella, Bhatia, Chei-roidea, Cubasina, Junewangia, Piricauda, Pithomyces, Pseudoacrodictys (for species with appendaged conidia), Rhexoacrodictys, Synnemacrodictys. refs: Ellis, Mycol. Pap. 79: 1–23, 1961 (n. spp.); ibid. 93: 24–25 (1–33), 1963 (n. sp.). — Sutton, Can. J. Bot. 47: 853–858, 1969 (n. spp., key). — Ellis, Demat. Hyphom., pp. 128–131, 1971 (illus.). ― Narendra & V.G. Rao, Kavaka 1: 47–49, 1973 (n. sp.). — Matsushima, Ic. Microf. Mats. lect., pp. 3–4, 1975 (Pl. 148–150). — Ellis, More demat. Hyphom.,


68


68

a-B pp. 101–103, 1976 (key). — Lunghini & Rambelli, G. bot. ital.

112: 175–195, 1978 (illus.). — Hughes, N. Z. Jl Bot. 17: 185 (139–188), 1979 (exclusions). — Matsushima, Mats. Mycol. Mem. 2: 5–6, 1981 (Figs 67, 68, synanamorph). — Wang & Sutton, Mycologia 74: 489–500, 1982 (n. sp.). — Matsushima, Mats. Mycol. Mem. 3: 2, 1983 (n. sp., Fig. 203). — Holubová-Jechová, Česká Mykol. 37: 12–18, 1983 (n. var.). — Holub-ová-Jechová & Mercado, Česká Mykol. 38: 96–120, 1984 (n. sp., in part as Piricauda). — Matsushima, Mats. Mycol. Mem. 4: 2, 7–8, 1985 (Figs 249, 250, synanamorph). — Holubová-Jechová & Mercado, Česká Mykol. 40: 143–146, 1986 (illus.). — Matsushima, Mats. Mycol. Mem. 5: 2, 1987 (Fig. 435). — Mel′nik & Popushoi, Nesoversh. Griby na drevesn. kustarnik, pp. 12–15, 1992 (illus.). — Matsushima, Mats. Mycol. Mem. 7: 3, 42, 1993 (n. sp., Figs 537–541, P-216, 217, 221). — Sutton, Mycol. Pap. 167: 8 (1–93), 1993 (descr.). — Whitton et al., Fungal Diversity 4: 159–169, 2000 (n. spp., key). — Mel’nik, Opredelitel’ Gribov Rossii, Kl. Hyphomycetes, Vyp. 1, pp. 31–34, 2000 (key). — Baker et al., Mycotaxon 81: 293–319, 2002 (emend.). — Cai et al., Nova Hedwigia 75: 525–532, 2002 (n. sp.). — Mercado et al., Mycol. Res. 109: 723–728, 2005 (n. comb.). — Manoharachary et al., Indian Phytopathology 59: 91–93, 2006 (n. spp.). — Kodsueb et al., Cryptog. Mycol. 27: 111–119, 2007 (n. sp.). — Castañeda et al., Mycotaxon 102: 91–99, 2007 (n. sp.). — T. Zhang et al., Fl. Fung. Sinicorum 31: 21 (1–231), 2009 (n. comb.).

aCRODONTIELLA U. Braun & Scheuer 1996 (‘1995’) –– Sydowia 47: 146 (146–149, Fig. 1) / A. fallopiae U. Braun & Scheuer 1996 CDM: none. SET: none. CPH: unbranched or sparing-ly branched, hyaline. CGC: sympodial, minutely denticu-late. CDA: amero or didymo, hyaline, single, dry, schizo. pl. 94B. On leaves (Fallopia): Europe. One species. notes: A hyaline counterpart to Acrodontium. Com-pare with Sporothrix. ref.: Braun, Monogr. Cercosporella, vol. 2, p. 414, 1998 (additions).

