Volume 1 Â· No. 2 Â· December 2010 V o lu m e 1 Â· N o ... - IMA Fungus

T h e G l o b a l 

M y c o l o g i c a l J o u r n a l 

Volume 1 · No. 2 · December 2010 

N E W S · R E P O R T S · R E S E A R C H N E W S · A R T I C L E S · C O R R E S P O N D E N C E 

S O C I E T Y A N D A S S O C I A T I O N N E W S · B O O K N E W S · f o r t h c o m i n g M E E T I N G S

Colofon 

IMA Fungus 

Compiled by the International 

Mycological Association for the 

world’s mycologists. 

Scope: All aspects of pure and 

applied mycological research and 

news. 

Aims: To be the flagship journal 

of the International Mycological 

Association. IMA FUNGUS is 

an international, peer-reviewed, 

open-access, full colour, fast-track 

journal. 

Frequency: Published twice per year 

(June and December). Articles are 

published online with final pagination 

as soon as they have been 

accepted and edited. 

ISSN 

E-ISSN 

2210-6340 (print) 

2210-6359 (online) 

Websites: www. imafungus.org 

www.ima-mycology.org 

E-mail: d.hawksworth@nhm.ac.uk 

Volume 1 · No. 2 · December 2010 

Cover: Sporophores of Armillaria 

mellea in Kirstenbosch Botanical 

Garden, Cape Town, South Africa 

EDITORIAL BOARD 

Editor-in-Chief 

Prof. dr D.L. Hawksworth CBE, Departamento de Biologia Vegetal II, Facultad de Farmacia, Universidad Complutense de 

Madrid, Plaza Ramon y Cajal, 28040 Madrid, Spain; and Department of Botany, Natural History Museum, Cromwell 

Road, London SW7 5BD, UK; E-mail: d.hawksworth@nhm.ac.uk 

Layout Editors 

M.J. van den Hoeven-Verweij & M. Vermaas, CBS-KNAW Fungal Biodiversity Centre, P.O. Box 85167, 3508 AD 

Utrecht, The Netherlands; E-mail: m.verweij@cbs.knaw.nl 

Associate Editors 

Dr T.V. Andrianova, M.G. Kholodny Institute of Botany, Tereshchenkivska Street 2, Kiev, MSP-1, 01601, Ukraine; 

E-mail: tand@darwin.relc.com 

Prof. dr D. Begerow, Lehrstuhl für Evolution und Biodiversität der Pflanzen, Ruhr-Universität Bochum, Universitätsstr. 

150, Gebäude ND 44780, Bochum, Germany; E-mail: dominik.begerow@rub.de 

Dr S. Cantrell, Department of Plant Pathology and Crop Physiology, Louisiana State University, Agricultural Centre, 455 Life 

Sciences Bldg., Baton Rouge, LA 70803, USA; E-mail: scantrel@suagm.edu 

Prof. dr D. Carter, Discipline of Microbiology, School of Molecular Biosciences, Building G08, University of Sydney, 

NSW 2006, Australia; E-mail: d.carter@mmb.usyd.edu.au 

Prof. dr P.W. Crous, CBS-KNAW Fungal Biodiversity Centre, P.O. Box 85167, 3508 AD Utrecht, The Netherlands; E- 

mail: p.crous@cbs.knaw.nl 

Prof. dr J. Dianese, Departamento de Fitopatologia, Universidade de Brasília, 70910-900 Brasília, D.F., Brasil; E-mail: 

jcarmine@unb.br 

Dr P.S. Dyer, School of Biology, Institute of Genetics, University of Nottingham, University Park, Nottingham NG7 2RD, 

UK; E-mail: paul.dyer@nottingham.ac.uk 

Dr M. Gryzenhout, Dept. of Plant Sciences, University of the Free State, P.O. Box 339, Bloemfontein 9300, South Africa; 

E-mail: Gryzenhoutm@ufs.ac.za 

Prof. dr L. Guzman-Davalos, Instituto de Botánica, Departamento de Botánica y Zoología, Universidad de Guadalajara, 

A.P. 1-139 Zapopan, 45101, México; E-mail: lguzman@cucba.udg.mx 

Dr K. Hansen, Kryptogambotanik Naturhistoriska Riksmuseet, Box 50007, 104 05 Stockholm, Sweden; E-mail: karen. 

hansen@nrm.se 

Prof. dr K.D. Hyde, School of Science, Mae Fah Luang University, Tasud, Chiang Rai, Thailand; E-mail: kdhyde3@gmail. 

com 

Prof. dr L. Lange, Dean of Research, Aalborg University, Denmark; E-mail: lla@adm.aau.dk 

Prof. dr L. Manoch, Department o Plant Pathology, Faculty of Agriculture, Kasetsart University, Bangkok 10900, Thailand; 

E-mail: agrlkm@ku.ac.th 

Prof. dr W. Meyer, Molecular Mycology Research Laboratory, CIDM, ICPMR, Level 3, Room 3114A, Westmead Hospital, 

Darcy Road, Westmead, NSW, 2145, Australia; E-mail: w.meyer@usyd.edu.au 

Dr D. Minter, CABI Bioservices, Bakeham Lane, Egham, Surrey, TW20 9TY, UK; E-mail: d.minter@cabi.org 

Dr L. Norvell, Pacific Northwest Mycology Service, LLC, 6720 NW Skyline Boulevard, Portland, Oregon 97229-1309, 

USA; E-mail: llnorvell@pnw-ms.com 

Dr G. Okada, Microbe Division / Japan Collection of Microorganisms, RIKEN BioResource Center, 2-1 Hirosawa, Wako, 

Saitama 351-0198, Japan; E-mail: okada@jcm.riken.jp 

Prof. dr N. Read, Fungal Cell Biology Group, Institute of Cell and Molecular Biology, Rutherford Building, University of 

Edinburgh, Edinburgh EH9 3JH, UK; E-mail: nick@fungalcell.org 

Prof. dr K.A. Seifert, Research Scientist / Biodiversity (Mycology and Botany), Agriculture & Agri-Food Canada, K.W. 

Neatby Bldg, 960 Carling Avenue, Ottawa, ON, K1A OC6, Canada; E-mail: seifertk@agr.gc.ca 

Prof. dr J. Taylor, Department of Plant and Microbial Biology, University of California, 111 Koshland Hall, Berkeley, CA 

94720, USA; E-mail: jtaylor@berkeley.edu 

Prof. dr M.J. Wingfield, Forestry and Agricultural Research Institute (FABI), University of Pretoria, Pretoria 0002, South 

Africa; E-mail: mike.wingfield@fabi.up.ac.za 

Prof. dr W.-Y. Zhuang, Systematic Mycology and Lichenology Laboratory, Institute of Microbiology, Chinese Academy of 

Sciences, Beijing 100080, China; E-mail: zhuangwy@sun.im.ac.cn 

 

i m a f U N G U S

IMA vision 

My first opportunity to speak to the world’s mycological community through this column must begin with sincere 

thanks to the Past President of the International Mycological Association, Pedro Crous of the Centraalbureau voor 

Schimmelcultures and the Universities of Utrecht and Wageningen in the Netherlands. Under President Crous the scope 

of the IMA was broadened well beyond its traditional focus on the quadrennial International Mycological Congress while, 

at the same time, sponsoring a most memorable IMC9 in Edinburgh. Regarding IMC9, thanks are especially due to the 

IMC9 organizing committee, consisting of chair Nick Read and members Simon Avery, Nick Clipson, Geoff Gadd, and 

Neil Gow. Thanks also are due the Elsevier group, led by Nina Cosgrove. I also want to thank Karen Hansen for agreeing 

to serve a second term as IMA Treasurer and Dominik Begerow for accepting the task of IMA Secretary. 

EDITORIAL 

Over the past four years, the 

foundation for a reinvigorated 

IMA was laid in the form of new 

statutes, which were ratified at the General 

Assembly of the IMC in Edinburgh and which 

can be viewed at the IMA website using this 

URL: . The new statutes are more explicit 

than the old about membership in the IMA, 

about election of IMA officers and members 

of the IMA Executive Committee, about 

proposals to host future IMCs, and about 

awards made by the IMA. 

Given that the mission of the IMA is “the 

promotion and encouragement of mycology 

throughout the world,” the IMA website, 

initiated by previous IMA officers, including 

Gioconda San Blas, Trond Schumacher and 

Meredith Blackwell, has assumed increased 

importance. The site 

has become essential for coordinating the 

activities of the six IMA Regional Mycological 

Member Organizations, which represent 

Africa, Asia, Australasia, Europe, North 

America and South America. 

A very significant contribution to the 

mission of global mycology is the transfer of 

responsibility for MycoBank from CBS to 

the IMA. MycoBank () 

aims to host nomenclatural, genetic and 

phenotypic data for all described fungi and 

to make that information freely available to 

all of science. The IMA membership clearly 

appreciates the worth of MycoBank, having 

voiced 86 % support that “Deposition of key 

nomenclatural information in one or more 

approved depositories (e.g., MycoBank) 

should be made mandatory for the valid 

publication of new fungal names.” 

A second, significant contribution is 

IMA Fungus, which you, dear reader, clearly 

appreciate. Deep thanks are due to Pedro 

Crous and IMA Fungus Editor David 

Hawksworth, and to the members of the IMA 

Executive Council, who now also wear the 

hats of Associate Editors of IMA Fungus. 

The third major contribution concerns 

IMA awards. IMC9 witnessed the resurrection 

of the two awards traditionally made by 

the IMA, but absent since 1996 -- the De 

Bary Medal for outstanding career research, 

awarded at IMC9 to Franz Oberwinkler, 

and the Ainsworth Medal for extraordinary 

service to world mycology, awarded at IMC9 

to both Emory Simmons and Richard Korf. 

In addition, 2010 saw the initiation of a new 

set of awards, the IMA Young Mycologist 

Awards, designed to honor the research 

accomplishments of young mycologists from 

each of the six regions. The first awards, which 

we had hoped to make in time for IMC9, will 

be determined over the next year and awarded 

in 2014, along with the second set of IMA 

Young Mycologist Awards to be determined 

early in 2014. The announcement about the 

IMA Young Mycologist Awards is in this 

issue of IMA Fungus and can also be found at 

. 

Mention of 2014 brings us to the final 

contribution, which is centered on selection 

of Thailand as the site for IMC10, with Leka 

Manoch as chair of the IMC10 Organizing 

Committee. To establish continuity between 

IMCs, the new statues tap the experience of the 

past chair of the IMC organizing committee 

by appointing this person as one of two IMA 

Vice-Presidents, the other being the chair of the 

next IMC organizing committee, in this case, 

Nick Read and Leka Manoch, respectively. 

To further encourage continuity, one to three 

members of the IMA executive council will 

serve on the IMC10 program committee along 

with mycologists from host nation and region. 

Although the focus of the IMA remains 

the quadrennial IMC, the meeting now 

shares the stage with the other contributions 

that the IMA makes to global mycology, 

through the IMA website, MycoBank, IMA 

Fungus, and support provided by the IMA to 

Regional Mycological Member Organizations. 

Clearly, the tasks and opportunities have 

grown beyond what can be attended to solely 

by the officers and Executive Committee of 

the IMA. Over the next four years, to ensure 

that the innovations made over the past four 

years succeed, I will be calling on the Regional 

Mycological Member Organizations and 

their members to work for the IMA in all 

of these areas. I certainly hope that you, the 

membership, will help us move the IMA and 

mycology forward to 2014. 

John Taylor President, IMA 

(Berkeley, CA, USA; jtaylor@berkeley.edu) 

v o l u m e 1 · n o . 2 

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News 

The home stretch for fungal barcoding 

In 2009, the plant research community published 

a paper (Hollingsworth et al. 2009) 

that designated two genes as a barcode for 

all plants. This means that among the major 

groups of eukaryotes, only the Fungi remain 

without a formal barcode. This may come 

as a surprise to many mycologists, who continue 

to loosely apply the term ‘barcode’ in 

numerous papers and projects. Importantly, 

barcodes are intended to be used as identification 

markers and may have properties, 

such as unalignable hypervariable regions, 

that make them inappropriate as phylogenetic 

markers. A strict set of criteria for barcodes 

has been established () 

and GenBank will only apply barcode tags 

to sequences of the genes or loci accepted as 

barcodes for a particular taxonomic group 

by the Consortium for the Barcode of Life 

(CBOL). Despite its long history of DNA 

sequencing and diverse ecological interests, 

the mycological community has not engaged 

with CBOL and associated organisations 

with much enthusiasm. At two major 

conferences in 2010 (the Mycological Society 

of America in Lexington, Kentucky, and 

the International Mycological Congress in 

Edinburgh), we suggested that it is in mycology’s 

best interest to engage and collaborate 

with the international barcode community, 

and judging by the response after these sessions, 

this message was received clearly by a 

significant number of mycologists. 

The road towards adoption of a fungal 

barcode has already had numerous twists 

and turns (Seifert 2009). An important 

milestone was a meeting of 37 mycologists 

from 12 countries at Front Royal, Virginia, 

in May 2007. At this meeting, participants 

were unanimous that the 5.8S nuclear ribosomal 

gene and its two flanking spacer 

regions (ITS) was the most likely candidate 

for a universal fungal barcode, despite some 

promising but mixed results for the default 

barcode sanctioned by CBOL, COI (Seifert 

et al. 2007). For a gene other than COI to 

be granted status as a barcode, CBOL has 

established a set protocol for validation. 

For mycologists, this process includes the 

publication of a proposal comparing the ITS 

against a set of other candidates. We intend 

to publish a paper in 2011 and together 

with an international group of interested 

mycologists, we are gathering and generating 

data for this purpose (Eberhardt 2010). 

Until now we received commitments from 

colleagues covering more than 40 clusters 

of five or more sibling species, covering 12 

of the main fungal lineages identified in 

phylogenies generated from ‘Assembling the 

Fungal Tree of Life’ (AFTOL) project. The 

final paper will probably involve more than 

50 coauthors from more than 12 countries 

and we intend to keep participation open to 

all interested parties until the data gathering 

deadline is passed in February 2011. More 

details can be found at the discussion group 

‘Fungi’ on connect.barcodeoflife.net and the 

project website(). 

As we set our sites on this publication, it 

is also important to look at projects and opportunities 

that may follow. We hope that the 

energy now building within the community 

will accelerate and improve collaborations 

between like-minded researchers across 

national borders. One pressing need is to 

coordinate and accelerate ongoing efforts to 

barcode authoritatively identified specimens 

and type cultures in herbaria and culture collections. 

Equally critical is coping with the 

vast amount of sequences generated by environmental 

sequencing projects. 

Making a dent in this mountain of 

Fig 1. A. Zygospores of Zygorhynchus heterogamous. B. The yeast Cryptococcus macerans. C. Basidiomata of Hygrocybe cantharellus. D. Basidiomata of Tremiscus 

helvelloides. E. Xanthoria parietina on Walter Gams’ roof in Bomarzo, Italy. F. Helicoconidia of Helicomyces roseus. G. Tar spot of sugar maple caused by Rhytisma 

acerinum. H. Morchella elata ready for further processing. Photos: Keith Seifert. 

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i m a f U N G U S

undescribed fungal diversity will require a 

monumental community effort to update 

and curate our sequence databases with accurate 

sequences tied to well validated samples; 

barcoding will be central to this. In addition, 

barcoding should provide excellent opportunities 

to expand the boundaries of fungal 

taxonomy by engaging teachers and amateur 

groups to help collect, locate and identify 

samples that could eventually be identified 

or validated by barcoding. Much is possible, 

but first we urgently need to formally declare 

a fungal barcode. We must not let this opportunity 

slip away to finally clear this hurdle. 

Conrad Schoch and Keith Seifert 

(schoch2@ncbi.nlm.nih.gov; and keith. 

seifert@agr.gc.ca) 

NEWS 

Eberhardt U (2010) A constructive step towards selecting a DNA barcode for fungi. New Phytologist 187: 266–268. 

Hollingsworth PM, Forrest LL, Spouge JL, Hajibabaei M, Ratnasingham S, et al. (2009) A DNA barcode for land plants. Proceedings of the National Academy of Sciences, 

USA 106: 12794–12797. 

Seifert KA (2009) Progress towards DNA barcoding of fungi. Molecular Ecology Resources 9: 83–89. 

Seifert KA, Samson RA, Dewaard JR, Houbraken J, Levesque CA, et al. (2007) Prospects for fungus identification using C01 DNA barcodes, with Penicillium as a test 

case. Proceedings of the National Academy of Sciences, USA 104: 3901–3906. 

Fungi and the Convention on Biological Diversity 

Recognizing that fungi are not plants, but need their own conservation strategies, is a first step to proper 

protection for species like critically-endangered Pleurotus nebrodensis, the only mushroom currently on the 

IUCN red-data list. Photo: David Minter. 

The tenth follow-up conference of the parties 

to the 1992 Rio Convention on Biological 

Diversity was held in Nagoya, Japan on 

18–29 October 2010. For the first time at 

these meetings, as a result of lobbying from 

the new International Society for Fungal 

Conservation (see Reports in this issue), and 

the Cup Fungi, Truffles and their Allies specialist 

group of the IUCN Species Survival 

Commission, fungi were explicitly considered 

independently from animals and plants. 

The focus of the lobbying was section 

E, paragraph 10 of the Global Strategy for 

Plant Conservation. The wording of that 

paragraph read, “accordingly the Strategy 

addresses the Plant Kingdom with main 

focus on higher plants, and other welldescribed 

groups such as Bryophytes and 

Pteridophytes. This does not imply that 

these lower groups do not have important 

ecological functions, nor that they are not 

threatened. Parties may choose on a national 

basis to include other taxa, including algae, 

lichens and fungi”. It was pointed out to 

delegates of the conference that the final 

sentence of this paragraph gave the mistaken 

impression that fungi are “lower” plants, 

that “lichens” are different from fungi, and 

that strategies for fungal conservation could 

be treated as an optional extra. 

At Nagoya, delegates discussed these 

concerns and agreed that the wording needed 

to be changed. The revised and agreed 

wording reads, “while the Strategy addresses 

the plant kingdom with main focus on higher 

plants, and other well-described groups 

such as bryophytes and pteridophytes; Parties, 

other governments and other relevant 

stakeholders may consider developing conservation 

strategies for other groups such as 

algae and fungi (including lichen-forming 

species)”. 

The revised wording recognizes that lichens 

are fungi, and that fungi are not plants. 

It also recognizes the possibility that fungi 

should have their own separate strategy. In 

1992, the Convention established that all 

groups of living organisms have the right to 

exist on this planet and to be protected. Up to 

now, the Convention has not lived up to that 

promise in respect of the fungi. By explicitly 

recognizing fungi as different from plants, this 

re-wording can be seen as a small but significant 

first step towards getting the convention 

to honour its promise for the fungi. 

The Convention also agreed key targets 

for 2020, which included commitments 

to: cut the rate of loss of natural habitats by 

at least half; to increase terrestrial nature 

reserves from 13 % to 17 % of the world’s 

land area; increase marine and coastal nature 

reserves from 1 % to 10 % of the world’s 

seas; restore at least 15 % of the areas where 

biodiversity is classed as ‘degraded’; and 

safeguard at least 75 % of plant species in 

collections. For further information see 

. 

David W Minter 

(d.minter@cabi.org) 

v o l u m e 1 · n o . 2 

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News 

Mycologists go political, with success 

A critically important message in this 

International Year of Biodiversity is to 

promote the missing “F” word at every 

opportunity – Fauna, Flora, AND 

Fungi. This was part of a rallying call to 

all mycologists delivered in a recent IMC 

paper by Dave Minter, and through the 

establishment of the International Society 

for Fungal Conservation during IMC9. 

Already, the Society’s founding members 

have demonstrated the power of global 

coordination to influence a bastion of public 

broadcasting – the BBC. 

On 15 October 2010, the BBC’s 

“The World Tonight” programme focused 

attention on the biodiversity crisis ahead 

of the COP 10 meeting in Nagoya. 

Although an engaging and well-produced 

45-min programme, it perpetuated the 

simplification of biodiversity to “fauna 

and flora”. A call to action went out from 

Dave, and mycologists globally responded 

by petitioning the BBC. Dave sent e-mails 

to 180 people, all the Founder Members of 

the new International Society for Fungal 

Conservation. At least 62 messages were 

sent to the BBC, which came from a 

minimum of 29 countries: Argentina, 

Australia, Brazil, Canada, China, Cuba, 

Ecuador, Egypt, France, Ghana, Greece, 

India, Italy, Malaysia, Mexico, New 

Zealand, Philippines, Poland, Puerto Rico, 

Russia (Novosibirsk oblast, St Petersburg 

oblast, Tula oblast; i.e. Russia in Asia and 

in Europe), Serbia, South Africa, Spain, 

Sweden, Ukraine, UK, United Arab 

Emirates, USA, and Zimbabwe. 

Is this the first ever global political 

action by mycologists? That appears to be 

the case. The result was a triumph; within 

hours the BBC producer Alistair Burnett 

by way of the BBC editors’ blog () 

apologised for the omission, stating: 

“The UN’s member states are getting 

together next week ...... to stop the loss 

of plants, animals and fungi species - 

or biodiversity - ......Just a note to our 

listeners who’ve e-mailed us in the past 

day pointing out that we failed to mention 

fungi when describing what biodiversity 

is, my apologies, we could have been more 

explicit.” 

The word “fungi” was hyperlinked to an 

excellent blog about the fundamental 

importance of fungi: . 

Peter Buchanan 

(BuchananP@landcareresearch.co.nz) 

Mushrooms – the new plastic? 

Styrofoam is one of the most egregious 

offenders in the disposable plastics category, 

US $ 20 Bn of this material is produced 

annually and used in products as diverse 

as building insulation, coffee cups, and 

packaging; it is said to occupy around 

25 % of landfill sites, and will stay there for 

100s of years. In the oceans it can break into 

smaller and smaller globules, but these do 

not really ever disappear. Gavin MacIntyre 

and Eben Bayer (Ecovative Design, Green 

Island, NY, USA), with financial support 

from the National Science Foundation 

(NSF), have been developing a way to 

transform agricultural wastes, especially 

seed husks, with basidiomycete mycelia to 

form an entirely new class of materials, 

which perform much like plastics, but as 

they are made from crop waste differ in 

being totally compostable when no longer 

required. The waste material is transformed 

into a chitinous polymer, Mycobond that 

can be moulded into different shapes and 

used in products ranging from packaging 

to insulation boards for the construction 

industry (Greensulate). The inventors 

have developed a continuous system 

which cleans, cooks, cools and pasteurizes 

the waste materials, while continuously 

inoculating them with the mycelium. The 

resultant stream of material can be made 

into almost any shape which self-assembles 

in a mould. This invention is particularly 

exciting as there is potential for different 

locally available materials to be used 

according to what agricultural wastes are 

available in a country. The basidiomycete 

fungi utilized are not, however, named in 

the released information. 

For further information on this 

most exiting and novel discovery, 

which has such enormous economic 

potential and environmental benefits, 

see the video on , and the NSF report 

on . 

White truffles still demand a high price 

In times of austerity, it might have been 

reasonable to expect that luxury and 

gourmet foods would drop in value. 

However, at the Fiera Nazionale del Tartufo 

Bianco d’Alba, the annual truffle fair in the 

town of Alba in the Piedmont district of 

northern Italy, which ran from 9 October to 

14 November 2010, a single 900 g specimen 

of Tuber magnatum, the Alba White Truffle, 

was sold to a Hong Kong buyer for a massive 

105 000 € (US $ 141 605), i.e. 117 € g -1 . The 

previous record paid for a single truffle was 

one of 1.5 kg which sold to a Macau casino 

owner for £ 165 000 (194 330 €, US $ 262 

255) in 2007, equating to 129 € g -1 . 

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i m a f U N G U S

Fungi on German TV: Focus on black fungi 

Sybren de Hoog and Bert Gerrits van den 

Ende of CBS (Utrecht) featured on Germany’s 

second TV network in an episode of 

the ZDF science program ‘Abenteuer Wissen’. 

The CBS homepage () 

has a link to the original program, as well as 

a version with English subtitles. De Hoog 

explained on the background of a brain 

infection after near-drowning of a child, 

published by Mursch et al. (2006). This 

concerned a 2-year-old boy who fell into a 

rainwater container and aspirated polluted 

water for several minutes. The child was reanimated 

and did well, but later he became 

sleepy and fell into a deep coma. Multiple 

fungal brain abcesses were observed and 

removed by surgery. The etiologic agent 

proved to be Scedosporium apiospermum, 

one of the very few species consistently associated 

with near-drowning fungal encephalitis 

(Guarro et al. 2006), which is almost 

always fatal. The boy was saved thanks to 

timely diagnostics and appropriate therapy. 

De Hoog and coworkers recently published 

a special issue of Medical Mycology 

on Scedosporium (47 (4), 2009: ‘Pseudallescheria 

and Scedosporium: emerging 

opportunists’, JP Bouchara et al., eds), 

stemming from a series of Workshops on 

this group of fungi. They are abundantly 

present in stagnant, nutrient-rich waters, 

as well as in poorly aerated industrial and 

agricultural soils. Alkane pollution is a promotive 

factor. Investigations are ongoing by 

the group of Johannes Rainer in Innsbruck 

to utilize S. dehoogii for bioremediation of 

diesel-polluted sites. The taxonomy of the 

genus is still under debate (Gilgado et al. 

Processing Scedosporium samples, recorded by a ZDF TV team for ‘Abenteur Wissen’, a science program on 

Germany’s second network. 

2005, Lackner et al. 2010), but it is clear 

that a complex of species is concerned each 

occupying different slots in an ecological 

continuum. Scedosporium apiospermum has 

a bias towards virulence, whereas S. dehoogii 

at the opposite end of the spectrum is almost 

strictly environmental. 

There are more fungi inhabiting oils 

and toxic pollutants. Hans-Michael Fürst 

demonstrated frequent isolation of Hormoconis 

resinae, not only from hydrocarbonpolluted 

soils, but also as a monoculture in 

the oil tanks and machinery of ships (Fürst 

2000). Biofouling by this fungus may cause 

engines to stop, which potentially is quite a 

threat to aviation. But authorities will never 

admit that there is a problem, said Dr Fürst. 

Growth of Hormoconis is also promoted by 

phenol-rich creosote used in wood protection, 

in which respect there is a striking resemblance 

to black yeasts (Prenafeta-Boldú 

et al. 2006). Chaetothyrialean black yeasts 

are massively isolated from treated wood by 

enrichment with phenolic aromates (Zhao 

et al. 2010). Chaetothyrialean black yeasts 

differ by combining this feature with oligotrophy 

(Satow et al. 2008), which explains 

their abundance in steam baths and dishwashers 

(Zalar et al. 2011). 

Black yeasts belonging to the orders 

Dothideales and Capnodiales are mainly 

extremophiles, surviving harsh conditions 

of heat, cold, dryness and solar irradiation. 

Their evolutionary origin lies far back, when 

they occupied micro-niches in crevices 

between crystals in bare rock of deserts, 

mountains or the Antarctic (Selbmann et al. 

2005). On German TV, Thomas Warscheid 

gave a more recent example of their rockinhabiting 

life style in the deterioration of 

the famous Dom of Cologne, Germany. 

Monuments are crumbling under the slow 

but effective force of black fungi. Earlier 

Warscheid observed the same thing happening 

with the temples of Angkor Wat in 

Cambodja, as he explained in The New York 

NEWS 

Blackish discoloured skin of toes and toe webs with scaling. [from Li et al. 

(2008) Studies in Mycology 61: 131–136]. 

Skin biopsy stained with HAE; some fungal elements are visible (arrows). Bar = 5 

µm. [from Li et al. (2008) Studies in Mycology 61: 131–136]. 

v o l u m e 1 · n o . 2 

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News 

Times of 24 June 2008. The fungi utilize 

hydrocarbon air pollutants and assimilation 

products of algae, acidify their environment, 

and finally turn all buildings black. Aspects 

of this behaviour were explained in a special 

issue of Studies in Mycology (61, 2008: 

‘Black Fungal Extremes’, GS de Hoog & M 

Grube, eds), while medically significant species 

were presented in one of Medical Mycology 

(47(1), 2009; ‘The Dark World of Black 

Fungi’, Vitale et al., eds). Fungal Biology will 

also devote a special issue to black yeasts 

shortly, entitled ‘The Emerging Potential of 

Black Yeasts’ (Gunde-Cimerman et al., eds). 

Black fungi are remarkable for their odd 

ecology, which we still have a long way to go 

towards understanding. In ‘Abenteuer Wissen’, 

Axel Brakhage explained his research on 

Aspergillus fumigatus, a ubiquitous compost 

fungus which is hypothesized to express factors 

enhancing escape from phagocytosis when 

confronted with human alveolar macrophages 

(Behnsen et al. 2007). Similar processes are 

to be expected when black yeasts invade our 

homes and our bodies from the specialized 

niches somewhere in nature (Sudhadham et 

al. 2008). They differentially adapt to a highly 

deviating environment using different combinations 

of the essential factors determining 

their natural habitat. Gerrits van den Ende et 

al. (2011) applied the term ‘ecological fitting’ 

to this process enabling rapid host and environmental 

shifts, eventually leading to sympatric 

speciation. Many extremophiles, however, 

already occupy their ultimate habitat and don’t 

have an adaptive space around them. They are 

likely to have entered a dead-end street and are 

likely to become extinct when climate changes 

dramatically, as it is expected to do. 

Sybren de Hoog 

(s.hoog@cbs.knaw.nl) 

Behnsen J, Narang P, Hasenberg M, Gunzer F, Bilitewski U, et al. (2007) Environmental dimensionality controls the interaction of phagocytes with the pathogenic fungi 

Aspergillus fumigatus and Candida albicans. PLoS Pathogens 3: e13. 

Fürst HM (2000) Ökologie des Hyphenpilzes Hormoconis resinae und Eigenschaften seines n-Alkan-induzierten P450-Monooxygenasesystems. Thesis. Technische Universität 

Berlin. 

Gilgado F, Cano J, Gené J, Guarro J (2005) Molecular phylogeny of the Pseudallescheria boydii species complex: proposal of two new species. Journal of Clinical Microbiology 

43: 4930–4942. 

Guarro J, Kantarcioglu AS, Horré R, Rodriguez-Tudela JL, Cuenca Estrella M, et al. (2006) Scedosporium apiospermum: changing clinical spectrum of a therapy-refractory 

opportunist. Medical Mycology 44: 295–327. 

Lackner M, Gerrits van den Ende AHG, Hoog GS de, Kaltseis J (2010) Barcoding of the therapy-refractory species of Pseudallescheria and Scedosporium. Medical Mycology: 

in press. 

Mursch K, Trnovec S, Ratz H, Hammer D, Horré R, et al. (2005) Successful treatment of multiple Pseudallescheria boydii brain abscesses and ventriculitis/ependymitis 

in a 2-year-old child after a near-drowning episode. Child’s Nervous System 22: 189–192. 

Prenafeta-Boldú FX, Summerbell R, Hoog GS de (2006) Fungi growing on aromatic hydrocarbons: biotechnology’s unexpected encounter with biohazard? FEMS 

Microbiology Reviews 30: 109–130. 

Satow MM, Attili-Angelis D, Hoog GS de, Angelis DF, Vicente VA (2008) Selective factors involved in oil flotation isolation of black yeasts from the environment. 

Studies in Mycology 61: 157–163. 

Selbmann L, Hoog GS de, Mazzaglia A, Friedmann EI, Onofri S (2005) Fungi at the edge of life: cryptendolithic black fungi from Antarctic desert. Studies in Mycology 

51: 1–32. 

Sudhadham M, Prakitsin S, Sivichai S, Chaiyarat R, Dorrestein GM, et al. (2008) The neurotropic black yeast Exophiala dermatitidis has a possible origin in the tropical 

rain forest. Studies in Mycology 61: 145–155. 

Zhao J, Zeng J, Hoog GS de, Attili-Angelis D, Prenafeta-Boldú FX (2010) Isolation and identification of black yeasts by enrichment on atmospheres of monoaromatic 

hydrocarbons. Microbial Ecology 60: 149–156. 

Executive Committee and Officers of the International Mycological Association (2006–2010). 

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REPORTS 

IMC9 Edinburgh: a selection of memorable moments featuring some of those who helped to make it such a success. 

v o l u m e 1 · n o . 2 

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REPORTS 

IMC9: The Biology of Fungi 

A personal reflection 

The idea of holding IMC9 in Edinburgh in 

2010 started five years ago when I received 

a phone call from Ellen Collingsworth at 

the Edinburgh Convention Bureau. She 

said that the last (and the first) International 

Mycological Congress (IMC) to be held 

in the UK was in 1971 in Exeter, and 

wouldn’t I like to organize the next one? 

My response was initially very negative (I 

will omit the expletives I used) because I 

was very conscious of the profound effect 

that organizing an IMC would have on 

both my academic and personal life over 

the next five years. However, after having 

many discussions with senior officers of 

the British Mycological Society (BMS), 

seeing the proposed Congress venue (the 

superb Edinburgh International Conference 

Centre, EICC), and giving the matter 

considerable thought, I strongly warmed to 

the idea of taking on this onerous job. I felt 

that it presented some exciting challenges 

which ultimately could have a big impact on 

global mycology. 

Early in 2006, the ECB, BMS, and 

myself put together and submitted a bid to 

the IMA to hold IMC9 in Edinburgh, with 

the BMS agreeing to act as host. I received 

extremely strong support from the BMS 

Council, who generously agreed to commit 

£ 100,000 to pump-prime the Congress. 

I took on the organization of IMC9 

with a clear vision of how I wanted it to 

be: (1) the whole of mycology had to be 

represented in all of its guises in a very 

balanced way across the immense breadth of 

the subject -- to try and give a flavour of this 

and provide the Congress with an up-todate 

image, I subtitled it The Biology of 

Fungi which I felt might appeal to a broader 

range of scientists working on fungi; (2) 

the Congress had to have a stellar scientific 

programme with a strong emphasis on where 

the excitement of the subject would be at in 

2010, and where its big areas of impact will 

be in the future -- it was essential that the 

conference programme and speakers should 

inspire young and old mycologists alike; (3) 

the scientific programme should evolve by 

a very carefully regulated process of natural 

selection in which only the best symposia 

proposed by the community would be 

Delegates being led from the Usher Hall to the EICC following the Opening Ceremony by Scottish pipers. 

chosen; (4) it would be compulsory for 

every symposium to have young researchers 

(postgrads and/or postdocs) giving talks; 

(5) the poster sessions would be given a 

high profile and be very accessible to the 

delegates in areas where they congregated 

in the Congress venue (e.g. at lunchtime); 

(6) delegates should be able to experience 

the delights of the City of Edinburgh and 

all that it has to offer as one of the world’s 

main cultural heritage sites -- here I felt 

that it was important that the timing of 

the Congress should be at the beginning of 

August just before the Edinburgh Festival 

when hotel prices reach their maximum; (7) 

the conference party should be at the end of 

the conference when people could properly 

relax and let their hair down. I openly 

declared that this should be the “conference 

party to end all conference parties”; and (8) 

I made it clear to everyone involved in the 

organization that we should try our hardest 

to make IMC9 the best IMC ever! From 

my perspective, even with my somewhat 

biased point of view, I felt that my vision for 

IMC9 was largely if not completely fulfilled. 

However, this only came about as a result of 

the team effort of hundreds of participating 

individuals. 

Once having won the bid and the dust 

from IMC8 in Cairns having settled, I 

set about finding a suitable Professional 

Conference Organizer (PCO) with the help 

of Geoff Robson and Nick Clipson on the 

Steering Group. To organize a Congress of 

the size of an IMC (our best guesstimate was 

we that we would attract between 1200 and 

2500 delegates to Edinburgh), it was absolutely 

essential to have an extremely good 

PCO to work closely with. Amongst other 

things, the PCO is responsible for most of 

the conference administration, invitation of 

speakers and poster presenters, organization 

of delegate registration, communication 

with delegates, interfacing with the venues, 

organizing the exhibition, obtaining 

sponsorship, marketing the conference, etc, 

etc. Any naïve notion that I could organize a 

conference on this scale without a PCO was 

quickly kicked into touch once it became 

clear how massive the task of organizing a 

conference on this size is. We short-listed 

three PCOs and finally took on board 

Elsevier who, as well being publishers, have 

a big PCO Department. There were several 

key issues in Elsevier’s favour over the other 

PCOs we interviewed. What was particularly 

significant for the BMS was that Elsevier 

were the only PCO to agree to take on 

the complete financial risk for the Congress 

if it went ‘belly up’ (e.g. due to volcanic dust, 

acts of terrorism, fears of epidemics). We 

were concerned that an ‘act of God’ could 

potentially result in bankrupting the BMS. 

The cost of organizing IMC9 approached a 

£ 1 million! Another aspect strongly in Else- 

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i m a f U N G U S

vier’s favour was that they published four 

journals for the BMS, and IMC9 presented 

various exciting publishing opportunities. 

We also felt that Elsevier, with its experience 

in publishing, would be able to market the 

Congress well and give it the image we felt 

that it should have. We did not want IMC9 

to have a boring clinical feel to it, which is 

so typical of many conferences. There is no 

question that we made the right choice, and 

the Elsevier team, led by the inimitable Nina 

Cosgrove, were outstanding and a joy for me 

to work with over the last four years. 

One of my aims was that the scientific 

programme should present the whole 

breadth of mycology in a very balanced way 

without any single topic dominating. In 

consultation with the Steering Committee, 

I divided the subject into five themes 

which I felt represented the main areas of 

the subject, and I gave these themes equal 

weighting. These five themes were: 

1. Cell Biology, biochemistry and 

physiology 

2. Environment, ecology and interactions 

3. Evolution, biodiversity and systematics 

4. Pathogenesis and disease control 

5. Genomics, genetics and molecular 

biology 

Across all five themes ran applied aspects of 

the subject (e.g. fungal biotechnology). 

I set up a number of Committees to 

bring the Congress organization, and particularly 

the scientific programme, to fruition. 

First, we had a Steering Group of six individuals 

chaired by me that had an advisory 

role and oversaw the Congress organization. 

Second, we had a Scientific Programme 

Committee comprising the chairs of the five 

scientific themes, and this committee was 

also chaired by me. And then finally we had 

the five scientific theme committees each 

containing five eminent scientists covering 

the breadth of each theme. 

We next invited the mycological 

community to propose symposia for the 

scientific programme. We had decided that 

it would be possible to hold 45 symposia, 

each 2.5 h long with seven speakers during 

five days of the Congress (Monday–Friday). 

This equated to nine symposia per theme. 

However, we made sure that many of the 

symposia were inter-thematic. Amazingly, 

and a tribute to the enthusiasm of the 

mycological community, we received over 

Delegates in session at the EICC. 

220 symposium proposals. The five scientific 

theme committees then set to work 

to prioritize these and the final selection 

was made by the Scientific Programme 

Committee. As you can imagine, this was 

an extremely difficult task because we had 

so many outstanding proposals for the 

45 symposium topics. Because we had so 

many excellent suggestions that didn’t make 

the cut as symposia, we decided that these 

should be converted into Special Interest 

Group meetings to be held on the Sunday 

before the Opening Ceremony. Finally, we 

were also able to hold three Nomenclature 

Sessions during the Congress because this 

was going to be a very hot topic in 2010 

with potentially major changes in fungal 

nomenclature afoot. 

About two years before the Congress, I 

had the idea of organizing an exhibition of 

fungi in the superb new John Hope Gateway 

exhibition centre that had just been built 

at the Royal Botanic Garden Edinburgh 

(RBGE). I went to see the Regius Keeper 

of the RBGE and his Deputy and said how 

cool it would be if the RBGE could hold 

an exhibition for 2 or 3 weeks around the 

time of the Congress which would not only 

appeal to IMC9 delegates but also to the 

general public. They warmed to the idea, 

but said that an exhibition on chocolate was 

planned for that time. I wasn’t defeated by 

their response and went ahead with extra 

determination and put forward a proposal 

for holding the fungal exhibition. The 

RBGE responded by saying that they didn’t 

like my idea of a 2–3 week exhibition but 

wanted this to be the main exhibition at 

RBGE in 2010, and for it to be four months 

long! At this point I got the BMS involved 

in its organization, and the exhibition 

became the main outreach experience of the 

congress, and one of the largest the BMS has 

ever been involved in. The exhibition was 

called ‘From Another Kingdom: the Amazing 

World of Fungi’ and was accompanied by 

a coffee-table book aimed at the general 

public as well as academics, edited by Lynne 

Boddy and Max Coleman. The John Hope 

Gateway exhibition centre, together with 

the exhibition, also provided a superb venue 

for two of the receptions held during IMC9, 

and which were sponsored by the BMS, 

Mycological Society of America and the 

British Society for Plant Pathology. 

We worked hard to keep the Registration 

costs of the Congress as low as possible, 

and certainly these costs were lower than 

most equivalent meetings covering six 

days held at the EICC. We also realized, 

however, that these fees would still be too 

high for the majority of potential delegates 

from low-middle income countries, so, for 

the first time in IMC history, we introduced 

a substantially reduced fee for them. We 

additionally realized that if we were going 

to attract the biggest stars in the field to 

speak at the Congress, then we would have 

to provide a significant financial incentive 

which was greater than has been provided 

for invited speakers at previous IMCs. As 

a result we were able to contribute over £ 

100,000 towards 220 invited speakers and 

Symposium Organizers. We also set up a 

REPORTS 

v o l u m e 1 · n o . 2 

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REPORTS 

Delegates enjoying the Conference Party at the completely transformed EICC. 

bursary scheme in which we able to provide 

£ 91,000 in bursaries to 296 delegates from 

80 countries as financial assistance to attend 

the Congress. The bursary scheme was 

primarily managed through the herculean 

efforts of Geoff Robson. 

Originally, the conference was going 

to be held entirely in the EICC because 

they had planned to build an extension 

which was due to be completed in 2009. 

However, two years before the congress 

was due to start, the contractors withdrew 

from building the extension and as a result 

we ended up holding part of the conference 

in the somewhat dramatic Usher Hall, 

Fortunately this worked well since the two 

venues were only a 5 min walk apart, and 

delegates were guided between them by fly 

agaric mushrooms adorning the pavement. 

The only major obstacle was the very busy 

Lothian Road which separated them. 

However, with the assistance of the police 

this potentially nightmarish problem was 

overcome. One of my long lasting memories 

of the congress was seeing 1200+ mycologists 

on each day of the conference bringing 

the traffic of Edinburgh to a standstill! 

A scary aspect of organizing any 

conference is not knowing how many people 

would actually register, and then on top of 

that, as indicated earlier, there is always the 

possibility that some ‘act-of-God’ might 

prevent delegates actually getting to the 

conference. Indeed, four months before 

the IMC9 started all flights in and out of 

the UK had come to a standstill because of 

volcanic dust drifting across from Iceland. 

At the end of the day this nightmare 

scenario did not happen. 

Ultimately, the success of any conference 

lies with the delegates, many of whom have 

to travel considerable distances. I am very 

proud to say that 1593 delegates from 83 

different countries finally registered for 

IMC9. The ‘I’ in IMC9 was thus fully 

deserved. About 330 delegates gave oral 

presentations in Symposia and Special 

Interest Group sessions, and some in the 

Nomenclature Sessions. In addition, there 

were ~ 1,200 posters presented at the 

meeting. I am extremely indebted to all of 

those who made such a big effort to attend 

and participate in the Congress. 

After the official opening of the congress 

and the handing over of the new IMC 

gavel, made of wood from every continent 

on the globe (including Antarctica), John 

Taylor (University of California at Berkeley) 

kicked off the scientific programme with 

an outstanding talk on the “The poetry of 

mycological accomplishment and challenge” 

whilst kitted out in full Scottish regalia. We 

couldn’t have had a better start. Besides integrating 

mycology with poetry, John’s major 

‘take home’ message was for mycologists 

to ‘think big’. Each successive day of the 

conference began with a Plenary Lecture by 

a mycological superstar, except on the last 

day when we were treated to two superstars. 

These mycological leading lights were: 

• Gero Steinberg (Exeter University, 

UK): Organelle transport in fungi - 

stochastic or controlled? 

• David Hibbett (Clark University, 

USA): Knowing and growing the 

fungal tree of life 

• Joe Heitman (Duke University, USA): 

Microbial pathogens in the fungal 

kingdom 

• Nick Talbot (Exeter University, UK): 

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Welcome to the pressure dome: investigating 

the molecular genetics of plant 

infection by the rice blast fungus 

• Alastair Fitter (University of York, UK): 

A forgotten phylum? 

• Nancy Keller (University of Wisconsin 

at Madison, USA): Unlocking the fungal 

treasure box 

Every one of their talks was truly inspirational 

and exceptional in their scope and in the scientific 

excitement they each generated. 

The Plenary Lectures in the morning were 

followed by five parallel sessions of symposia, 

with a long break in the middle of the day for 

lunch and viewing poster presentations. I was 

not able to attend as many of the symposia 

as I would have liked, but all the sessions I 

did attend were of outstanding quality. The 

feedback I received from those attending other 

symposia was excellent. 

The conference was brought to an 

official end on the Friday with the Closing 

Ceremony. During this session, the IMA 

General Assembly, a business meeting, was 

presided over by the President of the IMA, 

Pedro Crous, who was highly praised for the 

exceptional job that he has done for the IMA 

over the last four years. John Taylor was 

announced as the incoming IMA President 

for the next four years. The IMA presented 

two medals, the De Bary medal for outstanding 

scientific contributions to Franz Oberwinkler, 

and Ainsworth Medals for outstanding 

services to mycology to ‘Dick’ Korf and 

Emory Simmons. It was also announced that 

a new series of medals for younger mycologists 

in the countries covered by each of the 

five IMA Regional Committees was being 

established, and that the new IMA journal 

IMA FUNGUS was being launched. Twenty 

prizes for outstanding poster presentations, 

generously provided by Elsevier, were also 

made. The formal business closed with a 

short presentation by Lekha Manoch inviting 

mycologists to IMC10 which is to be held in 

Bangkok, Thailand, in 2014. 

However, IMC9 did not end there, as after 

the Closing Ceremony, we had few hours to 

kill and put on our glad rags and dancing shoes 

in preparation for the Conference Party. I had 

spent a lot of time organizing the party to be an 

experience that would be thoroughly enjoyed 

and remembered. About 700 delegates attended 

the party, which was also held in the EICC, 

which was completely transformed from a 

scientific venue into a party environment. The 

party-goers were entertained by four bands, 

ceilidh dancing, karaoke, salsa dancing, whisky 

tasting, and food from all over the world that 

was provided at different locations around the 

EICC. To my mind, the party provided the best 

way to finish what had been an amazing week of 

science and fun(gi). 

There is no question that the organization 

of IMC9 took a lot of hard work and 

commitment, but I have to say that I really 

enjoyed all of it. However, it would never have 

happened without extraordinary teamwork. I 

started to add up the number of people who 

had been involved in different aspects of its 

organization, and in making it an unquestionable 

success, and after getting up to 150 

individuals I gave up! I can’t thank all of these 

people anything like enough. It was clear that 

all of those attending the Congress all shared 

one thing in common – a passion for fungal 

biology. I was very struck during the meeting 

by the fact that no one was standing around 

looking bored. Everyone was either intensely 

engaged in the science or in lively communication 

with each other. We all know the 

importance of communication, interactions, 

and networking, not only for mycelia but also 

for the progress of any scientific discipline, 

including mycology. IMC9 provided that. My 

long lasting feeling about the congress was 

that fungal biology in 2010 is in a very healthy 

state and there has never been a more exciting 

time to be studying the subject. 

Nick D. Read 

(nick.read@edu.ac.uk) 

REPORTS 

International Commission on the Taxonomy of Fungi 

(ICTF) 

The ICTF held a General Meeting on 2 

August 2010 during IMC9. The ICTF is 

COMCOF of IUMS and a Commission 

of the IMA. A full record of the meeting 

appears on the ICTF website () and 

only a synopsis is presented here. 

Subcommissions and working groups 

Several taxon-specific Subcommissions (SC) 

and a Working group (WG) are associated 

with the ICTF: 

Fusarium SC – (chair: David Geiser). 