aCRODONTIUM de Hoog 1972 –– Stud. Mycol. 1: 23 (23–27, Fig. 9) / A. crateriforme (F.H. Beyma) de Hoog 1972 ≡ Chloridium crateriforme F.H. Beyma 1933 CDM: none. SET: none. CPH: unbranched or branched or verticillate, (sub)hyaline or = cgc. CGC: sympodial, minutely denticulate, rachis, (sub)hyaline. CDA: amero, hyaline, dry, single, schizo. pl. 94a. On wood, bark and leaves, fungi (rusts) and slime moulds (Stemonitis): Cosmopolitan. About 10 species. Teleomorphs: Amplistroma, Wallrothiella, fide Huhndorf et al. 2009; Ascocorticium, fide Oberwinkler et al. 1967. ITS barcode: AY843112 (TRN 267). notes: Anamorphic Ascomycota (Amplistroma-taceae). Similar to Tritirachium, but with brown coni-diophores, with a less sinuous rachis. Compare also to Acrodontiella, Beauveria, Tritirachium. For A. album see Engyodontium. refs: Oberwinkler et al., Nova Hedwigia 14: 283–289, 1967 (Fig. 98, 99, as Rhinotrichella, teleomorph). — de Hoog & Rao, Persoonia 8: 207–212, 1975 (n. sp.). — Cole & Samson, Patterns Dev. conid. Fungi, p. 36–37, 1979 (conidiogenesis). — von Arx, Gen. Fungi Sporul. Pure Cult. 3rd ed., pp. 314, 319, 1981 (Fig. 86 a). — Crane & Schoknecht, Trans. Br. mycol. Soc. 79: 345–347, 1982 (n. sp., SEM, cf. Engyodontium). — Kirk, Trans. Br. mycol. Soc. 81: 401–404, 1983 (n. sp.). — Ellis & Ellis, Microf. Land Plants, p. 592, 1987 (repr. 1997) (Fig.

2185). — Cabello, Mycotaxon 36: 91–94, 1989 (n. sp.). — Wang & Zabel, Fungi from Utility Poles, pp. 174–175, 1990 (illus.). — Mel’nik & Popushoi, Nesoversh. Griby na drevesn. kustarnik, pp. 14–16, 1992 (illus.). — Mel’nik, Opredelitel’ Gribov Rossii, Kl. Hyphomycetes, Vyp. 1, pp. 34–36, 2000 (illus.). — Kubátová et al., Czech Mycol. 53: 237–255, 2001 (illus.). — Fernando et al., Mycotaxon 92: 323–338, 2005 (illus.). — Huhndorf et al., Mycologia 101: 904–919, 2009 (teleomorphs, LSU).

aCROGENOSPORA M.B. Ellis 1971 –– Demat. Hyphom., p. 114 (114–115, Fig. 75 a) / A. sphaeroce-phala (Berk. & Broome) M.B. Ellis 1971 ≡ Monotospora sphaerocephala Berk. & Broome 1859 ≡ Monotospora Sacc. 1880, nom. illegit. Art. 53 [non Monoto-

spora Corda 1837], fide Mason 1941 ?= Monotosporella S. Hughes 1958, cf. Hughes 1979 CDM: none. SET: none. CPH: unbranched, brown. CGC: integrated, percurrent, brown. CDA: amero, brown, single, dry, schizo. pl. 113B. On wood, branches and other fungi (Graphium, rhizomorphs): Pantemperate and subtropical. Nine species. Teleomorph: Farlowiella, fide Mason 1941 (as Monotospora), Ellis 1971, 1976. Synanamorph (hypho): phialides, fide Arnold 1990. LSU: GU301791 (CBS 164.76). notes: Anamorphic Ascomycota (basal to Gloniaceae, Dothideomycetes). Compare with Domingoëlla, Monoto-sporella, Endophragmiella. refs: Mason, Mycol. Pap. 5 (Annot. Account, List II, Fasc. 4): 103–113, 1941 (as Monotospora). — Ellis, Mycol. Pap. 131: 5–6 (1–25), 1972 (n. comb.). — Hammill, Can. J. Bot. 50: 581–585, 1972 (TEM conidiogenesis, as Monotosporella). — Matsushima, Ic. Microf. Mats. lect., p. 4, 1975 (Pl. 114). — Ellis, Demat. Hyphom., pp. 74, 1976 (illus.). — Hughes, N. Z. Jl Bot. 16: 312–317 (311–370), 1978 (n. spp.). — Cole & Samson, Patterns Dev. conid. Fungi, p. 76–78, 1979 (illus. conidiogenesis). — Hughes, N. Z. Jl Bot. 17: 184–186, 1979 (discussion of Monotosporella). — Minter et al., Trans. Br. mycol. Soc. 79: 75–93, 1982 (conidiogenesis). — Sivanesan, Bitun. Ascomycetes, pp. 280–282, 1984 (teleomorph). — Ellis & Ellis, Microf. Land Plants, pp. 28, 47, 1987 (repr. 1997) (Figs 102, 168). — Arnold, Wiss. Z. Friedr.-Schiller-Univ. Naturw. R. 39: 353–357, 1990 (phialidic synanamorph). — Mel’nik & Popushoi, Nesoversh. Griby na drevesn. kustarnik, pp. 16–18, 1992 (illus.). — Muntañola-Cvetković et al., Revta catal. Micol. 20: 63–84, 1997 (illus.). — Goh et al., Mycol. Res. 102: 1309–1315, 1998 (revis., key, n. sp.). — Mel’nik, Opredeli-tel’ Gribov Rossii, Kl. Hyphomycetes, Vyp. 1, pp. 36–38, 2000 (key). — H. Zhu et al., Mycotaxon 92: 383–386, 2005 (n. sp.). — Hu et al., Sydowia 62: 191–203, 2010 (n. sp., key).