This group also works under the auspices 

of the International Society of Plant 

Pathology Commission on Fusarium, and 

holds meetings prior to the International 

Congress of Plant Pathology (ICPP). The 

EF1-alpha DNA sequence database created 

by David Geiser with much data from Kerry 

O’Donnell (USDA) was augmented with 

an RPB2 database to enable identification 

of Fusarium strains from a curated, barcodelike 

database, and was moved to a new web 

platform at . The list of current names 

of Fusarium continues to be available at 

, but has been integrated 

in the MycoBank database () and is maintained at that site. The 

Fusarium SC group met at the 10 th International 

Fusarium Workshop (Alghero, Italy, 

August 2008) after the Torino, Italy ICPP. 

Discussions were initiated to organize a 

specialist workshop on Fusarium taxonomy 

and molecular phylogenetics, to discuss a 

community oriented approach to solving 

some of the more pressing issues in this 

genus. The next meeting is planned for the 

ICPP in China in 2013, and the organization 

of the workshop is already underway 

by Ulf Thrane (Technical University of 

Denmark) , the chair of the ISPP Fusarium 

Subject Matter Committee. 

Trichoderma SC (ISTH) – (chair: 

Irina Druzhinina). The barcode identification 

system, TrichoKey2, continues to be 

maintained on the subcommission website, 

. This website also has extensive 

literature and additional information on 

Trichoderma and its sexual states, Hypocrea. 

This group has been active in developing 

and publishing collaborative, polyphasic 

projects such as the special issue of Studies 

in Mycology (56, 2006). They are also active 

v o l u m e 1 · n o . 2 

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REPORTS 

in organizing and making presentations 

at international meetings, and held an 

international workshop on Trichoderma in 

agriculture in Haifa, Israel, in October 2010. 

International Commission on Penicillium 

and Aspergillus (ICPA) – (chair: 

Robert Samson). This commission reports 

separately to the IUMS, but the chair also 

sits on the ICTF. It organized an international 

workshop on “Aspergillus systematics 

in the genomic era” in April 2007, and 

published the proceeding in Studies in 

Mycology (59, 2007). This Commission 

has been very active at IUMS meetings, 

and organized a session on “Advances in 

molecular phylogenetics/systematics of 

Penicillium and Aspergillus species” at the 

2008 Istanbul Congress, and plans a session 

at the 2011 Sapporo Congress. Their 

website is maintained at . 

Ceratocystis/Ophiostoma SC – 

(co-chairs: Keith Seifert, Michael Wingfield). 

A three day pre-congress symposium 

attended by 45 people was organized at 

IMC8. The editing of these proceedings is 

in progress, but establishment of a formal 

structure remains on hold pending potential 

members securing permanent positions. 

Mycosphaerella SC – (chair: Pedro 

Crous). This informal subcommission is 

centred around CBS and its collaborators. 

A one day symposium on Cercospora beticola 

was held at the American Phytopathological 

Society/Canadian Plant Pathological 

Society/Mycological Society of America 

meeting in Quebec City in August 2006. 

There will also be a specialist workshop on 

Mycosphaerella in Australia in April 2011. 

Fungal Barcoding WG (FunBOL) – 

This subcommission is a shared committee 

with the Consortium for the Barcode of 

Life (CBOL, Smithsonian Institution, 

Washington DC; ). 

Originally organized by Keith Seifert, 

Pedro Crous and John Taylor (University 

of California Berkeley) at the invitation of 

CBOL, it is now chaired by Conrad Schoch 

(GenBank). Membership and terms of reference 

are currently being finalized, and the 

group is expected to be active in initiating the 

official recognition of DNA barcode markers 

for fungi, lobbying for the inclusion of fungi 

in barcoding projects, and developing large 

scale fungal barcoding projects. 

Additional SCs – Discussions amongst 

interested mycologists had been held with a 

view to initiating additional SCs concerned 

with Colletotrichum (Peter Johnston) and 

Stachybotrys (Keith Seifert) but no formal 

arrangements had been made. It was recognized 

that most of the active workers in 

these groups were graduate students or postdocs 

who could not become committed at 

this time. 

Other activities 

In addition to the activities of the subcommissions, 

the ICTF has also been involved 

in the following activities: 

How to Describe a Fungal Species – A 

draft document presenting the procedures 

necessary to effectively describe a fungal 

species was written by Keith Seifert and 

circulated to the rest of the commission for 

comment, and after revision is published in 

this issue of IMA Fungus. The document is 

intended to provide guidance as to formal 

requirements and good practice for students 

or non-taxonomic mycologists who find 

that they need to describe new fungi. 

IUMS Congress in Istanbul in 2008 

– The ICTF organized a symposium on 

“Taxonomic developments in economically 

important fungal genera” at the IUMS 

Congress in Istanbul. Robert Samson was 

the principal organizer, with support from 

Irina Druzhinina. 

Future directions and activities 

New executive officers of the ICTF need 

to be elected, but it was felt that, a priori, 

the remit and future direction of the 

commission should be clarified, beyond 

the need to provide a structural framework 

for subcommissions and working groups. 

Scott Redhead (Agriculture and Agri-Food 

Canada) suggested that the ICTF could be 

active in developing subcommissions that 

could develop consensive nomenclature for 

specific taxonomic groups; noting that the 

“Names in Current Use in the Trichocomaceae” 

project initiated by ICPA had been 

granted special protected status at the International 

Botanical Congress in Tokyo in 

1993 (see the Preamble in the Tokyo Code). 

Michael Wingfield felt that problems with 

the Botanical Code were pressing issues for 

mycologists, as evidenced by the Nomenclature 

Session discussions during IMC9, 

and that the ICTF could have a role in the 

resolution of these problems, perhaps even 

taking responsibility for drafting an independent 

code of mycological nomenclature 

should that become necessary in the future. 

Rob Samson, stated that the ICTF could 

have a positive influence on microbiologists, 

by presenting information on changes of 

names and refinement of species concepts, 

which would help to make fungal taxonomy 

visible to applied scientists – something it 

had done through a series of publications in 

the early days of the ICTF. David Hibbett 

saw a need for a broader involvement of 

the taxonomic community in the ICTF. 

In order to progress matters, David Hawksworth, 

who had been the founding Chair 

of the ICTF, agreed to prepare a “vision 

paper” for discussion by a working group 

comprising Rob Samson, Pedro Crous, José 

Carmine Dianese, David Hibbett, Peter 

Johnston, Michael Wingfield, Keith Seifert, 

Gen Okada, and Scott Redhead. This 

“vision paper” is included under Correspondence 

in this issue of IMA FUNGUS. 

A suggestion by Robert Samson and Pedro 

Crous that CBS host a symposium with 

ICTF in April 2011 around the theme 

“One fungus: One name” (1F: 1N) where 

the working group’s proposals could also 

be considered was warmly accepted. The 

Edinburgh meeting also agreed to delegate 

to the April symposium the power to elect 

the new Commission, which would then 

elect its new officers. 

Against this background, Keith Seifert and 

Gen Okada agreed to continue to act as 

Chair and Secretary, respectively to facilitate 

any transition to a new executive. 

Keith Seifert and Gen Okada 

(keith.seifert@agr.gc.ca; okada@jcm. 

riken.jp) 

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REPORTS 

IMC9: Moments to reminisce on captured by the camera. 

v o l u m e 1 · n o . 2 

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REPORTS 

IMC9: Interludes of ambience, relaxation, and committee-work, amongst the hard-science. 

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i m a f U N G U S

IMA Medals awarded at IMC9 

The highest honour that the International 

Mycological Association can bestow on 

any mycologist, is embodied in two highly 

prestigious medals, namely the de Bary 

Medal for outstanding career research in 

mycology, and the Ainsworth Medal for 

extraordinary service to world mycology. The 

last time these awards were made was in 

1996, when the de Bary award was made 

to John Corner (for lifetime achievement 

in mycological research, particularly, 

contributions to ecology and the systematics 

of wood-decaying basidiomycetes), and 

Terence Ingold (for lifetime achievement in 

mycological research, particularly, contributions 

to our knowledge of fungal spore 

release and dispersal and the recognition of 

aquatic fungi as ecological specialists). The 

Ainsworth award was made to John Webster 

(for extraordinary service to international 

mycology through the IMA). 

In several meetings held by the Executive 

Committee (EC) during 2006–2010, 

these awards were discussed in detail, and 

finally a call was posted on the IMA website 

for proposals for potential candidates to 

receive the awards at the IMC9 in Edinburgh. 

An IMA Awards Committee was 

established to evaluate the proposals. The 

committee consisted of the Officers (Mike 

Wingfield, Susumu Takamatsu, John Taylor, 

Wieland Meyer, Karen Hansen and Geoff 

Robson), and up to three members from the 

EC (Keith Seifert, David Hawksworth and 

Dominik Begerow), and was chaired by the 

President (Pedro Crous). In a short period 

of time, several excellent applications were 

received, and then the more difficult part of 

the discussion started. 

Three candidates were proposed for 

each of the two medals. The committee 

debated for several weeks, and actually voted 

twice, as it first had to agree on how many 

awards could be made, and the secondly, 

on who the best candidates were to receive 

these awards. Several committee members 

felt that all six candidates deserved to be 

recognised, but eventually a compromise 

was made, and based on the fact that 

no awards were made since 1996, the 

committee agreed to (as an exception) make 

two awards for a lifetime contribution to 

mycology (Ainsworth), and one for career 

research in mycology (de Bary). 

AWARDS AND PERSONALIA 

De Bary Medal: Franz Oberwinkler 

Franz Oberwinkler receiving the De Bary Medal from IMA President Pedro Crous. 

Extracts from the nomination letter submitted by Micheal Weiß and 

Robert Bauer are the following: “Franz Oberwinkler has contributed to 

science for almost 50 years in various fields of fungal research. Besides 

his overwhelming scientific output he is a great teacher and many of 

his scholars have become influential mycologists as well. Franz Oberwinkler 

had a major impact on our understanding of the systematics of 

basidiomycetes. Inspired by a deep interest in detailed morphology and 

anatomy of basidia and basidiocarp ontogeny and their comparative 

analysis his work revolutionarised basidiomycete systematics in wide 

parts. He investigated not only morphology, but included all relevant 

characters ranging from physiology to genetics to understand the 

main evolutionary trends in basidiomycetes. Our modern view on 

Basidiomycota is highly influenced by his work. He was participant on 

all International Mycological Congresses. He was President of the IMA 

from 1994–1998 and holds memberships of many mycological societies 

worldwide. Therefore, we would like to nominate Prof. dr Franz 

Oberwinkler for the De Bary Medal of the International Mycological 

Association in 2010”. 

For me personally, the awards were the highlight of IMC9. It 

was truly a privilege to share the stage with such mycological greats, 

to congratulate Franz Oberwinkler, to see the movie clip of Dick 

Korf (after I upgraded to the Mac platform, thanks to John Taylor!), 

and to hear Emory’s parting words “Save your dollars, your Euros, 

your Pounds, save whatever currency you have and come to Bangkok 

for the IMCX.” See you there! 

Ainsworth Medal: Richard P Korf 

Extracts from the nomination letter 

submitted by Donald Pfister are the 

following: “Korf has focused his research 

on taxonomy of discomycetes and on 

nomenclature. Most of the more than 340 

papers he has published over his more than 

60 active years concentrate on the taxonomy 

of these fungi. His 1973 keys in The Fungi: 

an advanced treatise remain the only 

comprehensive keys for the inoperculate 

discomycetes and has been influential in 

training generations of systematists but 

also in pointing to problem areas. No less 

important than his taxonomic papers are 

his nomenclatural papers. Here he gives 

insight into the intricacies of the rules of 

naming and has sought to stabilize the use 

and application of names. In his series 

Nomenclatural notes, which now number 12 

contributions, he has outlined and solved 

various issues related to the proper application 

of the Code. In 1974 he was motivated 

by several realities of the publication scene 

to found a new independent, international, 

v o l u m e 1 · n o . 2 

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Richard P Korf as "Elias Fries" and Karen Hansen. Inlay: Don Pfister accepting the Award on behalf of Richard 

P Korf. 

mycological journal – Mycotaxon. Taking 

the model of a camera-ready journal of 

the time he improved the methods and 

established the journal that is now in its 

111 th volume accounting for about 60 000 

published pages. In the early volumes he did 

all the editorial work and related correspondence 

and at the same time was the business 

manager. Dick is at heart a teacher and 

devoted himself to instruction both in and 

out of the classroom. He has been honored 

as the State University of New York Chancellor’s 

Award for Excellence in Teaching, 

given the Distinguished Teaching Award 

by Gamma Sigma Delta, and the William 

H. Weston Distinguished teaching award. 

Teaching and outreach build on his interest 

in the theatre. He has had an active career as 

an actor participating in local performances 

and in professional productions. We mycologists 

know him best for his impersonation 

of our father in mycology, Elias Fries. He 

served the Mycological Society of America 

in many ways, as Councilor, as secretarytreasurer, 

as vice president and president 

in 1970–71. He attended nearly all of 

the International Mycological Congresses 

and before that he generally attended the 

International Botanical Congresses. At these 

he was an active participant and looked to 

as a leader. His is a long and distinguished 

career highlighted by engagement in his 

science, excellence in teaching, and service 

to the international mycological community. 

The Ainsworth medal would be a fitting 

recognition of this mycological great”. 

Ainsworth Medal: Emory G Simmons 

Emory G Simmons receiving the Ainsworth Medal from IMA President Pedro Crous. 

Extracts from the nomination letter 

submitted by Mary E. Palm are the following: 

“It is with great pleasure that I write this 

letter to nominate Dr Emory G. Simmons 

for the IMA Ainsworth Medal recognizing 

his truly extraordinary service to world 

mycology. Throughout his career he has 

directly and indirectly, officially and unofficially, 

intentionally and unintentionally 

been a strong voice and a driving force 

for international mycology. In this letter I 

will highlight his significant international 

mycological activities over the past half 

century, especially his tireless efforts in the 

development, maintenance and preservation 

of biological resource collections. I also 

will discuss his significant role in the 

establishment and success of the community 

to which I write this letter, the International 

Mycological Association (IMA). From 

1953–74 Emory was the Director of the US 

Army Quartermaster Culture Collection 

of Fungi. The collection was an essential 

component of U.S. studies on the resistance 

of fabric, glass, wood and other surfaces to 

degradation by fungi. Emory also was an 

active member, Secretary and part of the 

Executive Board of the World Federation 

for Culture Collections to which he was 

elected an Honorary Life Member. He is a 

Centennial Fellow of the British Mycological 

Society. He has served in an advisory 

capacity to the American Type Culture 

Collection, President of the U.S. Federation 

for Culture Collections, and numerous 

other positions in which he positively 

influenced the development of collections 

and educated administrators, politicians, 

and others on the importance of those 

collections for industrial, economic, and 

biological uses. Dr Simmons played a major 

role in the founding and development of 

the IMA. His involvement started in 1967, 

while he was President of the Mycological 

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i m a f U N G U S

Society of America, in the form of correspondence 

with G. C. Ainsworth whose 

brainchild became the IMA. As a testament 

to the esteem in which Dr Simmons is 

held by the IMA, he was elected in 2002 as 

Honorary President of the IMA for life. It 

is impossible to close this nomination letter 

Order of Canada: Stanley J Hughes 

The Order of Canada is the centrepiece of 

Canada’s system of public honours, which 

serves to recognize a lifetime of outstanding 

achievement, dedication, and service to the 

nation. Welshman Stanley ‘Stan’ J Hughes 

was amongst those made a Member of the 

Order in 2010. Stan needs little introduction 

to mycologists working with microfungi 

worldwide. Having cut his mycological teeth 

at the Commonwealth Mycological Institute 

in 1945–1952, where he came under the 

influence of Edmund W Mason, Stan moved 

to what is now Agriculture and Agri-Food 

Canada in Ottawa in 1952, ‘retiring’ in 1985. 

He is best known for his novel and groundbreaking 

scheme for distinguishing methods 

of conidiogenesis in asexual fungi published 

in 1953, the resolution of many uncertainties 

in the application of generic names of conidial 

fungi in 1958, and his masterly overview 

of sooty moulds and their asexual stages in 

1976. 

without mentioning the publication of his 

recent tome “Alternaria: An Identification 

Manual” in 2007 in the CBS Biodiversity 

Series, which he finished several years after 

his 80 th birthday. Thank you for considering 

this nomination of Dr. Emory Simmons 

for the Ainsworth Medal. This would be 

In addition he has produced meticulous 

accounts of numerous microfungi, especially 

ones arising from studies of New Zealand 

material, which he embarked on in 1963 

and continue . . . . Stan has received numerous 

honours and awards over the years, 

including election as a Fellow of the Royal 

Society of Canada in 1974, and continues 

to help and inspire aspiring mycologists, 

not least myself. Although still active at 92 

years, Stan donated his personal library to 

the National Botanic Garden of Wales and 

attended a reception in his honour at the 

Garden in April 2009. All mycologists will 

wish to not only congratulate Stan on this 

additional honour, but also trust he will 

continue to enjoy good health and elucidate 

further fascinating aspects of his beloved 

‘soots’. 

David L Hawksworth 

(d.hawksworth@nhm.ac.uk) 

the ultimate honor for the man who helped 

Ainsworth make an idea become a reality in 

the form of the IMA”. 

Pedro W. Crous 

Past-President, IMA (p.crous@cbs.knaw. 

nl) 

Stanley J Hughes. 


Obituary: C Terence Ingold (1905–2010) 

C Terence Ingold. 

It is with deep regret that we record the death 

of Cecil Terence Ingold on 31 May 2010 at the 

age of almost 105 years. Terence is best-known 

amongst mycologists for his pioneering work 

on spore discharge and dispersal in fungi, the 

five editions of his textbook The Biology of 

Fungi (1961–84) on which so many students 

cut their mycological teeth, and his discovery 

of the fascinating helicoid and tetraradiate 

spores of aquatic hyphomycetes – of which 

over 300 species are now known. After periods 

at the University of Reading and what is now 

the University of Leicester, from 1944 until 

retirement in 1972 he held a professorship 

at Birkbeck College of the University of 

London – where the laboratory he used to 

work in is now named after him. Birkbeck is 

unusual in concentrating on part-time mature 

students, and there he inspired or supervised 

numerous mycologists, including Richard 

Bailey, Hilda Canter, Steve Moss, David Pegler, 

Bryan Plunkett, and Guy Willoughby. In his 

‘retirement’, Terence conducted numerous 

studies from his home in Benson (Oxfordshire) 

on spore germination in diverse fungi until 

1998, mostly published in The Mycologist, 

aided and abetted by supplies organized by Jane 

Nicklin who is now responsible for mycological 

work at Birkbeck. He also continued to attend 

field meetings of the British Mycological 

Society whenever he could. In later life he 

moved to Wooler (Northumberland) where he 

lived with one of his daughters. His numerous 

honours and awards include appointment as 

a Companion of the Order of St Michael and 

St George (CMG) in 1970 for work in higher 

education, especially in Africa and Jamaica, but 

also in the UK where he was involved in establishing 

universities in Belfast and Canterbury. 

A special double-issue of the Botanical Journal 

v o l u m e 1 · n o . 2 

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of the Linnean Society was published as a tribute 

to Terence in 1985, and that includes a full list 

of his 174 publications to that date; about 100 

appeared after that date. The IMA is especially 

indebted to Terence for agreeing to serve as the 

New IMA Awards 

President of IMC1 in Exeter in 1971, and he 

was one of the two first recipients of the IMA’s 

de Bary medal in 1996. For further information 

on this remarkable man and his achievements, 

see the references listed below. 



Plunkett BE (1985) Professor Cecil Terence Ingold C.M.G., D.Sc., F.L.S., Hon. D.Litt. (Ibadan), Hon. DSc. (Exter), Hon. D.C.I. (Kent). 

Botanical Journal of the Linnean Society 91: vii–xv. 

Webster J (2005) Centenary of a mycologist: C. Terence Ingold. Mycological Research 109: 753–754. 

Webster J (2010) Obituary: Professor Terence Ingold. The Linnean 26 (3): 38–41. 

To honor the accomplishments of those who are the future of our field, the IMA Executive Committee has initiated IMA Young Mycologist 

Awards. Awards such as the IMA Young Mycologist Awards do more than honor the recipients, they also promote mycology by helping to 

advance young mycologists. The accomplishments that we celebrate can be no greater than the pool of nominees. Therefore, I encourage 

everyone in the IMA to take the time to consider candidates and initiate nominations. Note that there will be two sets of IMA Young Mycologist 

Awards, one for the past IMC9 and another for the upcoming IMC10. Note, also, that the award includes 500 euros. 

John W Taylor, President, IMA 

(jtaylor@berkeley.edu) 

IMA Young Mycologist Awards 

The IMA has established six IMA Young Mycologist Awards to mark outstanding research accomplishment by young mycologists from each 

of the six IMA Regional Mycological Organizations. 

• Ethel Mary Doidge Medal - African Regional Mycological Member Organization 

• Keisuke Tubaki Medal - Asian Regional Mycological Member Organization 

• Daniel McAlpine Medal - Australasian Regional Mycological Member Organization 

• Elias Magnus Fries Medal - European Regional Mycological Member Organization 

• Carlos Luis Spegazzini Medal - Latin American Regional Mycological Member Organization 

• Arthur Henry Reginald Buller Medal - North American Regional Mycological Member Organization 

Eligibility 

To be eligible to be nominated for an IMA 

Young Mycologist Award, a member of the 

IMA must, at the time of the next International 

Mycological Congress, be within ten years 

of his or her PhD degree, e.g. for IMC10 in 

2014, a nominee must have received his or her 

PhD degree no earlier than 2004. 

Establishment of Award Regional 

Committees 

The President of each IMA Regional Mycological 

Member Organization is asked to 

establish an IMA Young Mycologist Award 

Regional Committee consisting of at least 

two members (although the appointment of 

more members to represent the diversity of 

mycology in the region is encouraged) and a 

chair, who will serve from one International 

Mycological Congress to the next. To ensure 

a broad pool of nominees, nine months 

prior to the next International Mycological 

Congress, this committee will notify the 

members of the IMA Regional Mycological 

Member Organization of the award, solicit 

nominations and vote to select a candidate. 

To ensure a breadth of experience on the 

IMA Young Mycologist Award Regional 

Committee, the chair and members of this 

committee should be senior mycologists 

with distinguished records and should 

represent the diversity of Mycological 

Member Organizations and Sustaining 

Mycological Member Organizations from 

the region. The chair will vote only in the 

event of a tie vote by the other members. 

Responsibilities of the Award 

Regional Committees 

1. Nine months prior to the next International 

Mycological Congress and 

three months prior to the deadline for 

receiving nominations, members of each 

IMA Regional Mycological Member 

Organization shall be notified of the call 

for nominations for the relevant IMA 

Young Mycologist Award through the 

IMA website and the IMA Regional 

Mycological Member Organization 

website and through emails sent to 

delegates from the region who attended 

the preceding International Mycological 

Congress. 

2. Nomination of candidates for each IMA 

Young Mycologist Award shall consist 

of a letter of nomination, two letters of 

support from mycologists familiar with 

the candidate, and a current curriculum 

vitae of the candidate. Candidates 

are expected to have contributed 

appreciably to mycology and they 

should have achieved international 

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i m a f U N G U S

ecognition based on several criteria: (a) 

The quality, innovation, thoroughness, 

and impact on science of their published 

research, with consideration given to 

the contribution of the nominee to 

multi-authored publications. (b) Service 

as editors of journals or as officers of 

societies. (c) Membership on national 

or international policy committees. 

(d) Invitations to present research at 

national or international meetings. 

3. The committee will forward the nomination 

material for each nominee and 

the committee’s choice for the award 

to the IMA Young Mycologist Awards 

Committee convened by the IMA President 

no more than one month following 

the deadline for nominations and no less 

than five months prior to the next IMC. 

The IMA Young Mycologist Awards 

Committee is charged with ratifying the 

choices made by the IMA Young Mycologist 

Award Regional Committees. 

To reiterate, the timeline for the IMA 

Young Mycologist Awards, the notification 

to the members of the call for nominations 

shall be nine months prior to the IMC, the 

deadline for receipt of nominations by the 


Committees shall be six months prior to 

the IMC, and the receipt of the IMA Young 

Mycologist Award Regional Committee’s 

choice by the IMA Young Mycologist 

Awards Committee shall be five months 

prior to the IMC. In this way, recipients 

can be notified of their award at least four 

months prior to the IMC to encourage their 

attendance at the IMC. 

Presentation of the IMA Young 

Mycologist Awards 

The IMA Young Mycologist Awards for 

IMC9 and IMC10 will be presented by 

the President of the IMA at the Closing 

Ceremony of the International Mycological 

Congress. They shall consist of a certificate 

and 500 euros. 

Special instructions for the IMC9 

awards 

The IMA Executive Committee has voted 

to make awards for both IMC9 and the 

upcoming IMC10, both of which will be 

awarded at IMC10 in Thailand. To be 

eligible for the IMC9 awards, the nominee 

must have received his or her Ph.D. degree 

no earlier than 2000 and, for the IMC10 

award, no earlier than 2004. 

For the special case of the IMA 

Young Mycologist Awards for IMC9, 

each IMA Young Mycologist Award 

Regional Committee must be organized 

and announced to the IMA Executive 

Committee by May 1, 2011. The 

membership of each Regional Mycological 

Member Organization must be notified of 

the nomination procedure by June 1, 2011. 

Nominations must be received by each 


Committee by September 1, 2011. The 


Committees must communicate their 

nomination materials and their choice 

to the IMA Young Mycologist Awards 

Committee by October 1, 2011. The IMA 

Young Mycologist Awards Committee shall 

announce the awards by November 1, 2011. 

The awards will be presented at IMC10 in 

Thailand in 2014. 


Mycologists attending the general meeting and awards ceremony of the IMC9 in the Usher Hall, Edinburgh. 

v o l u m e 1 · n o . 2 

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CORRESPONDENCE 

A vision for the future of the ICTF 

David Hawksworth searching for a lichenicolous fungus on a Xanthoparmelia in Yosemite National Park, 

California, 8 November 2010. 

The International Commission on the 

Taxonomy of Fungi (ICTF) was established 

by the IUMS Division of Mycology at the 

XII International Microbiology Congress 

in Boston (MA, USA) in August 1982. At 

IMC5 in Vancouver in 1994, the ICTF decided 

that it it was appropriate for it to act 

as an inter-union body between IUMS and 

the International Union of Biological Sciences 

(IUBS) – the body in which the IMA 

is a scientific member; ICTF was accepted 

as an inter-union scientific member of IUBS 

at the IUBS General Assembly in Paris in 

September 1994. 

The ICTF was envisioned as a taxonomic 

complement to the nomenclatural 

Committee for Fungi which is established 

by each six-yearly International Botanical 

Congress, and which is concerned with formal 

nomenclatural proposals. The principle 

objectives of the new Commission were, in 

contrast, to foster interest and good practice 

in fungal taxonomy, to provide a mechanism 

to foster collaborative work on critical 

groups, and to identify areas of concern on 

nomenclatural matters amongst microbiologists 

for debate – and where appropriate 

transmit those views to the Committee for 

Fungi. 

The activities have included the production 

of a synopsis of changes in the International 

Code of Botanical Nomenclature 

relevant to fungi following the Sydney Congress 

(Hawksworth 1984), a list of genera in 

most need of taxonomic work (Hawksworth 

1986), a Code of practice for systematic mycologists 

(Sigler & Hawksworth 1987), and 

a series of papers drawing attention to (and 

explaining the reasons for) name changes 

in fungi of microbiological, industrial, and 

medical importance (Cannon 1986-90) 

– the short-lived name-changes series was 

made possible through financial support 

received from the IUMS Division of Mycology. 

The latest publication is the How to 

describe a new fungal species in this issue of 

IMA FUNGUS (Seifert & Rossman 2010). 

The ICTF has also been responsible for the 

organization of symposia during IMA and 

IUMS congresses on several occasions. The 

rôle of the ICTF as an umbrella organization 

for international collaborations on 

different groups of fungi, normally genera, 

has been especially successful with subcommissions 

working on various critical groups. 

Most productive have been those concerned 

with Aspergillus and Penicillium (formed 

1986; later an IUMS commission in its 

own right and produced a subsequently 

protected list of species names in current 

use; Pitt & Samson 1993), Fusarium (with 

the International Society for Plant Pathology), 

and Trichoderma (formed 1989). 

Less active have been those concerned with 

Ceratocystis/Ophiostoma, and Cladosporium/ 

Mycosphaerella. Suggestions for subcommissions 

on Alternaria, Colletotrichum, and 

Pythium sadly never progressed. However, 

the new subcommission on Fungal Barcoding 

(formed in 2009; FunBOL) has made a 

promising start. A report on recent activities 

is included in this issue of IMA FUNGUS 

(Seifert & Okada 2010). 

As to the future, the ICTF is currently 

considering its role, and how it could or 

indeed should develop in the coming years. 

As a start to this debate, and to open this 

discussion to a wider mycological community 

than those nominated at IMC9 to consider 

the matter, my vision is that the ICTF 

should in future be involved in: 

1. Overseeing existing, and being proactive 

in the formation of, subcommissions 

on a wider spectrum of fungi, including 

macromycetes and lichens – with a 

target of 10 being active by 2014, and 20 

by 2018. 

2. Providing annual summaries and explanations 

of the reasons for name changes 

in fungi of all groups which are of 

agricultural, ecological, environmental, 

industrial, or medical importance – in 

IMA FUNGUS and also on the IMA 

and IUMS Mycology Division websites. 

3. Developing a series of pragmatic how-to 

guidelines for good practice in systematic 

mycology – on the lines of How to 

describe a fungus. 

4. Developing a set of minimum standards 

for the description of new genera and 

species in particular groups – for distribution 

to all mycological journals. 

5. Organizing one or more Nomenclature 

Sessions during all future IMCs – to 

both debate and vote on formal proposals, 

and also to develop acceptable solutions 

to controversial issues. 

6. Organizing at least one symposium during 

all future IMCs and IUMS Division 

of Mycology congresses – these to be on 

exciting advances or controversial issues. 

7. Providing inputs of views on particular 

nomenclatural issues to the Committee 

for Fungi – when requested to do so or 

proactively. 

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i m a f U N G U S

8. Preparing agreed responses to International 

Botanical Congress Nomenclature 

Section meetings decisions on 

proposals related to mycological issues – 

for example the voting on governance of 

the Code and the mandatory deposit of 

nomenclatural information on new taxa 

at the 2011 Melbourne Congress. 

9. Developing a co-ordinated mycological 

response to the proposals for a BioCode 

to cover all groups of living and fossil 

organisms – a new draft (updated from 

1997), and prepared at the request of 

the IUBS General Assembly held in 

Cape Town in 2009, is due to be released 

by January 2011. 

10. Providing a mechanism to decide which 

of two or more competing generic 

names should be adopted for fungi 

with pleomorphic life-cycles – this may 

become necessary should proposals to 

move towards a single scientific name 

for each fungal species be formally 

adopted. 

11. Preparing reports of its activities annually 

to the IMA Executive Committee (for 

inclusion in IMA FUNGUS), and to 

each IMC, IUBS General Assembly, and 

IUMS Mycology Division congress – 

and making proposals for financial support 

whenever the opportunity arises. 

12. Advising the IMA Executive Committee 

on all matters concerned with the 

taxonomy of fungi – both at the request 

of the Executive Committee and proactively. 

13. Extending the number of ICTF members 

from 15 to 30 – to provide representation 

of a wider spectrum of fungal 

groups and with members delegated to 

be responsible for the furtherance of 

particular activities. 

This list is certainly daunting, and made 

without prejudice to the final decisions of 

the group of ICTF members mandated to 

address this issue, but 13 has always been a 

lucky number for me. In essence, the points 

represents a personal vision of the ICFT 

becoming the key over-arching body responsible 

for the furtherance of all aspects 

of fungal systematics that are not within the 

remit of the Committee for Fungi. To fully 

achieve this vision will take many years, but 

some points could be progressed in the short 

term. In this, the 13 th point is of critical importance. 

In my opinion, the principle reason 

for the ICTF not to have yet achieved 

its potential and realized its original vision 

has been that its officers and members have 

tended to be senior mycologists already 

over-committed with heavy personal and 

institutional duties. In consequence they, 

and not least me, have not been able to accord 

ICTF matters the priority they require. 

I personally feel guilty that I had not been 

able to do so much more during my term as 

ICTF Chair. 

The future of the ICTF, and the election 

of new members and officers, are scheduled 

to take place during the One Fungus: One 

Name (1F: 1N) in Amsterdam on 19–21 

April 2011. Do not be backward in coming 

forward to express your own views on 

the vision presented here, however critical 

or controversial you think they may be, to 

make new suggestions, or to volunteer to 

join or work in a rejuvenated Commission 

(or one of its less active or proposed Subcommissions). 

Responses should be sent 

to the current Chair (Keith Seifert; keith. 

seifert@agr.gc.ca) and Secretary (Gen 

Okada; okada@jcm.riken.jp) by 28 February 

2011 – in order to inform the working 

group established at IMC9 to develop 

its report on the future of the ICTF for the 

April meeting. 


ICTF Chair 1982–2000 


CORRESPONDENCE 

Cannon PF (1986–90) International Commission on the Taxonomy of Fungi (ICTF): Name changes in fungi of microbiological, industrial and medical importance. 

Parts 1–4. Microbiological Sciences 3: 168–171 (1986), 285–287 (1986), 5: 23–26 (1988); Mycopathologia 111: 75–83. 

Hawksworth DL (1984) Recent changes in the international rules affecting the nomenclature of fungi. Microbiological Sciences 1: 18–21. 

Hawksworth DL (1986) Fungal genera in urgent need of taxonomic work. Microbiological Sciences 3: 58. 

Pitt JI, Samson RA (1993) Species names in current use in the Trichocomaceae (Fungi, Eurotiales). Regnum Vegetabile 128: 14–57. 

Seifert KA, Okada G (2010) International Commission on the Taxonomy of Fungi (ICTF). IMA Fungus 1: (11)–(12). 

Seifert KA, Rossman AY (2010) How to describe a new fungal species. IMA Fungus 1: 109–116. 

Sigler L, Hawksworth DL (1987) International Commission on the taxonomy of Fungi (ICTF): Code of practice for systematic mycologists. Microbiological Sciences 4: 

83–86; Mycopathologia 99: 3–7; The Mycologist 21: 101–105; Acta Mycologica Sinica 8: 154–159 (1989). 

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RESEARCH News 

Carbonaceous spherules exposed as fungal sclerotia, 

are not evidence of the impact of a comet 

In the last few years, some palaeoecologists 

postulated that the onset of the 

Younger Dryas climate interval at ~ 

12,900 yBP was due to an air-bursting or 

impacting comet which caused intensive 

wildfires which raged across Europe and 

North America (Firestone et al. 2007; Kennett 

et al. 2008); this hypothesis sparked 

a controversial debate amongst palaeoclimatologists. 

Carbonaceous spherules (and 

similar “elongates”) found around the 

horizon have been interpreted as evidence 

of intensive wildfires that the postulated 

comet initiated. However, morphologically 

and anatomically identical spherules 

have subsequently also been discovered 

not only in earlier and later palaeoecological 

preparations, but further in modern 

heathland sites where there had been recent 

fires (notably Thursley Bog in Surrey, UK). 

When fractured and studied by scanning 

electron microscopy (SEM), the internal 

structures of the sphaerules appeared to be 

pseudoparenchymatous throughout with 

the exception of a cellularly differentiated 

outer cortex-like layer. The structures were 

consequently unquestionably biological 

in origin, but ascertaining what organism 

present today they belonged to proved elusive. 

Now, the question has been answered 

by the careful SEM studies of Scott et al. 

(2010) who compared the structure of the 

spherules with that of the sclerotia of species 

of Cenococcum and Sclerotium – both fresh 

and more significantly experimentally charred 

by fire. The superb illustrations in the 

paper and accompanying online material 

leave no doubt that the implicated carbonaceous 

spherules are fungal sclerotia, while 

the “elongates” were sclerotia or possibly in 

some cases coprolites. Especially illuminating 

was the discovery that at 350 0C the 

outer cortical cells tended to coalesce, and at 

450 0C voids developed inside the sclerotia. 

Structures that had previously been termed 

“nanodiamonds” were also evident in the 

charred sclerotia. 

The degree of surface reflectance of the 

structures and sclerotia was also measured 

as that is known to be indicative of the 

temperatures to which organic material 

has been subjected. As a consequence of 

the structural comparisons and reflectance 

data, Scott et al. conclude that these 

carbonaceous spaerules and many of the 

“elongates” are fungal sclerotia which have 

been subjected to low-intensity burning, 

and are not indicative of high-intensity fires 

as had previously been postulated; i.e. the 

putative evidence for the catastrophic explanation 

of the onset of cooling and associated 

megafaunal extinctions around 12,900 yBP 

is unsound and alternative explanations 

must be sought. 

Margaret E Collinson and Andrew C Scott 

kindly prepared the accompanying figure 

especially for this item. 

A. Modern charred fungal sclerotium (Thursley, Surrey, UK). B. SEM of section through charred fungal 

sclerotium from the Younger Dryas, 12 900 Cal BP (Santa Rosa Island, CA, USA). C. SEM of section through 

uncharred Cenococcum (Peace River Canada; photo: A Heiss). D. SEM of section through charred sclerotium 

of the genus Sclerotium, charred at 350 0C for 5min. Bars: A = 200 µm, B = 50 µm, C–D = 20 µm. 

Firestone RB et al. (2007) Evidence for an extraterrestrial impact 12,900 years ago that contributed to the megafaunal extinctions and the 

Younger Dryas cooling. Proceedings of the National Acedemy of Sciences, USA 104: 16016–16021. 

Kennett DJ, Kennet JP, West GJ, Erlandson JM, Johnson JR, Hemdry IL, West A, Culleton BJ, Jones TL, Stafford TW jr (2008) Wildfire 

and abrupt ecosystem disruption on California’s Northern Channel Islands at the Allerød—Younger Dryas boundary (13.0-12.9 ka). 

Q uarterly Science Review 27: 2350–2545. 

Scott AC, Pinter N, Collinson ME, Hardiman M, Anderson RS, Brain PR, Smith SY, Marone F, Stampanoni M (2010) Fungus, not 

comet or catastrophe, accounts for carbonaceous spherules in the Younger Dryas “impact layer”. Geophysical Research Letters 37: . 

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i m a f U N G U S

Physiological differences between wet- and drydistributed 

conidia 

The distinction between fungi which 

produce conidia in slimy heads 

(“wet-spored” and waterborne) and 

those which form them in powdery masses 

or chains (“dry-spored” and airborne) has 

been recognized as of fundamental importance 

in the characterization of genera since 

the early 20 th century. Now, some clues as 

to the basis of this distinction have been generated 

by van Leeuwen et al. (2010). The 

freshly harvested condia of two wet-spored 

(Fusarium oxysporum and Lecanicillium 

fungicola 1 ) and two dry-spored species (Aspergillus 

niger and Penicillium discolor) were 

compared physiologically using electron 

spin resonance (ESR) to test for cytoplasmic 

viscosity and staining with the fluorescent 

dye filipin to indicate the presence of ergosterol. 

Striking differences were found; the 

wet-spored species had lower cytoplasmic 

viscosity than the dry-spored pair, and the 

staining showed ergosterol was present in 

the plasma membrane only of the wetspored 

ones. Whether this correlation can 

be used as a generalization must, however, 

await studies on a much more diverse range 

of fungi. 

RESEARCH NEWS 

Leeuwen MR van, Doorn TH van, Golovina EA, Stark J, Dijksterhuis J (2010) Water- and 

air-distributed conidia differ in sterol content and cytoplasmic microviscosity. Applied and 

Environmental Microbiology 76: 366–369. 

1 

The authors used the name Verticillium fungicola, but that species has been transferred to 

Lecanicillium (Zare R, Gams W, Mycological Research 112: 818, 2008). 

Formation of conidia by Aspergillus niger (top) and Fusarium oxysporum (bottom) observed by scanning cryo-electron microscopy. Numerous conidia of A. niger are 

formed on erect conidiophores while conidia of F. oxysporum are formed within the mycelium. Bars = 10 µm. 

Mobile chromosomes: the clue to pathogenicity in 

Fusarium species 

Fungal chromosomes are notoriously 

difficult to visualize, although a few 

mycologists have been able to achieve 

this, including Punithalingam (1975) on 

Fusarium. Now it appears that fungal cytology 

could have provided the clue to why 

some fusaria are pathogenic and some are 

not. Ma et al. (2010) sequenced genomes in 

F. oxysporum f. sp. lycopersici and F. verticilliodes, 

and compared the results with those 

of the genome of F.graminearum that had 

previously been sequenced. The three species 

have different numbers of chromosomes: 

15 in F. oxysporum, 12 (11 mapped) in F. 

verticilloides, and only four in F. graminearum. 

It was found that the four additional 

chromosomes in F. oxysporum were the ones 

which were rich in transposons and genes 

related to pathogenicity. That these chromosomes 

were indeed the clue to enhanced 

pathogenicity was shown experimentally 

v o l u m e 1 · n o . 2 

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RESEARCH News 

within F. oxysporum by transferring two of 

these chromosomes from a pathogenic to 

a non-pathogenic strain by co-incubation; 

that whole chromosomes were involved 

was demonstrated by electrophoresis. 

The non-pathogenic strain then became 

pathogenic to tomato. Over 28 % of the 

entire F. oxysporum genome was composed 

of repetitive sequences, and while only 

20 % of those only found in this species 

could be functionally classified, they were 

evidently rich in ones considered related to 

secreted effectors and virulence factors. The 

results were also compared with the recently 

published genome of F. solani, which has 

three “supernumerary” chromosomes 

(Coleman et al. 2009) which lacked the 

suites of genes involved in the pathogenicity 

of F. oxysporum. As F. graminearum and F. 

verticilloides have Gibberella teleomorphs, F. 

solani one in Haematonectria (not Nectria 

as wrongly stated in the Nature paper, 

and also by Coleman et al. 2009), and F. 

oxysporum f. sp. lycopersici none (though 

one in Gibberella would be expected as it 

clusters with F. verticilloides), the difference 

with F. solani might have been no surprise 

to a taxonomist. Perhaps the most exciting 

aspect of this study, but one that should 

cause concern to those involved with food 

security, is the ease with which pathogenicity 

was transferred by whole chromosome 

movement between F. oxysporum isolates 

growing together. It also shows that when 

possible unexpected pathogenicity emerges 

in fungi that it could be advantageous to 

try pulse-gel electrophoresis to see whether 

extra chromosome transfer was involved as 

a prelude to embarking on similar complex 

molecular high cost studies – this one 

involved 63 co-authors distributed through 

25 laboratories . . . . 

Coleman JJ et al. (2009) The genome of Nectria haematococca: contribution of supernumerary chromosomes to gene expression. PLos 

Genetics 5: e10000618. 

Ma L-J et al. (2010) Comparative genomics reveals mobile pathogenicity chromosomes in Fusarium. Nature 464: 367–373. 

Punithalingham E (1975) Cytology of some Fusarium species. Nova Hedwigia 26: 275–303. 

Ectomycorrhizal symbiosis in basidio- and 

ascomycete fungi have different origins 

Whole genome studies are 

remarkable in what unexpected 

information can emerge. 

The sequencing of the whole genome of 

the Périgord truffle (Tuber melanosporum) 

by French and Italian researchers (Martin 

et al. 2010) is no exception, First, at ~ 125 

megabases it is the largest by far of all fungal 

genomes sequenced to date. Second, the size 

is a result of a proliferation of transposable 

elements that account for 58 % of the size 

and not a consequence of whole-genome 

duplication. Third, it lacks large sets of 

carbohydrate-cleaving enzymes, but has 

some involved with plant cell wall degradation 

that are induced in symbiotic tissues. 

And fourth, most fascinatingly, comparison 

with whole genome data from the ectomycorrhizal 

basidiomycete Laccaria bicolor 

revealed that strikingly distinct proteomes 

were encoded; i.e. that different “molecular 

toolkits” had evolved to enable these two 

fungi to be become ectomycorrhizal. This 

implies that the ectomycorrhizal habit 

has evolved independently in asco- and 

basidiomycete fungi – just as appears from 

phylogenenetic studies to have been the case 

with lichenization. 

Martin F et al. (2010) Périgord black truffle genome uncovers evolutionary origins and mechanisms of symbiosis. Nature 464: 1033–1038. 

A basal bryophilous fungus associates with 

cyanobacteria 

For almost 40 years, Peter Döbbeler 

has, almost single-handedly, been 

revealing the enormous variety and 

abundance of specialized ascomycetes that 

live on bryophytes. The phylogenetic position 

of many of the genera he discovered 

has been obscure, but now, in collaboration 

with Finnish mycological colleagues, a fivegene 

phylogeny has been constructed for 

an amazing 61 bryosymbionts. DNA was extracted 

from cultures prepared from freshly 

collected specimens – in itself a major achievement. 

As expected, the fungi were found 

to be dispersed through different ascomycete 

lineages, including Dothideomycetes, Eurotiomycetes, 

Lecanoromycetes, Leotiomycetes, 

Pezizomycetes, and Sordariomycetes. Most of 

the fungi are commensals, weak parasites, or 

saprobes, and “symbiont” is used here in the 

original sense of organisms living together 2 , 

whether mutualists or not. The results suggest 

that bryophily has arisen at different 

times in different lineages of ascomycetes. 

However, one of the fungi is of particular 

phylogenetic and biological interest: the newly 

described Trizodia acrobia which inhabits the 

living shoot apices of at least eight Sphagnum 

2 

The term “symbiosis” is attributed here to an 1879 publication of de Bary, but actually originated from one of AB Frank of 1877 who studied 

lichen anatomy; viz Hawksworth (Nature 374: 841, 1995) and Sapp (Evolution by Association: a history of symbiosis, 1994). Frank went on to 

coin the term “mycorrhiza” in 1887. 

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i m a f U N G U S

Trizodia acrobia, hyphae enveloping cyanobacterial 

colonies on the leaf of Sphagnum. Photo: Tomi 

Laukka. Bar = 30 µm. 

species in Finland. The fungus has minute 

convex ascomata which are translucent 

white to pale pink when fresh, inoperculate 

unitunicate asci, and single-celled colourless 

ascospores. Trizodia appeared as a basal clade 

in Leotiomycetes, and is of special interest 

biologically because it is associated with Nostoc 

cyanobacteria as well as Sphagnum. The genus 

is not “categorized as a lichen per se [as] it does 

not form organized thallus structures” (p. 16), 

but that is also the situation in some of the 

fungi traditionally studied by lichenologists, 

especially pyrenocarpous species on bark! 

Amongst other fungi that would fall in this 

category is Nectria phycophora (syn. Calonectria 

phycophora), which occurs on the moss 

Dawsonia grandis and has pockets of algal 

cells in the perithecial wall (Döbbeler 1978, 

Hawksworth 1988). Many fungi do not fit 

neatly into biological categories conceived by 

humans, and the authors describe this as “a 

primitive form of a bryosymbiotic pyrenolichen” 

– but does that matter? It is the fungi 

themselves and the diversity of their intimate 

associations that are so fascinating – evidently 

not least amongst the bryosymbionts. 

This is a most elegant study that now 

needs to be emulated for the obligately 

lichenicolous fungi – which sadly prove 

more recalcitrant to culture and yield their 

DNA than those that are bryophilous. 

RESEARCH NEWS 

Döbbeler P (1978) Moosbewöhnende Ascomyceten I. Die pyrenocarpen, den gametophyten besiedelnden Arten. Mitteilungenaus der botanischen 

Staatsammlung, München 14: 1–360. 

Hawksworth DL (1988) The variety of fungal-algal symbioses, their evolutionary significance, and the nature of lichens. Botanical Journal of 

the Linnean Society 96: 3–20. 

Stenroos S, Laukka T, Huhtinen S, Döbbeler P, Myllys L, Syrjänen, Hyvönen J (2010) Multiple origins of symbioses between ascomycetes and 

bryophytes suggested by a five-gene phylogeny. Cladistics 26: 281–300. 