aCROPHIALOPHORA Edward 1961 (‘1959’) –– Mycologia 51: 784 (782–784, Figs 1, 2) / A. nainiana Edward 1961 = A. fusispora (S.B. Saksena) M.B. Ellis 1971 CDM: none. SET: none. CPH: verticillate or branched, brown. CGC: phialides, (sub)hyaline. CDA: amero, (sub)hyaline, in basipetal chains, schizo. pl. 56c. On plants and soil: Cosmopolitan. Three spe-cies. ITS barcode: DQ648458/9 (IHEM 19209). notes: Anamorphic Ascomycota (Chaetomiaceae, Sordariales). Compare with Sagenomella, Phialosim-plex, Phialomyces, and genera of the Verticillium com-plex.


69

The Genera of Hyphomycetes a-B refs: Samson & Mahmood, Acta. bot. neerl. 19: 804–808,

1970 (n. sp., key). — Ellis, Demat. Hyphom., pp. 533–534, 1971 (illus.). — Subramanian, Hyphomycetes, pp. 605–606, 1971 (as Paecilomyces). — Domsch et al., Compendium of Soil Fungi, pp. 30–31, 1980; 2nd ed. p. 39, 2007 (docum.). — von Arx, Gen. Fungi Sporul. Pure Cult. 3rd ed., pp. 263–264, 1981 (Fig. 69f). — Kirk, IMI Descr., no. 1051, Mycopathologia 115: 131–132, 1991 (illus.). — Moubasher, Soil fungi in Qatar and other Arab countries, pp. 30–31, 1993 (illus.). — de Hoog & Guarro, Atlas clin. Fungi, pp. 386–387, 1995; 2nd ed., pp. 420–421, 2001 (medically important spp.).

aCROPHRAGMIS Kiffer & Reisinger 1970 –– Revue Écol. Biol. Sol 7: 16 (12–16, Fig. 1, Pl. i, Fig. 1) / A. coronata Kiffer & Reisinger 1970 CDM: none. SET: none. CPH: unbranched, brown. CGC: integrated, percurrent, brown. CDA: phragmo or stauro, with a darker central cell and hyaline polar cells or multiple protuberances, single, dry, schizo. pl. 205B. On branches and litter: Cosmopolitan. Three species. notes: Compare with Arachnophora, Uberispora. For A. laevispora see Endophragmiella. refs: Ellis, More demat. Hyphom. p. 107–108, 1976 (illus.). — Vasant Rao & Reddy, Indian J. mycol. Res. 16: 365 (361–369), 1978 (n. sp.). — Hughes, Fungi Can., no. 143, 1979 (n. sp.). — Hughes, N. Z. Jl Bot. 17: 180–181 (139–188), 1979 (discussion). — Vasant Rao & de Hoog, Stud. Mycol. 28: 10–11 (1–83), 1986 (illus.). — W. Wu & W. Zhuang, Sporidesmium, Endophragmiella and related genera from China, pp. 295–297, 2005 (illus.).