Molecular clocks and evolutionary rates 

The evolution of fungi is attracting 

increasing interest as the discovery 

of fossils and the production of 

phylogenetic trees with representatives of 

increasing numbers of families and orders 

accelerates – with earlier and earlier dates 

being suggested for major divergences. For 

example, Ascomycota and Basidiomycota 

could have diverged as early as 1.2 Byr ago 

(Heckman et al. 2001). Berbee & Taylor 

(2010) have now revisited this question, 

one that has fascinated them since the early 

1990s. They used the BEAST program to 

apply a relaxed lognormal clock analysis to 

a data set comprising 50 loci from 26 taxa 

and then endeavoured to ground-truth 

nodes on the basis of assumptions made as 

to the systematic position of some wellresearched 

fungal fossils of known age. The 

assumptions on position had a dramatic 

effect. In the case of Palaeopyrenomycites 

devonicus, if it can be placed in Ascomycota 

but no further the minimum age of the 

phylum (and its sister phylum) would be 

just 452 Myr. However, if is considered 

to be assignable to Pezizomycotina the 

probable date for the divergence to 843 

Myr. And most strikingly if it is in Sordariomycetes, 

as seems very likely, that pushes 

the Ascomycota/Basidiomycota divide back 

to 1489 Myr – a result not so different from 

the figure computed by Heckman et al. Some 

of the most potentially intriguing fossils that 

could be pertinent are unfortunately difficult 

to interpret in modern terms, for example 

the Lower Devonian Prototaxites and the 900 

Myr-old Proterozoic Tappania. Further, if 

the Silurian hyphae and especially conidumlike 

structures reported by Sherwood-Pike 

& Gray (1985) could be pinned-down to 

modern taxa, that could materially change 

the picture pushing orders even further back. 

Berbee & Taylor recognize that the rates 

of divergence, i.e. the speed of the molecular 

clock, could well vary in different fungal 

groups. This issue has been examined by 

Wang et al. (2010) who studied the rates of 

substitution in 21 protein-coding genes in 

the lichen-forming Rhizoplaca chrysoleuca 

and used them to test for discrepancies in 

rate between Ascomycota and Basidiomycota 

by comparing them with sequences in 

GenBank – in all 299 taxa were considered, 

13 of which had data on 105 protein-coding 

genes. Significant differences emerged 

between the phyla, the higher rate being 

in Ascomycota. Within Ascomycota, the 

fastest evolutionary rates detected were in 

Sordariomycetes. They speculate that the 

high level of speciation in the class is due to 

founder-effects rather than the development 

of mutualisms, ecological adaptations, 

shorter generation times or metabolic rates; 

v o l u m e 1 · n o . 2 

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RESEARCH News 

i.e. isolated populations undergoing rapid 

change as a result of genetic drift. 

This is becoming a fascinating area of 

enquiry which would benefit from much 

more involvement of palaeontologists with 

both morpho-mycologists and molecular 

systematists. For example, there are enormous 

numbers of fossil fungal propagules 

now documented and given scientific names 

(Kalgutkar & Jansonius 2000), which are 

rarely considered by mycologists . . . . . 

Berbee ML, Taylor JW (2010) Dating the molecular clock in fungi – how close are we? Fungal Biology Reviews 24: 1–16. 

Heckman DS, Geiser DM, Eudell BR, Stauffer RL, Kardos NL, Hedges SB (2001) Molecular evidence for the early colonization of land by 

fungi and plants. Science 293: 1129–1133. 

Kalgutkar RM, Jansonius J (2000) Synopsis of Fossil Fungal Spores, Mycelia and Fructifications. [AASP Contributions Series No. 39.] Dallas, 

TX: American Association of Stratigraphic Palynologists. 

Sherwood-Pike MA, Gray J (1985) Silurian fungal remains: probable records of the class Ascomycetes. Lethaia 18: 1–20. 

Wang H-Y, Guo S-Y, Huang M-R, Lumbsch HT, Wei J-C (2010) Ascomycota has a faster evolutionary rate and higher species diversity than 

Basidiomycota. Science China (Life Sciences) 53: 1163–1169. 

Numbers of fungi in China 

Dai & Zhuang (2010) have systematically 

scoured publications 

dealing with fungi recorded from 

China. They located reports of 16 046 species 

and 297 varieties from the Chinese territories 

as a whole; the mainland had 14 846 

species reported, Taiwan 6 207, and Hong 

Kong 2 122. The overall total included 300 

chromistan (straminipilous) fungi, and 340 

protozoan fungi (myxomycetes). However, 

as far as I can gather from the Chinese text 

and especially the literature cited, these 

totals omit the lichen-forming species. An 

unfortunate oversight which would have 

added at least 1766 species to the Chinese 

mainland total (Wei 1991), 261 to that for 

Hong Kong (Aptroot & Seaward 1999), 

and 559 to that for Taiwan (Lai 2000); allowing 

for species in common to these three 

regions the known lichens actually amount 

to some 2200 species (J Wei in litt.). This 

oversight is most unfortunate as the names 

applied to lichen associations strictly refer 

to the fungal partners alone! 

As to the number of fungi to be 

expected in a total inventory for China, as 

32 200 plants are known in China according 

to the World Resources Institute database 

portal (), the now 

often cited 6:1 model (Hawksworth 1991) 

would suggest the real figure could be as 

much as 193 200 species. This would imply 

that only around 9 % of the fungi of China 

have yet been recognized and catalogued. 

The authors note that 2849 species 

new to science and 5260 new records of 

non-lichenized fungi were added over the 

years 1978–2010, i.e. the rate of accrual 

was 89 and 164 species per year respectively. 

However, they suggest that the totals of new 

taxa be reduced because of possible synonymy, 

proposing a reduction of 10 % – much less 

than the 70 % that would be predicted by 

the 1:2.6 accepted species to synonym ratio 

derived from an analysis of 15 monographs of 

fungal genera (Hawksworth 1992). Such an 

estimate is perhaps pessimistic as the 1:2.6 ratio 

covered the whole period since 1753 when 

formal fungal nomenclature is now considered 

to have started, and does not allow for the 

diligence of mycologists in their endeavours 

to discover possible candidate names in often 

obscure literature. However, Yu-Chen Dai (in 

litt.) informs me that he used the 10 % figure 

because “many species are complex, and in fact 

more species inside a single name”. 

There has been an accelerating interest in 

fungal systematics in China since the 1980s, 

largely stimulated by the vigorous Systematic 

Mycology and Lichenology Laboratory of 

the Institute of Microbiology of Academia 

Sinica in Beijing. Nevertheless, it is evident 

that huge additional resources need to be 

devoted to inventorying fungi in China to 

raise the level of knowledge to a level approaching 

that in European countries. While such 

investment would be unthinkable in western 

countries today, perhaps that need not be 

dismissed out of hand so readily in China, 

considering the enormous resources now 

being placed on molecular sequencing in the 

country through the BGI genomics centre 

in Beijing () – established 

in 1999 and set to become the world leader 

(Cyranoski 2010). Dai & Zhuang’s synopsis 

of the numbers of fungi known in China 

deserves to prompt and stimulate discussions 

within China as to how the description of the 

mycobiota of the country can be accelerated. 

I am indebted to Yu-Cheng Dai and 

Jiang-chun Wei for the supplementary 

information they kindly supplied while this 

note was being prepared. 

Aptroot A, Seaward MRD (1999) Annotated checklist of Hongkong lichens. Tropical Bryology 17: 57–101. 

Cyranoski D (2010) The sequence factory. Nature 464: 22–24. 

Dai Y-C, Zhuang J-Y (2010) Numbers of fungal species hitherto known in China. Mycosystema 29: 625–628. 

Hawksworth DL (1991) The fungal dimension of biodiversity: magnitude, significance and conservation. Mycological Research 95: 641–655. 

Hawksworth DL (1992) The need for a more effective biological nomenclature for the 21 st century. Botanical Journal of the Linnean Society 

109: 543–567. 

Lai M-J (2000) Lichen Checklist of Taiwan. Taichung: M-J Lai. 

Wei J-C (1991) An Enumeration of Lichens in China. Beijing: International Academic Publishers. 

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i m a f U N G U S

International Society for Fungal Conservation 

At a special meeting on 6 August 

2010 at the Royal Botanic Garden, 

Edinburgh, mycologists from over forty 

countries established the International 

Society for Fungal Conservation. For 

the conservation movement, this was 

an event of enormous and historical 

significance. Almost unbelievably, this 

new Society appears to be the first 

anywhere in the world exclusively and 

explicitly devoted to protecting fungi. 

The present note is a short account of 

how the Society came to be formed, 

and what it hopes to achieve. For information 

about how to join the Society, 

please visit its website (). 

Fungi: the orphans of Rio 

The challenges for fungal conservation 

are daunting. As nature's recyclers, 

fungi are like municipal refuse 

collectors employed to take away our 

rubbish. We don't notice them until 

they go on strike. The well-being of 

fungi is necessary for sustainable life 

on this planet. Scientists have known 

for over 100 years that, like animals 

and plants, fungi too are affected by 

the destructive activities of mankind. 

The impact of air pollution on lichenforming 

fungi is one particularly well 

documented example. Although there 

is still insufficient information about 

the conservation status of fungi, there 

is no reason to suppose that fungi are 

any less vulnerable than other groups 

of organisms to habitat loss and climate 

change. The topic is far too important 

to ignore. 

Cryptomyces maximus: provisionally assessed globally 

as critically endangered. IUCN Species of the Day, 

20 September 2010 (). Photo: David 

Harries. 

Public awareness of their importance 

is, however, very low, not least 

because biodiversity - the full and 

wonderful diversity of life - is still 

widely portrayed as "flora and fauna" or 

"animals and plants". These are lazy and 

misleading descriptions. Those responsible 

include a range of major biological 

institutions and learned societies which 

should and do know better. The five 

kingdom classification of life, which 

recognizes fungi in a kingdom of their 

own, has been generally accepted by 

scientists since at least 1970. 

The broader conservation movement, 

as a result, remains largely 

unaware of the need to conserve fungi. 

Priority habitats for conservation, 

such as biodiversity hotspots, are 

almost always defined on the basis 

of bird, mammal and flowering plant 

diversity. This means that habitats 

rich in fungal diversity are missed 

and remain unprotected. Most nature 

reserve management plans do not take 

fungi into account. Fungi (for example 

host-specific species known only on 

rare endemic plants) are often treated 

as part of the problem (a threat to the 

plant) rather than recognized as themselves 

being in need of protection. In 

many countries there is no explicit legal 

protection for fungi. 

This failure to take fungi into 

account spilled over into the Rio 

Convention on Biological Diversity 

[CBD]. Laudibly, it claims to protect 

all forms of life, but its text defines 

biodiversity as "animals, plants and 

micro-organisms", i.e. two taxonomic 

kingdoms and a third category by size. 

Fungi belong in neither the animal nor 

plant kingdom and, as they include in 

their number the largest single living 

individual known on earth, far bigger 

than the blue whale or any of the great 

redwoods, they can hardly be described 

as micro-organisms. They simply do 

not fit any of the CBD categories. As 

a result, many national biodiversity 

action plans produced as a result of 

that convention fail to consider fungi 

at all. The few which do, treat them 

as "lower plants" - an obscure corner 

of botany. As David Hawksworth, 

Poronia punctata: provisionally assessed globally as 

vulnerable. IUCN Species of the Day, 21 August 

2010 (). 

Photo: David Minter 

one of the world's leading mycologists 

so eloquently put it, fungi are "the 

orphans of Rio". 

The start of the modern fungal 

conservation movement 

Fungal conservation is in its infancy. 

The European Council for Conservation 

of Fungi (now the fungal conservation 

group of the European Mycological 

Association) was established in Oslo in 

1985, and that event marked the start 

of the modern fungal conservation 

movement. Thereafter, specialist 

groups for "lichens" and "fungi" were 

set up in the Species Survival Commission 

of the International Union for 

Nature Conservation (IUCN), the 

Australasian Mycological Society formed 

a continental-level fungal conservation 

group, and the ground-breaking 

volume Fungal Conservation: issues and 

solutions (Moore et al., eds, Cambridge 

University Press, 2001) drew worldwide 

attention to the topic. 

Recent developments 

To date, mycologists have concentrated 

on gathering scientific evidence of 

population declines. But fungal conservation 

is more than just scientific 

evidence. It also has a political dimension: 

the movement must use such 

scientific evidence ("fungal populations 

are declining") in political activities 

("something needs to be done about 

it") to promote policy changes which 

will result in better protection for 

fungi. The political dimension makes it 

important to keep the science separate 

SOCIETY AND ASSOCIATION NEWS 

v o l u m e 1 · n o . 2 

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SOCIETY AND ASSOCIATION News 

Zeus olympius: provisionally assessed globally as critically endangered. IUCN Species of the Day, 21 February 

2010 (). Photo: David Minter. 

from the politics. The science needs to 

be protected from the unscrupulous 

verbal abuse which is so often an 

unfortunate side-effect of lobbying. To 

achieve this, the movement for fungal 

conservation must have some basic 

infrastructure, including components 

independent from learned societies. 

There must also be an awareness of 

how to work in the political arena. In 

this respect, important and exciting 

developments have occurred in fungal 

conservation over the past few years. 

• November 2005. A pioneering 

workshop was organized by the 

European Council for Conservation 

of Fungi in Córdoba, Spain. 

This was one of the earliest uses 

(perhaps the first) for fungi of the 

IUCN conservation status evaluation 

system. 

• September 2007. Three prototype 

specialist committees were 

established for conservation of 

fungi inadequately covered by the 

IUCN's specialist groups of that 

time: 

1. Mildews, Moulds & Myxomycetes 

(). 

2. Non-lichen-forming Ascomycetes 

(). 

3. Rusts & Smuts (). 

• November 2007. The Sociedade 

Brasileira de Micologia established 

a national fungal conservation 

group, perhaps the first in South 

America. 

• December 2007. At an international 

meeting in Spain on 

conservation and sustainable use of 

fungi, the Declaration of Córdoba 

() 

was 

published by over 150 mycologists 

from 35 countries - a first attempt 

to establish global principles for 

fungal conservation. 

• August 2008. The Mycological 

Society of America established a 

continental-level fungal conservation 

group for North America. 

• November 2008. The Asociación 

Latino-Americana de Micología 

established a working party to 

set up a continental-level fungal 

conservation group for South 

America. 

• January 2009. The African Mycological 

Association established a 


group for Africa. 

• February 2009. The Species 

Survival Commission of the IUCN 

formally recognized fungi as 

needing fully separate representation 

within the Commission's 

structure. The Commission also 

decided to increase the number 

of Specialist Groups representing 

fungi to five by adopting the 

prototype specialist committees 

described above, and re-defining 

the old "Fungal Specialist Group" 

so that it henceforth covered the 

larger basidiomycetes. 

• October 2009. At an international 

meeting in the UK (), the 

Chairs of the five IUCN Species 

Survival Commission fungal 

Specialist Groups, together with 

representatives from each continental 

level fungal conservation 

group, and mycologists from over 

20 countries appointed a Steering 

Committee to prepare proposals 

for a federation of fungal conservation 

groups. 

• November 2009. The Mycological 

Committee for Asia, at its Asian 

Mycological Congress established a 


group for Asia. 

Collectively, these events constituted 

huge progress for fungal conservation. 

In geographical terms within every 

continental-level mycological learned 

society, there is now a conservation 

group in existence or at least in preparation, 

and therefore an identifiable 

cadre of people interested in fungal 

conservation. Conservation groups 

have also started to appear in national 

mycological learned societies, and even 

in one or two places at a local level. 

In terms of taxonomic coverage the 

reform and enlargement of the IUCN 

Species Survival Commission fungal 

specialist groups mean that there are 

now identifiable teams responsible for 

promoting conservation of all fungal 

groups. By the start of 2010, the one 

missing component was a general 

society specifically dedicated to fungal 

conservation, which could engage 

support from the general public, draw 

all of these elements together, and take 

responsibility for the political lobbying 

which conservation always entails. 

The new society is established 

IMC9 in August 2010, gathered mycologists 

from all over the world. This 

provided an excellent opportunity for 

meetings beyond the formal and purely 

scientific congress programme. The 

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i m a f U N G U S

steering committee mentioned earlier, 

appointed in October 2009, produced 

proposals for a new society for fungal 

conservation, and a special meeting was 

convened to discuss those proposals. 

The Royal Botanic Garden Edinburgh 

generously hosted the event. Nearly 60 

mycologists from 21 different countries 

attended, with messages of interest 

and support from about ninety more, 

taking the total number of countries 

represented to over forty. In addition, 

there were messages of support from a 

range of learned societies, NGOs and 

national representatives of the CBD 

Subsidiary Body on Scientific, Technical 

and Technological Advice (the scientists 

who advise the Rio Convention). 

After animated discussion, there was 

overwhelming support to set up the 

International Society for Fungal Conservation. 

Time constraints made it impossible 

to agree all of the draft constitution 

which had been circulated, but 

enough was agreed to determine the 

name of the Society, its status, address, 

objectives and activities, and to establish 

English as the Society's working 

language. A new steering committee 

was appointed to carry out the remaining 

work needed to bring the Society 

to an operational state. The composition 

of that Steering Committee 

is as follows: Peter Buchanan (New 

Zealand), Sharon Cantrell (Puerto 

Rico), Marieka Gryzenhout (South 

Africa), Greg Mueller (USA), David 

Minter (UK, co-ordinator), and 

Tatyana Svetasheva (Russia). The 

Steering Committee is now occupied 

in revising the draft constitution so 

that it can be circulated to Founder 

Members for comment and, hopefully, 

eventual adoption. As soon as possible, 

the minimum officers necessary for the 

Society to function will be appointed, 

and activities will begin. Even at this 

early stage - particularly at this early 

stage - new members are very welcome. 

Likely activities of the new society 

Until the constitution is adopted, and 

a system of governing the Society has 

been set up, there can be no formal 

policy within the Society. The following 

comments are therefore merely 

speculative and tentative ideas on 

general directions. They are far from 

exhaustive, but it is already clear that 

the Society will need to work in at 

least four general areas: infrastructure, 

education, science and politics. 

Infrastructure. The Society will 

need to start fund-raising. Little can be 

achieved without money. The Society 

must promote and support efforts to 

establish a network of conservation 

societies for fungi working at different 

levels (continental, national and local). 

At present, fungal conservation is 

supported mainly by field mycologists 

and taxonomists. The Society needs to 

raise awareness among other mycologists, 

for example experimental mycologists, 

that their work is relevant to 

the movement, and that they too have a 

responsibility to promote conservation. 

The Society should also seek to raise 

awareness among curators of fungal 

culture collections of the important 

role these bodies have for ex situ 

fungal conservation. The Society will 

also seek to establish links with other 

organizations promoting conservation 

of overlooked and under-valued groups 

of organisms, so that experience and 

resources can be pooled. 

Education. The Society will work 

with learned mycological societies to 

raise general public awareness of the 

importance of fungi, to promote the 

teaching of mycology at all levels in 

education, and to develop educational 

websites and other resources appropriate 

for that objective. 

Science. The Society will work to 

identify, classify and publicise threats 

to fungi, and to identify important 

areas for fungi (fungal hotspots 

and coldspots), important fungal 

associated organisms, and impacts on 

human society which may occur as a 

result of fungal population declines 

and extinctions. The Society will 

furthermore promote the message that, 

without taking fungi into account, the 

ecosystem approach to conservation 

is so severely compromised as to be 

invalid. This will entail raising awareness 

that fungi are essential components 

of ecosystems. 

Politics. The Society will develop 

policy, and will develop political expertise 

where possible by learning from 

the experiences of other conservation 

societies. The Society will seek to raise 

awareness of fungi among the CBD 

National Focus Points, and will also 

seek to engage governments which are 

not signatories to the CBD, making 

them aware of the importance of fungal 

conservation. The Society will seek 

to raise the profile of fungi, in part 

through a campaign to encourage biological 

institutions and learned societies 

to ensure that the language used in 

their promotional material properly 

reflects the true importance of fungi. 

This will, for example, entail discouraging 

language which results in confusion 

of fungi with plants ("botany" does 

not include "mycology", fungi are not 

"lower plants", they are not part of a 

"flora" etc.). Use of "biodiversity" as 

shorthand for "animals and plants" will 

also be discouraged. 

The Society will also work to 

promote representation by mycologists 

on bodies concerned with biodiversity 

and conservation. If fungi are the 

"orphans of Rio", then mycology, 

like an orphan, enjoys little of the 

family wealth (mycologists are usually 

hidden away in obscure departments 

of botanical institutions, getting a very 

small share of resources), and mycology 

is rarely consulted on family matters 

by the biological sciences. Biodiversity 

initiatives should as a matter of course 

involve mycologists as equal playes 

from their inception. At present, in 

general, they don't. 

Conclusions 

As already stated, it is almost unbelievable 

that up to now there has been no 

society explicitly devoted to protecting 

fungi. Fungal conservation is far too 

important to be left to chance. The 

challenges are daunting. Establishing 

this new Society was an important 

and historic event in the conservation 

world. The Society needs enthusiastic 

support from all those who understand 

the pressing need to protect the 

"orphans of Rio". 

David W. Minter President, European 

Mycological Association 

(Whitby, UK; d.minter@cabi.org) 


v o l u m e 1 · n o . 2 

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SOCIETY AND ASSOCIATION News 

Mycological Society of America 

The Mycological Society of America had its 2010 annual meeting 

among the gently rolling hills of the inner Bluegrass Region of the 

eastern United States - Lexington, KY, the “horse capital of the 

world.” This was a joint meeting with the International Symposium 

on Fungal Endophytes of Grasses (ISFEG), a society that we had not 

met with in recent years. Both societies have many common interests 

and it was an excellent opportunity for the two groups to interact. 

The theme of the meeting was “Symbiosis.” 

The MSA pre-meeting foray took mycologists to the Bernheim 

Arboretum and Research Forest in Western Kentucky. This is a 

private educational facility dedicated to the restoration and preservation 

of Kentucky’s native ecosystems, including forests, wetlands and 

grasslands. A wide array of fungi was found despite the dry weather. 

It was wonderful to visit with friends and colleagues among the trees. 

The invitational Karling Lecturer was Sally Smith, an excellent 

speaker and powerful researcher in the field of mycorrhizal symbioses. 

She gave a charming overview of her work on phosphorous 

transfer in arbuscular mycorrhizae. The keynote speaker of ISFEG 

was Charles W. Bacon who gave a review of the basic biology of 

forage grass fungal endophytes. Symposia included “Impacts of 

endophytes on host plant ecology and biotechnology,” “The role of 

stress response mechanisms in symbiotic and pathogenic associations 

of fungi,” “Advances in DNA barcoding for fungi,” and “Molecular 

insights into the fungus:host plant interface.” An informal meeting 

was held for mycologists interested in the status of white-nose 

syndrome, a devastating fungal disease of bats in eastern North 

America. 

Several prominent mycologists received awards at the meeting. 

These included: Gary Samuels (USDA-ARS Systematic Mycology 

and Microbiology Laboratory, Beltsville, MD) – Distinguished 

Mycologist; Joseph Spatafora (Oregon State University) – Weston 

Teaching Award; Anne Pringle (Harvard University) – Alexopoulus 

Prize for Outstanding Early-Career Mycologist; Steven Stephenson 

(University of Arkansas) – MSA Fellow; and José Carmine Dianese 

(Universidade de Brasilia) – Honorary Member. Twenty-five 

students and young professionals received awards for travel and 

research excellence. Eight students received International Travel 

Awards to help defray expenses for their participation in IMC9. 

The annual banquet was held at the famous Kentucky Horse Park 

() and included local cuisine and entertainment. 

The annual auction featured a wide array of literature and 

mycological curiosities so that everyone left with something new! 

Preparations are already underway for next year’s annual meeting 

at the University of Alaska – Fairbanks. The theme of that meeting 

will be “High latitude fungi in a changing climate.” More information 

on this meeting can be found throughout 2011 at . MSA has not previously met in Alaska, so this 

should be an excellent opportunity to explore boreal fungi in the 

“Land of the Midnight Sun.” 

Jessie A. Glaeser 

(msasec1@yahoo.com) 

Silent auction at MSA 2010 annual meeting. Photo: Thomas J. Volk. 

MSA pre-meeting foray at the Bernheim Arboretum and Research Forest, near Bardstown, KY. Photo: Thomas J. Volk. 

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i m a f U N G U S

Iranian Mycological Society 

It gives me great pleasure to announce that the Iranian Mycological 

Society is now established. The calls for the establishment of such 

society first came some 15 years ago, but the major move was made 

two years ago at the 18 th Iranian Plant Protection Congress in 

Hamedan (Iran)in 2008 where about 70 mycologists from universities 

and research institutions in the country gathered and registered 

to become members of this society; the society was subsequently 

officially registered at the high commissions for scientific societies 

of the Iranian Ministry of Science, Research and Technology. With 

respect to the number of mycologists, the need for this society was 

approved by this commission. Consequently, the first general meeting 

was announced and took place on 15 September 2010. Members 

of the executive committee were elected by those present in the 

general meeting and five members were elected to officially follow 

up on various tasks for the Iranian Mycological Society. A website is 

dedicated to this society () with a permanent office at the 

Iranian Research Institute of Plant Protection, Tehran. The following 

mycologists were elected for the 3-year period 2010-2013 -- the first 

officers of the executive committee of the Iranian Mycological Society: 

President: Djafar Ershad (Emeritus Professor, Iranian Res. Inst. 

of Plant Protection) 

Vice-President: Mohammad Javan-Nikkhah (Associate Professor, 

University of Tehran) 

Secretary: Rasoul Zare (Professor, Iranian Research Institute of 

Plant Protection) 

Treasurer: Mehrdad Abbasi (Associate Professor, Iranian 

Research Institute of Plant Protection) 

Newsletter editor: Akbar Khodaparast (Associate Professsor, 

University of Guilan) 

Rasoul Zare 

(simplicillium@yahoo.com) 


Mycologists at the end of the first general meeting of the Iranian Mycological Society. Left to right: D. Ershad, S. Rezaee, A. Khodaparast, H. Azimi, M.J. Soleimani, A. 

Abbasi-Moghaddam, Gh. Hedjaroude, M. Abbasi, M. Javan-Nikkhah, V. Minassian, Z. Banihashemi, Gh. Balali, B. Sharifnabi, and R. Zare. 

v o l u m e 1 · n o . 2 

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BOOK NEWS 

From Another Kingdom: the amazing world of fungi. Edited by Lynne Boddy and Max Coleman. 2010. ISBN 978-1-906129-67-5. 

Pp. 176, illustr. Edinburgh: Royal Botanic Garden. Price: £ 20. 

What a superbly presented book! 

Should you ever require just one to 

capture the imagination, or to get 

friends or family to appreciate why 

you are a fungal fan, this is it. The 

large format, striking colour photographs 

on virtually every page (and 

in some cases a whole page), cannot 

but enchant. It really lives up to the 

subtitle, proclaiming that there is an 

“amazing world of fungi”. Aimed at 

the general public, this production 

has been a team effort, involving 21 

authors (all based in the UK) – and 

72 suppliers of photographs. 

The Introduction by Nick Read 

and Lynne Boddy, designed to catch 

the imagination, is followed by a 

characteristic ‘tour de force’ overview 

by Roy Watling (Ch 1). In this, 

amongst many other things, Roy illustrates 

the difficulty in inventorying 

fungi by his experience at Dawyck 

Botanic Garden; although recording 

started there in 1993, the rate of 

discovery of additional species still 

showed no sign of reduction in 2009 

(p. 30). The first group of chapters 

have an ecological and exploitive 

theme: ‘Recycling the World’ (Ch 

2), ‘Plant Pests and Perfect Partners’ 

(Ch 3) which covers symbiotic associations, 

‘Animal Slayers, Saviours and 

Socialists’ (Ch 4) with some stunning 

pictures of entomopathogens, and 

‘Amazing Chemists’ (Ch 5). The 

extent to which the potential benefits 

of fungi will be realized, however, is 

finance dependent; for example, who 

will pay for the prophylactic production 

and spraying of ‘Green Muscle’ 

for locust control in poor arid regions 

of the world (p. 77)? 

I really enjoyed the chapters 

on ‘Fungi and Humanity’ (Ch 6) 

and ‘Fungal Monsters in Science 

Fiction’ (Ch 7). However, I must 

admit to being surprised not to see 

mentions of Aldous Huxley’s last 

novel Island (1964) where the demeanour 

of the indigens is attributed 

to hallucinogenic mushrooms they 

cultivate, nor any reference to Eleanor 

Cameron’s much collected and 

delightful Mushroom Planet five-book 

series (1954-67); the first was The 

Wonderful Flight to the Mushroom 

Planet (1954) – which was appropriately 

named Basidium! ‘Growing Edible 

Fungi’ (Ch 8) includes commercial 

and personal approaches, but I do wish 

anyone coaxing Lentinula edodes plugs 

to produce on logs has a less frustrating 

and more productive experience 

than mine! Perhaps citing Coprinus 

comatus spawn kits for growth in 

lawns instead would have been 

preferable. ‘The Fungal Forager’ (Ch 

9) on collecting for food in the wild, 

rightly also highlights the issue of the 

mixed messages in current UK legislation 

designed to control collecting and 

promote conservation. ‘Safeguarding 

the Future’ (Ch 10), devoted to 

conservation, is politically charged 

and reiterates that it is first necessary 

to conserve mycologists with identification 

skills; this chapter should 

be compulsory reading for those in 

government agencies and departments 

responsible for science management 

and the environment. Then follows a 

section with six recipes using different 

wild mushrooms; the dishes look great, 

but I do wonder at the appropriateness 

of including one using Hygrophorus 

marzuolus as it is unlikely to be discovered 

north of Spain – though I have 

to admit it does have a rather special 

taste! What did seem something of an 

omission in this series of chapters, was 

any hints as to reliably identify fungi, 

whether for food or curiosity – or the 

wealth of information on distributions 

and ecologies now accumulated in the 

Fungal Records Database of Britain 

and Ireland (FRDBI) and the Association 

of British Fungus Group’s CATE. 

I was at first pleased to see lichens 

had due mention, but then a little 

disappointed to see them repeatedly 

referred to as “organisms” (pp. 16, 52, 

169) rather than associations, and also 

by the oxymoron “fungi and lichens” 

(p. 84); at least on p. 27 we read 

that they were “once thought to be 

independent organisms”! Some myths 

are also slain, for example presenting 

Amanita muscaria as a hallucinogen 

rather than a poisoner (p. 103), but 

others are perpetuated, such as that 

of spores being “spread in the wind” 

(p.16-17) – something that actually 

holds for only a rather limited number 

of species. There are inevitably small 

slips, for example, it is not Psilocybe 

semilanceata itself which is now 

categorized as a Class A drug (p.97), 

but rather the compounds psilocyin 

and psilocybin in whatever fungi they 

occur. And while the nomenclature 

is generally up-to-date, it would have 

been good to have see Gliocladium 

roseum replaced by Clonostachys rosea 

(p. 90). 

A feature of all the main chapters I 

particularly enjoyed was the separately 

authored half-page species profiles; 

these are wonderfully eclectic, for 

instance who would have expected to 

see features on Coniochaeta polymegasperma 

(p. 49) or Phellinus ferreus 

(p. 91)? – access the book to discover 

why! 

The one-page Bibliography unfortunately 

has a strong mushroom identification 

and cultivation bias, with 

not a single text on fungal biology 

cited, nor even a mention of Martin 

and Pam Ellis’ (1997) Microfungi 

on Land Plants. The few websites 

mentioned could usefully have been 

extended, and an opportunity to 

draw attention to Field Mycology, 

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i m a f U N G U S

The Forayer, and courses was missed. 

A suitably enthused reader could be 

frustrated and left floundering as 

to how to reveal even more amazing 

facts! Four pages are then devoted 

to portraits and biographies of 14 of 

the authors, and two to a glossary; 

the need for the latter could have 

been made superfluous by a policy of 

including definitions in the text and 

just avoiding unnecessary jargon such 

as ‘coprophile’ or ‘mycobiont’. 

The book was launched to accompany 

an extravaganza of an exhibition 

with the same name, which ran from 

31 July to 21 November 2010 at the 

Royal Botanic Garden in Edinburgh 

which many of the delegates to IMC9 

were able to enjoy. The modest cost 

was made possible by a generous donation 

from the Wellcome Trust, and 

makes the work easily accessible to the 

public at large. However, it is packed 

with tit-bits that are sure to also fascinate, 

or perhaps even amaze, the most 

hardened of mycologists. Secure a copy 

while you can, enjoy dipping into it, 

and do leave it lying around open to 

help ensnare the next generation of 

organismal mycologists. 

BOOK NEWS 

The Kingdom Fungi: the biology of mushrooms, molds, and lichens. By Steven L Stephenson. 2010. ISBN 978-0-88192-891-4. Pp. 

271, col. pl. 124. Portland, ORE: Timber Press. Price: US$ 34.95, £ 20. 

The flyleaf indicates that this book is aimed 

at “the general naturalist, amateur mycologist, 

or interested lay-person who simply 

wants to become more familiar with, and 

more appreciative of, the fascinating world 

of fungi”. The arrangement is primarily 

systematic, but with chapter titles designed 

not to deter. In the case of the agarics, there 

are entries even for families, which are 

grouped by spore colour, and throughout 

there is information on collecting and 

hands-on clues to identification intermixed 

with notes on ecology and biology. As 

might be expected from a specialist in myxomycetes, 

they have their own chapter, and so 

do lichens – although the trap of calling the 

latter “organisms” is not avoided (p. 21) and 

they are hardly “traditionally grouped with 

fungi” as the integration of lichen-forming 

fungi into the overall fungal system has 

only really been achieved over the last three 

decades. There are final chapters on the role 

of fungi in nature, interactions of fungi and 

animals, fungi and humans, and fossil fungi. 

I was pleased to see the myth that fungal 

species in general are widely distributed and 

that long-distance dispersal is a common 

phenomenon knocked down (p. 24). There 

are also numerous pieces of information that 

may be unfamiliar and sure to interest many 

professional mycologists. For example, that 

Pacific Northwest Northern flying squirrels 

rely on Bryoria species as their primary 

winter food source, the appreciation of the 

“Asa Gray Disjunction” between eastern 

Asia and North American closely allied 

fungi (e.g. Ciboria carunculoides and C. 

shiraiana), and a synopsis of what is known 

and postulated on regarding the enigmatic 

6 m tall fungal Lower Devonian fossil 

Prototaxites – including a hypothetical 

reconstruction in colour (pl 122). 

The book is well-presented and I did 

not detect many proofing slips, although 

the legend to the plate showing Arcyria 

sulcata in Baltic amber (pl 124) gives a 

date of “35 to 59 years old” having lost the 

million (which is correctly included on p. 

237). The illustrations comprise 124 superb 

colour plates on coated paper, arranged in 

two tipped-in signatures, which are sure to 

capture the interest of anyone chancing to 

open it. There are no line illustrations, but 

the writing style renders the whole most 

readable and engrossing – and the author’s 

enthusiasm for all things fungal permeates 

throughout. It is perfectly tailored to 

its intended target audience, and can be 

unhestitatingly recommended to the general 

naturalist and citizen scientist. 

Mycorrhizal Biotechnology. Edited by Devarajan Thangadurai, Carlos Alberto Busso & Mohamed Hijri. 2010. ISBN 978-1-57808- 

691-7. Pp. x + 216. Enfield, NJ: Science Publishers. Price: £ 57.99. 

Knowledge of the operation of plant-fungus 

interactions in arbuscular-mycorrhizal 

(AM) fungi at the molecular level has 

developed dramatically in the last decade, 

and on seeing this title I expected the focus 

to be some synthesis of our this new understanding 

and discussions of the potential 

for exploitation. The emphasis is indeed on 

AM fungi, as only one of the 14 chapters 

deals with ectomycorrhizas (Ch 2) -- and 

that concentrates on general issues and not 

technologies for exploitation. But the thrust 

of the AM fungal chapters is on pragmatic 

low technological exploitation rather 

than blue-skies visions, with some original 

studies thrown in. The interplay with 

bacteria, especially nitrogen-fixing rhizobia, 

is emphasized in Ch 1, and this theme is 

developed further in Ch 3. 

Low-cost production of enough 

inoculum is a limiting factor to commercial 

exploitation, especially in tropical countries 

where there is the greatest need to increase 

crop yields; the methods tried are covered 

in Ch 4, though no mass culture system 

seems yet to have been developed. There 

is an account of a research project carried 

out on a single grassland site in Argentina 

which was weeded and then planted with 

two grasses; AM-fungal colonization was 

assessed and biomass production measured 

(Ch 5). There was a negative correlation 

between the development of mycorrhizal 

associations and biomass production and it 

is concluded that the associations may not 

be beneficial where the soil has high levels 

v o l u m e 1 · n o . 2 

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BOOK NEWS 

of available phosphorous – I cannot but 

think that such a contribution would have 

been better-placed in a primary journal. In 

contrast, in the case of horticultural crops, 

an increased yield of 20–40 % is claimed (p. 

137) but not referenced and the information 

is stated to be limited (Ch 9). 

Mechanisms of reduced resistance to 

pathogens in plants are reviewed, and the 

importance of bacteria, including inoculated 

Pseudomonas strains, is stressed; although 

some promising results with Glomus species 

are mentioned, the incidence of leaf pathogens 

can be increased (Ch 6)! Rhizosphere 

management is clearly a complex issue, but 

bioaugmentation by AM-fungi is again 

frustrated by difficulties in securing enough 

biomass for field inoculations (Ch 8). 

Perhaps of greatest potential is “mycobization”, 

a term coined in 2005 for the novel 

process of co-inoculating plants with different 

functional groups of fungi, for example 

phosphate-mobilizing Glomus mosseae and 

solubilizer Aspergillus niger (along with 

rock phosphate) in tomato cultivation (Ch 

10). As to the future, the need to inoculate 

plants being used in bioremediation, 

actually so-called “phytoremediation”, of 

polluted soils devoid of AM fungi is pointed 

out (Ch 7); Glomus mosseae can apparently 

withstand 1200 mg kg -1 zinc. This theme 

is also the topic of a second chapter (Ch 

11), stressing enhancing the functionality 

of the 400 or so known “hyperaccumulator” 

plants in “phytoextraction” where polluting 

compounds are taken up into harvestable 

plant tissues. 

A survey of the actual diversity of 

AM-fungi, surprisingly, does not feature until 

almost the end of the book, where there is 

one chapter providing a general overview of 

the families and genera of Glomeromycota 

with a summary of the PCR-based studies 

conducted (Ch 12) – but without illustrations 

illustrating critical characters used in 

their morphological identification. More 

pragmatic for many of those working with 

these fungi is the survey of molecular tools 

for detection (Ch 13) which is especially to 

be commended in ranging from RFLP to 

sequence data, but more so in its focus on the 

composition of the different primers available 

which occupy seven tables. The final chapter 

(Ch 14) considers the topical and pertinent 

issue of possible effects of climate change on 

AM fungi, which respond to elevated soil 

temperatures; yet the experimental results 

to date are “contradictory and do not allow 

general conclusions” (p. 205). One complication 

is surely that a single patch of vegetation 

may support 30–40 AM fungal species, 

which I presume would not necessarily all 

respond in a regimented manner. 

The editors are clearly mycorrhizal 

chauvinists as they claim “mycorrhizae 

are the most common symbiotic species 

on earth” (p. vii).Yet a “mycorrhiza” is 

not a species but a mutualistic association 

between a plant root and a fungus, and 

I suspect that there would be bacterial 

contenders for the commonest association 

title, and possibly also claims from the lichenologists. 

I was amused to see that the oldest 

fossil evidence of a bryophyte-like land plant 

was stated to be from just 100 years ago (p. 

103). Five of the chapters are contributed 

by staff of the Tamil Nadhu Agricultural 

University in Coimbatore, India, where I 

was privileged to have lectured in 1991; I 

was much-impressed by the mycological 

work being carried out there at that time, 

and am so-pleased to see them feature so 

strongly here. I enjoyed the delightful and 

sometimes amusingly phrased Indian- 

English that features throughout but do feel 

more rigorous editing in content was needed 

– how many times do readers wish to be 

told that Frank introduced the term “mycorrhiza” 

in 1885? Further, that date was for 

what later became termed ectomycorrhizal 

and not endomycorrhizal fungi, two terms 

he introduced in 1887! Author citations 

appended to scientific names are of course 

quite inappropriate for such an applied 

work, but do crop up in two chapters (Chs 

10 and 12), and in one the citations do not 

even follow the recommended system. And 

there are some misspellings, such as “Dydimella” 

for “Didymella” which are difficult to 

excuse . . . . 

There are two main audiences that I feel 

would profit from digesting this book: (1) 

those in less-developed countries endeavouring 

to exploit AM-fungi in increasing crop 

yields; and (2) those researching AM-fungal 

mechanisms at the molecular level that 

need to be aware of the practical difficulties 

in exploitation. Sadly, the price is likely to 

prohibit access by the first group, and the 

lack of state-of the art molecular work deter 

the second from buying a copy. 

Oomycete Genetics and Genomics: diversity, interactions and research tools. Edited by Kurt Lamour and Sophien Kamoun. 

2009. ISBN 978-0-470-25567-4. Pp. xvii + 574. John Wiley & Sons, Hoboken, NJ, USA. Price: £100.00, € 120.00. 

Although this book was published in 2009, 

it only just came to my attention and I 

am much impressed by the coverage and 

caliber of the 67 contributors drawn from 

14 countries. No less than 27 chapters are 

accommodated, which leads to chapters with 

a clarity and pertinence too often absent in 

multi-authored works. The title understates 

the contents. As Francine Govers states in her 

Foreword, this “ . . . is the first time that the 

existing knowledge on oomycetes has been 

brought together in one volume” (p. ix). 

While it is true that genetic and 

genomic studies are authoritatively considered, 

and these are incorporated where 

appropriate, there is much basic information 

on these fungal analogues that is not otherwise 

easily accessible. This is especially true 

for the first five chapters which address the 

phylogeny, ecology, and life-cycles of these 

fungi. I found the first (by Gordon Beakes 

and Satoshi Sekimoto) on the overall phylogeny 

especially useful as these fungal analogues 

are rarely considered in fungal systems 

that embrace only the members of the 

kingdom Fungi. The major part of the work, 

14 chapters, are concerned with succinct 

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i m a f U N G U S

accounts of studies of particular genera and 

species. These include many groups of major 

economic importance as pathogens of crops, 

fish, and even humans, for example different 

species of Phytophthora, Saprolegnia on fish, 

various downy mildews on particular crops, 

and Pythium insidiosum in mammals (including 

humans!). This last chapter on P. insidiosum 

I found to be an especially valuable 

synopsis with fine SEMs of appressoria. The 

Phytophthora species selected for particular 

attention are, not unsurprisingly, P. brassicae, 

P. capsici, P. infestans, P. ramorum, and P. 

sojae – but I missed a chapter devoted to P. 

cinnammoni which causes such devastation 

of native forests in Australia in particular. 

The last six chapters focus on some of 

the cutting-edge molecular approaches to 

work on these fungi, which from the title 

might have been expected to dominate. 

These cover transformations, expression 

systems, the promise of gene silencing, 

proteomics, and the strategy towards 

genome sequencing (the last with a useful 

glossary for the non-geneticist). 

These fungal analogues tend to be increasingly 

not, or rather poorly, represented in 

mycological treatises and textbooks, since 

it became recognized that they were not 

real Fungi, but merely fungi (i.e. organisms 

studied by mycologists). This book would 

thus help address this situation in mycological 

libraries, and should be seen in that 

context and purchased for that reason. Its 

scope would have been better reflected in 

a title on the lines of “Oomycete diversity, 

interactions, and molecular biology” – and 

it is unfortunately that the actual title 

may deter many mycologists and plant 

pathologists that would actually find much 

to interest them here. 

The whole is extremely well-produced, 

and I especially liked the tipped-in signature 

of colour plates on coated paper comprising 

colour versions of eight half-tone figures 

from various chapters. It also seems as well 

up-to-date as can be expected in such multiauthored 

works, with many papers from 

2008 being cited. And the price is reasonable 

by current standards for a book of this 

quality. The editors are to be congratulated 

on marshalling such a work, which clearly 

merits wide circulation amongst the broader 

mycological community. 

BOOK NEWS 

Lichens. By William Purvis. 2010. ISBN 978-0-565-09153-8. Pp. 112, illustr. London: Natural History Museum. Price: £ 12.99. 

First published in 2000, and reprinted 

in 2007 with a different cover, this wellillustrated 

introduction to lichen ecology 

and biology has now been re-issued “with 

updates”. The updates have been rather 

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though there is a different illustration on 

the title page inside. Even the list of selected 

books (p. 111) has not been revised from 

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replacements; that would hardly have been 

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and revise sections such as that on pollution 

effects where the influence of nitrogenous 

compounds might have had a higher profile. 

Nevertheless, with no equivalent full-colour 

book aimed at the general naturalist now on 

the market, it is pleasing that this little book 

is available once more – but if you have one 

of the earlier editions, do not put acquiring 

this one high on your list of desiderata. 

v o l u m e 1 · n o . 2 

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i m a f U N G U S

IMA Fungus · volume 1 · no 2: 101–108 

The enigma of Calonectria species occurring on leaves of Ilex 

aquifolium in Europe 

Christian Lechat 1 , Pedro W. Crous 2 and Johannes Z. Groenewald 2 

ARTICLE 

1 

AscoFrance, 64 route de Chizé, F-79360, Villiers en Bois, France; corresponding author e-mail: lechat@ascofrance.fr 

2 

CBS-KNAW Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands 

Abstract: Species of Calonectria are common saprobes and plant pathogens on a wide range of hosts 

occurring in subtropical to tropical regions of the world. The aim of the present study was to resolve 

the status of new Calonectria collections obtained on Ilex leaves from France. Based on DNA sequence 

data of their b-tubulin and histone gene regions, as well as morphology, the new collections matched 

the ex-type strain of Cylindrocladium ilicicola. On the host and in culture, yellow to brownish-yellow 

perithecia were observed that did not strain red in 3 % KOH. Based on these results, C. ilicicola and 

its purported teleomorph, Ca. pyrochroa, were shown to represent two distinct species, as the latter 

has bright red perithecia that strain purple in KOH. A new combination, Ca. lauri, based on Tetracytum 

lauri, is subsequently proposed for C. ilicicola. Calonectria lauri is distinct from Ca. ilicicola, a pathogen 

commonly associated with Cylindrocladium black rot of peanut. Finally, Ca. canadiana is proposed as 

new name for Cy. canadiense, which is a nursery pathogen involved with root rot of several tree genera 

in Quebec, Canada. 

Key words: 

Hypocreales 

Calonectria 

Cylindrocladium 

Ilex aquifolium 

TUB 

HIS 

systematics 

Article info: Submitted: 1 September 2010; Accepted: 29 September 2010; Published: 2 November 2010. 

Introduction 

Species of Calonectria are members of Nectriaceae 

(Hypocreales, Ascomycetes) (Lombard 2010a–c). 

Nectriaceae are characterised by having uniloculate, 

orange to purple, superficial ascomata (Rossman 

et al. 1999). Calonectria is easily distinguished 

from other members of the family based on its 

Cylindrocladium anamorphs. Formerly Cylindrocladium 

also included members of Cylindrocladiella, a genus 

that accommodates Cylindrocladium-like species with 

small conidia (Boesewinkel 1982, Victor et al. 1998) and 

Nectricladiella teleomorphs (Schoch et al. 2000). Other 

morphologically similar genera that have also since been 

separated from this complex include Xenocylindrocladium 

(Decock et al. 1997), Curvicladiella (Crous et al. 2006a) 

and Dematiocladium (Crous et al. 2005). Following the 

approach of Crous et al. (2006b, 2008, 2009a, b) with 

other fungal groups, Lombard et al. (2009, 2010a–d) 

chose to use the older Calonectria name for the genus, 

irrespective whether the teleomorph or Cylindrocladium 

anamorph, unnamed microconidial, megaconidial, or 

chlamydospore-like synanamorph was observed. All 

taxa are since accommodated in Calonectria, which is a 

monophyletic genus (Lombard et al. 2010a–c). 

Most species of Calonectria occur commonly in 

soil, especially in subtropical to tropical regions of 

the world. Although the genus was originally regarded 

as saprobic (Graves 1915), taxa have since been 

proven to be important plant pathogens, associated 

with a wide host range of plants, causing disease 

symptoms ranging from leaf spots to stem cankers, 

damping off, cutting rot, root and fruit rot (Crous et al. 