aCROSPEIRA Berk. & Broome 1857 –– In Berkeley, Introduction to Cryptogamic Botany, p. 305 (305–306, Fig. 69) / A. mirabilis Berk. & Broome 1857 = Spirospora L. Mangin & Vincens 1920, fide Hughes 1958 CDM: none. SET: none. CPH: reduced, branched, (sub)hyaline. A-anamorph: CGC: monoblastic, (sub)hya-line. CDA: dictyo or helico, brown, single, dry, rhexo. B-anamorph: CGC: phialides, hyaline. CDA: amero, hyaline, in basipetal chains, dry, schizo. fig. 101, pl. 154c. On nuts (Castanea): Europe. One species. Synanamorph (hypho): proteophiala-like fide Barron 1968, von Arx 1981. notes: Anamorphic Ascomycota (Ceratostomataceae, Hypocreales). Compare with Chlamydomyces, Arachno-phora, Tetracoccosporium. refs: Wiltshire, Trans. Br. mycol. Soc. 21: 232–237 (211–239), 1938 (illus.). — Barron, Gen. Hyphom. Soil, pp. 87–88, 1968 (illus.). — Ellis, Demat. Hyphom., pp. 70–71, 1971 (illus.). — von Arx, Gen. Fungi Sporul. Pure Cult. 3rd ed., pp. 338–339, 1981 (Fig. 93b). — Ellis & Ellis, Microf. Land Plants, p. 106, 1987 (repr. 1997) (Fig. 443).

Acrospira Mont. 1857 –– Annls Sci. nat. Bot., Sér. 4, 8: 299 (299–301, no illus.) / A. crouanii Mont. 1857 notes: The protologue described helical conidia joined in spiral chains. Its identity in modern terms is unknown.

Acrosporella Riedl & Ershad 1978 (‘1977/78’) –– Sydowia 29: 166 (166–169, Fig. 4) / A. sarmentorum Riedl & Ershad 1978 = Cladosporium Link 1815 : Fr., fide U. Braun (pers. comm., W!)

Acrosporium Bonord. 1851 –– Handbuch allg. Mykol., p. 80 (Taf. iv, Fig. 91) / A. tenue Bonord. 1851 nom. illegit. Art. 53 [non Acrosporium Nees 1816 (hyphomycetes)]

Acrosporium Nees 1816 –– Das System der Pilze und Schwämme, p. 53 (Pl. 4) / A. monilioides Nees 1816 ≡ Oidium monilioides (Nees) Link 1824 = Acrosporium anamorph of Blumeria graminis (DC.) Speer 1975 nom. rej. = Oidium Link 1824, nom. cons. ref.: Subramanian, Hyphomycetes, pp. 832–841, 1971 (key in the sense of Oidium Link).

ACROSTALAGMUS Corda 1838 –– Ic. Fung. 2: 15 (Tab. x, Fig. 66) / A. cinnabarinus Corda 1838, lectotype fide Clements & Shear 1931 = A. luteoalbus (Link : Fr.) Zare, W. Gams & Schroers 2004 = Verticillium Nees 1816, fide Hughes 1951 but because of newly

conserved type, considered distinct by Zare et al. 2004 CDM: none, or pale, determinate synnemata. SET: none. CPH: verticillate or penicillate, brown or orange. CGC: phialides, hyaline or brown. CDA: amero, hyaline, slimy, schizo. pl. 51a. On plant debris and in soil: Cosmopolitan. Two species. ITS barcode: AJ292420 (IMI 182719). notes: Anamorphic Ascomycota (Plectosphaerella-ceae, Glomerellales). Compare with Phaeostalagmus and members of the Verticillium complex. Acrostalag-mus, long considered a synonym of Verticillium, is now recognized for phylogenetic and morphological reasons.

fig. 101. Acrospeira mirabilis, development of conidia.


70


70

a-B See notes under Verticillium.

refs: Hughes, Mycol. Pap. 45: 1–18 (1–36), 1951 (illus.). — Barron, Gen. Hyphom. Soil, pp. 38, 321–323, 1968 (illus.). — Ellis, Demat. Hyphom., pp. 535–536, 1971 (illus.). — Subra-manian, Hyphomycetes, pp. 648–650, 1971 (key to Indian spp.). — Domsch et al., Compendium of Soil Fungi, pp. 501–503, 1980; 2nd ed. pp. 39–40, 2007 (docum.). — Zare et al., Mycol. Res. 108: 576–582, 2004 (revis., ITS). — Pantou et al., Mycol. Res. 109: 889–902, 2005 (multigene). — Onofri et al., Conti-nental Antarctic Fungi, p. 95, 2007 (Antarctic sp.). — Réblová et al., Stud. Mycol. 68: 143–171, 2011 (multigene, n. comb.).