2004b, 2006a, Lombard et al. 2009, 2010a, d). Major 

diseases attributed to Calonectria infections include 

Cylindrocladium black rot of Arachis hypogea (peanut), 

and red crown rot of Glycine max (soybean) (Crous et 

al. 1993, Wright et al. 2010), as well as root rot and 

leaf diseases of numerous diverse hosts (Crous et al. 

2004b, 2006a). 

Over the past few years, a species of Calonectria 

was collected from leaves of Ilex aquifolium in 

France. Presently four species of Calonectria have 

been described from Ilex (Aquifoliaceae), namely 

Calonectria morganii on Ilex paraguayensis in 

Argentina, and Ilex vomitoria in Florida (USA); 

Calonectria avesiculata on Ilex spp. in Georgia and 

Florida (USA), Cylindrocladium ilicicola (as Calonectria 

pyrochroa) on Ilex aquifolium on Clare Island (Ireland), 

and Calonectria spathulata on Ilex paraguariensis in 

© 2010 International Mycological Association 

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v o l u m e 1 · n o . 2 

101

Lechat et al. 

ARTICLE 

Brazil (Crous 2002). Hawksworth & Sivanesan (1976) 

also reported a Calonectria species on Ilex aquifolium 

from Slapton, South Devon, England, which appears 

to be undescribed, with ascospores 3-septate, 14–22 

×3–4 µm. The collection obtained from France and 

treated in this study, is morphologically distinct from 

taxa presently reported from Ilex. 

In recent years there have been several revisions 

focused on either Calonectria or its anamorph 

genus, Cylindrocladium (Rossman 1979, Peerally 

1991, Crous & Wingfield 1994, Crous 2002). The 

first attempt to provide a molecular phylogeny of the 

genus was that of Schoch et al. (2001) based on 

b-tubulin DNA sequences. This gene region, however, 

proved insufficiently variable to reliably distinguish all 

species complexes in the genus (Kang et al. 2001a, 

b, Henricot & Culham 2002, Crous et al. 2004b, 

2006a). Since then, a concerted effort has been made 

to generate a multi-gene phylogeny for taxa in the 

genus, and identify the best suited gene for species 

delimitation (Lombard et al. 2009, 2010a–d). Based 

on these findings, a combination of b-tubulin DNA 

sequence data, supplemented with either calmodulin 

or elongation factor 1-a, proved the most effective in 

distinguishing all known taxa. 

The aim of the present study was to compare the 

new collections on Ilex from France to all species known 

in the genus, using morphology and DNA sequence 

analysis of their b-tubulin and histone gene regions in 

order to determine if it represented a novel taxon. 

Materials and methods 

Isolates 

Single ascospore isolates were obtained from leaves 

of Ilex aquifolium as explained in Crous & Wingfield 

(1994). Isolates were incubated on plates of 2 % malt 

extract agar (MEA), 2 % potato-dextrose agar (PDA) 

and oatmeal agar (OA) (Crous et al. 2009c) for 7 d 

at 25 °C under continuous near-UV light, to promote 

sporulation. Reference strains are maintained in the 

CBS-KNAW Fungal Biodiversity Centre (CBS) Utrecht, 

The Netherlands. 

DNA isolation, amplification and analyses 

Genomic DNA was isolated from fungal mycelium 

grown on MEA, using the UltraCleanTM Microbial DNA 

Isolation Kit (MoBio Laboratories, Inc., Solana Beach, 

CA, USA) according to the manufacturer’s protocol. 

Two loci were amplified and sequenced as explained 

in Crous et al. (2004b) and Lombard et al. (2010c), 

namely, part of the β-tubulin gene (TUB), amplified with 

primers T1 (O’Donnell & Cigelnik 1997) and CYLTUB1R 

(Crous et al. 2004b); and part of the histone H3 gene 

(HIS) using primers CYLH3F and CYLH3R (Crous et al. 

2004b). Part of the nuclear rDNA operon spanning the 

3’ end of the 18S nrRNA gene (SSU), the first internal 

transcribed spacer (ITS1), the 5.8S nrRNA gene, the 

second ITS region (ITS2) and the 5’ end of the 28S 

nrRNA gene (LSU) was amplified for some isolates as 

explained in Lombard et al. (2010c). The generated 

sequences were compared with other fungal DNA 

sequences from NCBI’s GenBank sequence database 

using a blastn search; TUB sequences with high 

similarity were added to the alignment and the result of 

sequences of the other loci were used as confirmation 

(not shown). The additional GenBank sequences were 

manually aligned using Sequence Alignment Editor 

v. 2.0a11 (Rambaut 2002). Phylogenetic analyses 

of the aligned sequence data were performed using 

PAUP (Phylogenetic Analysis Using Parsimony) v. 

4.0b10 (Swofford 2003) and consisted of neighbourjoining 

analyses with the uncorrected (“p”), the Kimura 

2-parameter and the HKY85 substitution models. 

Alignment gaps were treated as missing data and all 

characters were unordered and of equal weight. Any 

ties were broken randomly when encountered. For 

parsimony analyses, alignment gaps were treated as a 

fifth character state and all characters were unordered 

and of equal weight. Maximum parsimony analysis 

was performed using the heuristic search option with 

100 random (ITS) or simple (LSU) taxa additions and 

tree bisection and reconstruction (TBR) as the branchswapping 

algorithm. Branches of zero length were 

collapsed and all multiple, equally parsimonious trees 

were saved. The robustness of the trees obtained 

was evaluated by 1 000 bootstrap replications (Hillis 

& Bull 1993). Tree length (TL), consistency index 

(CI), retention index (RI) and rescaled consistency 

index (RC) were calculated. Sequences derived in this 

study were lodged at GenBank (), 

the alignment in TreeBASE (), and taxonomic novelties in MycoBank 

(; Crous et al. 2004a). 

Morphology 

Characteristics in culture were determined after 7 d on 

MEA, PDA and OA (Crous et al. 2009c). Morphological 

descriptions were based on sporulating cultures on 

synthetic nutrient-poor agar (SNA) (Nirenburg 1981, 

Lombard et al. 2009) and carnation leaf agar (CLA) 

(Crous et al. 2009c). Slide preparations were made 

from sporulating cultures (SNA for anamorph, CLA for 

teleomorph) in clear lactic acid, with 30 measurements 

determined per structure, and observations made with 

a Nikon SMZ1500 dissecting microscope, and with 

a Zeiss Axioscope 2 microscope using differential 

interference contrast (DIC) illumination. Colony 

characters and pigment production were noted after 

7 d of growth on MEA, PDA and OA (Crous et al. 

2009c) incubated at 25 ºC. Colony colours (surface 

and reverse) were rated according to the colour charts 

of Rayner (1970). 

102 

i m a f U N G U S

Calonectria lauri sp. nov. 

Table 1. Collection details and GenBank accession numbers of isolates of Calonectria lauri included in this study. 

Strain No. 1 Substrate Country Collector(s) GenBank Accession No. (TUB, HIS, ITS) 2 

CPC 15683 Leaves of Ilex aquifolium Netherlands W. Gams FR694682, FR694676, FR694679 

CBS 126269 = CPC 17978 Leaves of I. aquifolium France A. Gardiennet FR694683, FR694677, FR694680 

CBS 553.69 = IMI 299390 Root of Buxus sempervirens Belgium — FR694684, FR694678, — 

CBS 749.70 I. aquifolium Netherlands H.A. van der Aa FR694685, GQ267250, GQ280584 

1 

CBS: CBS Fungal Biodiversity Centre, Utrecht, The Netherlands; CPC: Culture collection of P.W. Crous, housed at CBS; IMI: International 

Mycological Institute, CABI-Bioscience, Egham, Bakeham Lane, UK. 

2 

TUB: partial beta-tubulin gene; HIS: partial histone H3 gene; ITS: Internal transcribed spacers 1 and 2 together with 5.8S nrDNA. 

ARTICLE 

Results 

Phylogeny 

Approximately 600, 480 and 680 bases were determined 

for the isolates indicated in Table 1 for TUB, HIS and 

ITS, respectively. Of the β-tubulin gene, 522 bases 

were used for phylogenetic analyses in the manually 

adjusted alignment containing 32 isolates (including 

the outgroup sequence). Of these 522 characters 

(including alignment gaps), 180 were parsimonyinformative, 

47 were variable and parsimonyuninformative, 

and 295 were constant. Neighbourjoining 

analysis using the three substitution models, 

as well as the parsimony analysis, yielded trees with 

exactly the same topologies. Parsimony analysis of 

the alignment yielded a single most parsimonious tree 

(TL = 381 steps; CI = 0.816; RI = 0.953; RC = 0.778), 

which is shown in Fig. 1. 

Taxonomy 

Calonectria lauri (Vanderw.) Lechat & Crous, 

comb. nov. 

MycoBank MB517423 

(Fig. 2) 

Basionym: Tetracytum lauri Vanderw., Parasitica 1: 

145. 1945. (as “laurii”). 

= Candelospora ilicicola Hawley, Proc. Roy. Irish 

Acad. 31: 11. 1912. [non Calonectria ilicicola Boedijn & 

Reitsma, 1950] 

= Cylindrocladium ilicicola (Hawley) Boedijn & Reitsma, 

Reinwardtia 1: 57. 1950. 

region, ostiole papillate, composed of palisade-like, 

cylindrical to narrowly ellipsoidal cells. Ascomatal wall 

50–65 µm thick of two regions; outer region comprising 

warts 50–55 µm thick, composed of globose to nearly 

angular, thick-walled cells, 10–30 × 5–16 µm, yellow, 

wall 1.5–2 µm thick; inner region 5–10 µm thick, 

composed of flattened, ellipsoidal cells, 12–18 × 3–5 

µm, hyaline; warts globose to subglobose 25–40 × 15– 

30 µm, yellow. Asci clavate, long stipitate, 110–130 × 

17–22 µm, 8-spored, multiseriate. Ascospores narrowly 

fusiform with rounded ends, lightly curved, guttulate, 

hyaline, smooth, (53–)60–86(–89) × 6.5–8(–9) µm, 

3-septate, not conctricted at the septa or constricted 

when overmature. Conidiophores consisting of a 

stipe bearing a penicillate arrangement of fertile 

branches, a stipe extension, and a terminal vesicle; 

stipe septate, hyaline, smooth, 40–150 × 3–5 µm; stipe 

extensions septate, straight to flexuous, 120–200 µm 

long, 2.5–3 µm wide at the apical septum, terminating 

in an obpyriform to ellipsoid vesicle, (5–) 7–8(–10) 

µm diam. Conidiogenous apparatus with primary 

branches aseptate or 1-septate, 15–20 × 4–5 µm; 

secondary branches aseptate, 8–15 × 4–5 µm; tertiary 

branches aseptate, 10–15 × 4–5 µm, each terminal 

branch producing 2–4 phialides; phialides doliiform 

to reniform, hyaline, aseptate, 6–12 × 2.5–4 µm; apex 

with minute periclinal thickening and inconspicuous 

collarette. Conidia cylindrical, rounded at both ends, 

straight, (45–) 55–68(–73) × (4–)5–6(–7) µm (av. = 60 

× 5.5 µm), (1–)3-septate, lacking a visible abscission 

scar, held in parallel cylindrical clusters by colourless 

slime. Megaconidia and microconidia unknown. 

Typus: Ireland, Clare Island, Ilex aquifolium, Hawley, K (M) 

61269!, holotype of Cy. ilicicola, IMI 76542 isotype. Netherlands, 

South-East Limburg, Vijlenerbos, Vijlen, Ilex aquifolium, Aug. 

1970, H.A van der Aa, epitype CBS H-15110, ex-epitype culture 

CBS 749.70. 

Ascomata perithecial, solitary, scattered, subglobose 

to ovoid, 450–550 µm high × 380–420 µm diam, 

superficial, not obviously stromatic but difficult to 

remove from the subtratum because basal cells of 

ascomata remain immersed in the substratum, yellow to 

brownish-yellow, dark-red at base, not changing colour 

in 3 % KOH or lactic acid, warted except at ostiolar 

Culture characteristics: Colonies on MEA sienna to 

brick on the surface, and sienna in reverse; sienna on 

OA (surface); sienna to umber on PDA (surface), and 

umber in reverse; chlamydospores on MEA moderate, 

occurring throughout the medium, with sparse to 

moderate sporulation on aerial mycelium. 

Additional specimens examined: Netherlands, Hilversum, 

on leaves of Ilex aquifolium, 11 Nov. 2008, W. Gams, CPC 

15683 = CBS 128031, CPC 15684, CPC 15685. France, 

Pressigny (52), on leaves of Ilex aquifolium, 05 Dec. 2009, A. 

Gardiennet, AG09308, CBS H-20476, culture CPC 17978 = 

CBS 126269; Forêt de Chizé, Villiers en Bois (79) on leaves 

v o l u m e 1 · n o . 2 103

Lechat et al. 

ARTICLE 

Cylindrocladiella lageniformis AY725652 

100 

10 changes 

66 

53 

90 

62 

77 

99 

100 

100 

62 

100 

76 

68 

DQ190564 

DQ190557 

DQ190560 

Calonectria pauciramosa AY078119 

51 

AF308466 

AF308465 

Calonectria citri AF333393 

CPC 15683 

CBS 749.70 

CPC 17978 

AY078122 

CBS 553.69 

84 

AY078120 

Calonectria “ilicicola” AY725641 

Calonectria colombiensis GQ267207 

Calonectria curvispora AF333395 

GQ267213 

AY725630 

GQ267209 

GQ267208 

AY725642 

GU256586 

GU256594 

AY725631 

AY725636 

AY725637 

AY725639 

AY725643 

AY725647 

AY725646 

AY725645 

100 

Calonectria colhounii 

100 

Calonectria pacifica 

Calonectria kyotensis 

Calonectria ilicicola 

Calonectria leucothoës 

Calonectria lauri 

Fig. 1. Single most parsimonious trees obtained from a heuristic search with 100 random taxon additions of the β-tubulin 

sequence alignment. The scale bar shows 10 changes and bootstrap support values from 1000 replicates are shown at the 

nodes. The tree was rooted to Cylindrocladiella lageniformis (GenBank AY725652). 

104 

i m a f U N G U S


ARTICLE 

Fig. 2. Calonectria lauri and its Cylindrocladium anamorph. A, B. Yellowish perithecia in vivo (A), and in vitro (B). C. 

Cylindrocladium anamorph. D. Vertical section through perithecium, showing wall anatomy. E. Ascospores. F–H. Conidiophores. 

I–L. Conidiogenous apparatus with phialides. M. Ellipsoid to obpyriform vesicles. N. Three-septate conidia. Scale bars: A, B = 

200 µm, C, E–H, M = 8 µm, D, J–L, N = 10 µm, I = 5.5 µm. 

v o l u m e 1 · n o . 2 105

Lechat et al. 

ARTICLE 

of Ilex aquifolium, 19 Sept. 2006, C. Lechat, CLL696. Belgium, 

Gent, on roots of Buxus sempervirens, July 1969, A. Roos, 

IMI 299390 = CBS 553.69. 

Notes: The name Calonectria ilicicola is already 

occupied, and thus the next available epithet for this 

species in Calonectria is that of Tetracytum lauri. 

Calonectria lauri is phylogenetically closely related to 

Ca. citri (known on Citrus from Florida). Morphologically 

the two species can be separated in that Ca. citri has 

ellipsoid to pyriform or obovoid vesicles, and 3-septate 

conidia that are slightly shorter and narrower, (25–)53– 

60(–65) × 3–4(–5) µm (Crous 2002). 

Discussion 

collection (on the host and on CLA in culture), forms 

yellow to brownish yellow perithecia that do not 

discolour in KOH (except at the perithecial base). 

The teleomorph of C. ilicicola could therefore not be 

Ca. pyrochroa as currently accepted (Lombard et al. 

2010c). Because the name Ca. ilicicola is already 

occupied by the pathogen causing Cylindrocladium 

black rot of peanut (Crous et al. 1993), a new name, 

Ca. lauri, is proposed for this species, which appears 

to occur commonly on Laurus, Ilex, as well as several 

other hosts in Europe (Brayford & Chapman 1987). 

Presently no cultures are available of Ca. pyrochroa, 

and further collections will have to be made from 

Platanus leaf litter in France to help clarify the 

morphology of its Cylindrocladium anamorph. 

The genus Calonectria is based upon Calonectria 

pyrochroa (on Platanus leaf litter, France, 

lectotype BPI), which Rossman (1979) found to be 

indistinguishable from Ca. daldiniana (on Magnolia 

grandiflora leaf litter, Italy, holotype RO). A separate 

collection from decaying leaves of Pittosporum 

undulatum collected in Madeira (CUP-MM 2407) 

produced a Cylindrocladium anamorph with clavate 

vesicles, which later led Rossman (1983) to conclude 

that the oldest anamorph epithet that could be linked 

to Ca. pyrochroa was C. ilicicola. 

Brayford & Chapman (1987) reported a wilting 

disease of Laurus nobilis in nurseries on the Isles 

of Scilly, and later on Arbutus andrachnoides and 

Gaultheria shallon in West Devon, UK. The causal 

organism was identified as C. ilicicola, but incorrectly 

linked to the teleomorph name, Ca. ilicicola. Based 

on a molecular comparison of ex-type strains, Crous 

et al. (1993) showed Ca. ilicicola was the teleomorph 

of C. parasiticum, a major pathogen associated with 

Cylindrocladium black rot of peanut. In a later study, 

Crous & Wingfield (1994) accepted the relationship 

between Ca. pyrochroa and C. ilicicola, as there 

were no cultures available at the time to refute this 

proposed link (Crous 2002). Following a revision of 

Cylindrocladium strains in the CBS culture collection, 

Crous et al. (2006a) discovered a strain linked to a 

specimen that closely matched the type of C. ilicicola, 

and subsequently designated CBS 749.70 (on Ilex 

aquifolium, the Netherlands) as ex-epitype strain for 

C. ilicicola. Sequence data derived from the ex-epitype 

strain, and morphology, proved to be identical to that 

of the new collection obtained from France (Figs 1–2), 

confirming it to be C. ilicicola. 

However, isolate CBS 126269 produced a 

Calonectria teleomorph in culture, which is clearly 

distinct from Ca. pyrochroa. The latter species (and 

its synonyms) have scarlet-red perithecia, which turn 

purple in 2 % KOH (Rossman 1979). The present 

APPENDIX 

In the recent treatment of the genus Calonectria, 

Lombard et al. (2010c) allocated the name 

Cylindrocladium canadense to Calonectria as Ca. 

canadensis (J.C. Kang, Crous & C.L. Schoch) L. 

Lombard, M.J. Wingf. & Crous, but overlooked the 

older existing name, Ca. canadensis (Ellis & Everh.) 

Berl. & Voglino. A new combination is required to 

resolve this homonym as follows: 

Calonectria canadiana L. Lombard, M.J. Wingf. 

& Crous, nom. nov. 


Basionym: Cylindrocladium canadense J.C. Kang, 

Crous & C.L. Schoch, Syst. Appl. Microbiol. 24: 210. 

2001. 

= Calonectria canadensis (J.C. Kang, Crous & C.L. 

Schoch) L. Lombard, M.J. Wingf. & Crous, Stud. Mycol. 

66: 56. 2010, non Calonectria canadensis (Ellis & Everh.) 

Berl. & Voglino, Addendum to Syll. Fung. 4: 212. 1886. 

Acknowledgements 

The authors thank the technical staff, Arien van Iperen 

(cultures), Marjan Vermaas (photo plates), and Mieke Starink- 

Willemse (DNA isolation, amplification and sequencing) for 

their invaluable assistance. Drew Minnis (USDA, Beltsville, 

USA) is also thanked for bringing the homonym associated 

with epithet “canadensis” to our attention. Finally, we thank 

Alain Gardiennet for the supply of specimens. 

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MJ, Groenewald JZ (2009a) Novel species of 

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GC, Burgess TI, Andjic V, Barber PA, Groenewald JZ 

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(2009c) Fungal Biodiversity. [CBS Laboratory Manual 

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Phylogeny and systematics of the genus Calonectria. 


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45: 45–62. 

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MJ, Alfenas AC (1998) Systematic appraisal of 

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How to describe a new fungal species 

Keith A. Seifert 1 and Amy Y. Rossman 2 

1 

Biodiversity (Mycology), Eastern Cereal and Oilseed Research Centre, Agriculture & Agri-Food Canada, Ottawa, Ontario K1A 

0C6 Canada; corresponding author e-mail: Keith.Seifert@AGR.GC.CA 

2 

Systematic Mycology & Microbiology Laboratory, USDA-ARS, Rm. 246, B0101A, 10300 Baltimore Ave., Beltsville, MD 20705, 

USA 

ARTICLE 

for the International Commission on the Taxonomy of Fungi, www.fungaltaxonomy.org 

Abstract: The formal requirements and best practices for the publication 

of descriptions of new fungal species are discussed. Expectations for DNA 

sequences and cultures are considered. A model manuscript offers one 

possible approach to writing such a paper. 

Key words: 

Culture Collections 

Herbaria 

International Code of Botanical Nomenclature 

Latin diagnoses 

Molecular phylogenetics 

Article info: Submitted: 9 October 2010; Accepted: 15 October 2010; Published: 2 November 2010. 

Introduction 

Every fungal species is unique. Therefore, every 

description of a fungal species is also unique. 

The morphological, physiological, ecological, and 

molecular diversity in fungi means that descriptions 

and illustrations differ from one taxonomic group 

to another. There are no formal standards for the 

description and illustration of species, but there are 

some formal (or ‘legal’) requirements for proposing 

names that are imposed by the International Code of 

Botanical Nomenclature (ICBN; McNeill et al. 2006). 

Furthermore, community standards of scientific rigor 

are enforced by editors and reviewers. 

For the beginner, it is useful to have models to assist 

with the preparation of descriptions and illustrations. In 

this paper, formal requirements and best practices that 

should be considered for any description are outlined, 

and a model manuscript for describing a new species 

is provided. Several ‘tricks of the trade’ and cautionary 

notes are also included. 

Additional hints can found in the now somewhat 

dated Code of Practice developed by the ICTF (Sigler & 

Hawksworth 1987), and the guidebook for mycologists 

by Hawksworth (1974). Although not exclusively 

concerned with fungi, the book by Winston (1999) also 

provides a valuable perspective. 

Formal requirements 

The International Code of Botanical Nomenclature 

(ICBN) governs the naming of fungi. This is a complex 

document, but you should read the relevant articles of 

the Code for exact wording of the regulations. The ICBN 

is updated every six years, after each International 

Botanical Congress, and is available on the World 

Wide Web (see references). The most recently 

published code must be followed, and previous Codes 

are considered obsolete. Although there have been 

discussions about a possible independent Mycological 

Code, or a BioCode covering all organisms, these are 

still in the dialogue stage. The Phylocode (Cantino & 

de Queiroz 2010) promotes phylogenetically based 

non-Linnaean nomenclature and is not relevant for the 

description of new species as presented here. 

In taxonomic language, species must be ‘effectively’, 

‘legitimately’ and ‘validly’ published. These three words 

have special meanings in taxonomic terminology (as 

do ‘illegitimate’ and ‘invalid’), and they should not be 

used in other ways in taxonomic manuscripts. 

1. To be effectively published (Arts. 29–31), i.e. to 

be made available, a description of a new species 

must be published in a journal that can be read 

by the scientific community. Species published 

in newspaper articles, or mentioned in oral 

presentations at scientific meetings, for example, 

are not considered effectively published. At present, 



Attribution: 






v o l u m e 1 · n o . 2 

109

Seifert & Rossman 

ARTICLE 

descriptions of new species cannot be published 

exclusively on electronic media such as CD-ROMs, 

DVDs or on the Internet. Effective publication is 

only accepted by the ICBN when at least two paper 

copies are archived in a scientific library or other 

despositary; however, we feel that many more 

printed copies should be deposited, preferably on 

each continent. Several mycological journals now 

publish articles including new species online and 

deposit printed copies in permanent libraries to 

meet effective publication requirements; the date 

of publication of the paper copy remains the official 

date, not the often earlier date of publication online. 

Academic PhD or other theses presented to 

universities as part of degree requirements are not 

considered effective publications, even if copies 

are distributed to other universities, unless they 

have an ISBN number or clearly state that they are 

to be intended effective publications (Art. 30.5). 

2. To be legitimately published (Art. 6), i.e. legally 

acceptable, a new species must have a unique 

binomial, i.e. it cannot have the same species 

epithet as another species validly published in the 

same genus. 

3. To be validly published (Arts 32–45), a new species 

must be clearly designated as a new species, have 

a Latin diagnosis, and a single, clearly designated 

and permanently preserved ‘type’, which fixes 

the application of the name. Usually, the type is 

preserved in a public herbarium (Holmgren et 

al. 1990) that will make the material available to 

interested scientists, has an on-line database of 

holdings, and that will assign a unique accession 

number, which you will then quote to clearly identify 

the type specimen. If there duplicates of the type 

specimen, or cultures derived from it, these must 

be explicitly distinguished from the holotype; the 

others are referred to as ‘isotypes’ or as ‘ex-type 

cultures’. To safeguard against loss and to facilitate 

access by other mycologists, isotypes should 

be deposited in several herbaria, on different 

continents if possible. 

There are many nuances to the concept of a type 

(Arts 7–10). For a new species, you will normally 

propose a ‘holotype’. The holotype is usually a 

dried, physiologically inert specimen (or a dried 

culture) that includes all diagnostic morphological 

characters of the species. For microscopic fungi, 

several separate individuals can be present as long 

as they are part of one sample collection, i.e. made 

at one time in a precise locality. Living cultures are 

now allowed as holotypes (Art. 8.4), but only if they 

are preserved in a metabolically inactive state (i.e. 

by lyophilization or in liquid nitrogen), ideally in an 

internationally recognized culture collection (see 

World Federation for Culture Collections, s.d.). 

This practice is not widely used in mycology except 

for yeasts. Cultures can be dried for use as type 

specimens (Constantinescu 1983); take care to dry 

an uncontaminated, optimally developed culture, 

not an old one that has started to degenerate. If 

you wish to designate a microscope slide as a type, 

or to include one with the type, it is worth the effort 

to make a permanent preparation using the method 

described by Kohlmeyer & Kohlmeyer (1972). 

Informal requirements 

To successfully describe a new species, the author 

must convince readers (especially reviewers and 

editors) that: 

1. The species is really undescribed. 

2. The species is being described in the most 

appropriate genus, and if molecular data are 

available, the genus including the new species 

remains monophyletic. 

3. The species is described, illustrated or otherwise 

characterized adequately so that it can be 

recognized again by subsequent workers. 

4. A sufficient number of cultures or specimens were 

examined. Ideally, new species should be described 

based on more than one specimen or culture, and 

some journals demand this. With limited material 

but clear taxonomic novelty, the author may be able 

to write a convincing argument for the proposal of 

a new species that is acceptable to editors and 

reviewers. 

Manuscripts that do not satisfy these criteria should 

not be published until they can be met. Normally, peer 

reviewers and editors assess whether these criteria 

are satisfied. 

In recent decades, partly as a result of the spirit 

of the UN Convention on Biological Diversity (CBD), 

taxonomists are encouraged, and sometimes legally 

required by national laws, to deposit type specimens 

in public reference collections in the country where 

the specimens were originally collected. If cultures 

were isolated, there may be a similar requirement 

or expectation from the originating country. Cultures 

of new species should be deposited in two or three 

internationally recognized public culture collections, 

which agree to make them available to other 

researchers. This latter practice is a condition for valid 

publication of new bacterial species (Lapage et al. 

1992), and is enforced as an editorial policy by some 

journals that publish new fungal species. 

It is critical that type specimens and cultures are 

available to other taxonomists who want to study 

and compare them with other material. The ICBN 

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recommends (Rec. 7A), but cannot enforce, that type 

specimens be deposited in public institutions with 

a policy to allow scientific researchers to examine 

material. A frequent problem is the unavailability 

of type or other specimens from under-resourced 

collections, or collections not curated by a mycologist. 

Some historical collections may never be sent on loan 

because of their fragility and extreme importance. 

Some nations forbid specimens or cultures from being 

sent abroad, under their interpretations of the CBD. A 

parallel situation is the reluctance of some industrial 

researchers (e.g. pharmaceutical companies) to 

allow access to cultures that they own. Balancing the 

question of open access to specimens or cultures 

against the legal or proprietary interests attached to 

that material is complicated, but must be considered 

when depositing type specimens. The scientific process 

demands reproducibility, and if this cannot be assured, 

responsible journals will not allow publication. The risk 

for the authors of species that cannot be re-examined 

or studied by other taxonomists is that the species 

will not be accepted by future scientists, and that the 

efforts and work of the authors of such species will be 

wasted and ignored. 

Almost all mycological journals now require 

that names and certain nomenclatural information 

for all newly proposed fungal taxa, including new 

combinations, be deposited in MycoBank (Crous et al. 

2004), and that the MycoBank accession number be 

included as part of the description. While the minimum 

requirements are the deposit of the Latin diagnosis and 

information on the type specimen, it is good practice 

to include as much information as possible, including 

illustrations, the English description, and links to 

molecular data, because this critical information will 

then be freely available to all scientists. 

genera should not be described in the absence of 

cultures or sequences. For other ascomycetes, singlespore 

cultures may yield unexpected anamorphs that 

will allow the description of a more complete life cycle. 

DNA sequences can be usually obtained from all 

but the most recalcitrant materials (such as fossils). 

There is a growing expectation that descriptions of 

all new species should be accompanied by molecular 

data, driven in part by the need for DNA sequence data 

to integrate new species into molecular phylogenies. 

The growth of molecular ecology, which relies on 

databases of reference sequences for identification 

of environmental sequences, has also highlighted the 

importance of sequencing all newly described species. 

Therefore, an increasing number of journals, editors 

or reviewers insist on cultures or DNA sequences 

before a manuscript is accepted for publication. If 

you do not have cultures or DNA sequences, your 

new species can only be published in a journal with 

different policies. We encourage mycologists who 

lack resources for culturing or DNA sequencing to 

collaborate with colleagues who can assist with this, 

often in return for co-authorship. 

ARTICLE 

Requirements for cultures and 

molecular data 

At present, there are no formal requirements that you 

must have cultures or DNA sequences of a fungus 

before you can describe a new species. Nevertheless, 

DNA sequences and cultures significantly enhance the 

value of a species description and you should make 

every effort to generate these resources. 

Mycologists describing new species should 

indicate whether they have tried to obtain cultures 

and what methods were attempted. Not all species 

can be cultured using currently available methods, 

but for most groups, culturing should be relatively 

straightforward after consulting the literature on related 

species. Cultures are essential for some groups where 

the modern morphological taxonomy is based entirely 

on in vitro characters, especially hyphomycetes such 

as Alternaria, Aspergillus, Cladosporium, Fusarium, 

Penicillium and Trichoderma; new species in these 

v o l u m e 1 · n o . 2 

111


ARTICLE 

Model Manuscript 

Title: Genus species sp. nov., an undescribed fungus (Taxonomic 

group) from habit in country with interesting biological properties 

Abstract: If your title is sufficiently engaging, a prospective reader will probably look next at 

the Abstract. The abstract should include all details necessary for the reader who does not 

have access to the whole article (i.e. someone looking at the abstract only on-line or in an 

abstract journal) so that they will know whether it is worth their time or money to obtain the 

full article. When describing a new species, you should include a summary of the diagnostic 

characters of the new species, especially the spore characters and dimensions. Make sure 

to include information about where your fungus was found and what it was growing on. If you 

have molecular data, it is useful to mention what genes you have sequenced, and what this 

information tells us about the fungus, such as what family or order it belongs to, and what are 

the most closely related species. Mention if a key to related species or comparative synoptic 

table is included, a feature that will increase potential readership. 

Key words: 

These should not 

reproduce words in the 

title. It is useful to list 

special techniques used 

in the description, e.g. 

electron microscopy, DNA 

sequencing, or chemotaxonomic 

methods. 

Article info: Submitted: dd month yyyy; Accepted: dd month yyyy; Published: dd month yyyy. 

Introduction 

The paragraphs of the introduction should be 

presented in a logical order, i.e. how they tell the best 

story. Remember, most people reading a scientific 

paper will only read the Introduction and Discussion, 

so the account of your discovery should be complete 

and complementary between these two sections. 

Normally, you will tell the reader about the larger 

projects (if any) that led to the discovery of the new 

species, provide information on its ecological niche 

and associated organisms, and give references to 

complementary publications where appropriate. 

Information should be provided about the taxonomy 

of genus in which the species is being described, 

such as the number of species already known, a brief 

review of recent revisions or monographs, perhaps 

discussion of controversies about the generic concept, 

and something about the biology of the species. Be 

diligent about citing all relevant literature. 

If you have DNA sequence data, usually one 

paragraph will provide a brief review of the existing 

state of molecular knowledge for the group your 

species belongs to, and explain to the reader what 

experiments you have done with your own species to 

fit it into the existing context. 

Another paragraph gives information about why 

the new species is suspected to be undescribed. 

This should be basic information that leads into the 

formal part of the paper. Some of this information 

may be repeated and presented in more detail in the 

discussion. 

Some papers may require a longer introduction. 

Manuscripts including molecular or physiological data 

are often longer. Situations where the new fungus 

could be described in one of several different genera 

also may require a longer introduction. Sometimes 

this will include a more extensive review of historical 

literature, or discussions of taxonomic characters of 

particular significance. You must judge whether this 

information is most suitable in the introduction, which 

the reader will read before the description itself, or if it 

is more logical to place it in the discussion. 

Usually the Introduction concludes with a statement 

like, “therefore, we decided that our fungus represents 

an undescribed species, which is described and 

illustrated here as Genus species sp. nov.” 


This section is often omitted from taxonomic papers 

that include only morphological data, but it is preferable 

to include as many details as possible. Some of the 

following subheadings and paragraphs might be 

appropriate. 

Collecting and field sites 

How the specimens were collected and transported 

to the laboratory and preserved or incubated prior to 

examination may be relevant. Information about specific 

field sites is usually given in the ‘Material examined’ 

section, but it might be appropriate to provide more 

details here if they are relevant to understanding the 

species. 

Cultures and media 

Give recipes for the isolation media employed, or cite a 

reference for the media. Give brand names for extracts 

used, such as malt extract, yeast extract, and the agar 

used for the media. Describe the inoculation methods, 

incubation conditions such as temperature and lighting 

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i m a f U N G U S


regime, and length of incubation before examination. 

Determine the cardinal growth temperatures (minimum, 

optimum and maximum) if possible. List the culture 

collections where the cultures are maintained, with 

accession numbers, either here or in a table. 

Isolation methods 

Explain all isolation methods used, such as explants 

from sporophores or infected host tissues, removal of 

spores directly from sporulating structures, transfer 

of actively discharged spores from Petri dish lids or 

spore prints, etc. If the substrate was treated before 

isolation, e.g. by some form of surface sterilization, 

these methods should be explained. Give the recipes 

for the isolation media employed including any 

antibacterial compound added, or cite a reference 

for those media. Describe the incubation conditions 

such as temperature and lighting conditions. If singlespore 

cultures were prepared, a good practice to be 

undertaken when possible (Choi 1999, Crous 2002), 

describe how this was done. 

Microscopy 

Give details of the kind of microscope used, including 

the illumination systems (e.g. phase contrast, 

differential interference contrast), the mounting media 

and stains employed for routine examination and for 

making measurements, and how many structures 

of each microscopic character were measured. A 

similar section, with details on dehydration protocols, 

fixation, staining, etc. should be given for electron 

microscopy methods, if these were used. Permanent 

microscopic preparations should be deposited with 

the type if possible, which will make your observations 

reproducible to later taxonomists, and limit the amount 

of material they might use on specimens in future 

studies. 

Techniques used for illustrations are often given 

here, but may also be briefly mentioned in the figure 

legends. For example, whether drawings were made 

with a drawing tube, a camera lucida, or by freehand, 

and the type of camera used for photography, may 

be relevant. It is essential to describe techniques 

used for image enhancement of digital photographs, 

such as the sharpening filters of PhotoShop or other 

imaging software, whether separate photographs were 

combined into one image, or whether colours have 

been altered (Microscopy Society of America, 2003). 

Modification of contrast has always been standard 

practice in photography and need not be mentioned. 

Physiological tests or chemotaxonomic 

methods 

For yeasts, substrate utilization and other physiological 

tests are standard parts of taxonomic descriptions. For 

some lichen groups, spot-tests with a standard set of 

chemical reagents are essential. These methods must 

be described carefully. If chemotaxonomic methods, 

such as isozyme analysis or secondary metabolite 

profiling, were employed, complete methods should be 

given to allow the resulting data to be reproduced. 

DNA extraction, PCR amplification, DNA 

sequencing, and phylogenetic analysis 

Note what kind of material was used for DNA 

extraction, such as cultures, single spores, or naturally 

occurring tissues, and give the details of the kits and 

methods used for DNA extraction. For DNA isolations 

from natural tissues, note whether procedures were 

repeated to reduce the chance of sequencing a 

contaminant or associated organism. Give the details 

of PCR amplification profile used, the concentrations of 

reagents used in the reactions, and information on the 

brand and model of thermocycler used. Provide details 

of any methods used to clean or otherwise process 

the PCR products before sequencing. If PCR products 

or other DNA fragments were cloned for sequencing, 

provide the relevant information for this procedure. 

For the DNA sequencing, provide details of the cycle 

sequencing profile used, the concentration of reagents 

used, the brand and model of the thermocycler, the 

relevant information about the sequencing chemistry 

used, and the brand and model of DNA sequencer 

employed. If you used a DNA sequencing service, list 

it here. Cite the literature where PCR and sequencing 

primers were first published. If you designed the 

primers yourself, give details of how you did this. 

Include details about how you did your phylogenetic 

analyses, including literature citations for sequences 

originating from published or unpublished work of 

colleagues, the software used for analysis, and the 

details of the parameters used for the analysis. Diverse 

methods of phylogenetic analysis are available. While 

the choice of methods is largely a matter of preference, 

there is a general agreement that it is critical to employ 

measures of confidence, such as the bootstrap, 

Bayesian posterior probabilities, the ‘decay index’ or 

congruence among independent data sets. If you have 

used several sequences from a previously published 

study, you should cite that study so that it is properly 

credited (Seifert et al. 2008). 

Results 

Many descriptions of new species will not have a 

Results section because all of the data are included in 

the Taxonomy section. However, if some experiments 

were done with the fungus, such as physiological tests 

or tests of antibiotic resistance, then these data should 

be presented in the Results section, in the same order 

as the methods are described in the Materials and 

Methods section. Often, the results of physiological 

tests are given in a Table. 

ARTICLE 

v o l u m e 1 · n o . 2 

113


ARTICLE 

General results of DNA sequencing analyses are 

usually given in this section. These can include details 

of the length and composition of the DNA fragments, and 

the results of comparisons with other sequences (e.g. 

BLAST searches). If phylogenetic trees are presented, 

the tree statistics may be given in this section, or 

otherwise in the figure legends. Describe what the data 

shows, e.g. that the sequence is similar to those in a 

particular genus, family or order, or that the sequence 

is apparently unique, but leave the conclusions derived 

from this for the Discussion section. Mention support 

values for the critical nodes in your tree based on 

bootstrap frequencies or other measures of confidence. 

If you have done analyses using different phylogenetic 

methods, or have analyzed several genes, then 

comparison of the results is appropriate here, but leave 

the conclusions for the Discussion section. 

TAXONOMY 

Genus species Authors 1 , sp. nov. 

MycoBank: MBxxxxxxx. 

Figs xxx–xxx 

The Latin diagnosis comes first and is essential for 

valid publication. It should list the diagnostic characters 

only, i.e. those that separate it from similar ones, and 

not be a complete translation of the description. Many 

journals now restrict the Latin to a few lines. Use 

published Latin diagnoses for models, then if possible 

have yours checked by a mycologist or botanist 

competent in scientific Latin. If there is no such expert 

in your own department, consult with colleagues in 

other institutions. Stearn’s Botanical Latin (1992) is a 

valuable resource for preparing Latin diagnoses. 

Holotypus: Collection acronym, accession number. 

Immediately after the Latin diagnosis, clearly and 

explicitly indicate the details of the single accession 

that will serve as holotype. If you wish to list isotypes 

or extype cultures here, be certain that they are clearly 

distinguished from the holotype or you may have 

problems with the validity of your new name (Art. 37.7). 

and moves towards the finest. Macroscopic characters 

are described next. Use a colour standard, such as 

Ridgway (1912), Rayner (1970), Kornerup & Wancsher 

(1984), or Munsell (1905 and many subsequent editions) 

to accurately describe colours. Most details will be 

visual, but sometimes texture and odour are useful 

additions. When describing microscopic characters, be 

as complete as possible about shape, colour, texture 

and size for every component of the fungus. There are 

standard terminologies for shape; check Ainsworth & 

Bisby’s Dictionary of the Fungi (Kirk et al. 2008, and 

earlier editions, see under ‘shapes’) as a starting 

point. Be aware that the terminology for describing 

three dimensional shapes sometimes differs from the 

terminology used to describe two dimensional shapes. 

The full range of observed dimensions should be given 

for all structures. Means should be calculated for all 

dimensions in the description (at least the spores), 

along with a statistical measure of variation in these 

measurements, such as standard error, standard 

deviation, confidence intervals or percentile ranges. 

If you have isolated a culture, its features are 

usually included in a separate paragraph. For some 

fungi, colony characters are described first; for others, 

this information follows the morphological description. 

At a minimum, give the growth rates on a specified 

medium and explain the temperature and light regime, 

and give a general impression of the colour and texture 

of the colonies. If the fungus sporulates in culture, it 

can be very helpful to compare the sizes and shapes 

of the microscopic structures to what occurred on 

the natural specimen. The detail employed in culture 

descriptions varies considerably from one taxonomic 

group to another, and you should consult published 

descriptions for the group you are working with. 

If you have done any physiological tests, or 

determined cardinal temperatures, this information is 

normally put in a separate paragraph in the description. 

It can also be put into a table or in a graph, which may 

be easier for a reader to follow. 

Substrate or Host: Provide a summary of the known 

hosts or substrates as a separate paragraph, especially 

if you have more than one specimen. 

A full description follows. Think of the descriptions and 

illustrations together as providing a blue-print for your 

new species. If someone wanted to build an exact scalemodel 

of the fungus, they should be able to do so using 

your paper. Where both asexual and sexual states (i.e. 

anamorph and teleomorph) occur, the description of 

the sexual state is traditionally given first. In general, 

a taxonomic description begins at the broadest scale 

1 

Names of author or authors, and their abbreviations, should 

follow the standards of Kirk & Ansell (1992). 

Distribution: Summarize the known disttibution, by 

continent, country (and by province or state for larger 

countries), along with relevant information on the 

biome, climactic or geological conditions. 

Etymology: Explain the meaning or derivation of the 

species epithet, and note the language of origin of the 

word(s) used for constructing the epithet. Avoid species 

epithets with more than five syllables and those similar 

to other epithets in the same genus. Epithets that are 

descriptive are most helpful, but names can be derived 

from any source, including acronyms or the name of a 

114 

 

i m a f U N G U S


ARTICLE 

Fig. 1. Genus species (specimen or culture number, noting whether it is type). A series of photographs of the fungus, showing the field habit, 

the appearance under the dissecting microscope, and microscopic photographs showing taxonomically relevant structures and preferably some 

developmental sequence. All illustrations should include scale bars. In this particular example: 

Fig. 1. Sarcinella questierii (DAOM 235813). A. Black growth on living leaves of Cornus sp. B. Black conidia on leaf surface. C. Development of 

dictyoconidia from conidiogenous hyphae, with hyphopodia (h) arising from the same hyphae (differential interference contrast). Bars: A = 1 cm, 

B = 25 µm, C = 10 µm. B–C, composite images created with CombineZ (Hadley 2006). 

person, usually someone involved in the discovery of 

the fungus or a mycologist who has made a significant 

contribution to the subject. 

Additional material examined: Most journals have 

a specific format for this part of the paper. For all 

specimens and cultures, including isotypes and extype 

cultures, list Country: Province/State/Territory/ 

County/Township, City/Town/Park, Specific location 

details (GPS coordinates, including altitude), Substrate 

or host, Date of collection and/or isolation, Collector’s 

name, Collector’s number (if any), Herbarium or culture 

collection abbreviations and accession numbers 

where the material is preserved. Often, some of this 

information is instead provided in a table including 

GenBank accession numbers for DNA sequences. 

Discussion 

The discussion completes the story that began in the 

Introduction. There are many ways to write this section, 

but one rule is not to introduce new data that should 

have been introduced in the ‘Results’ or ‘Taxonomy’ 

sections. Mixing of the Results and Discussion in one 

section is generally frustrating for the reader, unless 

the section is very short. 

It is often useful to start the Discussion by 

summarizing the diagnostic features of the fungus you 

have described. 

In a separate paragraph, you should compare your 

fungus to other similar species of the same genus, 

stating clearly how they differ. Many papers will include 

either a diagnostic key or synoptic table (or both), either 

including all species of a smaller genus, or only the 

most similar species of a larger genus, to assist the 

reader in understanding why the new species is distinct. 

If you have DNA sequence data, there are often 

several paragraphs of discussion relating to what they 

show or do not show. Discuss how the data support 

the classification of your fungus and its recognition as 

a distinct species. If you have done analyses using 

different phylogenetic methods, or have analyzed 

several genes, compare the results and explain your 

conclusions carefully, especially if contradictory 

evidence occurs in the different data sets or analyses. 

It is often useful to conclude the paper with 

discussion of the biology of the new species, either 

demonstrated by the experiments done in the paper or 

as an extension of field observations. A limited amount 

of speculation on this topic is usually tolerated by 

reviewers and editors. 

Illustrations 

The ICBN does not require that new species descriptions 

have illustrations (Fig. 1), but few journals allow 

the description of a new species without them, with 

the exception of yeasts. Usually, at least some of 

the illustrations will be of the holotype specimen or 

culture. The package of illustrations should present the 

complete concept of the species to the reader so that 

they can confidently identify your fungus. Provide visual 

information at several different size scales, from general 

habitat to the most detailed microscopy. Expectations vary 

among different taxonomic groups, but often a mixture of 

photographs and line drawings are included. Individual 

photographs are visual data that are proof of observations. 

Electron micrographs are generally unhelpful to facilitate 

identification, but scanning electron micrographs may 

be necessary for documenting spore ornamentation or 

tissue types, and transmission electron micrographs 

may be necessary to prove ultrastructural observations 

v o l u m e 1 · n o . 2 

115


ARTICLE 

of spore production (especially types of conidiogenesis). 

Line drawings are interpretations; they are not proof, 

but can present a complex concept in one image and 

be extremely helpful for someone trying to identify your 

species. With colour photographs now published with 

increasing frequency, and the relative economy and 

ease of digital photography and computerized imaging, 

the preparation of informative illustrations is one of the 

most exciting aspects of describing a new species. Find 

the best model illustrations for the group of organisms 

where your species fits, and then do better! 


In addition to the usual acknowledgements for a scientific 

paper (e.g. mentors, sources of funding), it is traditional 

to acknowledge any taxonomic specialists that you 

have consulted during your decision to describe a new 

species. Similarly, it is customary to acknowledge the 

curators of any collections who have provided specimens 

or cultures that was used in your study. For this paper, 

we wish to thank the other members of the International 

Commission on the Taxonomy of Fungi, especially David 

Hibbett and David Hawksworth, and the peer reviewers, 

for valuable comments on earlier drafts of this paper. The 

final sentence usually identifies funders of the research. 

References 

Do your best to cite the relevant historical and 

modern literature, including revisions, monographs, 

identification keys, and molecular studies that you 

have consulted. Reviewers often see submitted papers 

in some fields of taxonomy that only cite literature 

more than 25 years old. Study the guidelines of the 

journal carefully for citation formats. 

Cantino PD, Queiroz K de (2010) International Code of 

Phylogenetic Nomenclature. Version 4c. 

Choi Y-W, Hyde KD, Ho WH (1999) Single spore isolation of 

fungi. Fungal Diversity 3: 29–38. 

Constantinescu O (1983) Dried reference fungal cultures. 

A review and a simpler technique. Bulletin of the British 

Mycological Society 17: 139–143. 

Crous PW (2002) Adhering to good cultural practice (GCP). 


Crous PW, Gams W, Stalpers JA, Robert V, Stegehuis G 

(2004) MycoBank: an online initiative to launch mycology 

into the 21st century. Studies in Mycology 50: 19–22. 

Hadley A (2006) CombineZ. Version 5. Published by the 

author. 

Hawksworth DL (1974) Mycologist’s Handbook: an introduction 

to the principles of taxonomy and nomenclature in the fungi 

and lichens. Kew: Commonwealth Mycological Institute. 

Holmgren, PK, Holmgren NH, Barnett LC (1990) Index 

Herbariorum. Part I. The Herbaria of the World. 8 th edn. 

[Regnum Vegetabile vol. 120.] Utrecht: Bohn, Scheltema 

& Holkema. 

Kirk PM, Ansell AE (1992) Authors of Fungal Names: 

A list of authors of scientific names of fungi, with 

recommended standard forms of their names, including 

abbreviations. [Index of Fungi Supplement.] Wallingford, 

UK: CAB International. 

Kirk PM, Cannon PF, Minter DW, Stalpers JA, eds (2008) 

Ainsworth and Bisby’s Dictionary of the Fungi. 10 th edn. 

Wallingford, UK: CAB International. 

Kohlmeyer J, Kohlmeyer E (1972) Permanent microscopic 

mounts. Mycologia 64: 666–669. 

Kornerup A, Wanscher JH (1984) Methuen Handbook of 

Color. 3 rd edn. London, UK: Methuen. 

Lapage SP, Sneath PHA, Lessel EF, Skerman VDB, Seeliger 

HPR, Clark WA (eds) (1992) International Code of 

Nomenclature of Bacteria and Satutes of the International 

Committee on Systematic Bacteriology and Statutes 

of the Bacteriology and Applied Microbiology Section 

of the International Union of Microbiological Societies. 

Bacteriological Code (1990 Revision). Washington, DC, 

USA: American Society for Microbiology. 

McNeill J, Barrie FR, Burdet HM, Demoulin V, Hawksworth 

DL, Marhold K, Nicolson DH, Prado J, Silva PC, Skog 

JE, Wiersema JH, Turland NJ, eds (2006) International 

Code of Botanical Nomenclature (Vienna Code). [Regnum 

Vegetabile vol. 146.] Liechtenstein: Gantner Verlag, 

Ruggell. 

Microscopy Society of America (2003) MSA policy on digital 

imaging. www.microscopy.org/resources/digital_imaging. 

cfm 

Munsell AH (1905) A Color Notation. Boston: G. H. Ellis. 

(for a pop-up computer 

version). 

Rayner RW (1970) A Mycological Colour Chart. Kew: 

Commonwealth Mycological Institute. 

Ridgway R (1912) Color Standards and Color Nomenclature. 

Washington, DC, USA: published by the author. 

Seifert KA, Crous PW, Frisvad JC (2008) ACT: Appropriate 

Citation of Taxonomy. Inoculum 59(3): 4; Persoonia 20: 

105. 

Sigler L, Hawksworth DL (1987) International Commission on the 

Taxonomy of Fungi (ICTF): Code of practice for systematic 

mycologists. Mycologist 1: 101–105; Mycopathologia 99: 

3–7; Microbiological Sciences 4: 83–86. 

Stearn WT (1992) Botanical Latin: history, grammar, syntax, 

terminology and vocabulary. 4 th edn. Newton Abbot: David 

& Charles. 

Winston JE (1999) Describing Species: practical taxonomic 

procedure for biologists. New York: Columbia University 

Press. 

World Federation for Culture Collections. s.d. 

116 

i m a f U N G U S


What is Johansonia? 

Pedro W. Crous 1 , Robert W. Barreto 2 , Acelino C. Alfenas 2 , Rafael F. Alfenas 2 and Johannes Z. Groenewald 1 

1 

CBS-KNAW Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands; corresponding author e-mail: 

p.crous@cbs.knaw.nl 

2 

Departemento de Fitopatologia, Universidade Federal de Viçosa, 36.570 Viçosa, MG, Brazil 

ARTICLE 

Abstract: The bitunicate ascomycete genus Johansonia is presently treated as a member of 

Saccardiaceae, a family regarded as incertae sedis within the Ascomycota. Recent collections on 

leaves of a leguminous host, Dimorphandra mollis, in Mato Grosso, Brazil, led to the discovery of a 

new species of Johansonia, described here as J. chapadiensis. Based on DNA sequence data of the 

nuclear ribosomal DNA (LSU), Johansonia is revealed to represent a member of Dothideomycetes, 

Capnodiales. Although its family could not be resolved, it clustered basal to Schizothyriaceae and 

Mycosphaerellaceae, and could well represent a species of Saccardiaceae. DNA sequence data of 

other members of Saccardiaceae would be required, however, to confirm this classification. 

Key words: 

Dothideomycetes 

Johansoniella 

ITS 

LSU 

systematics 

Article info: Submitted: 19 October 2010; Accepted: 24 October 2010; Published: 2 November 2010. 

Introduction 

The genus Johansonia is based on J. setosa (Saccardo 

1889), a species known from leaves of Sapindaceae 

collected in South America (Müller & von Arx 1962). 

Due on its superficial discoid ascomata, bitunicate asci 

and hyaline, 1-septate ascospores, Müller & von Arx 

(1962) were of the opinion that the genus belonged 

to Schizothyriaceae. In a later study, however, von 

Arx & Müller (1975) again placed it in Saccardiaceae, 

suborder Dothideaceae in Dothideales, based on 

the ascomata having an epithecium of branched 

hyphal elements. Barr (1993) again placed it in 

Phillipsiellaceae in Loculoascomycetes, while 

Lumbsch & Huhndorf (2007) concluded that it was 

a member of Saccardiaceae, a family they regarded 

as incertae sedis in Ascomycota. In recent studies 

on Dothideomycetes (Schoch et al. 2006, 2009), no 

mention is made of Johansonia. As there are presently 

no DNA sequence data represented for any species 

of Johansonia in GenBank, its taxonomic position 

remains obscure. 

During a recent visit to Brazil, we collected fresh 

material of a species of Johansonia on leaves of a 

legume. The aims of the present study were, therefore, 

to identify the species of Johansonia, and at the same 

time to see if the taxonomic position of the genus could 

not be resolved. 


Isolates 

Leaves bearing ascomata were soaked in water for 

approximately 2 h, after which they were placed in 

the bottom of Petri dish lids, with the top half of the 

dish containing 2 % malt extract agar (MEA; Crous 

et al. 2009c). Ascospore germination patterns were 

examined after 24 h, and single ascospore and conidial 

cultures established as described earlier (Crous et 

al. 1991, Crous 1998). Colonies were subcultured 

onto potato-dextrose agar (PDA), oatmeal agar (OA), 

MEA (Crous et al. 2009c), and incubated at 25 °C 

under continuous near-ultraviolet light to promote 

sporulation. Reference strains are maintained in the 

CBS-KNAW Fungal Biodiversity Centre (CBS) Utrecht, 


DNA isolation, amplification and 

analyses 

Genomic DNA was isolated from fungal mycelium 

grown on MEA, using the UltraCleanTM Microbial DNA 

Isolation Kit (MoBio Laboratories, Inc., Solana Beach, 

CA, USA) according to the manufacturer’s protocols. 

The primers V9G (de Hoog & Gerrits van den Ende 1998) 

and LR5 (Vilgalys & Hester 1990) were used to amplify 

part of the nuclear rDNA operon spanning the 3’ end 

of the 18S rRNA gene (SSU), the internal transcribed 

spacer 1, the 5.8S rRNA gene, the internal transcribed 

spacer 2 (ITS) and the first 900 bases at the 5’ end of 

the 28S rRNA gene (LSU). The primers ITS4 (White et 



Attribution: 






v o l u m e 1 · n o . 2 

117

Crous et al. 

ARTICLE 

Phaeobotryosphaeria visci DQ377868 

10 changes 

100 

56 

86 

64 

81 

70 

52 

89 

74 

98 

100 

56 

97 

63 

92 

93 

68 

50 

Devriesia strelitziicola GU214417 

Devriesia strelitziae EU436763 

Devriesia lagerstroemiae GU214415 

Devriesia hilliana GU214414 

Teratosphaeria knoxdavesii EU707865 

Pseudoramichloridium henryi GQ303320 

91 

Staninwardia suttonii DQ923535 

Devriesia americana EU040227 

Devriesia pseudoamericana CPC 16174 

Penidiella pseudotasmaniensis GQ852625 

Penidiella tasmaniensis DQ246233 

Pseudocercosporella fraxini GU214682 

100 

Penidiella kurandae EU040214 

Devriesia thermodurans EU040229 

Devriesia shelburniensis EU040228 

100 Devriesia acadiensis EU040226 

95 Devriesia staurophora EF137359 

Penidiella strumelloidea EU019277 

100 

Microcyclospora malicola GU570549 

Microcyclospora sp. 1 FJ147168 

Microcyclospora malicola GU570550 

Microcyclospora tardicrescens GU570552 

Microcyclospora pomicola GU570551 

Microcyclospora sp. 2 FJ147169 

Johansonia chapadiensis CPC 18474 

Zygophiala cryptogama FJ147157 

Zygophiala wisconsinensis FJ147158 

Zygophiala sp. FJ147159 

Zygophiala sp. FJ147156 

Schizothyrium pomi GU570554 

Schizothyrium pomi EF134949 

Schizothyrium pomi EF134948 

Mycosphaerella intermedia DQ246248 

Mycosphaerella marksii GQ852614 

Mycosphaerella madeirae DQ204756 

63 

61 

73 

93 

74 

72 55 

76 

Pseudocercosporella sp. FJ031991 

Microcyclosporella mali GU570547 

Microcyclosporella sp. FJ031995a 

Microcyclosporella sp. FJ031995c 

Microcyclosporella sp. FJ031995b 



Microcyclosporella sp. FJ031990 





90 

100 

100 

54 

55 

100 

62 

76 

54 

62 

74 

Pseudocercospora vitis DQ073923 

Pseudocercospora natalensis DQ267576 

Pseudocercospora paraguayensis GQ852634 

Cercosporella virgaureae GQ852585 

Ramulispora sorghi GQ852653 

Pseudocercosporella capsellae GU214662 

Pseudocercosporella sp. GU214683 

83 

65 

Septoria leucanthemi GQ852677 

Septoria senecionis GQ852678 

Septoria apiicola GQ852674 

Septoria convolvuli GQ852675 




Mycosphaerella endophytica DQ246255 

Mycosphaerella pseudoendophytica DQ246253 

Mycosphaerella endophytica GQ852603 

Passalora daleae EU040236 

Mycosphaerella pini GQ852597 

Passalora fulva DQ008163 

Dothistroma pini GQ852596 

Pseudocercosporella bakeri GU570553 

Passalora bellynckii GQ852618 

98 

Mycosphaerella microsora EU167599 

Passalora brachycarpa GQ852619 

Mycosphaerella africana DQ246258 

Mycosphaerella ellipsoidea GQ852602 

Mycosphaerella buckinghamiae EU707856 

Mycosphaerella aurantia DQ246256 

Mycosphaerellaceae Schizothyriaceae 

Teratosphaeriaceae 

Incertae 

sedis 

Fig. 1. The first of 1000 equally most parsimonious trees obtained from a heuristic search with 100 random taxon additions of the LSU sequence 

alignment. The scale bar shows 10 changes, and bootstrap support values from 1000 replicates are shown at the nodes. The novel sequence 

generated for this study is shown in bold. Branches present in the strict consensus tree are thickened and important lineages are colour-coded. 

The tree was rooted to a sequence of Phaeobotryosphaeria visci (GenBank accession DQ377868). 

118 

 

i m a f U N G U S

Johansonia 

al. 1990) and LSU1Fd (Crous et al. 2009b) were used 

as internal sequence primers to ensure good quality 

sequences over the entire length of the amplicon. The 

PCR conditions, sequence alignment and subsequent 

phylogenetic analysis followed the methods of Crous 

et al. (2006, 2009a). Sequences were compared with 

the sequences available in NCBI’s GenBank nucleotide 

(nr) database using a megablast search and results 

are discussed in the relevant species notes where 

applicable. Based on the Blast results, the novel 

sequence was added to the alignment of Frank et al. 

2010 (TreeBASE study S10547). Alignment gaps were 

treated as new character states. Sequences derived in 

this study were lodged at GenBank, the alignment in 

TreeBASE (), and 

taxonomic novelties in MycoBank (; Crous 

et al. 2004). 

Morphology 

The morphological description is based on preparations made 

from host material in clear lactic acid, with 30 measurements 

determined per structure, and observations made with a Nikon 

SMZ1500 dissecting microscope, and with a Zeiss Axioscope 

2 microscope using differential interference contrast (DIC) 

illumination. Colony characters and pigment production were 

noted after 2 wk of growth on MEA, PDA and OA (Crous et 

al. 2009c) incubated at 25 ºC. Colony colours (surface and 

reverse) were rated according to the colour charts of Rayner 

(1970). Growth characteristics were studied on MEA plates 

incubated for 2 wk in the dark at 25 °C. 

RESULTS 

Phylogeny 

Approximately 1700 bases, spanning the ITS and LSU 

regions, were obtained from the sequenced culture. The 

LSU region was used in the phylogenetic analysis for the 

generic placement (Fig. 1) and ITS to determine specieslevel 

relationships (see notes under species descriptions). 

The manually adjusted LSU alignment contained 77 taxa 

(including the Phaeobotryosphaeria visci outgroup sequence) 

and, of the 731 characters used in the phylogenetic analysis, 

171 were parsimony-informative, 96 were variable and 

parsimony-uninformative and 464 were constant. Only the 

first 1000 equally most parsimonious trees were retained 

from the heuristic search, the first of which is shown in 

Fig. 1 (TL = 776, CI = 0.485, RI = 0.839, RC = 0.407). The 

phylogenetic tree of the LSU region (Fig. 1) show that the 

obtained sequence clusters basal to the Schizothyriaceae. 

Etymology: Named after the location where the holotype was 

collected, Chapada dos Guimarães, Mato Grosso, Brazil. 

Johansoniae brasiliensis morphologice similis, sed ascosporis 

minoribus, (13–)15–19(–24) × (5–)6–7 mm, discernitur. 

Typus: Brazil: Mato Grosso, Chapada dos Guimarães, on leaves 

of Dimorphandra mollis (Leguminosae; False Barbatimao), 18 Aug. 

2010, P.W. Crous, A.C. Alfenas & R. Alfenas, (CBS H-20484 – 

holotypus, cultures ex-holotype CPC 18475, 18474 = CBS 128068). 

(GenBank accession numbers: ITS, HQ423449; LSU, HQ423450). 

Leaves with brown spots, but ascomata also occurring on 

dead and green leaf areas. Mycelium superficial, consisting of 

septate, branched, medium brown, verruculose to warty, 2–5 

µm wide hyphae. Ascomata on lower leaf surface, superficial, 

situated on a hyphal stroma (occurring loosely on surface), 

discoid, dark brown, up to 300 µm diam, 200 µm high. Exciple 

15–20 µm diam, consisting of 3–6 layers of brown textura 

angularis to textura globulosa. Asci in parallel layer, bitunicate 

with ocular chamber, sessile, narrowly ellipsoid to subcylindrical 

or clavate, 8-spored, 32–45 × 11–19 µm. Paraphyses 

intermingled among asci, hyaline, branched, septate, 1.5–2.5 

µm wide, becoming somewhat darkened and branched towards 

the apical region, forming an epithecium. Ascospores hyaline, 

thick-walled, medianly 1-septate, thick-walled, constricted at the 

septum, prominently guttulate, (13–)15–19(–24) × (5–)6–7 µm. 

Ascospores after 24 h on MEA germinating from both ends, with 

germ tubes parallel to the long axis of the spore, developing 

lateral branches; ascospores remaining hyaline, prominently 

constricted, not distorting, 5–7 µm wide. Setae brown, erect, 

straight to curved, separate and surrounding ascomata, thickwalled, 

brown, smooth, with basal T-cell devoid of rhizoids, with 

slight taper towards apical cell, which is thin-walled, pale brown, 

and acutely to obtusely rounded, 5–10-septate, 130–260 × 4–5 

µm; 2.5–3 µm wide at apical septum. 

Culture characteristics: Colonies spreading, erumpent, with 

sparse aerial mycelium and diffuse, submerged margins. 

On PDA surface pale mouse-grey (centre), olivaceous-grey 

(middle) with smoke-grey to cream outer region; reverse 

olivaceous-grey; colonies reaching 5 mm diam. On OA 

smooth, somewhat slimy, surface umber to dark mousegrey; 

margin diffuse, reaching 8 mm diam. On MEA, surface 

smoke-grey; reverse greyish-sepia, reaching 10 mm diam 

after 2 wk. 

Additional specimen examined: Brazil: Pernambuco: Poço do 

Macaco, on Inga sp., 18 Sept. 1960, Osvaldo Soares de Silva (CBS 

H-5029 – isotype of Johansonia brasiliensis). 

ARTICLE 

Taxonomy 

Johansonia chapadiensis Crous, R.W. Barreto, 

Alfenas & R.F. Alfenas, sp. nov. 


(Fig. 2) 

Notes: The generic name Johansonia is based on J. setosa, 

a species described from living leaves of Sapindaceae 

collected in South America. The genus is characterised by 

having loose, superficial, discoid ascomata situated on a 

hyphal stroma, and an exciple covering the bitunicate asci. 

Paraphyses, which are intermingled among asci, are hyaline, 

v o l u m e 1 · n o . 2 

119

Crous et al. 

ARTICLE 

Fig. 2. Johansonia chapadiensis (CBS H-20484 – holotype). A. leaves colonised with J. chapadiensis. B–E. Ascomata on leaf surface from 

above (B, C), below (D), and a vertical section though an ascoma (E). F, L. Germinating ascospores. G. Vertical section through ascoma. H–J. 

Asci. K. Ascospores. Bars: B, C = 300 µm; D, E = 150 µm; G, H = 20 µm; F, I–L = 10 µm. 

branched, septate, and become somewhat darkened and 

branched towards the apical epithecium. Ascospores are 

hyaline and 1-septate. Ascomata are surrounded by brown, 

erect, straight to curved, septate setae (Müller & von Arx 

1962). Based on these features, J. chapadiensis is a typical 

member of the genus Johansonia. 

Morphologically, J. chapadiensis closely resembles J. 

brasiliensis (Fig. 3). The two species can be distinguished in 

that ascospores of J. chapadiensis are smaller, (13–) 15–19(– 

24) × (5–)6–7 µm, than those of J. brasiliensis, (18–24 × 6–7 

µm). Furthermore, asci of J. chapadiensis are narrowly ellipsoid 

to subcylindrical or clavate, 32–45 × 11–19 µm, while those of 

J. brasiliensis are broadly ellipsoid, obovoid to subcylindrical, 

never clavate, and larger, 40–58 × 15–23 µm. Finally, setae 

in J. chapadiensis are more acutely rounded, 2.5–3 µm diam 

at the apical septum, while those of J. brasiliensis are bluntly 

rounded, and wider at the apical septum, 4–6 µm diam. 

Discussion 

Although there are only 12 species of Johansonia listed in 

Index Fungorum, von Arx & Müller (1975) were of the opinion 

that Johansoniella maranhensis represented a further species 

of Johansonia. Batista et al. (1966) introduced the monotypic 

generic name Johansoniella (Schizothyriaceae), based 

on J. maranhensis, which they regarded as closely related 

to Johansonia. Morphologically, the description appears 

120 

 

i m a f U N G U S

Johansonia 

ARTICLE 

Fig. 3. Johansonia brasiliensis (CBS H-5029 – isotype). A. Ascoma on leaf. B–D. Asci and ascospores. Bars: A = 300 µm; B–D = 10 µm. 

somewhat different, as the ascomata are described as having 

an upper wall layer (though this may be an epithecium), and 

setae around the ascomata, as well as on top of the ascomata. 

Regardless of these supposed differences, von Arx & Müller 

(1975) treated Johansoniella in synonym with Johansonia. A 

re-examination of the holotype specimen (URM 47621) found 

it to be depauperate, and hence the status of Johansoniella 

could not be resolved in the present study. 

An attempt to make a key to the species described to date 

based on published descriptions has not proven feasible, as 

too many species either have similar ascospore dimensions, 

or are insufficiently known. Based on published descriptions, 

most taxa only seem distinct if aspects such as dimenions 

of the ascospores, asci and setae are combined with host 

and distribution. However, as most taxa have been recorded 

once only, the value of these characters seems unreliable, 

and hence a key would only be feasible once the specimens 

of all described taxa have been re-examined to help resolve 

possible species synonymies. 

Recent studies focused on elucidating the higher order 

phylogeny of Dothideomycetes (Schoch et al. 2009) and 

Capnodiales (Crous et al. 2009b) did not treat Johansonia, as 

the present collection represents the first known cultures of 

this genus. Von Arx & Müller (1975) were of the opinion that 

Johansonia belonged to Saccardiaceae, a treatment accepted 

by Lumbsch & Huhndorf (2007), though they regarded it as 

a family incertae sedis within Ascomycota. Based on the 

DNA phylogeny generated in the present study (Fig. 1), we 

can reveal that Johansonia belongs to the Dothideomycetes 

(Capnodiales), and is closely related to Schizothyriaceae and 

Mycosphaerellaceae. However, whether it is a member of the 

Saccardiaceae (von Arx & Müller 1975, Lumbsch & Huhndorf 

2007), could not be confirmed, as presently there are no 

known cultures of this family available for comparison. Parts 

of Saccardiaceae have been transferred to Schizothyriaceae 

von Arx & Müller (1975), and thus its close relationship to the 

Schizothyriaceae suggests Saccardiaceae a likely family for 

this genus, pending further collections and study. 


We thank the technical staff, Arien van Iperen (cultures), Marjan 

Vermaas (photo plates), and Mieke Starink-Willemse (DNA isolation, 

amplification and sequencing) for their invaluable assistance. The 

curator of URM (Recife, Brazil), Dr Leonor M. Maia, is acknowledged 

for making the type species of Johansoniella available for study. 

References 

Arx JA von, Müller E (1975) A re-evaluation of the bitunicate 

ascomycetes with keys to families and genera. Studies in 


Barr ME (1993) Redisposition of some taxa described by J.B. Ellis. 

Mycotaxon 46: 45–76. 

Batista AC, Bezerra JL, Cavalcanti AAAS da Silva (1966) 

Johansoniella, um novo gênero de Schyzothyriaceae. Atas do 

Instituto de Micologia, Universidade Federal de Pernambuco, 

Recife 3: 84–90. 

Crous PW (1998) Mycosphaerella spp. and their anamorphs 

associated with leaf spot diseases of Eucalyptus. Mycologia 

Memoir 21: 1–170. 

Crous PW, Gams W, Stalpers JA, Robert V, Stegehuis G (2004) 

MycoBank: an online initiative to launch mycology into the 21st 

century. Studies in Mycology 50: 19–22. 

Crous PW, Groenewald JZ, Risède J-M, Simoneau P, Hyde KD (2006) 

Calonectria species and their Cylindrocladium anamorphs: 

species with clavate vesicles. Studies in Mycology 55: 213–226. 

Crous PW, Groenewald JZ, Summerell BA, Wingfield BD, Wingfield 

MJ (2009a) Co-occurring species of Teratosphaeria on 

Eucalyptus. Persoonia 22: 38–48. 

v o l u m e 1 · n o . 2 

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Crous PW, Schoch CL, Hyde KD, Wood AR, Gueidan C, Hoog 

GS de, Groenewald JZ (2009b) Phylogenetic lineages in the 

Capnodiales. Studies in Mycology 64: 17–47. 

Crous PW, Verkley GJM, Groenewald JZ, Samson RA (eds) 

(2009c) Fungal Biodiversity. [CBS Laboratory Manual Series 1.] 

Centraalbureau voor Schimmelcultures, Utrecht. 

Crous PW, Wingfield MJ, Park RF (1991) Mycosphaerella nubilosa a 

synonym of M. molleriana. Mycological Research 95: 628–632. 

Frank J, Crous PW, Groenewald JZ, Oertel B, Hyde KD, Phengsintham 

P, Schroers H-J (2010) Microcyclospora and Microcyclosporella: 

novel genera accommodating epiphytic fungi causing sooty 

blotch on apple. Persoonia 24: 93–105. 

Hoog GS de, Gerrits van den Ende AHG (1998) Molecular diagnostics 

of clinical strains of filamentous basidiomycetes. Mycoses 41: 

183–189. 

Lumbsch HT, Huhndorf SM (eds) (2007) Outline of Ascomycota - 

2007. Myconet 13: 1–58. 

Müller E, Arx JA von (1962) Die Gattungen der didymosporen 

Pyrenomyceten. Beitrage zur Kryptogamenflora der Schweiz 11 

(2): 1–992. 

Rayner RW (1970) A Mycological Colour Chart. Commonwealth 

Mycological Institute, Kew. 

Saccardo PA (1889) Johansonia setosa. Sylloge Fungorum (Abellini) 

8: 785. 

Schoch CL, Shoemaker RA, Seifert KA, Hambleton S, Spatafora JW, 

Crous PW (2006) A multigene phylogeny of the Dothideomycetes 

using four nuclear loci. Mycologia 98: 1041–1052. 

Schoch CL, Crous PW, Groenewald JZ, Boehm EWA, Burgess 

TI, et al. (2009). A class-wide phylogenetic assessment of 

Dothideomycetes. Studies in Mycology 64: 1–15. 

Vilgalys R, Hester M (1990) Rapid genetic identification and 

mapping of enzymatically amplified ribosomal DNA from several 

Cryptococcus species. Journal of Bacteriology 172: 4238–4246. 

White TJ, Bruns T, Lee J, Taylor J (1990) Amplification and direct 

sequencing of fungal ribosomal RNA genes for phylogenetics. 

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Protocols: a guide to methods and applications: 315–322. 

Academic Press, San Diego. 

122 

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The history, fungal biodiversity, conservation, and future 

perspectives for mycology in Egypt 

Ahmed M. Abdel-Azeem 

ARTICLE 

Botany Department, Faculty of Science, University of Suez Canal, Ismailia 41522, Egypt; e-mail: zemo3000@yahoo.com 

Abstract: Records of Egyptian fungi, including lichenized fungi, are scattered through a wide array 

of journals, books, and dissertations, but preliminary annotated checklists and compilations are not 

all readily available. This review documents the known available sources and compiles data for more 

than 197 years of Egyptian mycology. Species richness is analysed numerically with respect to the 

systematic position and ecology. Values of relative species richness of different systematic and 

ecological groups in Egypt compared to values of the same groups worldwide, show that our knowledge 

of Egyptian fungi is fragmentary, especially for certain systematic and ecological groups such as 

Agaricales, Glomeromycota, and lichenized, nematode-trapping, entomopathogenic, marine, aquatic 

and coprophilous fungi, and also yeasts. Certain groups have never been studied in Egypt, such as 

Trichomycetes and black yeasts. By screening available sources of information, it was possible to 

delineate 2281 taxa belonging to 755 genera of fungi, including 57 myxomycete species as known from 

Egypt. Only 105 taxa new to science have been described from Egypt, one belonging to Chytridiomycota, 

47 to Ascomycota, 55 to anamorphic fungi and one to Basidiomycota. 

Key words: 

checklist 

distribution 

fungal diversity 

lichens 

mycobiota 

species numbers 

Article info: Submitted: 10 August 2010; Accepted: 30 October 2010; Published: 10 November 2010. 

INTRODUCTION 

Biological diversity (biodiversity) encompasses the variety 

of life forms occurring in nature, from the ecosystem to the 

genetic level, as a result of evolutionary history (Wilson 

1992). The idea that fungi form a kingdom distinct from plants 

and animals gradually became accepted only after Whittaker 

(1969). Presently, the “fungi” as a mega-diverse group span 

three kingdoms, most belonging to the Fungi (Eumycota), 

while others are classified in the Protozoa and Chromista 

(Straminipila) (Cavalier-Smith 1998, James et al. 2006b). 

The word “fungi”, lower case and not in italics, is commonly 

used as a collective term for organisms traditionally studied 

by mycologists from all three kingdoms (Hawksworth 1991). 

The myxomycetes have also been traditionally studied by 

mycologists (Everhart & Keller 2008, Rojas & Stephenson 

2008), and are included here. 

Estimates for the number of fungi in the world range up to 

ca. 13.5 M species (McNeely et al. 1990, Hawksworth 1991, 

2001, Hawksworth & Kalin-Arroyo 1995, Hyde 1996, Hyde et 

al. 1997, Tangley 1997, Groombridge & Jenkins 2002, Brusca 

& Brusca 2003, Rossman 2003, Crous et al. 2006, Adl et al. 

2007, Kirk et al. 2008). It might be expected that the predicted 

numbers of fungi on Earth would have been considerably 

greater than the 1.5 M suggested by Hawksworth (1991), 

which is currently accepted as a working figure although 

recognized as conservative (Hawksworth 2001). 

The 10 th edition of Ainsworth & Bisby’s Dictionary of the 

Fungi (Kirk et al. 2008) provided a total of 98 998 for the 

number of fungal species accepted to date (excluding taxa 

treated under Chromista and Protozoa). Kirk et al. (2008) 

reported 1 039 species chromistan fungal analogues and 

1 165 as protozoan in which 1 038 are regarded as protozoan 

fungal analogues: Percolozoa (Acrasida), Amoebozoa 

(Dictyostelia, Myxogastria, Protostelia), Cercozoa 

(Plasmodiophorida) which were previously treated as 

Myxomycota and Plasmodiophoromycota. 

Egypt’s geographical position at the junction between two 

large continents (Africa and Asia), and its inclusion as part 

of the Mediterranean basin, has indelibly influenced both the 

people and the biota of the country socially, economically 

and biologically. Egypt is part of the Sahara of North Africa, 

it has an area of about 1 M km 2 , divided by the River Nile 

into a western part including the Libyan Desert (681 000 

km 2 ) and an eastern part comprising the Eastern Desert (223 

000 km 2 ), and the Sinai Peninsula (61 000 km 2 ). The Nile 

basin, comprising the valley in the south (Upper Egypt) and 

Nile delta in the north (Lower Egypt), forms a riparian oasis 

(40 000 km 2 ) that constitutes the densely inhabited farmlands 

of Egypt. 



Attribution: 






v o l u m e 1 · n o . 2 

123


ARTICLE 

Kassas (2002) mentioned four gaps related to biodiversity 

knowledge: the number of species on Earth; the diversity 

of the less conspicuous organisms such as fungi, bacteria, 

algae, and protozoa; the role played by each species among 

biotic elements of ecosystems; and the human ability to 

assess and forecast bio-ecological degradation. 

Documentation of the Egyptian fungi may be dated 

back to 4500 B.C., when ancient Egyptians produced a 

number of hieroglyphic depictions of plants (many of which 

are psychedelic) on walls and within texts throughout 

Egypt. Temples with countless pillars are shaped like huge 

mushrooms with tall stems, umbrella caps, and mushroom 

engravings distributed all over the country (Fig. 1). These 

are shaped like Amanita sporophores, and some like 

Psilocybe. Others look like bracket fungi and are decorated 

with pictures of an incredible variety of plants (Arthur 2000). 

In the Egyptian Book of the Dead, the Papyrus of Ani 

(Budge 1967), mushrooms are called “the food of the gods,” 

or “celestial food” and “the flesh of the gods.” 

Studies on fungi in Egypt started at the beginning of the 

19 th century on lichens (e.g. Delile 1813a, b, Nylander 1864, 

1876, Müller 1880a–c, 1884, Stizenberger 1890, 1891). In 

the early 20 th century, Sickenberger (1901) and Steiner 

(1893, 1916) provided information for collections of lichens 

from Egypt in the 19 th and early 20 th Century. In the Flore 

d’Egypte, Delile (1813a) presented a scientific study of 

Egyptian fungi into the early19 th century (Mouchacca 2008), in 

which he described the gastromycete now known as Itajahya 

rosea (syn. Phallus roseus; Fig. 1) which he had collected in 

Damietta and Assiut in 1798 and 1799, respectively. It should 

be noted that some early works repeat previous records, 

sometimes ambiguously as a result of the misinterpretation 

of synonyms and erratic use of infraspecific ranks; further, 

in the case of Sickenberger, misspellings of scientific names 

(Seaward & Sipman 2006). 

By the beginning of the 20 th century, special attention 

was being given to phytopathogenic fungi on wild and 

domesticated plants of economic importance (e.g. Fletcher 

1902, Reichert 1921, Fahmy 1923, Shearer 1924, Briton- 

Jones 1922, 1923, 1925, Bishara 1928, Melchers 1931, Sirag 

El-Din 1931, Abdel-Salam 1933). 

Fig. 1. A. Giant mushroom-like pillars (upper part), which are 

common in Egyptian temples. B. Description of Phallus roseus by 

Delile (1813a). 

Both Reichert and Melchers are considered the pioneer 

scientists in the documentation of Egyptian fungi. Israel 

Reichert (1891–1975) went to study in Germany. Here he 

obtained his doctorate on Die Pilzflora Ägypten in which 237 

species were recognized, of which 42 were new to science. 

Unfortunately, none of his specimens were retained in 

Egypt, or if they were, there is no record of their whereabouts 

today. However, earlier material collected before 1914 was 

present in the Botanisches Museum in Berlin-Dahlem, 

which Reichert used when compiling his list of 1921, but it is 

not known if these specimens survived World War II. 

In 1927 Leo E. Melchers went to Egypt at the invitation of 

the Egyptian Minister of Agriculture as chief mycologist for 18 

months. He met a series of difficulties such as there being no 

records available on the occurrence, distribution, or dates of 

any mycological observations conducted previously by any 

investigator in Egypt, and no mycological reference collection 

existing in the country. His checklist, however, included 345 

species of fungi, especially those causing plant diseases 

(Melchers 1931). 

No studies were carried out on the soil fungi until the 

1930s, yet it was to be expected that, in such a country with 

rich agricultural traditions, knowledge of these fungi should 

have attracted considerable interest. Research on Egyptian 

soil fungi was probably commenced by Younis Salem Sabet 

(1898–1977). Sabet graduated in 1921 from the High School of 

Agriculture (now the Faculty of Agriculture of Cairo University), 

and soon after was sent to England to study botany at the 

University of London, where he obtained a BSc (Hons) in 1925. 

After his return, he joined the Ministry of Agriculture in the Plant 

Breeding Section. In 1927 he was appointed lecturer in Botany 

in the faculty of Science of the newly established Egyptian 

University, and in 1935 published his pioneering study, which 

was followed by many other publications (Sabet 1936, 1938, 

1939a). His exploration led to the discovery of three taxa which 

were described later as new to science. 

Sabet took the initiative in the establishment of some 

scientific organisations, and served as a member and 

president for several years in some others. Particularly of note 

were the Egyptian Academy of Sciences, Egyptian Botanical 

Society, Egyptian Science Union, Egyptian Association for 

Scientific Culture, Society of Applied Microbiology, Egyptian 

Phytopathological Society, Society for the History of Science, 

and Society of Atomic Energy. 

Near the end of the 1930s, new aspects of mycological 

research were introduced into Egypt by several investigators 

such as mycorrhizal fungi (Mostafa 1938, Sabet 1939b, 

1940, 1945, Yousef 1946); biocontrol (Mostafa & Gayed 

1953), rhizosphere (Montasir et al. 1956, Naim et al. 1957), 

air (Saad 1958, Zaki 1960), and stored seeds and grains 

(Assawah & El-Arosi 1960). 

In 1956 late Magdy A. Ragab (Department of Botany, 

Faculty of Agriculture, University of Cairo) isolated 16 new 

species for the first time from soil, water and some plant 

hosts (Ragab 1956). 

However, the credit for initiating real research concerned 

with Egyptian fungi must be given to Abdel-Al H. Moubasher 

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Mycology in Egypt 

ARTICLE 

Fig. 2. A selection of prominent mycologists who have contributed greatly to our knowledge of mycology in Egypt. A. Abdel-Al H. Moubasher. B. 

Samy M. El-Abyad. C. Jean Mouchacca. D. Abdul-Wahid F. Moustafa. E. Farida T. El-Hissy. F. Younis S. Sabet. G. Israel Reichert. H. Youssef 

A. Youssef. 

(Botany Department, Faculty of Science, Assiut University; 

Fig. 2). In the early 1960s, with colleagues and students, he 

broadened the scope of mycological research in Egypt by 

conducting many studies on fungi. These included aspects 

such as: cellulose-decomposition, thermophily, osmophily, 

seed and grain mycobiota, phylloplane fungi, mycotoxins, and 

aquatic fungi. Moubasher, with his colleagues and students, 

have published more than 150 scientific papers to date, and 

in 1993 he published his major contribution to mycology in 

the Arabic World, the lavishly illustrated Soil fungi of Qatar 

and other Arab Countries (Moubasher 1993). He also invited 

outside specialists to run courses from the 1980s and trained 

many PhDs students. Specialists included Colin Booth and 

David Hawksworth in the 1980s. 

El-Abyad & Abu-Taleb (1993) summarized the habitat 

diversity of Egyptian fungi, and in 1997 the late Samy M. 

El-Abyad (Botany Department, Faculty of Science, Cairo 

University; Fig. 2) presented his pioneering attempt to update 

the checklist of Egyptian fungi: 1 246 species were recorded 

of which 173 were referred to Mastigomycotina, 41 to 

Zygomycotina, 222 to Ascomycotina, 143 to Basidiomycotina, 

and 667 to Deuteromycotina. Different ecological and 

taxonomic groups were not separated cited in the checklist, 

such as protozoan fungal analogues (Myxomycota, 

Plasmodiophoromycota), lichens, yeasts, aquatic and marine 

fungi, entomopathogenic fungi, nematophagous fungi, and 

mycorrhizal fungi. A large numbers of taxa, either reported 

in routine isolations or as novel taxa, are completely absent 

from this list. This may be due to his inability to trace the 

majority of references, which is actually the main reason 

why updated information documenting the fungi of Egypt 

was needed today. Amongst records lacking in the El-Abyad 

(1997) checklist are seven Podaxis species (Melchers 1931), 

Chaetomium gelasinosporum and C. uniporum (Aue & Müller 

1967), C. mareoticum (Besada & Yusef 1969), Zygopleurage 

faiyumensis (Lundqvist 1969), Podospora aegyptiaca 

(Lundqvist 1970), Thermoascus aegyptiacus (Udagawa & 

Ueda 1983), and Gelasinospora hippopotama (Krug et al. 

1994). 

In addition to the previous efforts of Reichert (1921), 

Melchers (1931), El-Abyad & Abu-Taleb (1993), and El- 

Abyad (1997), several other studies have added to the 

documentation of Egyptian fungi: Moubasher (1993), Lado 

(1994), Mouchacca (1995, 1999, 2001a, b, 2003a, b, 2004, 

2005, 2008, 2009a, b; Fig. 2), Moustafa & Abdel-Azeem 

(2005a, b, 2006, 2010), Moustafa (2006), and Seaward & 

Sipman (2006). 

The late Abdel Razak Abo-Sedah organized the 

Second African Regional Mycological Congress, in Cairo 

in 1992, under the auspices of the IMA Committee for the 

v o l u m e 1 · n o . 2 

125


ARTICLE 

Development of Mycology in Africa. Then in 1993 he founded 

the Regional Center for Mycology and Biotechnology (RCMB) 

in Al-Azhar University, Cairo. The major tasks of this centre 

were the establishment of a fungal culture collection, the 

application of fungi in public health, agriculture, environment 

and industry, and supporting researchers as well as research 

projects. The centre actively participated in organizing further 

African regional and international conferences and meetings 

in Cairo in 1994, Vancouver in 1994, Zimbabwe in 1995, 

Cairo in 1996 on “Regulations of fungal activities”, and again 

in Cairo in 1999 on “Fungi and the Environment”. The center 

had collaborative agreements with the former International 

Mycological Institute (IMI) in the UK, and collaborative 

activities with Egyptian universities as well as with others 

in the UK, South Africa, Mauritius, Zimbabwe and Austria. 

The centre also initiated and published The African Journal 

of Mycology and Biotechnology from 1993 to 2001, which 

contained numerous contributions by Egyptian authors, and 

also a mycological newsletter in Arabic. 

From the beginning of 2005 to the end of 2007, the 

Biodiversity Monitoring and Assessment Project (BioMap) had 

as its primary objective to develop and strengthen biodiversity 

research, monitoring and assessment across Egypt. In this 

project an extensive e-database was established to map the 

distribution of species across Egypt, and document up to 

50 % of the Egyptian fungi (). 

As mentioned above, the information concerning the fungi 

of Egypt is still incomplete and cannot be fully documented 

without an updated checklist of all taxa reported for the 

country. The present contribution assesses the diversity of 

fungi in Egypt. In addition, major groups of fungi are discussed 

briefly to highlight the extent of their diversity, followed by 

examples of habitats that are unique and deserve greater 

attention. These data show that the present contribution is 

a preliminary one concerning the diversity of Egyptian fungi, 

and therefore this summation is intended to enhance our 

knowledge of, and stimulate research into, the fungi of Egypt. 

MATERIALS AND METHODS 

The present contribution is based on an exhaustive revision of 

the available literature and sources of the Egyptian fungi reported 

from the 19 th century to the present, including dissertations, 

published papers, compilations and checklists. Name corrections, 

authorities, and taxonomic assignments of all taxa reported in 

this article were checked against the Index Fungorum database 

(). In addition, websites of international 

mycological centres such as the ATCC (USA) (), 

CABI (UK) (), CBS (The 

Netherlands) (), MUCL (Belgium) () and the catalogue of the culture 

collection of the Assiut University Mycological Center (AUMC 

2010) were also consulted. The systematic arrangement in the 

present article follows Kirk et al. (2008). 

This study extended to more than eight years in 

documenting and updating the information on Egyptian 

fungi. All results of the present study can be checked against 

the last updated checklist (El-Abyad 1997). 

RESULTS 

General features of Egyptian fungi 

The number of fungi recorded in Egypt is 2 281 species, 

out of which 105 taxa have been described from Egypt as 

new to science: one in Chytridiomycota, 47 in Ascomycota, 

56 in anamorphic fungi, and one in Basidiomycota. Reichert 

introduced 24 of the new taxa, representing 24.7 % of the 

novel taxa, followed by Jean Mouchacca and his colleagues 

(Laboratoire de Cryptogamie, Muséum National d’Histoire 

Naturelle, Paris), who described 18 new species (17.1 % 

of the total), and Abdul-Wahid F. Moustafa (Fig. 2) and his 

colleagues and students at the Suez Canal University who 

contributed 11 new taxa. 

Protozoan fungal analogues 

The kingdom Protozoa contains 115 000 known species. 

They are extremely diverse in their cell structure, patterns 

of nutrition, metabolic needs, reproduction, and habitat. This 

kingdom contains a grab-bag of organisms that do not fit 

into the other kingdoms. Protozoa are extremely difficult to 

classify so for the purpose of this survey, they are grouped 

by their nutritional patterns. Protozoan fungal analogues 

are heterotrophic and most are decomposers that feed on 

dead plants and animals by endocytosis (Kendrick 2000). 

According to Kirk et al. (2008) there are about 1 165 fungal 

protozoan analogues described. Slime moulds are a small 

and relatively homogenous group of eukaryotic organisms, 

and these are referred to as Myxomycota (Mycetozoa). In 

Egypt the slime moulds have never been the target of any 

widescale study (Lado 1994, Stephenson & Stempen 1994), 

except for the pioneer study of Abdel-Raheem (2002) on 

those of Upper Egypt (Ndiritu et al. 2009). 

Abdel-Raheem (2002) reported 20 species belonging 

to 17 genera in his first inventory of the protozoan fungal 

analogues (Myxomycota) of Upper Egypt from wood, 

bark of living and dead trees and leaf litter. Exhaustive 

examination of all available literature concerning protozoan 

fungal analogues in Egypt led to the discovery of reports of 

Protostelium irregulare (as “irregularis”; Olive & Stoianovitch 

1969) and Eidamella spinosa (Kowalik & Sadurska 1973). 

The protozoan fungal analogues occurring on decaying 

wood, bark, leaf litter and papyrus papers presently amount 

to 57 species belonging to 25 genera. For more details 

refer to the PBI: Global Biodiversity of Eumycetozoans 

() and 

Farghaly (2008). In addition, three species representing three 

genera of Cercozoa (previously Plasmodiophoromycota) 

have been recorded: Plasmodiophora, Spongospora, and 

Woronina. No dictyostelid cellular slime moulds are so far 

known from Egypt (Cavender et al. 2010). 

126 

 

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Chromistan fungal analogues 

The kingdom Chromista (Straminipila) is a collection of 

eukaryotic, walled microorganisms that produce heterokont, 

wall-less cells in their life-cycles, including some fungallike 

groups that are not considered to be ancestors of 

any members of the Fungi (Lutzoni et al. 2004). Kirk et al. 

(2008) estimated the Chromistan fungal analogues as 1 039 

known species and included the phyla Hyphochytriomycota, 

Labyrinthista, and Oomycota along with some taxa of 

uncertain position (incertae sedis). 

The late Farida T. El-Hissy (Botany Department, Faculty of 

Science, Assiut University; Fig. 2) was the founder of aquatic 

mycology research in Egypt. El-Hissy and her students 

published more than 60 papers on this topic. However, the 

plant pathogenic Oomycota have been the target of many 

research investigations since 1921, and in the present study, 

186 taxa of chromistan fungal analogues were recorded, of 

which 172 belong to 40 genera of Oomycota. Four species 

and two genera of Labyrinthista were recorded, while 

Hyphochytriomycota are represented by six species within 

three genera. For more details refer to El-Helaly et al. (1963, 

1966), Ali Hassanein et al. (1972), Khallil et al. (1995), and 

El-Hissy et al. (1990, 1992, 1997, 2004) 

Fungi (Eumycota) 

Blastocladiomycota 

This phylum was once considered part of the chytrids. 

However, most of the true chytrids (Chytridiomycota) 

produce a limited mycelium while Blastocladiomycota 

usually make extensive mycelia. Thus, they superficially 

resemble the water moulds to which they were thought to 

have been affiliated. Like the chytrids, Blastocladiomycota 

and Neocallimastigomycota are the only members of the 

fungi in which motility has been retained. In overall growth 

habit, the blastocladiomycetes tend to be eucarpic, in which 

there is an extensive vegetative growth habit in which some 

part of the organism participates in reproduction (asexual 

and sexual). Members of this phylum do exhibit a complete 

alternation of generation between a haploid gametophyte 

and a diploid sporophyte (Barr 1990, James et al. 2006a). 

Kirk et al. (2008) give a world total for Blastocladiomycota of 

179 species. In Egypt, 27 species and one variety belonging 

to seven genera of Blastocladiomycota were found in this 

study. For more details see Ragab (1956), Yusef (1964), Gad 

et al. (1967), Gad & Sadek (1968), El-Hissy (1974), El-Hissy 

et al. (1997), El-Abyad (1997), Shoulkamy et al. (2001), and 

Abdel-Moneim (2010). 

Chytridiomycota 

Chytridiomycota are a phylum of fungi that reproduce 

through the production of motile spores (zoospores), typically 

propelled by a single, posteriorly directed flagellum. These 

organisms, often referred to as chytrid fungi or chytrids, 

have a global total of approximately 1 000 described species 

(James et al. 2006a). Based on biochemical characteristics, 

including chitin in cell walls, the α-aminoadipic acid lysine 

synthetic pathway, and storage carbohydrates (i.e. glycogen), 

Bartnicki-Garcia (1970) classified Chytridiomycota as true 

fungi and this is supported by current molecular studies 

(Hibbett et al. 2007). In the past some authors considered the 

chytrids as a transitional group between protists and fungi 

because of their production of motile zoospores (Barr 1990). 

Kirk et al. (2008) give the number known Chytridiomycota as 

706. 

The study of Gaertner (1954) on Chytridiomycota of 

Africa is considered one of the pioneer mycological studies 

in Egypt. However, the real start of research on chytrids in 

Egypt must be credited to Samy Kamel Mohamed Hassan 

(Minia University) who obtained his PhD from the University 

of Warsaw for work on chytrids and aquatic fungi in 1982. 

Later, Hassan and Mohamed Abdel-Wahab El-Naghy (Minia 

University) made a series of studies on chytrids in Egypt. 

Intensive revision of the nomenclature showed that 84 

species belonging to 32 genera of Chytridiomycota were 

recorded in Egypt. For more details see El-Naghy et al. (1985, 

1987), Hassan (1991a-d, 1993), Hassan & Fadl-Allah (1991), 

Hassan & Shoulkamy (1991), and Hassan & Shaban (1991). 