Acrostaphylus G. Arnaud ex Subram. 1956 –– J. Indian bot. Soc. 35: 483 (483–486, no illus.) / A. hypoxyli G. Arnaud 1954, nom. inval. Art. 36 = Nodulisporium Preuss 1849, fide Barron 1968, Petrini & Müller 1986 refs: Arnaud, Bull. trimest. Soc. mycol. Fr. 69: 269–272, 1954 (nom. inval., Fig. 1n). — Subramanian, Hyphomycetes, pp. 418–420, 1971 (key to Indian spp.).

aCROSTAURUS Deighton & Piroz. 1972 –– Mycol. Pap. 128: 94 (94–96, Fig. 49) / A. turneri Deighton & Piroz. 1972 CDM: none. SET: none. CPH: unbranched, brown. CGC: percurrent, brown. CDA: stauro, brown, single, dry, schizo. pl. 368a. On fungi (Asterina, Cirsosia): Africa, Asia. One species. notes: Compare with Acrophragmis, Uberispora. ref.: Ellis, More Demat. Hyphom., pp. 108–109, 1976 (illus.).

aCROSTROMA Seifert 1987 –– Can. J. Bot. 65: 2196 (2196–2201, Figs 12–20) / A. annellosynnema Seifert 1987 = Acrostroma anamorph of Batistia annulipes (Mont.) Cif. 1958 CDM: determinate synnemata, dark, annellated, stro-mata. SET: none. CPH: branched or penicillate, brown. CGC: phialides, brown. CDA: amero, pale brown, basi-petal chains, dry, schizo. pl. 89c. On bark: Neotropics. One species. Teleo-morph: Batistia, fide Samuels & Rodrigues 1989. Syn-anamorph (hypho): phialophora-like, fide Samuels & Rodrigues 1989. LSU: AY346262 (G.J.S. 6059). notes: Anamorphic Ascomycota (Batistiaceae). Com-pare with Pesotum, Stromatographium. refs: Samuels & Rodrigues, Mycologia 81: 52–56, 1989 (teleomorph). — R.K. Verma et al., Forest Fungi of Central India, pp. 121–124, 2008 (n. spp., probably misclassified fide Seifert).

Acrothamnium Nees 1816 : Fr. –– Das System der Pilze und Schwämme, p. 74 (74–75, Pl. 5) : Fries, Syst. mycol. 1: xlvi, 1821 / A. violaceum Nees 1816, lectotype fide Donk 1962 nomen dubium, fide Donk, Taxon 11: 103–104, 1962

Acrotheca Fuckel 1860 –– Jb. Nassau. Ver. Naturk. 15: 42 (42–43, no illus.) / A. gei Fuckel 1860 = Acrotheca anamorph of Depazea geicola Fuckel 1870 = Ramularia Unger 1833, fide Hughes 1958, Braun 1990

notes: For A. acuta see Myrmecridium, for A. aquaspersa see Rhinocladiella.

Acrotheciella Koord. 1907 –– Verh. K. Ned. Akad. Wet. Amsterdam, Sect. 2, 13: 250 (250–251, Fig. 61) / A. javanica Koord. 1907 ?= Curvularia Boedijn 1933, cf. Mason 1928 notes: Mason (1928) considered Acrotheciella a pos-sible home for the fungus now known as Curvularia lunata, but rejected it because the protologue lacked criti-cal details. Boedijn (1933) also considered Acrotheciella before describing Curvularia. refs: Mason, Mycol. Pap. 2: 8 (1–43), 1928 (discussion). — Boedijn, Bull. Jard. Bot. Buitenz. III, 13: 123–132, 1933 (discussion).

Acrothecium (Corda) Corda 1851 –– apud Preuss In Sturm, Deutschl. Fl. III (Pilze), Bd 6, Heft 30: 85 (Tab. 43) / A. parasitans (Corda) Corda ex Streinz 1862 ≡ Tri-chothecium parasitans Corda 1838 ≡ Trichothecium subgen. Acrothecium Corda, Ic. Fung. 2: 10,

1838 nomen dubium, fide Hughes 1958 notes: In the sense of Preuss, perhaps similar to Dac-tylaria, and in the sense of Corda, perhaps similar to Fusarium solani. For A. globosum see Sterigmatobotrys, for A. lunatum see Curvularia. ref.: Preuss, Linnaea 24: 110–111, 1851 (n. sp.).