Zygomycota 

Zygomycota are a particularly ecologically diverse group 

of fungi, occurring as saprobes (Mucorales), harmless 

inhabitants of arthropod guts (Harpellales), plant mutualists 

forming ectomycorrhizas (Endogonales), and pathogens of 

animals, plants, amoebae, and especially other fungi (all 

Dimargaritales and some Zoopagales are mycoparasites) 

(James & O’Donnell 2007). Conversely, some Mucorales 

have a negative economic impact as they cause storage rots 

or plant diseases, while others can cause life-threatening 

opportunistic infections in diabetic, immuno-suppressed, and 

immuno-compromised patients. In addition, several species 

of Microsporidia cause serious human infections (de Hoog et 

al. 2000, James & O’Donnell 2007). 

According to Kirk et al. (2008) the total world number of 

Zygomycota is 1 065 species. Data collected from previous 

studies show the Zygomycota in Egypt to be fragmentary 

because members belonging to this group either have long 

been overlooked or simply reported as rare taxa during 

routine isolations. 

Abdel-Kader (1973) carried out a pioneering study in 

which he was able to isolate 11 species from a range of soils 

collected from various Egyptian localities. The second most 

relevant study is probably that of Al-Alfy (1995) who reported 

21 species from various substrates, including soil, dung, 

stored seeds and grains, and the phyllosphere. In his recent 

contribution on Zygomycetes in Egypt, Moustafa (2006) 

reported 33 species, out of which nine were considered new 

Egyptian records. 

Revision of all available data showed that Zygomycota in 

Egypt comprises 70 taxa including eight varieties and seven 

special forms within 35 genera. In addition, Absidia aegyptiaca 

(Sartory et al. 1939) is omitted from the list, as no living or 

other type of authentic material is apparently preserved; 

furthermore the name was not validly published as it lacked a 

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Latin diagnosis (Mouchacca 1995). For more information on 

Egyptian Zygomycota refer to Kharboush (1969a, b), Besada & 

Yusef (1968), Abdel-Rahman et al. (1990), Moubasher (1993), 

El-Abyad & Abu-Taleb (1993), Swelim et al. (1994), Mouchacca 

(1995), El-Abyad (1997), Abdel-Azeem (2003), Moustafa 

(2006), Ali & Ibrahim (2008), Afify et al. (2009), and Moubasher 

et al. (2010). 

Glomeromycota 

The Glomeromycota currently comprises 169 described 

species (Kirk et al. 2008). The phylum is not as diverse as 

other phyla of fungi with only three families and such a modest 

number of species. However, they make up for this uniformity 

by being among the most abundant and widespread of all 

fungi. As far as we know, all species of Glomeromycota are 

mutualistic with plants, forming endomycorrhizas. Although 

there are various types of mycorrhizas, involving different 

fungal and plant symbionts, the arbuscular mycorrhiza type 

is the most widespread occurring in around 80 % of plant 

species (Redecker & Raab 2006). 

The pioneering work of Mostafa (1938) and Sabet (1939b, 

1940, 1945; Fig. 2) is now accepted as the starting point of 

research on Egyptian Glomeromycota (Kelley 1950, Abdel- 

Moneim & Abdel-Azeem 2009). These studies were followed by 

many other investigations concerned mainly with the ecology 

and physiology of endomycorrhizas in Egypt, viz. Fares 

(1986), Ishac et al. (1986), Abdel-Fattah (1991), Aboulkhair & 

El-Sokkary (1994), Mankarios & Abdel-Fattah (1994), Abdel- 

Fattah & Mankarios (1995), Abdel-Fattah & Rabie (1995), 

Abdel-Fattah et al. (1996), Abdalla & Abdel-Fattah (2000), 

Abdel-Fattah (2001), and Abdel-Azeem et al. (2007). 

However, surveys of Egyptian Glomeromycota are limited, 

and had never been the sole target of any study until Fares 

(1986) conducted a survey of vesicular arbuscular mycorrhizas, 

followed by Agwa (1990) on mycorrhizas and nodulation in 

some Egyptian plants. After 10 years, Agwa (2000) studied the 

arbuscular mycorrhizal fungi associated with medicinal plants as 

Glomales in Egypt (I)”. Agwa & Abdel-Fattah (2002) followed up 

their work “Glomales in Egypt (II)” as an ecological view of some 

saline affected plants in the delta of the Mediterranean coast. A 

study of the distribution of Glomales in the Egyptian Protectorates 

was published by Agwa & Al-Sodany (2003) as “Glomales in 

Egypt (III)”, which surveyed the distribution and ecology in some 

plants in the El-Omayed Biosphere Reserve. Later, other relevant 

studies were carried out by several investigators such as El- 

Zayat et al. (2007) and Abdel-Moneim & Abdel-Azeem (2009) on 

the Wadi Allaqi and Saint Katherine Protectorate, respectively. 

Recently, Mansour (2010) screened 71 soil and root samples for 

endomycorrhizas in North Sinai and adopted some of them as 

biocontrol agents against fusarium-wilt of tomato. 

Eight genera and 19 species have been recorded in 

Egypt since 1938: Acaulospora, Entrophospora, Gigaspora, 

Glomus, Paraglomus, Sclerocysti, Scutellospora and 

Rhizophagus. Both Paraglomus occultum and Rhizophagus 

were recorded and never cited in any publication related 

to Egyptian Glomeromycota. For more details see Sabet 

(1939b) and Morton & Redecker (2001). 

Lichen-forming fungi 

Lichens are unique associations composed of two to three 

different organisms living together in a mutualistic relationship 

in which the fungal partner forms the external structure. The 

name used is that of the fungal parter, and the photosynthetic 

partner or partners have independent scientific names. 

Estimates for the number of lichen fungi worldwide vary, but 

a draft global checklist has 18 882 names of lichen-forming 

and allied fungi (Feuerer & Hawksworth 2007). 

Egyptian lichens have received the attention of many 

researchers since the early 1800s (Delile 1813a, b, Nylander 

1864, 1876, Müller 1880a–c, 1884, Stizenberger 1890, 

1891, Sickenberger 1901, Steiner 1893, 1916, Werner 1966, 

Galun & Garty 1972, Temina et al. 2004, 2005, Seaward & 

Sipman 2006). Egyptian investigators have participated in a 

few studies of lichens, namely in North Sinai (Khalil 1995) 

and on trees (Koriem 2003), and there have also been some 

physiological studies on the bionts (Koriem 2006). Khalil 

(1995) recorded 43 species belonging to 18 genera, all of 

which are ascolichens without any basidiolichens at all, and 

only one of these had a perithecioid ascoma (Gonohymenia 

sinaica). 

Seaward & Sipman (2006) reported 157 taxa of 

lichenized fungi (149 species and 8 infraspecific taxa) and 

six lichenicolous fungi (fungi obligately growing on lichens). 

Foliose lichens are very scarce, only being represented by the 

genera Xanthoria (7 species) and Physcia (1 species). The 

fruticose growth form is better represented, with members of 

the genera Ramalina, Roccella, Seirophora and Tornabea. 

At the family level, Teloschistaceae accommodated the most 

taxa (39), followed by Roccellaceae (16), and Physciaceae 

(12). For more information concerning Egyptian lichens 

please see: check-lists of Lichens and Lichenicolous 

Fungi (), the Tel Aviv University 

Herbarium (TELA) (), 

Galun & Garty (1972), Khalil (1995), Koriem (2003, 2006), 

Temina et al. (2004, 2005), and Seaward & Sipman 2006). 

Ascomycota (non-lichenized) 

Numerically Ascomycota constitute by far the largest group 

of fungi so far known, accommodating a relatively large 

assemblage of taxa estimated to be 65 % of all described 

fungi (Kirk et al. 2008) occurring in various habitats; aquatic 

or terrestrial, under moderate or stress conditions (Kodsueb 

et al. 2008a, b, Kruys & Ericson 2008, Thongkantha et 

al. 2008). A large number of Ascomycota species are 

economically important (e.g. Fusarium spp., Kvas et al. 

2009; Colletotrichum spp., Damm et al. 2009, Hyde et al. 

2009; Mycosphaerella spp., Crous 2009), while few are 

edible (morels and truffles), and some are used also in 

the production of food (including bread), drinks, organic 

acids, mycofungicides, fungal biofertilizers, cosmetics and 

hormones (Kaewchai et al. 2009, Hyde et al. 2010). 

This phylum encompasses biologically diverse forms. 

Many are free-living saprobes including species which may 

be cellulose decomposers, chitinolytic, keratinolytic, or 

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coprophilous, others are parasitic forms including species 

which cause very serious plant diseases like powdery-mildew, 

wood-canker, ergot, rot, blight, scab, leaf curl, and leaf-spots 

(e.g. Alves et al. 2008, Aveskamp et al. 2008, Simonis et al. 

2008, Wulandari et al. 2009). Others that are considered 

symbiotic forms contain species which live in association with 

insects or algae (lichens) or roots of plants (mycorrhizas). 

Ascomycota characteristically, when reproducing sexually, 

produce non-motile spores (ascospores) in a distinctive 

“ascus”. However, some members of the Ascomycota do 

not reproduce sexually and do not form asci or ascospores 

(anamorphic Ascomycota). These asexual members are 

assigned to Ascomycota based upon morphological and/ 

or physiological similarities to ascus-bearing taxa, and in 

particular by phylogenetic comparisons of DNA sequences. 

In old classification systems these were often placed in 

a separate artificial phylum, the deuteromycota (or “fungi 

imperfecti”). Molecular analyses can now place these genera 

and species among ascus-bearing taxa, or more rarely in 

other phyla such as Basidiomycota. 

The first reports of mutualistic non-lichenized ascusforming 

fungi from Egypt were those of Terfezia, Tirmania, 

and Morchella by Reichert (1921), and then Melchers (1931) 

who recorded six species. Later, Sabet (1935) recorded some 

saprobic Chaetomium species. The saprobic Ascomycota 

did not receive attention, and therefore information remained 

limited until the early 1970s, when some research on the group 

was initiated by Moubasher and his co-workers during their 

studies on soil fungi. Since then, fragmentary information has 

been accumulating, but these fungi had never been the main 

objective of any Egyptian study focusing on their ecology, 

distribution, and substrate preferences, untill the study of 

Abdel-Azeem (2003). 

Three hundred and three species of teleomorphic 

saprobic Ascomycota (including ascosporic yeasts) have 

been recorded from all terricolous substrates of Egypt 

(Moustafa & Abdel-Azeem 2010, and unpubl.). In their 

studies, 10 species of edible Ascomycota were recorded from 

Egypt within the genera Morchella, Terfezia, and Tirmania. In 

total, 328 taxa were recorded in this survey, of which only 

32 species are ascosporic yeasts. Binyamini (1973) reported 

Peziza vesiculosa as a coprophilous fungus from occupied 

Palestine, and some samples were even collected from north 

Sinai during the occupation in 1967, but never cited as an 

Egyptian record in any checklist. In addition, Byssonectria 

tetraspora was recorded for the first time in Egypt by El- 

Saadawi & Shabbara (1999) as an association between a 

fungus and a moss. 

For more details see Sabet (1936, 1939a), Binyamini 

(1973), El-Saadawi & Shabbara (1999), Abdel-Hafez et al. 

(1995), Ibrahim (1995), El-Abyad (1997), Zaki et al. (2005), 

Moustafa & Abdel-Azeem (2005a, b, 2006, 2008, 2010), and 


Records of phytopathogenic fungi in Egypt were scattered 

through the literature until 1921, when Israel Reichert (Fig. 

2) carried out his pioneer study of Egyptian fungi. This was 

followed by a comprehensive checklist of plant diseases 

and fungi occurring in Egypt by Melchers (1931). Records 

concerning aspects of plant pathology in Egypt continued to 

be accumulated during many decades until El-Helaly et al. 

(1963, 1966) started to update the information, and another 

updated bibliography of agricultural studies conducted 

in Egypt between the period 1900 to 1970 appeared (Ali 

Hassanein et al. 1972). This revealed records of 82 species 

of teleomorphic plant pathogenic Ascomycota. For more 

details please check, Natrass (1933), Abou El-Seood (1968), 

Ghoniem (1985), El-Desouky & El-Wakil (2003), Phillips et al. 

(2006), and Hafez (2008). 

Anamorphic genera are gradually disappearing into the 

overall ascomycete system, though it will take many years 

before anamorph genera have fully integrated. 

A school of medical mycology in Egypt was formed at the 

beginning of 1967, when the late Youssef A. Youssef (Ain 

Shams University, Faculty of Science; Fig. 2) published two 

papers on fungus infection of the human ear. Youssef and 

his students and colleagues became interested in medical 

mycology, serology and fungi affecting human health. For 

more details see Youssef & Abdou (1967a, b), Hassan et al. 

(1980a–e, 1981), Youssef & Karam El-Din (1988a, b), Karam 

El-Din et al. (1994 a–c, 1995, 1996), and Youssef et al. (1989, 

1992, 1993). 

In 1979 Ismail Abdel-Razak M. El-Kady received credit 

as the Egyptian mycologist working on mycotoxin producing 

fungi in Egypt. El-Kady and his coworkers studied the majority 

of aspects related to toxinogenic fungi, e.g. factors affecting 

mycotoxin production, toxinogenic taxa in food and feed, and 

mutagenic effects of fungal toxins. For more details see El- 

Kady & Moubasher (1982 a, b), and El-Kady et al. (1989, 

1994). 

In 1987, Mamdouh S. Haridy (Minia University, Faculty of 

Science) became the pioneer Egyptian mycologist in yeast 

identification and taxonomy, having completed his PhD thesis 

on the taxonomy of yeasts (“Taxonomie milchwirtschaftlich 

wichtiger Hefen”, Technical University, Munich). He conducted 

a series of extensive studies on the Egyptian saprobic yeasts 

from different ecological habitats and sources (Haridy 1992a, 

b, 1993a, b, 1994a, b, 2002). 

Recently, other areas of Egyptian mycology have 

been established, such as on the identification of human 

and plant pathogens by molecular techniques. Youssuf A. 

Gherbawy (Botany Department, Faculty of Science, South 

Valley University) focused on the identification of plant 

pathogens and saprobic fungi of food by means of molecular 

techniques (Gherbawy 2004, Gherbawy & Abdelzaher 2002, 

Gherbawy & Farghaly 2002, Gherbawy & Voigt 2010), 

and Sherif M. Zaki (Microbiology Department, Faculty of 

Science, Ain Shams University) extended the research 

of Youssef A. Youssef using the molecular techniques in 

species identification of human pathogens (Zaki et al. 2005, 

2009, Zaki 2008). 

About 905 filamentous or yeast-like anamorphic fungi 

have been reported from Egypt. These taxa colonize, survive 

and multiply in air, litter, soil, plant surfaces, the human 

body and other substrates. Of these, only 28 are species 

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of anamorphic ascomycetous yeasts, which belong to three 

genera. Furthermore, five genera of basidiomycetous yeasts 

and 18 species are recorded from all habitats in Egypt. 

For more information consult Al-Doory (1968), Abdel- 

Fattah (1985), Sherief (1985), Bagy & Abdel-Hafez (1985), 

Khater (1989), Shalouf (1989), Shindia (1990), Abdel-Mallek 

et al. (1995), Abdul Wahid et al. (1996), Hamdi & Hassanein 

(1996), El-Tanash (1997), Shalaby (1999), Mahmoud (1999), 

Ismail & Sabreen (2001), Teramoto et al. (2001), Abdel- 

Wahab (2002), Farghaly et al. (2004), Nofal & Haggag (2006), 

Haggag et al. (2007), Abdel-Hamed (2008), and Kottb (2008). 

Aquatic and marine fungi 

Marine fungi form an ecological, and not a taxonomic group 

(Raghukumar 2008, Jones et al. 2009, Hyde et al. 2000). 

Among these, the obligate marine fungi grow and sporulate 

exclusively in seawater, and their spores are capable of 

germinating in seawater (Hyde et al. 1998). On the other hand, 

facultative marine fungi are those obtained from freshwater 

or a terrestrial milieu, and have undergone physiological 

adaptations that allow them to grow and possibly also sporulate 

in the marine environment (Kohlmeyer & Kohlmeyer 1979). 

These fungi belong mostly to ascomycetes, their anamorphs, 

and a few basidiomycetes. Among the straminipilan fungi, 

those belonging to Labyrinthulomycetes, comprising the 

thraustochytrids, aplanochytrids, and labyrinthulids are 

obligate marine fungi (Raghukumar 2002), and those 

belonging to the oomycetes are also fairly widespread in the 

marine environment. 

About 3000 fungi (exclusive of yeasts) have been reported 

from aquatic habitats of which Ascomycota (1 527 spp.) and 

anamorphic taxa (785 spp.) are the most diverse groups, 

followed by Chytridiomycota (576 spp.) with Basidiomycota 

(21 spp.) as the least diverse group (Vijaykrishna et al. 2006, 

Shearer et al. 2007). 

Anwar Abdel Aleem (Faculty of Science, University of 

Alexandria), or Peripatetic Aleem as he was known among his 

colleagues, is one of the most brilliant Arab marine botanists 

and oceanographer extraordinaire. He is considered one of 

the pioneer marine Egyptian mycologists, with studies on 

marine fungi dating back to 1950 (Aleem 1950a–c, 1952a–c, 

1953, 1962, 1974, 1975, 1978, 1980a, b, Aleem & Mailbari 

1981). 

In Egypt, obligate and facultative marine fungi are 

considered as forgotten fungi (Jones 2001) because they 

never featured in research topics until 1993, which is 

considered the starting point of marine mycology research 

in Egypt. This provided Mohamed Abdel-Wahab (Botany 

Department, Faculty of Science, South Valley University, 

Sohag, Egypt) the possibility to publish his pioneering study 

on the Egyptian obligate mangrove-inhabiting fungi of the 

Red Sea in 1996. Three contributions of El-Sharouny et al. 

(1998, 1999) and Abd-Elaah (1998) shed light on the ecology 

and taxonomy of mangrovicolous, algicolous and aquatic 

fungi of the Red Sea in Upper Egypt. Abdel-Wahab (2000) 

obtained his PhD on the biodiversity of fungi in subtropical 

mangroves; he recorded 25 fungi on intertidal wood of 

Avicennia marina collected from three mangrove stands of 

the Red Sea coast of Egypt. Abdel-Wahab et al. (2001a, 

b) published three new species, Halosarpheia unicellularis, 

Swampomyces aegyptiacus and S. clavatispora, from Red 

Sea mangroves. Pang et al. (2002) erected Jahnulales as a 

new lignicolous freshwater ascomycete order with the new 

species Patescospora separans from Egypt. Abdel-Raheem 

(2004) studied the effect of different techniques on diversity 

of freshwater hyphomycetes in the River Nile (Upper Egypt). 

Abdel-Wahab (2005) examined the diversity of marine fungi 

on intertidal decayed wood of A. marina and on decayed 

prop roots of Rhizophora mucronata in mangrove stands in 

the southern part of the Egyptian Red Sea coast; 39 species 

were identified on decayed wood of A. marina, of which 19 

were new records for Egypt and the Red Sea. Freshwater 

fungi are those relying on freshwater for at least part of their 

life-cycle (Wong et al. 1998, Raja et al. 2009). Abdel-Aziz 

(2008) studied the diversity of aquatic fungi in Lake Manzala, 

which was the first report of aquatic fungi from the lake. Sixty 

taxa including 26 ascomycetes and 34 anamorphic fungi 

were recorded, of which 19 species were new records for 

Egypt. El-Sharouny et al. (2009) studied the fungal diversity in 

brackish and saline lakes in Egypt; 97 fungi (40 ascomycetes, 

55 anamorphic fungi and 2 basidiomycetes) were identified 

from 764 collections, obtained from 545 samples, of which 70 

were new records for Egypt. 

The revision of all available data sources reveals that the 

total number of marine and aquatic fungi known in Egypt is 

207 taxa (87 Ascomycota, 117 anamorphic taxa, and 3 Basidiomycota). 

There is no checklist of aquatic Egyptian fungi so 

far. For more details on these fungi see the website (), search mangrove fungi (), and check relevant studies 

(Khallil 2001, Abdel-Aziz 2004, Abdel-Wahab et al. 2009, 

2010). 

Entomopathogenic fungi 

The taxonomy of the entomopathogenic fungi has received 

much attention since the 1970s. More than 700 species of 

fungi are associated with insects, spiders, and mites (Samson 

et al. 1988, Hajek & St. Leger 1994, Sung et al. 2007, Aung 

et al. 2008). 

The invertebrate pathogenic fungi can be classified 

in the Mastigomycota, Zygomycota, Ascomycota, and 

allied anamorphic fungi; no truly entomopathogenic 

basidiomycetes have been documented (Samson et al. 

1988). Entomopathogenic fungi range from commensals 

or mutualists, through ectoparasites which do not seriously 

affect their hosts, to pathogens which are lethal and include 

representatives of all the groups of fungi (Hawksworth et al. 

1995). 

Few records appeared reporting the occurrence of 

entomogenous fungi in Egypt until Natrass (1932) published 

preliminary notes on some of these fungi in Egypt. He 

recorded five species: Empusa grylli (= Entomophaga grylli), 

Beauveria bassiana, Aspergillus flavus, Mucor racemosum, 

and Metarhizium anisopliae. In the beginning of the 1960s, 

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Egypt started to apply biocontrol methods to insects by 

entomopathogenic fungi, and Gad et al. (1967) studied the 

occurrence of Coelomomyces indicus in Egypt. 

There are several studies on this ecological group of fungi 

in Egypt, such as Badran & Aly (1995), Shoulkamy et al. (1997), 

Shoulkamy & Lucarotti (1998), Hafez et al. (1997), Sewify 

(1997), Abdel-Baky (2000), Sewify Hashem (2001), Abdel-Sater & Eraky (2002), 

Ali (2003), Abdel-Mallek 

et al. (2003 a, b), El-Hady (2004), Mourad et al. (2005), 

Abdel-Mallek & Abdel-Rahman (2006), El-Maraghy et al. 

(2006), and Moubasher et al. (2010). As a result of these 

only 18 species belonging to 13 genera were recorded as 

entomopathogenic fungi of Egypt. For more details please 

refer to this site (). 

Nematophagous fungi 

In Egypt, the study of nematophagous fungi dated back to 

1963 when Hamdy Aboul-Eid (Department of Plant Pathology, 

Nematology Laboratory, National Research Centre, Dokki, 

Cairo) isolated and illustrated four species belonging to two 

genera. Various studies on the biocontrol of nematodes by 

fungi have been the target of many studies in Egypt; the most 

relevant are: Ali (1994, 1995), Ali & Barakat (1994), Aboul-Eid 

et al. (1997a, b, 2006), Ashour & Moustafa (1999), and Amin 

& Moustafa (2000). Out of these various data and information 

only 10 species belonging to seven genera were recorded as 

nematophagous fungi of Egypt. 

Basidiomycota 

The Basidiomycota contains about 31 503 described species, 

which represents 31.8 % of the known species of true Fungi 

(Kirk et al. 2008). This group includes mushrooms, puffballs, 

bracket fungi and some yeasts (Petersen et al. 2008, 

Wannathes et al. 2009). Many Basidiomycota decay dead 

organic matter, including wood and leaf litter symbiotic lifestyles 

(intimate mutually beneficial or harmful associations with other 

living organisms) are well developed in the Basidiomycota. They 

include major plant pathogens, such as “rusts” (Uredinales) and 

“smuts” (Ustilaginales), which attack wheat and other crops, 

and some human and animal pathogens. Not all symbiotic 

Basidiomycota cause harm to their partners. Indeed, some form 

ectomycorrhizas with the roots of plants, principally forest trees 

such as oaks, pines, dipterocarps, and eucalypts (Smith & Read 

1997, Rinaldi et al. 2008). Other symbiotic Basidiomycota form 

associations with insects, including leaf-cutter ants, termites, 

scale insects, wood wasps, and bark beetles (Wheeler & 

Blackwell 1984, Mueller et al. 1998). 

Macro-Basidiomycota 

The first information on hyphenate macro-basidiomycota 

(phytopathogenic or saprobic) in Egypt dates back to Delile 

(1813a), Melchers (1931), and Morse (1933). In her study on 

the genus Podaxis, Morse referred to some samples collected 

from Egypt. After six decades more information about macrobasidiomycota 

came to light through a series of studies 

carried out by several investigators, such as Mouchacca 

(1977), Zakhary (1979), Salem & Michail (1980), Zakhary et 

al. (1983), MalenÇon (1984), Assawah (1991), Chen (1999), 

Abu El-Souod et al. (2000), El-Fallal (2003), El-Fallal & Khedr 

(n. dat.), El-Fallal & El-Diasty (2006), Kim et al. (2006), and 


An exhaustive revision of all the available literature and 

sources mentioned since 1931 shows that 108 taxa belonging 

to 65 genera, 104 species, and 4 varieties of Egyptian macrobasidiomycotese 

had been recorded up to the present time. 

Plant pathogenic Basidiomycota 

Though many basidiomycetes are saprobes or wood-rotters, 

the Basidiomycota contains two common and destructive 

groups of plant pathogens: rusts and smuts. Rust fungi 

are the largest group of fungal plant pathogens, containing 

7 000 species that possess the most complex life-cycles in 

the kingdom fungi (Sert 2009). They are obligate biotrophs 

and cause disease on most crops, ornamentals, and many 

other plants (Hawksworth et al. 1995). In addition to basidia 

and basidiospores, rusts produce other types of spores such 

as teliospores spermatia, aeciospores, and uredospores. 

Rusts that produce all five types of spores are referred to as 

macrocyclic, while rusts that lack one or more spore type are 

referred to as microcyclic. Unlike rusts, smuts produce only 

basidiospores and teliospores which can survive in the soil 

away from a host plant. Smuts commonly infect the ovaries 

of grains and are easily recognized by the formation of galls 

which contain masses of black spores (Agrios 2005). 

The initial research and documentation of rust and smut 

diseases in Egypt was by Reichert (1921), Briton-Jones 

(1922), Philp & Selim (1941), Abdel-Hak & Abdel-Rehim 

(1950), Ragab & Mahdi (1966), and Assawah (1969). Later, 

in-depth research was carried out by Egyptian and other 

investigators, with different targets such as taxonomy, 

pathogenicity, biocontrol and serology. The most relevant 

studies are: Sherif et al. (1991), El-Shamy (1996), Baka & 

Gjaerum (1996), Mennicken et al. (2005), Abd El Fattah et al. 

(2009), Abd EL-Ghany (2009) and Ismail et al. (2009). Baka 

& Gjaerum (1996) gave the first serious modern taxonomic 

treatment of local rusts, reporting 23 rust species on various 

monocotyledonous and dicotyledonous plants in the Nile 

Valley (see Mouchacca 2003b). As a result of these studies, 

112 species of plant pathogenic Basidiomycota belonging to 

21 genera were recorded from Egypt. 

Total recorded species 

After the omission of duplicate names, name correction, 

allowance for synonyms and taxonomic assignments of all 

reported taxa from Egypt, the number of the Egyptian fungi 

recorded is 2 281 taxa belonging to 755 genera (Table 1). 

At the generic level, some genera exhibit an extraordinary 

high species richness such as Aspergillus (100 spp.) and 

Penicillium (83). Other genera show moderate richness such 

as Chaetomium (53 spp.), Fusarium (49), Puccinia (41), 

Pythium (30) and Alternaria (27). 

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Table 1. Numbers of recorded Egyptian fungi. 

Groups and Phyla El-Abyad (1997) Present survey 

Amoebozoa 0 25 

Protozoan fungal analogues 

Cercozoa 0 3 

Hyphochytriomycota 1 3 

Labyrinthista 0 2 

Chromistan fungal analogues 

Oomycota 25 40 

Incertae sedis 0 1 

Blastocladiomycota 3 7 

Chytridiomycota 21 32 

Zygomycota 17 35 

Glomeromycota 0 8 

Teleomorphic genera 80 251 

Ascomycota 

Anamorphic genera 181 261 

Basidiomycota 32 87 

Total no. of genera recorded in Egypt 360 755 

Total no. of species recorded in Egypt 1246 2281 

DISCUSSION 

It is generally accepted that only about 7 % of all fungi 

have so far been discovered, and about 93 % still wait to 

be discovered. Fungi are neglected organisms and they 

are not well protected, but like animals and plants, they 

are endangered by human activities. Although the 1992 

Convention on Biological Diversity extends protection to 

all groups of organisms, it is worded in terms of “animals, 

plants and microorganisms” and fungi do not fit well into 

these categories. In Egypt and up to now fungal biodiversity 

and conservation topics have been overlooked. As a 

result, countries which signed the Convention have almost 

universally overlooked fungi in preparing their biodiversity 

conservation plans: fungi are truly the orphans of Rio (Minter 


Threats to fungi throughout the globe are of concern 

since they are not only beautiful but also play a significant 

role in human welfare. Three steps were suggested by Moore 

et al. (2001) for fungal conservation: (1) conservation of 

habitats; (2) in situ conservation of non-mycological reserves/ 

ecological niches; and (3) ex situ conservation especially for 

saprobic species growing in culture. To help collections of 

fungal cultures to maintain appropriate standards, the World 

Federation for Culture Collections (WFCC) has formulated 

guidelines which outline the necessary requirements 

(Hawksworth 1991, Smith et al. 2001, Smith 2003). There are 

573 microbial culture collections in 68 countries registered 

in the World Directory of Collections of Microorganisms 

(DCM) (). In Egypt only two 

centers are recorded: EMCC (WDCM583) Egypt Microbial 

Culture Collection, Cairo Microbiological Resources Centre 

(Cairo MIRCEN), Ain Shams University, and NODCAR 

WDCM822 Marwa Mokhtar Abd Rabo, National Organization 

of Drug Control and research. However, Moubasher and his 

colleagues founded the Assiut University Mycological Centre 

(AUMC) in 1999 where more than 6 000 fungal isolates 

belonging to more than 500 species are being preserved 

under low temperature (5 °C), deep-freezed (-80 °C), and 

lyophilized; this is the biggest reference culture collection in 

the Arab countries. The centre also has a collection of dried 

specimens (i.e. a fungarium) which is rare in Arab countries. 

In spite of this the AUMC is not yet registered with the WFCC. 

The number of habitats that potentially support 

specialized fungi is enormous. The fungi described as new to 

science during 1981 to 1990 were associated with 1 982 host 

genera or substrata (Hawksworth & Rossman 1997). Some 

unexplored substrata and habitats from which these fungi 

were found include the rumens of herbivorous mammals, 

algae, lichens, mosses, marine plants, including mangroves 

and driftwood, rocks and insect scales. 

The Egyptian fungi are presently represented by 2 281 

taxa (1 035 species and 395 genera) out of the 101 202 

world estimate. In comparing the fungal diversity recorded 

in Egypt with other countries, it is important to mention that 

some ecological groups of fungi are completely ignored or 

have never been studied in a comprehensive way in Egypt, 

such as Trichomycetes (a group of enigmatic fungi occurring 

in the hindguts of insects and other invertebrates; Lichtwardt 

2002), in addition to hypersaline and black yeasts. Other 

groups needing more exploration such as algicolous fungi, 

invertebrate associated fungi, mycorrhizas, endophytic fungi, 

lichens, wood deteriorating, and coprophilous fungi. The 

potential fungal resources of Egypt are globally important 

and there are vast areas that are still unexplored. At present, 

Egypt needs more investigators and funds to explore and 

develop this research field and, therefore, the extensive 

collection of fungi in unexplored areas remains a priority. 

This review will be followed by an updated checklist of 

all recorded Egyptian fungi up to the present, a bibliographic 

study of Egyptian mycological research, and a book on the 

fungi of Egypt, supplemented with provisional keys to all 

species listed. 

132 

 

i m a f U N G U S


ACKNOWLEDGEMENTS 

All my thanks to the late Samy M El-Abyad (Botany Department, 

Faculty of Science, Cairo University) for his courage in documenting 

the Egyptian fungi in 1997. I express my appreciation to Paul M Kirk 

(CABI Europe, UK) for data on species names in the Index Fungorum 

database; the late John C Krug (Centre for Biodiversity and 

Conservation Biology, Royal Ontario Museum, Toronto) for providing 

some unavailable papers. I am further indebted to Jean Mouchacca 

(Laboratoire de Cryptogamie, Muséum National d’Histoire Naturelle, 

Paris), Yaacov Katan (Department of Plant Pathology and 

Microbiology, Faculty of Agriculture, Food and Environmental Quality 

Sciences, The Hebrew University of Jerusalem, Israel) and Hussien 

M Rashad (Ashtoum El-Gamil Protectorate, Port Said, Egypt) for 

their unfailing help during this work. I also owe thanks to Robert A 

Blanchette (Forest Pathology and Wood Microbiology Research 

laboratory, Minnesota University) and El-Sayeda M Gaml El-Din 

(Botany Department, Faculty of Science, Suez Canal University) for 

critical reading the manuscript. I also thank Pedro W Crous (CBS- 

KNAW Fungal Biodiversity Centre, Utrecht) who worked closely with 

me in preparing and editing the manuscript and photoplates, and the 

two anonymous reviewers for their constructive comments. 

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IMC9 Edinburgh Nomenclature Sessions 

Lorelei L. Norvell 1 , David L. Hawksworth 2 , Ronald H. Petersen 3 and Scott A. Redhead 4 

1 

Pacific Northwest Mycology Service, Portland OR 97229-1309, USA; corresponding author e-mail: llnorvell@pnw-ms.com 

2 

Departamento de Biología Vegetal II, Facultad de Farmacia, Universidad Complutense de Madrid, Plaza Ramón y Cajal, Madrid 

28040, Spain and Department of Botany, Natural History Museum, Cromwell Road, London, SW7 5BD, UK 

3 

Department of Biology, University of Tennessee, Knoxville TN 37920, USA 

4 

Biodiversity (Mycology and Botany), Agriculture & Agri-Food Canada, Ottawa, Ontario K1A 0C6, Canada 

ARTICLE 

Abstract: The proceedings of the 3–5 August 2010, IMC9 Edinburgh Nomenclature 

Sessions are briefly summarized. The final resolution approved by the General Assembly 

endorses the recommendations by the Nomenclature Sessions regarding transfer of the 

governance of fungal nomenclature from botanical to mycological congresses, mandatory 

pre-publication deposit of nomenclatural information for valid publication of new fungal 

names, and the acceptability of English as an alternative to Latin in the valid publication 

of fungal names. Complete results from the IMC9 nomenclature questionnaire are also 

provided. 

Key words: 

dual nomenclature 

electronic publication 

governance 

International Code of Botanical Nomenclature 

Latin 

MycoBank 

Article info: Submitted: 25 October 2010; Accepted: 7 November 2010; Published: 12 November 2010. 

Introduction 

Three successive groundbreaking two-hour long 

nomenclatural sessions were held August 3–5, 2010, 

during this summer’s International Mycological Congress 

(IMC9) in Edinburgh, Scotland. Convener/Rapporteur David 

Hawksworth (Spain/UK), who supervised preparation of the 

IMC9 nomenclatural booklet + questionnaire, was assisted 

by Chair Ron Petersen (USA), Vice-Chair Scott Redhead 

(Canada), Nomenclature Committee for Fungi (NCF) 

Secretary Lorelei Norvell (USA), and Advisor & International 

Botanical Congress Rapporteur-général John McNeill 

(UK). IMC delegates attending each day’s session voted 

on nomenclatural proposals to recommend actions to next 

year’s International Botanical Congress (IBC) Nomenclature 

Section in Melbourne. Attendance was relatively high, 

particularly in view of the conflict caused by scheduling the 

three nomenclature and three (of four) poster sessions for 

the same 2–4 pm time periods. As each poster session 

presented authors and posters for only one day, this was an 

unfortunate conflict that influenced attendance numbers at 

the nomenclatural sessions. However, the questionnaires, 

distributed to all IMC9 delegates for return to the registration 

desk by the end of the Congress, permitted each delegate a 

chance to express an opinion, even if unable to attend any or 

all of the Nomenclature Sessions. 

Originally the entire proceedings, which proved to be 

lively, informative, and often amusing, were to be recorded. 

Due to an unfortunate communications failure, no recordings 

survive. The overly brief summary below has therefore been 

extracted from secretarial notes, the nomenclature booklet, 

and the returned questionnaires. 

Background 

When initially formed in 1971, the International Mycological 

Association (IMA) established a Nomenclature Secretariat to 

address issues of concern to mycologists. This led to a series 

of proposals on starting points and other matters that were 

adopted by the International Botanical Congress in Sydney 

in 1981, after which it was disbanded, having completed its 

tasks. Since that time, discussions of nomenclatural issues at 

IMCs have been confined to occasional debates on particular 

topical issues. However, at IMC8 in Cairns in 2006, some 

delegates spoke strongly in favour over a separate Code 

for fungi. Subsequently, proposals that could fundamentally 

change aspects of fungal nomenclature have been published; 

these are to be voted on at the forthcoming International 

Botanical Congress (IBC) in Melbourne in July, 2011. As 

IBCs occur only every six years, and decisions made there 

generally come into force 1–2 years later, any issues not 

decided in 2011 would have to wait until 2018 or 2019 to be 

implemented. The Nomenclature Sessions at IMC9 were 

convened to: (1) enable a broad spectrum of mycologists to 

express their views on a wide range of topics and also to vote 

on proposals already made; and (2) establish that IMCs can 

incorporate effective Nomenclatural Sessions. 



Attribution: 






v o l u m e 1 · n o . 2 

143

Norvell et al. 

ARTICLE 

Session 1: Governance of fungal 

nomenclature 

Approximately 100 delegates attended the first session 

convened by Hawksworth at 2 pm on August 3. After Chair 

Petersen set forth the “rules of engagement” for audience 

participation during all sessions, two introductory background 

presentations were given. Vincent Demoulin (Belgium, 

Chairman of the Committee for Fungi) spoke in defense of 

retaining governance of fungi within the Botanical Code and 

Hawksworth reported on the progress being made toward 

one unified code for all organisms. (See Appendix 1, below.) 

The floor was then opened to discussion of the formal 

proposals for the governance of fungal nomenclature, the 

composition of the Nomenclature Committee for Fungi, 

and a (very) brief discussion of the proposed exclusion of 

Microsporidia from the ICBN. At the close of the two-hour 

session, those remaining in the auditorium were polled as to 

their preferences, summarized as follows: 

Props. 016–020 (Hawksworth et al. 2009) all passed. 

Votes were actually counted for the first two proposals: both 

Prop. 016 (to amend the current Botanical Code to establish 

more clearly that it covers fungi, including changing the 

name to the “International Code of Botanical and Mycological 

Nomenclature”) & Prop. 017 (to replace “plants” by “plant(s) 

or fungus/fungi” throughout) passed with 87 yes and 4 no 

votes. Thereafter, due to time pressures, only the ‘no’ votes 

(out of 91 total) were counted, with 3 voting against Prop. 018 

(to provide for the election of the Permanent Nomenclature 

Committee for Fungi by an International Mycological 

Congress), 3 voting against Prop. 019 (to relegate decisionmaking 

on proposals relating solely to organisms treated as 

fungi to an IMC), and 1 against Prop. 020 (to insert a new Div. 

III.5 requiring the presence of the Secretary for the Committee 

for Fungi or Committee alternate on the Editorial Committee). 

Unanimous support was given to retaining the current 

members of the Committee for Fungi until the 2014 IMC10 

in Bangkok, provided that the 2011 International Botanical 

Congress in Melbourne accepts the fungal governance 

proposals above. 

Props. 048–051 (to exclude the governance of the 

phylum Microsporidia from the Code; Redhead et al. 2009) 

passed with only one dissenting vote, but as the vote 

was held as delegates were leaving the session, it may 

not accurately reflect the wishes of the majority. Demoulin 

has since submitted Prop. 190 to limit Art. 45.4 (Demoulin 


Session 2: Mandatory pre-publication deposit 

in a nomenclatural repository, electronic 

publication, type cultures, and illustrations 

After opening introductions, Paul Kirk (UK) provided an 

overview of the current strides made in data-basing taxonomic 

names of all organisms worldwide. (See Appendix 1, below.) 

A fluctuating audience (estimated at 97 total for the 2-hour 

session) discussed at length and eventually recommended 

Props. 117–119 (Hawksworth et al. 2010). Prop. 117 (to 

require deposition of names and required nomenclatural 

information in a recognized repository (such as MycoBank) for 

valid publication) received 58 yes, 5 no, and 1 abstaining votes. 

Props. 118 (to recommend deposit of minimal information 

elements, accession identifiers, and bibliographical details for 

valid publication) and 119 (to require citation of a repository 

identifier for valid publication) received almost universal 

support, with 1 and 2 abstentions respectively. Kirk also 

announced that it would be possible to deposit names via 

Index Fungorum, although the mechanism (still in progress) 

was not detailed. 

An informal poll showed no clear consensus for or against 

valid electronic publication of names. 

Prop. 138 (Nakada 2010), which seeks to add Rec. 8B.3, 

including the phrase “permanently preserved in a metabolically 

inactive state” or its equivalent when designating a culture as 

a type) likewise showed no clear consensus with the majority 

abstaining. 

The session concluded with a second informal poll 

(showing 4 for, 25 against, and the majority abstaining) 

regarding the addition of illustrations as a requirement for 

valid publication. 

Session 3: Moving to one name for one 

fungus and ending the requirement of Latin 

diagnoses for valid publication 

Approximately 145 delegates attended the final (and most 

controversial) “Article 59” session on August 5. Background 

on attempts to modify dual nomenclature was provided by 

Redhead (Secretary for the Special Committee on Names 

of Fungi with a Pleomorphic Life History), followed by a 

presentation by Walter Gams (Netherlands), who spoke on 

the limitations of “teleotypifying” fungal names according to 

Art. 59.7. (See also Appendix 1, below.) 

Emotions ran high in this session, and discussion was 

lively, entertaining, lengthy — and inconclusive. No formal 

proposals were before the Session, so no vote was scheduled 

on Art. 59. It was assumed that Congress participants would 

mark their opinions on their questionnaires. 

Due to the lengthy Art. 59 debate, the scheduled 

discussion and vote on whether to end the requirement of 

a Latin diagnosis for the valid publication of scientific names 

(also to be considered in 2011 at Melbourne) became a side 

issue. Entrants crowding the doors for the next scheduled 

mycological session dictated Chair Petersen’s decree for 

adjournment, which drowned out the plaintive cry from the 

back of the hall, “Why can’t we vote to abolish Latin?” and a 

call to hold a vote on Art. 59. 

Final resolution approved by the General 

Assembly — and a note of caution 

At the close of the first Nomenclature Session, 103 

questionnaires had already been returned. By the evening of 

the final session, Hawksworth and Norvell had tabulated 167 

results and identified three clear preferences for presentation 

to the delegates during the IMC9 closing ceremonies on 

August 6. The General Assembly voted by acclamation to 

approve the resolution below: 

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i m a f U N G U S


This General Assembly of the IMA endorses the decisions 

of the Nomenclature Session convened during IMC9 with 

respect to 

— the transference of the governance of the nomenclature 

of fungi from the International Botanical to International 

Mycological Congresses, 

— the mandatory pre-publication deposit of nomenclatural 

information in a recognized depository for the 

valid publication of new fungal names, 

— the acceptability of English as an alternative to Latin in 

the valid publication of fungal names, 

and requests the permanent Nomenclature Committee for 

Fungi, the special Committee on the names of Pleomorphic 

Fungi, the International Commission on the Taxonomy of 

Fungi, and the next International Botanical Congress to 

take note of the results of the questionnaire completed by 

delegates of IMC9. 

In summary, we must emphasize that these are 

recommendations and not approved changes. Currently 

fungal names are still governed by the International Code 

of Botanical Nomenclature, and — until changed — a 

Latin description or diagnosis is still required, as are other 

established requirements for valid publication as set forth in 

the current International Code of Botanical Nomenclature 

(McNeill et al. 2006). Nonetheless the interest shown in 

nomenclature at IMC9 was gratifying, and we are optimistic 

that many of the innovations supported by most mycologists 

will be made. 

Appendix 1: IMC9 Nomenclature Session 

presentation abstracts 

Fewer nomenclatural codes, not more, is what we need 

(Demoulin): At the first IMC (Exeter, 1971) the idea of a 

nomenclature code especially for fungi was discussed and a 

nomenclature committee was created under the auspices of 

the IMA. This committee reported at the 2 nd IMC in Tampa, Fl. 

1977. At that congress, the idea of a mycological code was 

abandoned in favour of more involvement by mycologists in 

the elaboration of the Botanical Code, which has ruled the 

nomenclature of fungi since its origin. A consequence was 

the important change in the starting point system adopted 

at the 13 th International Botanical Congress (Sydney, 1981). 

Progress towards a BioCode (Hawksworth): In October 

2009, the General Assembly of the International Union of 

Biological Sciences (IUBS) decided to re-activate the initiative 

to produce a unified Code of nomenclature for all organisms, 

by updating the Draft BioCode (Greuter et al.1998). This 

is being taken forward by the International Committee for 

Bionomenclature of the IUBS/IUMS (International Union of 

Microbiological Societies). The need for, and route towards, 

a revised and agreed BioCode is reviewed as a background 

to the Session’s deliberations. 

A web of data for fungal biology research — the 

registration question (Kirk): Why do we give names to fungi? 

It’s a simple question with a simple answer - to allow us to 

effectively communicate about the fungi, for the name is the 

link to all that is known about the organism. But in this answer 

the word ‘us’ is already of secondary importance. The web is 

the primary means of communication today and increasingly 

that means computer to computer communication. In addition, 

the current version of the web - a web of information - is 

rapidly being replaced by a web of data (the Semantic Web, 

especially Linked Data using RDF triples of entity-attributevalue) 

which will allow more rapid (real time) advances in 

synthesis, analysis, hypothesis, etc. The founder of the web 

Tim Berners-Lee, amongst others, is pushing for this to happen 

and we can be part of this effort. This short presentation will 

describe how name registration can operate, how associated 

data can be made available, what the barriers are, and how 

it all fits into existing and developing major global initiatives. 

It will indicate how fungal taxonomist and nomenclaturalists 

can be part of this with respect to the names we give to fungi. 

How do mycologists wish to treat names based on 

anamorphs? (Redhead): Fungal nomenclature dates back to 

Linnaeus (1753) when the use of microscopes was limited 

and the existence of sexual life cycles amongst them was 

unknown. Nearly 200 years later (1935) mycologists realized 

they had been naming different parts of fungal life-cycles as 

new species or genera, and formalized nomenclature rules 

giving priority to names for pleomorphic fungi based upon 

perfect states. Exceptions and refinements were instituted in 

1950 and continue today. Many fungi only produce anamorphs, 

many generic names are based upon anamorphs, and many 

fungi are better known under anamorph names. However, 

complications in merging and then prioritizing names have 

created a nightmare situation that has divided the mycological 

community and now acts as a roadblock. Proposals to block 

the deliberate generation of alternative names and smooth the 

transition to normal nomenclature were partially approved for 

Article 59 in the International Code of Botanical Nomenclature 

(McNeill et al. 2006) while remaining issues were referred to a 

Special Committee by the IBC. After >4 years this Committee 

was unable to reach consensus upon changes. Some 

mycologists have decided to ignore existing rules or to take 

nomenclatural risks. Genetic sequence phylogenetic analyses 

have revealed many new relationships leading to binomial 

recombinations and even a PhyloCode. Having reached an 

impasse it can be asked if mycologists wish to eliminate dual 

nomenclature? If the answer is yes, it may be asked how to 

resolve conflicts, and then to create a process or body capable 

of dealing with such conflicts. 

Teleotypification of fungal names and its limitations 

(Gams): This presentation was submitted without a formal 

abstract and too late to be included in the printed program. 

Gams discussed the effects of ‘teleotypification,’ which 

permits — after a teleomorph discovered for a fungus 

previously known only as an anamorph (and for which 

there is no existing legitimate name for the holomorph) — 

designation of an epitype exhibiting the teleomorph stage for 

the hitherto anamorphic name, even when there is no hint 

of the teleomorph in the protologue of that name. Several 

examples were forwarded to show that teleotypification is not 

the same as ordinary epitypification. For further information, 

see Props. (172–174; Gams et al. 2010). 

ARTICLE 

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Appendix 2: IMC9 Nomenclature 

questionnaire results 

From August 1–10, IMC9 delegates returned questionnaires 

in which they were to circle a Y (yes) or N (no) to 24 questions 

on 4 topics. We discovered during our first tabulation that one 

number (#19) appeared twice, bringing the actual number 

of questions to 25, and have renumbered the text below 

accordingly. Of the 174 questionnaires received, 7 were 

declared ‘spoiled’ as the respondents had placed an X over 

an option so that we could not determine whether agreement 

or rejection was intended. Both raw numbers and majority 

percentages are shown. We note that protocols followed at the 

2005 International Botanical Congress in Vienna with respect 

to the preliminary mail-in ballots decreed that proposals 

receiving 60 % or higher support merited further discussion 

by the attending Nomenclature Section, while 75 % support 

virtually ensured passage for all but the most controversial 

proposals. In the results reported below, opinions showing 

60 % (or greater) support are highlighted in bold. 

A. Codes of nomenclature 

(Fungal names are now governed by the International Code of Botanical Nomenclature) 

1 One code for the future nomenclature of all organism names would be ideal 

y-72 n-71 ...............................................................................................................................................................50 % (tie) 

2 Fungi should continue to be covered under the Botanical Code (ICBN) 

y-54 n-76 ................................................................................................................................................................ 58 % no 

3 Fungi should continue to be covered under the ICBN provided it is renamed the “Botanical and Mycological Code” 

y-97 n-40 ............................................................................................................................................................... 71 % yes 

4 Fungi should be covered by a separate mycological Code (ICMN) 

y-51 n-91 ................................................................................................................................................................ 61 % no 

5 Under either ICBN or ICMN, decisions on fungal nomenclature should be voted at an International Mycological 

Congress (and not an International Botanical Congress), guided by a secure advanced web publication and 

mail/email votes 

y-133 n-21 ............................................................................................................................................................. 86 % yes 

B. Language requirements for valid publication of names 

6 Latin diagnoses/descriptions should continue to be required 

y-49 n-91 ................................................................................................................................................................ 65 % no 

7 English diagnoses/descriptions rather than Latin should be required 

y-69 n-69 .............................................................................................................................................................. 50 % (tie) 

8 Either Latin or English diagnoses/descriptions should be required 

y-88 n-56 ............................................................................................................................................................... 61 % yes 

9 Diagnoses/descriptions in any language should be permitted 

y-4 n-135 ................................................................................................................................................................ 97 % no 

C. Nomenclatural information databasing 

10 Deposition of key nomenclatural information in one or more approved depositories (e.g. MycoBank) should be made 

mandatory for the valid publication of new fungal names 

y-134 n-21 ............................................................................................................................................................. 86 % yes 

11 Historic names not included in Index Fungorum (after a set date) should no longer be treated as validly published 

y-55 n-68 ................................................................................................................................................................ 55 % no 

12 Deposited names should be automatically protected against any unlisted names after a date to be agreed 

y-90 n-39 ............................................................................................................................................................... 70 % yes 

13 An accurate and free list should be prepared of names in use or available for use 

y-126 n-19 ............................................................................................................................................................. 87 % yes 

14 Names with key information deposited (e.g. in MycoBank) should be automatically available provided other Code 

requirements are met 

y-105 n-22 ............................................................................................................................................................. 83 % yes 

15 Electronic on-line only publication should be accepted without restriction 

y-24 n-126 .............................................................................................................................................................. 84 % no 

16 Electronic on-line only publication should be accepted only when key nomenclatural information has been deposited 

(e.g. in MycoBank) 

y-113 n-36 ............................................................................................................................................................. 76 % yes 

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17 For journals publishing online and printed copies, the dates of names should be those when the works are available in 

final form on-line 

y-101 n-40 ............................................................................................................................................................. 72 % yes 

18 For journals publishing online and printed copies, the dates of names should be those when the works are distributed 

in printed form 

y-63 n-73 ................................................................................................................................................................ 54 % no 

19 Special Group Committees should be empowered to create lists of acceptable and rejected names in particular groups 

(e.g. Fusarium, Trichocomaceae, yeasts) 

y-102 n-31 ............................................................................................................................................................. 77 % yes 

ARTICLE 

D. Names for pleomorphic fungi (anamorphs, teleomorphs) 

20 The established system allowing dual nomenclature for anamorphs and teleomorphs should continue via Art. 59 

y-67 n-71 ................................................................................................................................................................ 51 % no 

21 Article 59 should revert back to its status prior to changes in the 2006 Vienna Code, i.e. keeping separate anamorph 

and teleomorph names 

y-43 n-82 ................................................................................................................................................................ 66 % no 

22 A system of progressively establishing one name for each fungus should be enacted via modification of existing 

Articles (e.g. Art. 59) 

y-101 n-38 ............................................................................................................................................................. 73 % yes 

23 The historical practice of allowing valid names for different morphs of a species should be prohibited in the future via 

modification of existing Articles 

y-74 n-45 ............................................................................................................................................................... 62 % yes 

24 The ability to select a “teleotype” (a type of epitypification) with a sexual state for a fungus previously only known in 

the asexual state should be continued 

y-88 n-31 ............................................................................................................................................................... 74 % yes 

25 Article 59 (that permits the dual system) should be deleted provided other changes ensure this would not retroactively 

invalidate existing names 

y-66 n-47 ............................................................................................................................................................... 58 % yes 

Acknowledgments 

We thank John McNeill (Royal Botanic Garden Edinburgh) for his 

perennially wise counsel and cheerful guidance. We further thank 

special presenters Vincent Demoulin, Paul Kirk, and Walter Gams; 

José Dianese (Brazil) for assisting in tabulating questionnaire results 

on 3 August; and all those who participated in the nomenclatural 

discussions at IMC9 Edinburgh and/or completed questionnaires. 

References 

Demoulin V (2010) Proposals to amend Articles 15, 36, and 45. 

Taxon 59: 1627–1628. 

Gams W, Jaklitsch WM, Kirschner R, Réblová M (2010) Three 

proposals to amend Article 59 of the Code concerning 

teleotypification of fungal names. Taxon 59: 1297. 

Greuter W, Hawksworth DL, McNeill J, Mayo MA, Minelli A, Sneath 

PHA, Tindall BJ, Trehane P, Tubbs P (eds) (1998) Draft BioCode 

(1997): the prospective international rules for the scientific 

names of organisms. Taxon 47: 127–150. 

Hawksworth DL, Crous PW, Dianese JC, Gryzenhout M, Norvell 

LL, Seifert KA (2009) Proposals to amend the Code to make it 

clear that it covers the nomenclature of fungi, and to modify the 

governance with respect to names of organisms treated as fungi. 

Taxon 58: 658–659; and Mycotaxon 108: 1–4. 

Hawksworth DL, Cooper JA, Crous PW, Hyde KD, Iturriaga T, Kirk PM, 

Lumbsch HT, May TW, Minter DW, Misra JK, Norvell L, Redhead 

SA, Rossman AY, Seifert KA, Stalpers JA, Taylor JW, Wingfield 

MJ (2010) Proposals to make the pre-publication deposit of 

key nomenclatural information in a recognized repository a 

requirement for valid publication of organisms treated as fungi 

under the Code. Taxon 59: 660–662; Mycotaxon 111: 514–519. 

McNeill J, Barrie FR, Burdet HM, Demoulin V, Hawksworth DL, 

Marhold K, Nicolson DH, Prado J, Silva PC, Skog JE, Wiersema 

JH, Turland NJ (eds) (2006) International Code of Botanical 

Nomenclature (Vienna Code) adopted by the Seventeenth 

International Botanical Congress Vienna, Austria, July 2005. 

[Regnum Vegetabile no. 146.] Ruggell: A.R.G. Ganter Verlag. 

Nakada T (2010) A proposal on the designation of cultures of fungi 

and algae as types. Taxon 59: 983. 

Redhead SA, Kirk PM, Keeling PJ, Weiss LM (2009) Proposals to 

exclude the phylum Microsporidia from the Code. Mycotaxon 

108: 505–507; Taxon 58: 669. 

[Reproduced with minor amendments from Mycotaxon 113: 503–511 

(2010).] 

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Fungal phoenix rising from the ashes? 

Michael J. Wingfield 1 , Martin P.A. Coetzee 1 , Pedro W. Crous 2 , Diana Six 3 and Brenda D. Wingfield 1 

1 

Department of Genetics, Forestry and Agricultural Biotechnology Institute (FABI), University of Pretoria, Pretoria, 0002, South Africa; 

corresponding author e-mail: mike.wingfield@fabi.up.ac.za 

2 

CBS-KNAW Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands 

3 

Department of Ecosystem and Conservation Sciences, College of Forestry and Conservation, University of Montana, Missoula, MT 59812, USA 

ARTICLE 

Abstract: During May 2010, sporocarps of what appeared to be an Armillaria sp. were found in large clumps in historic 

Kirstenbosch Botanical Gardens on the foot of Table Mountain, Cape Town, South Africa. These sporocarps could be 

physically linked to the roots of unidentified dead trees and Protea spp. The aim of this study was to identify the Armillaria 

sp. found fruiting in Kirstenbosch. To achieve this goal isolates were made from the mycelium under the bark of dead roots 

linked to sporocarps. The ITS and IGS-1 regions were sequenced and compared to sequences of Armillaria spp. available 

on GenBank. Cladograms were generated using ITS sequences to determine the phylogenetic relationship of the isolates 

with other Armillaria spp. Sequence comparisons and phylogenetic analyses showed that the isolates represented A. 

mellea. They were also identical to isolates of this species previously discovered in the Company Gardens in South Africa 

and introduced from Europe apparently by the early Dutch Settlers. Armillaria mellea is alien and apparently invasive in 

Cape Town, fruits profusely and has the potential to spread to sensitive native forests on the foothills of the City. 

Key words: 

Armillaria mellea 

Armillaria root rot 

fungal introduction 

Proteaceae 


INTRODUCTION 

Species of Armillaria are some of the most important 

pathogens of woody plants in the world. These fungi have 

been known as tree pathogens since their first discovery by 

Danish botanist Martin Vahl. While the taxonomy of these 

Armillaria spp. has been controversial and widely debated 

over an extended period of time, application of the biological 

species concept (Korhonen 1978, Anderson & Ullrich 1979, 

Ota et al. 1998, Qin et al. 2007) and more recently DNA 

sequence comparisons (Coetzee et al. 2000a, 2003a, 2005, 

Gezahgne et al. 2004, Keča et al. 2006, Mwenje et al. 2006, 

Hasegawa et al. 2010) have resolved many problems relating 

to the delineation of species. At least 40 species are now 

recognised in Armillaria (Volk & Burdsall 1995, Lima et al. 

2008, Pildain et al. 2010) and it is likely that other species will 

emerge from under-sampled areas in the future. 

Armillaria root rot, the disease caused by pathogenic 

Armillaria spp. can result in serious losses to productivity in 

tree plantations, fruit tree orchards and in gardens (Gregory 

et al. 1991, Hood et al. 1991). In native forests, Armillaria 

spp. cause disease but this is most typically a natural process 

(Kile et al. 1991). Interestingly, species of these fungi exist as 

clones covering huge areas of land and in these situations 

they are considered to be amongst the largest and oldest 

living organisms (Gould 1992, Smith et al. 1992). 

Only a single native Armillaria sp. occurs in South 

Africa (Coetzee et al. 2000a). This fungus, A. fuscipes is 

occasionally found on native trees (Kotzé 1935, referred 

to as A. mellea). In contrast, it can be a serious pathogen 

in plantations of non-native Pinus spp. and on fruit trees 

planted in moist areas that have been cleared of native 

forest (Lundquist 1986, 1987, Coetzee et al. 2000a). A more 

intriguing Armillaria sp. in South Africa is A. mellea that was 

discovered in the Company (Dutch East India Company) 

Gardens in the centre of Cape Town (Coetzee et al. 2001). 

The fungus in that situation represents a single genetic entity 

that was shown to be at least 358 years old. It was most 

likely introduced into the city when gardens were established 

to provide sailors travelling to the East with fresh produce 

(Coetzee et al. 2001). 

Some years after the discovery of the A. mellea clone in 

Cape Town, Armillaria root rot was found killing Protea plants 

in the historic Kirstenbosch Botanical Gardens () on the foothills of South Africa’s iconic Table Mountain 

(Coetzee et al. 2003b). The fungus in that situation was never 

seen fruiting but isolates were identified as those of A. gallica, 

and it was suggested that the fungus was introduced into the 

gardens with plants brought from Asia (Coetzee et al. 2003b). 

A few of the isolates collected on the Protea plants were also 

thought to represent A. mellea, but the identification was 

tentative and based only on RFLP comparisons, without 

comparison of DNA sequence data against other Armillaria 

spp. 

During May 2010, sporocarps of what appeared to be an 

Armillaria sp. were found in large clumps in the upper corner 



Attribution: 






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Fig. 1. Armillaria root rot in Kirstenbosch Botanical Gardens. A. Native woody shrubs and forest deeper in the valleys common on the Cape 

Peninsula. B, C. Clusters of fruiting bodies found on a stump. D–G. Robust fruiting bodies of Armillaria sp. showing a yellow cap, prominent 

annulus and stipe tapering down to the base. H. Rhizomorphs produced in culture. 

of Kirstenbosch Botanical Gardens and close to Rycroft’s 

Gate. These sporocarps could be physically linked to the 

roots of unidentified dead trees and Protea spp. (Fig. 1B, 

C). Upon removal of the bark from the dead roots, sheets 

of white mycelium typical of Armillaria root rot were found. 

The aim of this study was to identify the Armillaria sp. found 

fruiting in Kirstenbosch using data that were not available at 

the time of the discovery of Armillaria root rot in Cape Town 

(Coetzee et al. 2003b). 


Isolates 

Isolation and purification of isolates followed the methods 

outlined in Coetzee et al. (2003). Cultures were maintained 

on malt extract yeast agar (MYA) (15 g/L malt extract, 2 

g/L yeast extract, 15 g/L agar). Cultures are stored in the 

culture collection (CMW) of the Forestry and Agricultural 

Biotechnology Institute (FABI), University of Pretoria and with 

the Centraalbureau voor Schimmelcultures (CBS), Utrecht, 

Netherlands. 

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Fungal phoenix 

Molecular methods 

Isolates for DNA extractions were grown at 25 °C in the dark for 

3 wk in conical flasks containing liquid malt extract yeast (MY). 

Mycelium was harvested using a tea strainer, freeze-dried and 

lyophilised. DNA extractions followed the protocol of Coetzee 

et al. (2000b). A NanoDrop spectrophotometer (Thermo Fisher 

Scientific, USA) was used to quantify the DNA. The IGS-1 and 

ITS regions of the rDNA operon were amplified using primer 

pairs P-1 / O-1 and ITS-1 / ITS-4, respectively. PCR reaction 

mixture and conditions were the same as those published 

by Coetzee et al. (2000b), except that FastStart Tag DNA 

polymerase was used instead of an Expand High Fidelity PCR 

System. The PCR products were purified prior to sequencing 

using a MSB® Spin PCRapace kit (Invitek, Germany). DNA 

sequences for the IGS-1 and ITS-1 regions were obtained 

in both directions using the same primers employed for their 

amplification. The sequence reactions were carried out using 

an ABI PRISM Dye Terminator Cycle Sequencing Ready kit 

with AmpliTaq DNA Polymerases FS (Applied Biosystems) 

following the manufacturer’s instructions. Chromatographs 

were analysed and contigs assembled in CLC Main Workbench 

v. 5.7 (CLC bio, Denmark). 

DNA sequence comparisons and phylogenetic 

analyses 

DNA sequences were compared against those available in 

the NCBI GenBank database using a BlastN search. IGS-1 

and ITS sequences generated in this study were aligned 

against those of isolates CMW 3975 and CMW 3978 available 

on GenBank and originating from the Company Gardens in 

Cape Town. This was done to determine nucleotide variation 

between the isolates from the Company Gardens and those 

from Kirstenbosch Botanical Gardens. 

Phylogenetic analysis was conducted with a sub-set of 

the ITS-1 dataset generated by Coetzee et al. (2003). The 

dataset was amended with DNA sequences for A. fuscipes 

from South Africa and A. mellea from Europe, Asia, western 

USA, eastern USA and the Company Gardens, South Africa. 

Sequences were re-aligned using MAFFT v. 6 (Katoh & Toh 

2008). Cladograms were generated using a heuristic tree 

search algorithm in PAUP v. 4 with branch swapping set 

to TBR and random addition of sequences (10 replicates). 

Trees were rooted to A. fuscipes. Bootstrap analysis (1000 

replicates) was done to gain support for the grouping of 

taxa using the same settings as above but with addition of 

sequences set to nearest. 

RESULTS 

Isolates 

The macro-morphology of basidiocarps produced by the 

fungus was similar to that described for A. mellea (Watling et 

al. 1982) (Fig. 1D–G). The cap colour of the basidiocarps was 

distinctly yellow and they had thick annuli and stipes tapering 

towards the base. The basidiocarps also had a caespitose 

growth habit typical of A. mellea. 

Two isolates (CMW 36264 and CMW 36265) were 

retrieved from infected roots and these produced rhizomorphs 

typical of Armillaria spp. in culture (Fig. 1H). The rhizomorphs 

displayed a dichotomous growth habit and were produced 

in abundance. White aerial mycelium was observed on the 

surface of the rhizomorphs at areas that had grown out of 

the medium. 

DNA sequence comparisons and phylogenetic 

analysis 

The IGS-1 and ITS DNA sequences of isolates from 

Kirstenbosch were most similar to sequences of A. mellea 

in GenBank. Comparisons of IGS-1 and ITS sequences 

revealed the absence of nucleotide variation between 

isolates from Kirstenbosch and A. mellea from the Company 

Gardens. 

The ITS dataset included 925 characters of which 183 

characters were parsimoniously informative. A heuristic 

search generated 6 trees with tree lengths of 250 steps (Fig. 

2). The consistency index was 0.864, and retention index 

0.941. The isolates from Kirstenbosch formed a monophyletic 

group with A. mellea from the Company Gardens with strong 

bootstrap support and together these were placed in a clade 

that included sequences of A. mellea from Europe (99 % 

bootstrap support). 

DISCUSSION 

Sporocarps linked to infected roots from which cultures were 

made in this study were morphologically similar to those of A. 

mellea previously found in the Cape Town city centre. DNA 

sequence comparisons also showed that the cultures were 

those of A. mellea and the sequences were identical to those 

from the Company Gardens. Although vegetative compatibility 

tests were not done to test whether these represent the same 

clone as those in the City Centre, there was no IGS-1 or ITS 

nucleotide variation between isolates from the two locations 

and they most likely are the same. 

There are three possible means of introduction of A. 

mellea into Kirstenbosch Botanical Gardens, via air-dispersed 

basidiospores, on infected plant material or on infested wood 

mulch. Armillaria mellea in the Company Gardens fruits 

profusely every year at the onset of the first rains in autumn. 

Although the fungus clone is entirely surrounded by roads 

and buildings, the basidospore cloud is likely to easily spread 

within the city and at least up the foothills of Table Mountain, 

on which Kirstenbosch is situated. While A. mellea might 

have been introduced into Kirstenbosch separately to that of 

the clone found in the Company Gardens and as A. gallica 

must have been, it would perhaps more easily have spread to 

this nearby location via basidiospores. One further possible 

route of introduction to consider relates to the cultivation 

practices used in the garden. Flowerbeds and paths are 

frequently covered with wood and bark mulch. As Armillaria 

spp. are common wood rotting fungi, it is possible that A. mellea 

was introduced into the Gardens through this substrate. 

ARTICLE 

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ARTICLE 

Fig. 2. Cladogram generated from ITS DNA sequence data. Bootstrap values are indicated above the tree branches. The year during which 

isolates from Kirstenbosch and the Company Gardens were reported are indicated next to the taxon name. Isolates obtained during this study 

are shown in the rectangle. 

Armillaria sporocarps have not previously been found in 

Kirstenbosch. This may simply be related to the fact that they 

are ephemeral and have not been present when mycologists 

or plant pathologists might have been visiting the botanical 

garden. When these sporocarps were discovered, they were 

relatively widespread and all were morphologically similar. 

The infected roots from which isolates were made were 

also from a number of locations, none of which had been 

associated with the infection by A. gallica. It is possible that 

A. gallica also fruits in the garden, but at a different time to A. 

mellea, or it is less prone to fruiting. Regular observations will 

be needed to resolve this question. 

Peripheral surveys of the native forest on the foothills of 

Table Mountain and that extending out of the Kirstenbosch 

Botanical Gardens have not revealed evidence of Armillaria 

root rot. The fact that A. mellea is able to fruit profusely in 

the gardens suggests that it may spread to native forests 

in the vicinity (Fig. 1A) and more careful surveys should be 

undertaken to determine whether this is already occurring. 

Certainly this invasive alien fungus has the capacity to result 

in serious disease problems in the native environment as 

has been true with the introduced invasive Phytophthora 

cinnamomi on Leucodendron argenteum (Silver Trees) in 

Kirstenbosch (van Wyk 1973, Linde et al. 1997). This potential 

risk to Kirstenbosch and the native forest associated with it 

deserve consideration. 

The Dutchman Jan Van Riebeeck was the founder and first 

commander of Cape Town between 1652 and 1662. One of his 

tasks was to establish a vegetable and fruit garden to provide 

ships of the Dutch East India Company sailing between The 

Netherlands and East Asia with fresh produce and to offset 

serious problems due to vitamin C deficiency [for a fascinating 

account of the ship’s surgeons of the Dutch East India Company 

see Bruijn (2009)]. This is the origin of the Company Gardens 

and the historic avenue of oak (Quercus robur) trees that 

line Government Avenue, the death of which prompted the 

discovery of A. mellea in Cape Town (Coetzee et al. 2001). 

At the time of this discovery, popular press took an interest in 

the problem () and referred to the 

tragic death of historic trees as “Van Riebeeck’s curse”. The 

appearance of A. mellea fruiting profusely in Kirstenbosch, 

another historic garden of great national importance, suggests 

that the fungal “Van Riebeeck’s curse” remains not only present 

but is growing in importance. It further illustrates the devastating 

impact that invasive alien pathogens can have on natural woody 

ecosystems many years after their introduction. 

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Fungal phoenix 

ACKNOWLEDGEMENTS 

We thank the Department of Science and Technology (DST)/ National 

Research Foundation (NRF) Centre of Excellence in Tree Health 

Biotechnology (CTHB) for funding that made this study possible. 

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Modelling fungal colonies and communities: challenges and opportunities 

Ruth E. Falconer 1* , James L. Bown 2 , Eilidh McAdam 1 , Paco Perez-Reche 3 , Adam T. Sampson 2 , Jan van den Bulcke 4 and Nia A. 

White 1 

1 

SIMBIOS Centre, University of Abertay Dundee, DD1 1HG, Dundee Scotland UK; corresponding author e-mail: r.falconer@abertay.ac.uk 

2 

Institute for Arts, Media and Computer Games, University of Abertay Dundee, DD1 1HG, Dundee Scotland UK 

3 

Department of Chemistry, University of Cambridge, Lensfield Road, Cambridge, CB2 1EW, UK 

4 

Laboratory of Wood Technology, Department of Forest and Water Management, Ghent University and UGCT, University Ghent Centre for X-ray 

Tomography, Ghent University, Coupure Links 653, BE-9000 Gent, Belgium 

ARTICLE 

Abstract: This contribution, based on a Special Interest Group session held during IMC9, focuses on physiological based 

models of filamentous fungal colony growth and interactions. Fungi are known to be an important component of ecosystems, 

in terms of colony dynamics and interactions within and between trophic levels. We outline some of the essential components 

necessary to develop a fungal ecology: a mechanistic model of fungal colony growth and interactions, where observed 

behaviour can be linked to underlying function; a model of how fungi can cooperate at larger scales; and novel techniques 

for both exploring quantitatively the scales at which fungi operate; and addressing the computational challenges arising from 

this highly detailed quantification. We also propose a novel application area for fungi which may provide alternate routes for 

supporting scientific study of colony behaviour. This synthesis offers new potential to explore fungal community dynamics 

and the impact on ecosystem functioning. 

Key words: 

foraging 

fungal growth 

interactions 

invasions 

mycelia 

networks 


Introduction 

Fungi are a central component of the biosphere, essential for 

the growth of over 90 % of all vascular plants (Allen 1993), 

play an essential role in ecosystem services (Boumans 

2002) and it is estimated that there may be as many as 1.5 

million species of fungi globally (Hawksworth 1991). Fungi 

can impact on the outcome of plants and their enemies 

(Bennett et al. 2006) and are a life support network for most 

plants (Bardgett et al. 2006). Van der Heijden et al. (1995) 

stress the importance of understanding the structure and 

function of fungal communities is an important contributor to 

the maintenance of plant biodiversity. Development of such 

a fungal ecology requires an understanding of the spatiotemporal 

growth and interaction dynamics both within fungal 

communities and between fungi and plant systems. We 

therefore propose that an essential step in understanding the 

ecology of fungi is to combine: (1) a physiologically-based 

model of fungal community dynamics, capturing colony 

growth and interactions, in heterogeneous environments; 

(2) non-destructive quantification of community growth 

patterns through instrumentation; (3) models linking fungal 

communities to plant systems; and (4) next-generation 

computational approaches to simulate complex systems at 

scales consistent with that instrumentation. Here, we report 

on a Special Interest Group meeting held during IMC9 that 

considered this agenda and the four components needed to 

progress our understanding of fungal ecology. Additionally 

the Group considered the notion of fungi as a biological 

metaphor for complex management problems. 

Modelling fungal community 

dynamics 

In Falconer et al. (2005, 2007, 2008) we demonstrated the 

use of a physiologically-based model to explore the factors 

that influence the nature of fungal community diversity and 

the link between individual behaviour and the structure and 

function of fungal communities. The model is individual-based 

and incorporates the essential physiological processes of 

nutrient absorption, within colony biomass transport and 

recycling, inhibitor production and growth, and these occur 

differentially within a single mycelium as a consequence 

of local and non-local context. This differential behaviour 

permits different parts of the mycelium to expand and senesce 

concurrently. This framework was developed to capture the 

minimal set of physiological processes required to reproduce 

the observed range in phenotypic response in real colonies: 

uptake, redistribution of biomass, remobilisation of biomass, 

and growth which are known to be important for vegetative 

growth of fungi but have not collectively been incorporated 



Attribution: 






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Falconer et al. 

ARTICLE 

into previous modelling frameworks (Falconer et al. 2005). We 

have also investigated the consequences of environmental 

heterogeneity for biomass distribution (Falconer et al. 2007), 

identifying which trait sets allowed individuals to persist in 

given environmental contexts. The model has been used 

to explore the effect of different soil management strategies 

on fungal invasion and interactions (Fig. 1; Kravchenko et 

al. 2010). The enhancement of this model to incorporate 

inhibitor production that impacts inter-colony interactions is 

described in Falconer et al. (2008). The model was used to 

generate mycelial distribution maps that emerge from fungal 

interactions among a community of intrinsically different 

individuals (Falconer et al. 2010). This is the first attempt to 

model (physiologically) the dynamics of a fungal community 

in terms of a fungal ecology. We introduced the concept of 

a biomass-based abundance distribution function, described 

the form of that curve, and made the first attempt to identify the 

traits that affect the form of that curve. Ongoing developments 

are to apply the model to soil systems to understand the 

effect of physical and chemical processes on fungal diversity. 

It has been shown by experiments that the fungal colony 

exhibits a remarkably complex cooperative behaviour (Ritz 

1995, Hughes & Boddy 1996), and a linked experimentaltheoretical 

approach by Bown et al. (1999) demonstrated 

that community scale dynamics are a consequence of nonindependent 

local interactions. In our development of fungal 

ecology, we must also consider such cooperation, and here 

we consider as an example fungal pathogen invasion. 

Linking fungal communities to plant 

systems 

In Perez-Reche et al. (2010), we used probabilistic models 

to determine how cooperation at the individual scale led to 

epidemic spread at the community scale. To investigate this 

phenomenon we constructed a model of fungal invasion that 

is spatially explicit and considers heterogeneous, discrete 

resource distribution. In general, the transmission of the 

fungus from a colonised donor host (d) to a healthy recipient 

host (r) does not only depend on the d-r pair but it is influenced 

by the environment of the d-r pair. On the one hand, the rate of 

transmission of the pathogen may be enhanced (constructive 

synergy) if the fungus is using resources from several 

colonised hosts. On the other hand, the rate can be diminished 

(interfering synergy) because of several factors arising from 

the competition between different parts of the colony. We have 

addressed the question of whether synergistic effects occurring 

at the individual scale play an important role in epidemics 

spreading at the community-scale. We have also investigated 

the effect of synergy on properties such as the foraging strategy 

followed by the pathogen, the probability of epidemic invasion, 

and the efficiency of invasion. The approach is based on an 

extension of a spatial model for SIR (susceptible-infectedremoved) 

processes (Grassberger 1983) to incorporate 

synergistic effects in the transmission rate between pairs of 

hosts depending on the number of infected neighbours to the 

pair. Analysis of the model by means of numerical simulations 

has shown that synergy at the host level has non-trivial 

consequences at the population level. The foraging strategy 

of the pathogen changes from being explorative for interfering 

synergy to being exploitative for constructive synergy. The 

invasion in the exploitative regime is temporally more efficient, 

i.e., it is quicker, than invasion using an explorative strategy. 

However, explorative epidemics are spatially more efficient 

than exploitative epidemics because they can lead to invasion 

by infecting fewer hosts. The modelling carried out so far is 

based on simple assumption such as equal intrinsic infectivity 

and susceptibility for all the hosts in the population. Extensions 

of the model to account for heterogeneity in transmission of 

infection and perhaps other factors will be essential to provide 

quantitative predictions for possibly invasive epidemics in real 

populations. While it is possible to validate models of infection 

spread because the domain may be directly observed, such 

as in infected plants where the number of lesions can be 

determined via direct or indirect methods (Jeger 1987), this 

is much more challenging for opaque soil and wood systems. 

For these systems, in order to obtain the experimental data 

for model calibration and validation information regarding 

the spatial distribution and biomass amounts is required. 

One technique that has been used to determine the physical 

architecture of the soil and wood systems is X-ray Computed 

Tomography and progress is being made in quantifying and 

visualising fungal biomass in situ. 

Non-destructive quantification of 

community growth patterns 

Fig. 1. Effect of soil structure on fungal invasions and interactions 

for a single soil management practice. Blue and red isosurfaces 

correspond to the two boundaries of different fungal species. 

X-ray computed tomography enables a non-destructive 

view of the internal structure of an object and is therefore 

an extremely valuable technique in many research fields. 

The continuously improving performance of equipment, 

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rapidly increasing computing power, and faster algorithms 

for reconstruction and data processing make large volume 

scanning at high resolutions feasible. The state-of-the-art 

equipment at the UGCT (Centre for X-ray Tomography at 

Ghent University) is highly flexible, with in-house developed 

software for scanner control, sample reconstruction, 

analysis, and visualisation. This set-up allows scanning 

with a resolution of 0.2 mm for samples of 37 cm in 

diameter down to approximately 400 nm for objects about 

the size of a splinter. As such, apart from visualisation, 3D 

quantitative information can be retrieved from objects with 

a broad range of sizes. Sub-micron resolution scanning 

should enable the visualisation of fungal hyphae and by 

using time-lapse tomography the growth of these tubular 

structures could be monitored (van den Bulcke et al. 2009). 

The latter procedure however has associated challenges. 

First, fungal growth can interfere with scanning during 

moderately long scan times. Second, with lab-based X-ray 

sources, polychromatic X-rays, scattering, fluorescence and 

noise disturb the ideal acquisition (Vidal et al. 2005). Third, 

at sub-micron resolution phase contrast emerges especially 

at sharp edges, complicating thresholding and segmentation. 

Fourth, tube shift during long scans at sub-micron resolution 

can reduce image quality. Fifth, hyphal tubes are hollow 

thin-walled structures, as such having a very low X-ray 

attenuation. A drastic solution to some of the problems is the 

use of synchrotron radiation, having a monochromatic X-ray 

bundle, allowing faster scanning with less heating of the 

samples, but access to such facilities is a major bottleneck. 

Especially the available beam time is limited and as such 

this is not an option for long-running experiments, of the 

order of days to weeks, and for repeated experiments. Many 

of the aforementioned problems are handled at the UGCT 

facility. Post-processing can contribute to the enhancement 

of image quality; the phase contrast phenomenon can 

be solved using dedicated filtering (Boone et al. 2009, De 

Witte et al. 2009); and tube shift can be counteracted with 

correction software. Proper scanning and processing can 

result in the visualization of fungal hyphae as illustrated in 

Fig. 2, obtained after scanning of a piece of Pinus sylvestris 

subjected to white-rot. In order to study pigmented species 

with rather large hyphal structures, such as Aureobasidium 

pullulans (van den Bulcke et al. 2008), visualization is easier 

due to X-ray interference of the pigment. Apart from individual 

hypha tracking, processing of X-ray volumes should enable 

the quantification of the effects of material degradation on 

different spatial scales, which might be an important concept 

to implement a degradation monitoring system. With the 

existing scanners, frequent scanning and quantification 

of degradation or hyphal biomass on a larger spatial scale 

will be a very valuable tool for non-destructive time-lapse 

analysis. Advanced algorithms implementing X-ray physics 

during reconstruction will increase image quality, whereas 

more advanced image processing code will improve 

quantitative results. The field of X-ray tomography, both hardand 

software, is rapidly evolving and therefore is promising 

for in situ fungal monitoring and quantification in wood and 

perhaps soil systems in the near future, in addition to other 

modalities such as confocal laser microscopy (Hickey et al. 

2005) and magnetic resonance imaging (Müller et al. 2002). 

Next-generation computational 

approaches 

Our ability to exploit the experimental advantages described 

above is currently constrained by the limited scales at which 

existing simulation technologies are able to operate. For 

example, in spite of data at larger scales, in Falconer et al. 

(2010) we use a domain size for the soil/fungal interactions 

of approx 1 cm 3 with a voxel resolution of 30 microns; for 

predictions to be useful we need to work at, at the very 

ARTICLE 

Fig. 2. (a) Three dimensional 

rendering of hyphal tubes of a white 

rot fungus winding around a small 

piece of Pinus sylvestris in contact 

with malt extract agar (filling some 

cell lumina). (b) Cross-sectional and 

(c) longitudinal view illustrating the 

high anatomical detail. Bar = 200 

µm; voxel size 0.79 µm. 

v o l u m e 1 · n o . 2 

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Falconer et al. 

ARTICLE 

least, core scale (10 cm 3 ). We need simulations to operate 

at realistic scales in order to accurately reproduce the 

observed, often emergent behaviours we wish to study. 

Emergent behaviours are often scale-dependent, a simulation 

that includes only a restricted region of the system may 

demonstrate different emergent behaviours, and we may not 

know in advance what scale is appropriate. One approach to 

scaling up simulations is to simplify the underlying model, for 

example the homogenisation approach proposed by Roose 

& Schnepf (2008), where the environmental heterogeneity is 

carefully coarse-grained, and by model reduction, for example 

Gibbons et al. (2009), where stripping out unnecessary model 

components reduces computational demands. Both raise 

considerable difficulties. In each case, simplification requires 

identification of the important model components, which are 

unlikely to be obvious in a complex system. An alternative 

approach is to increase the computational power available 

to the simulation by taking advantage of multicore CPUs, 

GPUs and clusters. Parallelisation is widely regarded as an 

experts-only programming problem, and one that is strongly 

tied to the particular computational platform in use. However, 

complex biological systems simulation is a problem with an 

inherently high degree of concurrency in the natural system: 

a complex system fundamentally consists of a large number 

of independent, but interacting, agents and processes. Most 

approaches to parallel programming focus on highly regular 

numerical problems, and make building such a simulation 

difficult. Simulation becomes much more straightforward 

with the use of concurrent programming techniques, in which 

the concurrent activities in a system and their relationships 

are specified, and the execution of those activities in the 

most efficient manner across the available resources is 

managed automatically by software. With careful design, 

a simulation built this way can be truly scalable, meaning 

that its complexity can be increased in a near-linear fashion 

by dividing it across more computational resources. This 

approach has become particularly interesting with the rise of 

grid and cloud computing: researchers can now gain access 

as required for a short period of time to a very large number 

of nodes upon which to execute their simulation, rather than 

relying upon in-house resources. Using cloud resources, 

we can potentially scale up by a few orders of magnitude. 

Approaches to scalability and validation in complex systems 

simulation are currently being investigated by CoSMoS 

(www.cosmos-research.org), drawing on expertise in 

modelling, highly-dependable software engineering and 

concurrent programming to develop and document reusable 

techniques for complex systems modelling and simulation. 

CoSMoS techniques for parallel, distributed simulation of 

agent-based spatial models have been successfully applied 

to problems including those in the fields of immunology (insilico 

experimentation with lymphocyte migration (Andrews et 

al. 2008) and granuloma formation (Flügge et al. 2009)) and 

mycology (scaling up of the Falconer et al 2005 model, in 

preparation). 

Fungi as a metaphor 

Fungal colonies are a highly successful organism, 

demonstrating pervasive growth through harsh environments. 

They achieve this through their capacity to operate in a 

decentralised manner, reacting locally to changes in context 

while interoperating at the colony scale and with other 

organisms. Fungi have the capacity to recycle biomass, 

effecting dynamic reallocation of investment, capitalise on 

fluxes in available resource and self-heal. These properties 

make them attractive metaphors for managing large, 

complex, distributed artificial networks. Other researchers 

have used other biological metaphors for similar problems. 

For example, ant colonies have been used as a metaphor for 

telecommunication routing algorithms. Typically, stigmergic 

pheromone trails are used to profile rates of flow of ants 

and other network traffic and the local strength of the trail 

is coupled to the values in local routing tables (e.g. Di Caro 

& Dorigo 1998). Similarly, in the field of artificial immune 

systems, the properties of the immune system have been 

used to inspire solutions in the areas of anomaly detection 

and data mining (Timmis et al. 2008). We have been 

exploring the potential for fungi as a metaphor for protecting 

society’s critical infrastructures. Modern societies are heavily 

dependent upon a number of critical infrastructure networks 

that allow our societies to function, including water, power 

and transportation. These networks are open to failure 

through a range of processes including shortage of essential 

resources, breakdowns at key nodes and surges in demand 

and this means that effective management of such networks 

is challenging (Schulman et al. 2004). Bebber et al. (2007) 

recognise that understanding how fungi grow may inform the 

design of man-made networks and through image analysis, 

have characterised fungal colony growth patterns in terms of 

nodes and edges of a graph. They show that fungal colonies 

are efficient transport networks that are robust to damage 

and react to local variation in resource. In order to translate 

the concept into a working solution, we have developed a 

graph-based implementation of Falconer et al. (2005). We 

are now exploring the capacity of our model to provide robust 

and resilient management solutions to resource limitations, 

node failures, and demand surges. 

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Colletotrichum: species, ecology and interactions 

Ulrike Damm 1 , Riccardo Baroncelli 2 , Lei Cai 3 , Yasuyuki Kubo 4 , Richard O’Connell 5 , Bevan Weir 6 , Kae Yoshino 7 and Paul F. 

Cannon 8 

1 

CBS-KNAW Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands; corresponding author e-mail: u.damm@cbs. 

knaw.nl 

2 

School of Life Sciences, University of Warwick, Wellesbourne, Warwick, CV35 9EF, UK 

3 

Key Laboratory of Systematic Mycology & Lichenology, Institute of Microbiology, Chinese Academy of Sciences, No. 10, North 4 th Ring Road 

West, Beijing 100190, P.R. China 

4 

Graduate School of Life and Environmental Sciences, Kyoto Prefectural University, 1-5 Hangi-cho, Shimogamo, Sakyo-ku, Kyoto, Japan 

5 

Department of Plant Breeding and Genetics, Max Planck Institute for Plant Breeding Research, 50829 Köln, Germany 

6 

Landcare Research, Private Bag 92170 Auckland, New Zealand 

7 

Graduate School of Agriculture, Kyoto University, Sakyo-ku, Kyoto 606-8502, Japan 

8 

CABI Europe-UK, Bakeham Lane, Egham, Surrey TW20 9TY, UK; and Royal Botanic Gardens, Kew, Surrey TW9 3AB, UK 

ARTICLE 

Abstract: The presentations of the Special Interest Group meeting Colletotrichum: species, ecology and 

interactions, held on 1 August 2010 during IMC9 in Edinburgh, UK, are outlined. Seven research projects, 

ranged from systematics and population genetics to host-pathogen interactions and genome projects were 

presented. The meeting revealed that currently major species complexes in the genus Colletotrichum are 

being revised and the identities of many pathogens clarified on the basis of molecular phylogenies, and that 

the genomes of four species are sequenced and decoded providing an enormous amount of data that are 

used to increase our understanding of the biology of Colletotrichum species. 

Key words: 

genome sequencing 

host-pathogen interaction 

identification 

pathogenicity 

population genetics 

systematics 


INTRODUCTION 

A Special Interest Group meeting Colletotrichum: species, 

ecology and interactions, was held on 1 August 2010 at the 

9 th International Mycological Congress (IMC9) in Edinburgh, 

UK. The meeting, organised by Paul Cannon (UK) and Ulrike 

Damm (The Netherlands), brought together 23 scientists 

from 12 countries working in different fields of mycology, but 

with a common interest in the genus Colletotrichum. Seven 

presentations, covering a wide range of topics, ranged 

from systematics and population genetics to host-pathogen 

interactions and genome projects. This contribution provides 

a synopsis of the presentations made at that meeting. 

Systematics and identification 

The first four presentations dealt with systematics and 

identification of major Colletotrichum species complexes 

containing various important anthracnose pathogens 

worldwide. Four of these species are illustrated in Fig. 1. 

While the identity of many important species still require 

revision (Hyde et al. 2009), molecular techniques improve the 

delimitation of species that are hard to distinguish based on 

morphology alone and reveal their phylogenetic relationships 

(Cai et al. 2009, Crouch et al. 2009, Damm et al. 2009). This 

will inevitably result in name changes but has implications 

for everyone working with this genus, especially plant 

pathologists, and will improve our understanding of the role 

these species play in nature. 

Ulrike Damm gave an overview of her ongoing 

collaboration with Paul Cannon about the phylogeny of three 

species complexes. The aim of this project is to delimitate 

species within these complexes, characterise known and 

new species and designate epitypes to provide the basis for 

accurate identifications of Colletotrichum species. This goal 

has so far been achieved for species with curved conidia from 

herbaceous hosts (Damm et al. 2009), which in the past were 

mostly identified as C. dematium. Multi-gene analyses and 

morphological characterisation revealed several diverse and 

distantly related species, including four new species. Seven 

species were epitypified, including C. dematium and the type 

species of the genus, C. lineola. A second study confirmed 

most of the previously recognised groups (Sreenivasaprasad 

& Talhinhas 2005) within the C. acutatum species complex. 

Most of these could be defined on the basis of type strains or 

strains suitable for epitypification. Literature reports (Lubbe et 

al. 2004, Johnston et al. 2005) and preliminary studies using 

ITS sequence data indicated that C. boninense represents a 

species complex as well. A multilocus molecular phylogenetic 



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Fig. 1. Conidiomata and conidia of four Colletotrichum species. A, E. C. acutatum (CBS 112996, ex-paratype strain). B, F. C. lineola (CBS 

125337, ex-epitype strain). C, G. C. truncatum (CBS 151.35, ex-epitype strain). D, H. C. gloeosporioides (CBS 112999, ex-epitype strain). A, C, 

F, H. Conidia on SNA. B, G. Conidiomata on SNA. D, E. Conidiomata on Anthriscus stem. Scale bars: A = 10 μm, B = 100 μm, E = 200 μm. A 

applies to A, C, F, H. B applies to B, D, G. 

analysis of strains previously identified as C. boninense 

resulted in clades that could be recognised as separate 

species with differences in host range, distribution and 

morphology, including C. boninense sensu stricto, Glomerella 

phyllanthi, C. hippeastri and several presumably new species. 

Most of the species in the C. boninense complex and some 

in the C. acutatum species complex form teleomorph states 

in culture. Publications on the C. acutatum and C. boninense 

species complexes will appear in a 2011 issue of Studies in 

Mycology. 

Colletotrichum gloeosporioides sensu lato is a species 

complex with broad genetic and biological diversity grouped 

together by similar conidial morphology and ITS sequences. 

Bevan Weir and Peter Johnston (Landcare Research, 

Auckland, New Zealand) presented their research on this 

species complex and possible approaches to species 

delimitation through the Genealogical Concordance 

Phylogenetic Species Recognition (GCPSR). It was shown 

that that the taxa C. musae, C. kahawae, C. xanthorrhoeae, 

C. nupharicola, C. fragariae, C. gloeosporioides sensu 

stricto, C. horii, C. theobromicola, C. ignotum, C. tropicale, C. 

asianum, C. siamense, C. fructicola and C. hymenocallidis, 

as well as many putative undescribed species are part of 

the C. gloeosporioides sensu lato complex. They recently 

characterised and neotypified one of these species, C. horii 

(Weir & Johnston 2010). The GCPSR concept was used 

to delimit taxa within C. gloeosporioides sensu lato. This 

concept considers that phylogenetic trees of different genes 

show discordance within a species due to gene flow between 

individuals. The common node where different gene trees 

show concordance is considered the speciation point. 

They applied the GCPSR with eight genes using recently 

developed Bayesian analysis tool, BUCKy (Ané et al. 2007). 

The GCPSR concept worked well for species delimitation 

along currently recognised lines, except for C. kahawae, 

which was insufficiently distinct from several genetically 

similar non-Coffee Berry Disease causing taxa. It was 

suggested that this may be due to the recent emergence 

(1920) of C. kahawae as a pathogen and that insufficient 

time had passed for ecological niche specialisation to show 

as mutations in the genes used. They suggested that C. 

kahawae be recognised at the subspecific rank. A publication 

on their work on the C. gloeosporioides sensu lato species 

complex will appear in the 2011 Studies in Mycology issue 

on Colletotrichum as well. 

Colletotrichum acutatum causes economically significant 

losses of temperate, subtropical and tropical crops. Globally, 

C. acutatum populations display considerable genotypic 

and phenotypic diversity. Riccardo Baroncelli (University 

of Warwick, Wellesbourne, UK) presented his research on 

evolutionary relationships in C. acutatum populations in 

collaboration with Charles Lane (FERA, Sand Hutton, York, 

UK) and Prasad Sreenivasaprasad (University of Warwick, 

Wellesbourne, UK). The overall objective is to understand the 

evolutionary relationships within the species with particular 

reference to the pathogen populations associated with the 

strawberry production systems in the UK. More than 150 

C. acutatum isolates related to different hosts worldwide 

have been assembled. Phylogenetic analysis of sequence 

data from the rDNA block, Mat1-2 and βtubulin-2 genes 

shows eight distinct genetic groups within C. acutatum. The 

subsets of isolates represented within these genetic groups 

corresponded to the previously identified groups A1 to A8. 

Almost all of the homothallic isolates capable of sexual 

reproduction comprise a single genetic group, A7. Isolates 

representing populations capable of heterothallic sexual 

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reproduction belong to two distinct genetic groups A3 and 

A5. Molecular characterisation of C. acutatum populations 

representing the introduction and spread of the pathogen 

in the strawberry production systems in the UK showed the 

presence of three genetic groups (A2, A3 and A4). Their 

results suggest the existence of C. acutatum populations 

potentially undergoing speciation processes, related to their 

reproductive behaviour and host association patterns. Further 

molecular and phenotypic characterisation is in progress. 

Lei Cai (Key Laboratory of Systematic Mycology & 

Lichenology, Institute of Microbiology, Beijing, China) 

and Kevin Hyde (School of Science, Mae Fah Luang 

University, Chiang Rai, Thailand) presented their research 

on Colletotrichum species from Asian fruits and leaves. 