Actiniceps Berk. & Broome 1876 –– J. Linn. Soc., Lon-don 15: 85 (Tab. ii, Fig. 3) / A. thwaitesii Berk. & Broome 1876 = Dimorphocystis Corner 1950 (Basidiomycota), fide Boedijn 1959, Persoonia 1: 12–13, 1959

Actinochaete Ferro 1907 –– Nuovo Giorn. bot. ital. 14: 232 (232–234, Tav. iii, Fig. 8) / A. arachnoidea Ferro 1907 = nomen confusum, fide M.B. Ellis in Dict. Fungi, 10th ed., based on a mixture of Aspergillus and Septobasidium

aCTINOCLADIUM Ehrenb. 1819 : Fr. –– Jb. Ge-wächsk. 1: 52 (51–53, Fig. 1d) : Fries, Syst. mycol. 3: 352, 1832 / A. rhodosporum Ehrenb. 1819 : Fr. CDM: none. SET: none. CPH: unbranched, brown. CGC: monoblastic or percurrent, brown. CDA: stauro, 2–3 arms up, brown, single, dry, schizo. pl. 346B. On wood and bark: Cosmopolitan. Six spe-cies. notes: Compare with Triposporium, Ceratosporella. refs: Ellis, Demat. Hyphom., pp. 137–138, 1971 (illus.). — Hughes, N. Z. Jl Bot. 16: 317–318 (311–370), 1978 (illus.). — Ellis & Ellis, Microf. Land Plants, p. 48, 1987 (repr. 1997) (Fig. 169). — Castañeda, Fungi Cub. 3: 2, 1988 (n. sp., Fig. 1, as Ceratosporella). — Mel’nik & Popushoi, Nesoversh. Griby na drevesn. kustarnik, pp. 17–19, 1992 (illus.). — Matsushima, Mats. Mycol. Mem. 7: 43, 1993 (n. sp., Figs 733, 735, P-262). — Sutton, Mycol. Pap. 167: 8, 1993 (notes). — Castañeda et al., Mycotaxon 60: 275–281, 1996 (n. comb. from Cerato-sporella). — Muntañola-Cvetković et al., Revta catal. Micol. 20: 63–84, 1997 (illus.). — Zhao & Li, Mycosystema 16: 270–273, 1997 (n. sp.). — Mel’nik, Opredelitel’ Gribov Rossii, Kl. Hyphomycetes, Vyp. 1, pp. 39–40, 2000 (illus.). — Wu &


853

Plate 369. A. Stauriella aquatica. B. Phaeocandelabrum elegans.C. Candelabrum spinulosum. D. Glomopsis corni.

DC

Intermediate betw

een stauro and bulbils

BA

cf. 30B

Scf. 274A

cf. 284C, 358D


856

Plate 372. A. Burgoa verzuoliana. B. Minimedusa polyspora. C. Badarisama sojae. D. Papulaspora sepedonioides.

A

C

B

Bul

bils

, ± d

ark,

glo

bose

D

cf. 361D

Glycine


857

Plate 373. A. Beverwykella pulmonaria. B. Ramicephala sphaerospora. C. Aegerita candida. D. Aegeritina tortuosa. C, D with habit.

AB

Bulbils, (sub)globose, spaces betw

een interior cells

B

C D

D


858

D

Plate 374. All aeroaquatic (exc. A). A. Clathroconium arachnicola (with monilioid resting cells). B. Clathrosphaerina zalewskii. C. Clathrosporium intricatum. D. Dendroclathra caeruleofusca.B

ulbi

ls, c

lath

roid

, bra

nche

d fr

om b

ase

A B

C

spiders


859

Plate 375. All aeroaquatic (exc. D). A. Pseudoclathrosphaerina evamariae. B. Sympodioclathra globosa. C. Polyancora globosa. D. Pseudogaster singularis (with habit).

DC

D

B

Bulbils, clathroid, branched from

centre

BA

Scf. 329C


866

A

Plate 382. A. Peyronelina glomerulata. B. Nidulispora quadrifida. C. Akenomyces costatus (with habit and anatomical details). D. Megacapitula villosa.

B

C D

Bul

bils

, out

er h

ypha

e cu

rved

cf. Kalamarospora


867

Plate 383. A. Tretopileus sphaerophorus (hyphae with clamp connections). B. Decapitatus flavidus. C. Mycopappus alni. D. Piricaudilium lobatum (with hyaline cells from interior). A, B with habit.

C

A B

Bulbils or ‘gem

mae’, m

isc.

B

D

D

Coffea

cf. 358B, D