Fruit rots (anthracnose) were previously often attributed to 

C. gloeosporioides and C. acutatum. Identifications were, 

however, based on morphological characters or, if gene 

sequence data were used, comparisons were often made 

with wrongly applied names. Colletotrichum gloeosporioides 

was recently epitypified (Cannon et al. 2008) so that living 

cultures and sequence data are, for the first time available 

for comparison with fresh collections. Analysis of sequence 

data of 25 isolates (selected from 140 obtained strains based 

on diversity of host and morphology) from eight tropical fruits 

are compared with the C. gloeosporioides epitype. Contrary 

to previous assumptions, none of these isolates from tropical 

fruits was C. gloeosporioides sensu stricto (Phoulivong et 

al. 2010). The five gene regions used in this study resolved 

C. asianum, C fructicola, C. horii, C. kahawae and C. 

gloeosporioides in the C. gloeosporioides species complex 

as distinct phylogenetic lineages with high statistical support. 

Many tested strains could not be assigned to any known taxa 

in this analysis. They also reported Colletotrichum species 

from Amaryllidaceae, Orchidaceae, Cordyline fruticosa and 

Jasminum sambac, with the latter including two new species 

(Wikee et al. 2010), and updated the typifications of C. 

coccodes, C. falcatum and C. musae. 

Genomic studies 

Colletotrichum species provide excellent models for studying 

fungal-plant interactions (Perfect et al. 1999). Several largescale 

genome projects are in progress for Colletotrichum 

species aiming to produce high-quality assemblies of the 

genome sequences to provide resources for comparative 

genomics and the molecular analysis of fungal pathogenicity, 

which allows the identification of genes and proteins relevant 

to each stage of plant infection. 

Colletotrichum graminicola is a destructive pathogen of 

maize, causing stalk rot and leaf blight, while C. higginsianum 

attacks many cultivated forms of Brassica as well as 

Arabidopsis thaliana, providing a model pathosystem in which 

both partners can be genetically manipulated (O’Connell et 

al. 2004). Both pathogens employ a hemibiotrophic infection 

strategy, but while the biotrophic phase of C. graminicola 

extends into many host cells, that of C. higginsianum is 

confined to single epidermal cells. Richard O’Connell (Max 

Planck Institute for Plant Breeding Research, Cologne, 

Germany) gave an overview of the C. higginsianum and C. 

graminicola genome research, which he is conducting in 

collaboration with Lisa Vaillancourt (University of Kentucky, 

USA), Li-Jun Ma (MIT-Broad Institute, USA) and Mike Thon 

(CIALE-University of Salamanca, Spain). Comparing the 

genomes of two species with contrasting pathogenic lifestyles 

and host specificities will allow them to study lineage-specific 

expansions and contractions of gene families and identify 

genes undergoing rapid evolution (diversifying selection), 

which may be involved in interactions with the host plant, 

e.g. those encoding secreted effector proteins. The 57.4 Mb 

genome of C. graminicola comprises 13 chromosomes and 

was sequenced at the Broad Institute (8X Sanger, 11X pairedend 

454) giving an assembly of 1,151 contigs in 653 scaffolds 

(). The 52.5 Mb genome of C. 

higginsianum comprises 11 chromosomes and was 

sequenced with a combination of 454 (24X), Illumina pairedend 

(60X) and Sanger fosmid sequencing (0.2X). These data 

assembled into 8,301 contigs in 367 scaffolds (). For 

genome annotation, 890,000 ESTs were generated from C. 

higginsianum by 454-sequencing cDNA libraries representing 

appressoria formed in vitro, appressoria penetrating leaf 

epidermis, biotrophic hyphae and necrotrophic mycelium. 

In addition, ~22,000 Sanger ESTs were obtained from the in 

vitro mycelium of C. graminicola. Both genomes have now 

been annotated by the Broad Institute, with 12,006 proteincoding 

genes predicted for C. graminicola and ~15,900 for 

C. higginsianum. 

Colletotrichum orbiculare is an anthracnose fungus which 

infects Cucurbitaceae. Yasuyuki Kubo (Kyoto Prefectural 

University, Japan) gave an update of the C. orbiculare 

genome project conducted in collaboration with Yoshitaka 

Takano (Kyoto University, Japan) and Ken Shirasu (RIKEN 

Plant Science Center, Yokohama, Japan). Pathogenicity, 

morphology and the hemibiotrophic infection strategy of 

strain 104-T (=MAFF 240422) are well studied, some of the 

rationales to select this strain for the genome sequencing 

project. Moreover, strain 104-T has been very stable for 

pathogenesis and morphogenesis for many years and a 

variety of insertional mutants are available. Gene manipulation 

techniques such as Agrobacterium tumefaciens-mediated 

transformation or protoplast transformation are established 

(Tsuji et al. 2003) and for host parasite interaction studies, 

a model plant Nicotiana benthamiana is being used as a 

susceptible host. So far, several factors involved in infection 

related morphogenesis have been identified in C. orbiculare 

104-T. Signal transduction pathways, such as cAMP 

dependent pathway and MAP kinase pathway are essential 

for germination, appressorium development, infection 

hyphae formation and invasive growth, while the melanin 

biosynthesis pathway is essential for appressorium function. 

Several structural and regulatory genes involved in these 

pathways were identified (Takano et al. 1997, Tanaka et al. 

2009). Peroxisome function is essential for pathogenesis as 

well; and two peroxisome biogenese gens, PEX6 and PEX13 

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were functionally analysed (Kimura et al. 2001, Fujihara et 

al. 2010). Recently, it was reported that pexophagy factor 

ATG26 is essential for appressorium function (Asakura et al. 

2009). The genome analysis of C. orbiculare is now going to 

be completed, which allows comparisons of genomic data of 

three Colletotrichum species, C. higginsianum, C. graminicola 

and C.orbiculare that belong to different phylogenetic clades 

within the genus. These data will provide comprehensive 

basis for studying the biology of the different Colletotrichum 

species. 

A study on host-pathogen interaction between C. 

orbiculare and N. benthamiana was presented by Kae 

Yoshino (Graduate School of Agriculture, Kyoto University, 

Kyoto, Japan). This project is conducted together with Fumie 

Sugimoto (Graduate School of Agriculture, Kyoto University, 

Kyoto, Japan), Hirofumi Yoshioka (Graduate School of 

Bioagricultural Sciences, Nagoya University, Nagoya, Japan), 

Tetsuro Okuno, and Yoshitaka Takano (both Graduate School 

of Agriculture, Kyoto University, Kyoto, Japan). By functional 

screening of C. orbiculare cDNAs using N. benthamiana-A. 

tumefaciens in planta expression system they identified a 

novel effector gene NIS1 whose product induces cell death 

in N. benthamiana. Deletion of the signal peptide of Nis1 

abolished its cell death activity, indicating Nis1 recognition 

in apoplasts of N. benthamiana. The GFP-based localization 

study together with biochemical analysis revealed that Nis1 

with the inducting activity is secreted from C. orbiculare cells at 

fungal invasion phase inside plant tissue. Nis1-mediated cell 

death was cancelled by virus-induced gene silencing (VIGS) 

of SGT1 or HSP90 in N. benthamiana, whereas VIGS of 

RAR1 had no effects. These indicate that SGT1 and HSP90, 

the components of R-gene mediated defences, are involved 

in the Nis1-inducing cell death. Surprisingly, the deletion of 

NIS1 has no significant effects on the infection behaviour of 

C. orbiculare on N. benthamiana. Overexpression of Nis1 in 

Arabidopsis thaliana caused growth reduction, implying that 

the NIS1-encoded effector has an activity to interfere plant 

development. These data indicate that C. orbiculare secretes 

the effector that can be recognized by N. benthamiana, but 

this potential effector-triggered immunity might be repressed 

in this susceptible interaction. 

Alan Buddie (CABI Europe-UK, Egham, UK) announced 

a further genome project for C. gloeosporioides. The project 

is a collaboration between CABI and The Genome Analysis 

Centre (TGAC), Norwich, UK (), to carry 

out a whole genome sequence of the ex-epitype strain of C. 

gloeosporioides (IMI 356878). The DNA sequencing phase is 

nearing completion and they are going to start soon with the 

assembly. 

The meeting provided good evidence of the rapidity 

with which our understanding is improving of Colletotrichum 

genomics, and nicely complemented the outputs of another 

Colletotrichum workshop that was held earlier in 2010, in 

conjunction with the 10 th European Conference on Fungal 

Genetics in Leeuwenhorst, the Netherlands. It was particularly 

exciting to witness the increasing power of genomic research 

tools, and their potential impact on our understanding of fungal 

systematics and speciation. It provided good opportunities to 

coordinate research programmes, exchange data and share 

experiences. 


Alan Buddie (CABI Europe-UK, Egham, UK) is thanked for sharing 

the news about the new Colletotrichum genome project. 

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T, Sakai Y, Takano Y (2009) Atg26-Mediated Pexophagy Is 

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Cryptic species in lichen-forming fungi 

Ana Crespo 1 and H. Thorsten Lumbsch 2 

1 

Departamento de Biología Vegetal II,Facultad de Farmacia, Universidad Complutense de Madrid, E-28040 Madrid, Spain; corresponding author 

e-mail: acrespo@farm.ucm.es 

2 

Department of Botany, The Field Museum, 1400 South Lake Shore Drive, Chicago, Illinois 60605, USA 

ARTICLE 

Abstract: This contribution provides a synopsis of the presentations and discussions during the SIG session on 

cryptic speciation in lichen-forming fungi held during IMC9. In several cases, a re-examination of morphology against 

the background of molecular phylogenetic evidence revealed, sometimes subtle, morphological and/or chemical 

characters, supporting the distinction of particular clades at species level. However, there are also examples of cryptic 

species in which no morphological characters could be identified to distinguish between lineages. Several cases were 

presented in which distinct lineages are correlated with biogeographical patterns. When and how to name cryptic 

species was debated, and the use of terms such as “complex” or “aggregate” commended where the taxa formed part 

of a single lineage. 

Key words: 

Ascomycota 

biogeography 

lichens 

Parmeliaceae 

phylogeny 

species concepts 

Article info: Submitted: 5 November 2010; Accepted: 15 November 2010; Published: 23 November 2010. 

Introduction 

The presentations at the session demonstrated that 

current species recognition in lichen forming-fungi vastly 

underestimates the true number of species. Based on 

phylogenetic and population studies, many cases were 

presented showing that numerous distinct lineages are 

hidden under a single species name. The issues raised 

can be grouped under the following headings: naming 

cryptic species, numbers of cryptic species, recognition 

of cryptic species, supporting species separations, 

and phylogeographic correlations. Collectively, these 

presentations provide a synopsis of the current state of 

knowledge of cryptic speciation in lichen-forming fungi. 

Naming cryptic species 

The recognition and naming of cryptic species from cryptic 

lineages was discussed and approaches and options were 

suggested. Hawksworth (2010) examined different groups 

as foraminifera, plant-pathogenic fungi, insects, and plants. 

The main species concepts were reviewed, and a pragmatic 

concept was proposed, defining a species as “groups of 

individuals separated by inheritable discontinuities and 

which it is useful to give a species name to” (Hawksworth 

1996, 2010). The term cryptic species was circumscribed 

as “populations which are phylogenetically distinct and able 

to reproduce themselves, by sexual means or otherwise, 

but which are distinguished by molecular or other features 

that are either not evident macroscopically or generally 

overlooked” (Hawksworth 2010). 

An increasing number of lichen-forming species are used 

as biomonitors or bioindicators of pollutants, environmental 

disturbance, or ecological continuity. Consequently there was 

the issue of how to proceed when cryptic species or lineages 

are found in taxa used in such studies where identifications 

need to be made quickly during field assessments – and 

access to a modern molecular laboratory is impractical. An 

acceptable way of referring to such groups of species was 

commended by Hawksworth (2010). The term “complex” or 

“aggregate” was supported as used when the populations are 

closely related, i.e. have a recent shared common ancestor. 

This practice is already familiar to and regularly used by 

botanists, citizen scientists, and ecologists dealing with 

complexes in plants, for example the Rubus fruticosus aggr. 

and the Taraxacum officinale aggr. 

In some situations, however, the option of recognizing 

subspecies was suggested as perhaps the most appropriate 

solution, for example in paraphyletic populations (Figs 1 and 

2) such as that of Parmelina pastillifera and P. tiliacea s. str. 

(Núñez-Zapata et al. 2010) (Fig. 1). In contrast, in cases 

where the cryptic taxa are not closely related but a result of 

convergence, i.e. they do not either occupy the same clade or 

have a recent common ancestor, it has to be recognized that 

the “complex” approach could give a misleading impression 

of affinity, as in Parmelina cryptotiliacea (Núñez Zapata et al. 

2010) or lineages in Parmelia saxatilis (Divakar et al. 2010b). 



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Fig. 1. Phylogenetic tree of Parmelina (Parmeliaceae). Majority rule consensus tree based on 18000 trees from B/MCMC tree sampling 

procedure from a combined data set of nuITS rDNA and mtLSU rDNA sequences. Posterior probabilities ≥0.95 in the Bayesian analysis are 

indicated above the branches and MP boostrap values ≥0.75 below branches. Branches with significative support in both analyses are in 

bold. (AU=Austria, CI=Canary Islands, FR=France, GE=Germany, IN=India, IT=Italy, MO=Morocco, SP=Spain, SV=Slovenia, TK=Turkey, TN= 

Tunisia, USA=United States of America). Figure provided by Nuñez-Zapata et al. 

Numbers of cryptic species 

There is a growing body of evi dence that the approach to 

current species recognition in lichenized fungi, which is largely 

based on morphology and chemistry, vastly underestimates 

the number of phylogenetic species. Phylogenetic studies 

repeatedly indicate that numerous distinct lineages can be 

hidden under a single species name (Arguello et al. 2007, 

Baloch & Grube 2009, Grube & Kroken 2000, Kroken & Taylor 

2001, Molina et al. 2004, Wirtz et al. 2008). In a number of 

cases, morphological or chemical differences have been 

inter preted as intraspeci fic variability. Re-examination of 

morphology against the background of a molecular phylogeny 

often reveals, sometimes subtle, and previously overlooked 

or viewed as unimportant, morphological and/or chemical 

characters, supporting the dis tinction of these clades at 

species level (Arguello et al. 2007, Divakar et al. 2005a, 

2005b, Molina et al. 2004, Wirtz et al. 2008). However, there 

are also cases of cryptic species in which no morpho logical 

characters have yet been identified to distinguish distinct 

lineages. In several cases, distinct lineages are correlated 

with distinct bio geographical patterns (Arguello et al. 2007, 

Crespo et al. 2010, Molina et al. 2004, Wirtz et al. 2008). 

Phylogenetic studies identified distinct lineages that occur in 

different geo graphic regions, such as continents. 

Recognition of cryptic species 

The large and increasing number of cryptic lineages detected 

in fungi means that the recognition of these lineages as 

separate taxa is a major issue of current fungal taxonomy 

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Cryptic species in lichen-forming fungi 

those taxa are common. Thus, morphological identification 

of a lichen-forming species, sometimes even a genus, can 

be difficult. Therefore, cryptic taxa have been recognised 

historically in lichens, although not necessarily by that term. 

“The recognition and characterization of cryptic species is 

a burgeoning and exciting activity in current systematics, 

and a major challenge for mycologists of all kinds, not least 

lichenologists” (Hawksworth 2010). Suggestions for when to 

formally recognise species within cryptic lineages that are 

found in molecular studies were discussed (Muggia 2010, 

Pérez-Ortega & Printzen 2010), and a consensus of the session 

was to recognise species formally when the phylogeny was 

unequivocal and other evidence supported their separation, 

whether ultramicroscopic, “new” morphological, ecological 

(Muggia 2010) or geographical (Parnmen et al. 2010) were 

discussed as examples for complementary evidence. 

ARTICLE 

Supporting species separations 

Fig. 2. Parmelina pastillifera (MAF 16473; upper) and P. tiliacea 

(MAF 16632; lower) both showing isidia, but in P. pastillifera they are 

peltate while in P. tiliacea are cylindrical. Bars = 5 mm. 

(Crespo & Pérez-Ortega 2009, Hawksworth 2001). However, 

cryptic species in lichen-forming fungi may be compared to 

fungi with other biologies where morphological characters 

are almost absent, thus the pertinence of using this concept 

in lichens was discussed (Hawksworth 2010, Pérez-Ortega & 

Printzen 2010). Unlike many microscopic fungi, some groups 

of lichens form distinctive macroscopic structures, frequently 

with a foliose or fruticose form, and with easily observable 

phenotypical differences. Despite these structures, the 

plasticity of morphological and chemical characters in these 

fungi results in a relatively high number of lichens, species or 

genera, being “difficult” for identification, often accompanied 

by a frequent lack of generative characters (Divakar et al. 

2010b) or the frequency of homoplasy and convergence of 

characters (Grube & Hawksworth 2007, Muggia 2010, Muggia 

et al. 2010, Parnmen et al. 2010, Divakar et al. 2010b). 

Although only relatively few lichens have yet been 

identified as comprising cryptic species using molecular data 

(Grube & Kroken 2000, Kroken & Taylor 2001, Crespo et al. 

2002, Feuerer & Thell 2002, Printzen et al. 2003, Molina et 

al. 2004, Argüello et al. 2007, Wirtz et al. 2008, Fehrer et al. 

2009, Divakar et al. 2010a), assemblages of morphologically 

similar species where identification remains dubious due to 

variability or ambiguity of key characters used to distinguish 

Recent molecular phylogenies have supported some species 

separations that were previously based on subtle characters: 

for example, Parmelina carporrhizans and P. quercina 

(Argüello et al. 2007, Divakar et al. 2010b), Caloplaca alociza 

and C. albopruinosa (Muggia 2010). It is also frequently found 

that distantly related major lineages show a surprising degree 

of morphological convergence. Examples of this phenomenon 

can be found within large families such as Parmeliaceae . 

For example, Parmelina and Austroparmelina were recently 

separated as independent genera based on geography and 

phylogeny. However, all species of Austroparmelina were 

previously included in concept of the genus Parmelina (Crespo 

et al. 2010, Divakar et al. 2010b). Also there are examples in 

microlichens, as in Capnodiales where the morphologically 

similar genera Racodium and Cystocoleus belong to 

independent lineages in recent phylogenetic studies (Muggia 

et al. 2008, Muggia 2010). 

Phylogeographic correlations 

A number of lichen-forming species were historically thought 

to have wide distributions, including cosmopolitan and 

pantropical species. However, while that may be so for some 

species, molecular analyses have repeatedly demonstrated 

that many lineages can be hidden under a similar morphology. 

Several examples were discussed in the symposium (Divakar 

2010, Muggia 2010, Parnmen et al. 2010). Divakar et al.. 

(2010) also found a correlation between reproductive modes 

and distribution patterns. In fertile species, cryptic lineages 

were frequently found, and geographically disjunct populations 

were discovered to represent different lineages (Divakar et 

al. 2010a). Several examples of this type were presented, 

including Melanelixia glabra and Parmelina quercina, two 

species distributed in areas with winter rain (Mediterranean 

climate) in North Africa, Europe and North America (Argüello 

et al. 2007, Divakar et al. 2010a, b). In sorediate taxa, cryptic 

v o l u m e 1 · n o . 2 

169

Crespo and Lumbsch 

ARTICLE 

lineages have also been found, but in this case the lineages 

can include specimens from different geographical regions; 

examples include Flavoparmelia caperata, Parmotrema 

reticulatum, and P. tinctorum (Divakar et al. 2005, 2010). 

Acknowledgments 

We acknowledge the participation of David L. Hawksworth (University 

Complutense of Madrid, Spain) in finalizing this article; his revision 

enriched the manuscript with important suggestions improving the 

text. This work was supported by the Spanish Ministerio de Ciencia 

e Innovación (CGL 2010-21646/BOS, CGL2007-64652/BOS) and a 

grant of the National Science Foundation to H.T.L. (“Hidden diversity 

in parmelioid lichens”, DEB-0949147). 

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Sex in Penicillium series Roqueforti 

Jos Houbraken 1 , Jens C. Frisvad 2 and Robert A. Samson 1 

1 

CBS-KNAW Fungal Biodiversity Centre, Uppsalalaan 8, NL-3584 CT Utrecht, the Netherlands; corresponding author e-mail: j.houbraken@cbs. 

knaw.nl 

2 

Department of Systems Biology, Building 221, Søltofts Plads, Technical University of Denmark, DK-2800 Kgs. Lyngby, Denmark 

ARTICLE 

Abstract: Various fungi were isolated during the course of a survey in a cold-store of apples in the Netherlands. One 

of these fungi belongs to the genus Penicillium and produces cleistothecia at 9 and 15 °C. A detailed study using a 

combination of phenotypic characters, sequences and extrolite patterns showed that these isolates belong to a new 

species within the series Roqueforti. The formation of cleistothecia at low temperatures and the inability to produce 

roquefortine C, together with a unique phylogenetic placement, make these isolates a novel entity in the Roqueforti 

series. The name Penicillium psychrosexualis sp. nov. (CBS 128137 T ) is proposed here for these isolates. 

Key words: 

Penicillium 

roqueforti 

P. carneum 

P. paneum 

taxonomy 

phylogeny 


Introduction 

Penicillium species are commonly occurring worldwide, and 

have been isolated from various substrates including air, soil, 

various food and feed products and indoor environments (Pitt 

1979, Samson et al. 2010, Houbraken et al. 2010). Penicillium 

roqueforti is a member of this genus is and this species has both 

adverse and beneficial properties. The main beneficial property 

of this species is its role in the production of blue-veined cheeses, 

such as Roquefort, Danish blue, and Gorgonzola (Nichol 

2000). However, this species is also frequently encountered 

as a spoilage organism, and is able to damage a vast array of 

food and feed products, due to its ability to grow under harsh 

conditions. These conditions include growth at low oxygen and 

high carbon dioxide levels, in the presence of preservatives and/ 

or at low temperatures (Samson et al. 2010). 

The taxonomy of series Roqueforti was studied by Samson 

& Frisvad (2004) using a polyphasic approach, combining 

partial β-tubulin sequences, extrolite patterns, phenotypic 

and physiological data. They showed that P. paneum and P. 

carneum are closely related to P. roqueforti, together forming 

the series Roqueforti. This series shares certain characters, 

such as a fast growth rate on agar media, the ability to grow 

on malt extract agar supplemented with acetic acid and the 

production of the extrolite roquefortine C. Despite the various 

shared characters, also various features are known to 

differentiate between these species (Frisvad & Samson 2004, 

Karlshøj & Larsen 2005, O’Brien et al. 2008). These include 

the growth rate at 30 °C, reverse colours on Czapek yeast agar 

and yeast extract agar, extrolite patterns and Ehrlich reaction 

(Samson & Frisvad 2004, Samson et al. 2010). 

Various fungi were isolated during the course of a 

survey in a cold-store of apples in The Netherlands. The 

apples were stored in wooden crates, which were covered 

by a white fungal growth of Fubulorhizoctonia psychrophila. 

The latter species only grows at temperatures below 

20 °C, and during the isolation of this species growth of an 

ascospore-forming Penicillium species was detected. This 

species appeared to be related to the series Roqueforti and 

a detailed study was performed on these isolates using a 

polyphasic approach. For the phylogenetic analysis, ITS, 

partial β-tubulin and calmodulin sequences were used, and 

these data were combined with extrolite analysis and macroand 

microscopical characteristics. The combination of these 

datasets show that this species is new and is here described 

as Penicillium psychrosexualis. 

Material and methods 

Strains and morphological examination 

All examined strains belong to the Penicillium series 

Roqueforti. The strains (Table 1) were grown for 7 d as three 

point inoculations on Czapek yeast agar (CYA), malt extract 

agar, yeast extract sucrose agar (YES), creatine sucrose 

agar (CREA) and oatmeal agar (OA). The effect of various 

incubation temperatures (9–36 °C with intervals of 3 °C) on 

the growth was studied on CYA and OA. 

Molecular analysis 

Genomic DNA was isolated using the Ultraclean Microbial 

DNA Isolation Kit (MoBio, Solana Beach, CA, USA) according 



Attribution: 






v o l u m e 1 · n o . 2 

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Houbraken et al. 

ARTICLE 

Table 1. Overview of Penicillium strains used in this study. 

CBS no. Other no. Name Substrate, locality 

449.78 IBT 21509=IBT 3473=IBT 6753 P. carneum Cheddar cheese 

466.95 ATCC 46837=IBT 6885 P. carneum Cured meat, Germany 

467.95 IBT 3466 P. carneum Hotwater tank, North Sealand, Denmark 

112297 T IBT 6884 P. carneum Type, mouldy rye bread, Denmark 

463.95 IBT 12392 P. paneum Chocolate sauce, Norway 

464.95 IBT 11839 P. paneum Rye bread (non preserved), Odense, Denmark 

465.95 IBT 13929 P. paneum Mouldy baker’s yeast, Vangede, Denmark 

101032 T IBT 21541=IBT 12407 P. paneum Type, mouldy rye bread, Denmark 

112296 IBT 21729 P. paneum Cassava chips, Africa 

112294 IBT 16402=NRRL 1168 P. paneum Unknown substrate, Ottawa , Canada 

128137 T DTO 70G9 = IBT 29551 P. psychrosexualis Type, wooden crate in cold-store of apples, the Netherlands 

128136 DTO 70H7 P. psychrosexualis Wooden crate in cold-store of apples, the Netherlands 



135.67 IBT 19475=MUCL 8491 P. roqueforti Blue veined cheese, Germany 

221.30 NT ATCC 10110=ATCC 1129=CECT P. roqueforti Neotype, French Roquefort cheese, USA 

2905=IBT 6754=IFO 5459=IMI 

024313=NRRL 849 

234.38 IBT 19781=IMI 291202 P. roqueforti Blue Cheshire cheese 

479.84 IBT 21543 P. roqueforti Mouldy baker’s yeast, Denmark 

498.73 ATCC 24720=FRR 1480=IBT 19476=IMI 

174718=IMI 291199=VKM F-1748 

P. roqueforti Fruit of Malus sylvestris (apple), Russia 

the manufacturer’s instructions. The ITS regions (ITS), a part of 

the β-tubulin (BenA) or calmodulin (Cmd) gene were amplified 

and sequenced according the method described in Houbraken 

et al. (2007). Each dataset was aligned using the Clustal W 

program in MEGA5 (Tamura et al. 2007), and subsequently 

manually optimised. The evolutionary history was inferred 

by using the Maximum Likelihood (ML) method based on 

the Tamura-Nei model (Tamura & Nei 1993). The bootstrap 

consensus tree inferred from 1 000 replicates is taken to 

represent the evolutionary history of the taxa analysed (Tamura 

et al. 2007). The percentage of replicate trees in which the 

associated taxa clustered together in the bootstrap test (1 000 

replicates) is shown next to the branches. Initial tree(s) for the 

heuristic search were obtained automatically as follows. When 

the number of common sites is < 100 or less than one fourth of 

the total number of sites, the maximum parsimony method was 

used; otherwise BIONJ method with MCL distance matrix was 

used. The tree is drawn to scale, with branch lengths measured 

in the number of substitutions per site. All positions containing 

gaps and missing data were eliminated. Evolutionary analyses 

were conducted in MEGA5 (Felsenstein 1985, Tamura et al. 

2007). All phylograms were rooted with Penicillium egyptiacum 

CBS 244.32 NT . The newly obtained sequences were deposited 

in GenBank under accession numbers HQ442319–HQ442359. 

Extrolite analysis 

Plugs with mycelium and agar were extracted from 7 d old 

cultures grown on CYA and YES. Extracts were prepared 

using the method described by Smedsgaard (1997). 

Each extract was filtrated through a 0.45 PTFE filter and 

subsequently analysed using HPLC with diode array 

detection (DAD) detection. The UV spectrum and the RI 

value, and comparison with authentic chemical standards, 

were used to characterise the extrolites produced (Frisvad 

& Thrane 1987). 

Results 

Phylogeny 

The ITS regions and parts of the β-tubulin (BenA) and 

calmodulin (Cmd) gene were sequenced and analysed. The 

BenA alignment included 432 positions, and 35 positions 

were parsimony informative. The bootstrap consensus tree 

based on the results of the maximum likelihood analysis 

of this alignment is shown in Fig. 1. The total length of the 

calmodulin alignment was 500 positions long, and 27 sites 

were parsimony informative. The bootstrap consensus tree 

derived from the maximum likelihood analysis is shown in 

Fig. 2. The phylogram in Fig. 3 is based on the ITS regions 

(incl 5.8S rDNA), and 585 bases were used in the maximum 

likelihood analysis. Of these 585 characters, 16 were 

parsimony informative (including alignment gaps). 

The result of the analysis of the three datasets shows 

that P. psychrosexualis belongs to the series Roqueforti. The 

species is related to P. carneum and P. roqueforti in all three 

analysed loci, and P. paneum is basal to these three species. 

Penicillium carneum is the closest relative of P. psychrosexualis 

in the tubulin phylogram (99 %, Fig. 1), and P. roqueforti is 

basal to these two species. However, this relationship is not 

supported in the phylograms based on the calmodulin and ITS 

sequences. In these datasets, P. carneum and P. roqueforti are 

172 

i m a f U N G U S


93 

86 

CBS 467.95 P. carneum 

CBS 112297 T P. carneum 


CBS 128137 T P. psychrosexualis 

ARTICLE 

87 

CBS 128136 P. psychrosexualis 

CBS 128035 P .psychrosexualis 

CBS 449.78 P .carneum 

CBS 234.38 P. roqueforti 

49 



85 


CBS 112294 P. paneum 

CBS 101032 T P. paneum 

CBS 221.30 NT P. roqueforti 

0.01 

92 

CBS 464.95 P. paneum 



CBS 244.32 P. egyptiacum 

Fig. 1. Bootstrap consensus tree from a maximum likelihood 

analysis of partial β-tubulin sequences. The bootstrap values 

from 1 000 replicates are shown at the nodes, the branches in 

bold have a bootstrap support higher than 95 %. The tree was 

rooted with Penicillium egyptiacum CBS 244.32 NT . 


64 






77 

69 

63 




CBS128136 P. psychrosexualis 



94 



0.01 



CBS 244.32 NT P.egyptiacum 

Fig. 2. Bootstrap consensus tree from a maximum likelihood 

analysis of partial calmodulin sequences. The bootstrap values 

from 1 000 replicates are shown at the nodes, the branches in 

bold have a bootstrap support higher than 95 %. The tree was 

rooted with Penicillium egyptiacum CBS 244.32 NT . 

v o l u m e 1 · n o . 2 

173


ARTICLE 

72 






85 



69 






86 

68 






0.01 

CBS 244.32 P. egyptiacum 

Fig. 3. Bootstrap consensus tree from a maximum 

likelihood analysis of ITS sequences. The bootstrap values 

from 1 000 replicates are shown at the nodes, the branches 

in bold have a bootstrap support higher than 95 %. The 

tree was rooted with Penicillium egyptiacum CBS 244.32 NT . 

sister species and in both cases P. psychrosexualis is basal to 

these two species. Two isolates (CBS 449.78 and CBS 112296) 

warrant further attention. Penicillium carneum CBS 449.78, 

an isolate from cheddar cheese, has a unique position in the 

tubulin and calmodulin phylograms (Figs 1, 2). In addition, this 

strain is morphologically slightly deviating from the majority of 

examined P. carneum isolates. Isolate CBS 449.78 is creambrown 

in reverse on CYA, more restricted colonies on creatine 

agar and slightly slower growth rate at 30 °C. The other isolate 

which warrants attention is P. paneum CBS 112296. This strain 

has a unique β-tubulin, calmodulin and ITS sequence. However, 

extrolite analysis shows that this strain produces a typical array 

of P. paneum extrolites. More strains of these two types should 

be collected and examined to determine whether these strains 

should be raised to species level. 

Taxonomy 

Penicillium psychrosexualis Houbraken & Samson, 

sp. nov. 


(Fig. 4) 

In Penicillium subgenus Penicillium sect. Roqueforti ser. 

Roqueforti 

Coloniis in MEA cum 0.5 % acore acetica crescentibus et item in agaro 

MEA, CYS et YES celeriter crescentibus, et formatione cleistotheciorum 

ad temperationem exiguam. Roquefortino C haud producenti. 

Typus: The Netherlands: wooden crate in cold-store of apples 

covered by growth of Fubulorhizoctonia psychrophila, 3 Apr. 2008, 

J. Houbraken & F. van der Geijn (CBS H-20501 holotype; cultures ex 

type – CBS 128137 = IBT 29551 = DTO 70G9). 

Colony diameter at 7 d (in mm): CYA, 25 °C, 47–55; CYA, 15 

°C, 35–46; CYA, 30 °C, 14–27; no growth on CYA at 37 °C; 

MEA >60; YES >60; DG18, 40–50; ratio CYAS : CYA 1.2– 

1.4; creatine agar 15–25, good growth and no or weak acid 

production (under colony), delayed base production. 

Strong sporulation on CYA, velvety, slightly floccose 

in centre, dull green or dark dull green conidia, mycelium 

inconspicuous, exudates absent, soluble pigment absent, 

radial sulcate, reverse warm brown. Good sporulation on 

YES, conidia dull-green, soluble pigments absent, reverse 

mustard-yellow, none sporulating edge 6–10 mm. Good 

sporulation on DG18, conidia dull-green, reverse pale. 

Colonies on MEA dull-green towards pure-green, velvety, 

soluble pigments absent. No reaction with an Ehrlich test. 

Cleistothecia on OA at 25 °C sparsely produced and not 

visible due to the presence of a layer of conidia, formation 

of cleistothecia induced and sporulation reduced at low 

temperatures (9–15 °C, Fig. 5), cleistothecia white, soft and 

sterile when young, maturing slowly and becoming pale 

orange-brown after 3–4 mo of incubation, (50–)100–175 µm 

diam. Ascospores ellipsoidal, 4–5 × 3–4 µm, with two distinct 

equatorial ridges, often with additional secondary ridges, one 

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ARTICLE 

Fig. 4. Penicillium psychrosexualis (CBS 128036 , ex wooden crate in cold-store of apples, the Netherlands). (A–C) Colonies grown at 25 °C for 

7 d on (A) CYA, (B) MEA, and (C) YES; (D) cleistothecium; (E–F) ascospores; (G) conidiophores on DG18 with warted stipes; (H) conidiophore 

with smooth stipe; (I) conidia. Bar = 10 μm, except (F) = 1 μm. 

v o l u m e 1 · n o . 2 

175


ARTICLE 

Fig. 5. Growth of Penicillium psychrosexualis CBS 128036 on oatmeal agar at various incubation temperatures. A–F: 9, 12, 18, 24, 27 and 33 °C. 

on either side of the main ones, suggesting the presence 

of four ridges when observed with light microscopy, valves 

slightly roughened when viewed with SEM. Conidiophores 

terverticillate, slightly reduced conidiophores with smooth 

walled stipes on MEA and other agar media (PDA, PCA), 

on DG18 robust conidiophores with warted stipes, 3–4 µm. 

Metulae 10–15 × 3–4 µm. Conidiogenous cells (phialides) 

ampulliform, 8–10 × 3–4 µm. Conidia globose, smooth, 3.5–4 

µm. 

Extrolites: Penicillium psychrosexualis produces the 

extrolites andrastin A, mycophenolic, patulin, roquefortine C 

and the uncharacterized extrolite tentatively named “fumu”. 

Furthermore, P. psychrosexualis produces the same odour 

as P. roqueforti. 

Diagnostic features: The growth on MEA containing 0.5 % 

acetic acid, the formation of cleistothecia at relatively low 

temperatures for the genus (9 °C) and fast growth rate on MEA, 

CYA and YES are diagnostic features of P. psychrosexualis. 

An overview of characteristics of P. psychrosexualis in 

comparison with other members of the series Roqueforti is 

shown in Table 2. 

Similar species and taxonomy: Phylogenetically P. 

psychrosexualis belongs to series Roqueforti. This species 

shares a fast growth rate on agar media, the ability to grow 

on MEA supplemented with 0.5 % acetic acid and forms 

conidiophores with warted stipes on DG18. This species 

produces the extrolites andrastin A, mycophenolic, patulin and 

roquefortine C and is chemically close to P. carneum. However, 

P. carneum also produces penitrem A, isofumigaclavine A 

and cyclopaldic aicd, while P. psychrosexualis produces the 

uncharacterised extrolite “fumu”. Penicillium psychrosexualis 

produces the same odour as P. roqueforti, and is thus very 

different from the strong odour of P. carneum. Another 

difference between P. psychrosexualis and the other 

members of the Roqueforti series is the production of 

cleistothecia by the former species. The growth rate on 

CYA at 30 °C is a diagnostic tool to differentiate between P. 

roqueforti and P. carneum on one hand and P. paneum on the 

other. Penicillium psychrosexualis has similar growth rates 

at 30 °C as P. roqueforti and P. carneum. This observation is 

concordant with the phylogeny, which also shows that these 

three species are closely related and that P. paneum is basal 

to these species. An overview of growth rates on CYA at 

various temperatures is shown in Fig. 6. 

Nomenclature: Although the new species produces cleistothecia, 

we decided to describe the taxon in Penicillium rather than 

Eupenicillium in accordance with the recommendations of 

Hawksworth (2010) on best-practice in such instances in a 

period when the rules of nomenclature that permit the dual 

naming of pleomorphic fungi are under revision. 

176 

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ARTICLE 

Fig. 6. Overview of growth rates of the members of Penicillium series Roqueforti on CYA at various temperatures. Row, top to bottom: 9, 12, 18, 

24, 24 (reverse), 30 °C; columns, left to right: P. roqueforti DTO 81D6, P. paneum DTO 28G8, P. carneum DTO 128A9 and P. psychrosexualis 

CBS 128036. 

v o l u m e 1 · n o . 2 

177


ARTICLE 

Table 2. Overview of selected characters of members of Penicillium series Roqueforti (after Frisvad & Samson 2004, Sumarah et al. 2005, 

Nielsen et al. 2006, O’Brien et al. 2006, Månsson et al. 2010). See Karlshøj & Larsen (2005) for further differences in volatiles. 

Species Ehrlich reaction Reverse on YES Cleistothecia/ Growth rate on Extrolites** 

sclerotia CYA30 °C (mm) 

P. carneum Violet Cream-beige - 15–30 Roquefortine C, isofumigaclavine A&B, 

mycophenolic acid, patulin, cyclopaldic acid, 

penitrem A, andrastin A, (penicillic acid in 

CBS 449.78) 

P. paneum Negative Cream yellow/ 

beige* 

- 30–45 Roquefortine C, marcfortin A, patulin, 

andrastin A, citreoisocoumarin, 

(botryodiploidin) 

P. psychrosexualis Negative Mustard-yellow + 15–25 Andrastin A, mycophenolic, patulin and 

roquefortine C and the uncharacterized 

extrolite tentatively named “fumu” 

P. roqueforti Violet Blackish green -/(+) (0–)5–15 Roquefortine C, isofumigaclavine A&B, 

PR-toxin, andrastin A, citreoisocoumarin, 

(mycophenolic acid) 

*Often turning strawberry-red with age; with colour diffusing into the medium. 

**The extrolites mentioned between brackets are not produced by the majority of isolates. 

Distribution and ecology: This species has been isolated 

from wood and apples (Elstar) stored in a cold-store in the 

Netherlands. The conditions in the cold-store were 1.5–2.0 

°C in combination with an oxygen level of 1.0–1.5 %, a 

carbon dioxide level of 2.0 % and a relative humidity of 92–95 

%. These conditions strongly inhibit the growth of most fungi; 

however, a low temperature and microaerophilic conditions 

do not prevent growth of members of the Roqueforti series 

(Samson et al. 2010). 

Discussion 

The taxonomy of Penicillium series Roqueforti has been 

studied extensively in the past, mainly due to its role in cheese 

manufacture. These studies were based on phenotypic 

characters (Thom 1906, 1910, Raper & Thom 1949, Pitt 1980, 

Samson et al. 1977), extrolite patterns (Frisvad & Filtenborg 

1989, Boysen et al. 1996, Samson & Frisvad 2004, Smedsgaard 

et al. 2004) and/or molecules (Boysen et al. 1996, Skouboe et 

al. 1999, Samson et al. 2004). This is the first study using a 

multigene approach to determine the relationship of species 

belonging to the Roqueforti series. All three studied loci are 

suitable for species recognition. Even the ITS regions, normally 

not recommended for species identification in Penicillium, 

have enough variation in this series (Skouboe et al. 1999, 

Houbraken et al. 2010, Samson et al. 2010). Incongruence was 

detected during the phylogenetic analysis of the calmodulin, 

β-tubulin and ITS loci. Penicillium psychrosexualis was, with 

high bootstrap support, basal to P. carneum and P. roqueforti in 

the ITS and calmodulin dataset, while P. roqueforti was basal 

to P. carneum and P. psychrosexualis in the β-tubulin dataset. 

The use of β-tubulin in taxonomy was debated by Peterson 

(2008) and he excluded this locus in his study due to his doubt 

about the homology of this locus between members of sections 

in Aspergillus. Furthermore, Hubka & Kolařík (2010) showed 

that the commonly used primers Bt2a and Bt2b could amplify 

the β-tubulin paralog tubC in Aspergilli. The interpretation of 

paralogous genes with non-homologous function in the same 

phylogenetic analysis posses a great risk and might create 

incongruence within and between datasets (Hubka & Kolařík 


A limited number of penicillia are able to produce 

cleistothecia and ascospores, and these species were 

referred to the genus Eupenicillium in a number of studies. 

Only a limited number of penicillia known to reproduce 

sexually belong in subgenus Penicillium. Samson & Frisvad 

(2004) omitted these species in their monograph of this 

subgenus, and they recommended that a multigene study 

needs to be conducted to resolve the placement of these 

teleomorphic penicillia within the subgenus Penicillium. 

Peterson (2000) included various Eupenicillium species in 

his phylogenetic study of Penicillium, and showed that E. 

crustaceum, E. egyptiacum, E. baarnense, E. tularense, 

and Hemicarpenteles paradoxus belonged to Group 6. This 

group largely corresponds with the subgenus Penicillium 

as circumscribed by Samson & Frisvad (2004). Until now, 

only homothallic species are described in this subgenus; 

however, recent studies indicated that various species 

belonging to this subgenus are heterothallic. Hoff et al. (2008) 

178 

i m a f U N G U S


showed that P. chrysogenum is heterothallic, and analysis 

of 12 P. chrysogenum isolates showed an equal mating 

type distribution, indicating the potential of this species to 

reproduce sexually. In addition, Eagle (2009) detected either 

MAT1-1-1 or MAT1-2-1 gene fragments in isolates of P. 

camemberti, P. roqueforti and P. verrucosum, also indicating 

heterothallism. Although various trials were undertaken to 

inducing mating in P. chrysogenum (Hoff et al. 2008, Eagle 

2009, Houbraken unpubl. data) none of them have been 

successful. In addition, mating trials with P. roqueforti under 

conditions known to induce sex in Aspergillus fumigatus 

were unsuccessful and no cleistothecia were detected after 

6 mo of incubation (Eagle 2009). Various growth factors 

induce formation of cleistothecia, such as temperature, 

light, nutrients and oxygen levels (Han et al. 2003). In this 

study, we show that P. psychrosexualis, a species related 

to P. roqueforti, produces cleistothecia abundantly at 9 °C. 

The production of a sexual stage at low temperatures might 

be more widespread in Penicillium, and mating experiments 

with P. roqueforti at this temperature might result in a sexual 

stage. Furthermore, P. psychrosexualis might be a good 

model species for comparison purposes in sex induction 

experiments or expression studies of genes required for 

sex in P. roqueforti. There are also indications of a sexual 

stage in P. roqueforti. Sclerotia were observed in cultures 

in P. roqueforti (Samson et al. 1977, Shimada & Ichinoe 

1998) and it was postulated that similar structures have 

a dual function in the life-cycle in Aspergillus sect. Flavi. 

Survival of adverse conditions is one of them; the other is 

providing genetic variation in populations through sexual 

reproduction as a cleistothecium (McAlpin & Wicklow 2005, 

Horn et al. 2009). The possible discovery of the sexual 

stage in P. roqueforti could have consequences for the 

stability of starter cultures and might have advantages in 

strain improvement programs using conventional genetical 

approaches. 

The effect of temperature on sexual reproduction in species 

belonging to the subgenus Penicillium is poorly studied. Many 

of these species are capable to grow at low temperatures and 

are therefore common spoilage organisms in refrigerators. 

McCulloch & Cain (1928) found an effect of the temperature on 

the formation of sclerotia of Penicillium gladioli. This species 

produces blue-green conidial structures abundantly when 

incubated at 14–15 °C, but produced comparatively a high 

number of sclerotia and only a few conidial structures, when 

incubated at 22 °C or higher. This observation is opposite to 

the results reported here, if the assumption is followed that 

sclerotia are immature cleistothecia. On the other hand, large 

white sclerotia are occasionally seen in P. italicum, a species 

related to P. psychrosexualis and also belonging to the 

subgenus Penicillium. These structures have been observed 

in cultures incubated in darkness at 0 °C for 3 mo (Raper & 

Thom 1949, Samson & Frisvad 2004), also suggesting the 

induction of a sexual cycle at low temperatures. 

Members of series Roqueforti have a worldwide 

distribution, mainly related to human environments, and 

occur on various substrates. Penicillium roqueforti, P. 

paneum, and P. carneum occur on (preserved) food and 

silage, and only P. roqueforti has been frequently isolated 

as a saprobe in nature. Reports of the occurrence of P. 

carneum and P. paneum in nature are rare, and recently P. 

paneum has been found in stone tombs in Japan (An et al. 

2009). Penicillium psychrosexualis is the second saprobic 

species in this series and has also been isolated from wood. 

Several reports are made on the occurrence of P. roqueforti 

on woods such as sawn wood (logs), wood stakes in soil, 

wood in sea, cut lumber, Quercus robur, and very wet wood 

in indoor environments (Picci 1966, Pitt 1980, Land et al. 

1985, Kubátová 2000, Seifert & Frisvad 2000, Sumarah et 

al. 2005). 


We thank Frank van de Geijn (Agrotechnology & Food Innovations 

BV, Wageningen, the Netherlands) for collecting the wood and apples 

samples. Dae-Hoo Kim is greatly acknowledged for the preparations 

of the SEM images of the ascospores, and we thank Uwe Braun for 

providing the Latin diagnosis. 

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Anaerobic fungi: Neocallimastigomycota 

Gareth W Griffith 1 , Scott Baker 2 , Kate Fliegerova 3 , Audra Liggenstoffer 4 , Mark van der Giezen 5 , Kerstin Voigt 6 and Gordon 

Beakes 7 

1 

IBERS, Aberystwyth University, Aberystwyth SY23 3DD, Wales; corresponding author e-mail: gwg@aber.ac.uk 

2 

Chemical and Biological Process Development Group, Energy and Environment Directorate, Pacific Northwest National Laboratory, 902 

Battelle Boulevard, P.O. Box 999, MSIN P8-60 Richland, WA 99352 USA 

3 

Institute of Animal Physiology and Genetics, Academy of Sciences of Czech Republic, Videnska 1083, Prague 4 - Krc, 142 20 Czech Republic 

4 

Department of Microbiology and Molecular Genetics, Oklahoma State University, Stillwater, OK, USA 

5 

Centre for Eukaryotic Evolutionary Microbiology, Biosciences, College of Life & Environmental Sciences, University of Exeter, Stocker Road, 

Exeter EX4 4QD, UK 

6 

Friedrich Schiller University Jena, Institute of Microbiology, Department of Microbiology and Molecular Biology, Jena Microbial Research 

Collection, Neugasse 25, 07743 Jena, Germany 

7 

School of Biology, Newcastle University, Newcastle upon Tyne NE1 7RU, UK 

ARTICLE 

Abstract: This contribution is based on the six oral presentations given at the Special Interest Group 

session on anaerobic fungi held during IMC9. These fungi, recently elevated to the status of a separate 

phylum (Neocallimastigomycota), distinct from the chytrid fungi, possess several unique traits that make 

their study both fascinating yet challenging to mycologists. There are several genome sequencing programs 

underway in the US but these are hampered by the highly AT-rich genomes. Next-generation sequencing 

has also allowed more detailed investigation of the ecology and diversity of these fungi, and it is apparent 

that several new taxa beyond the six genera already named exist within the digestive tracts of mammalian 

herbivores, with others potentially inhabiting other anaerobic niches. By increased collaboration between 

the various labs studying these fungi, it is hoped to develop a stable taxonomic backbone for these fungi 

and to facilitate exchange of both cultures and genetic data. 

Key words: 

chytrid 

basal fungi 

genome sequencing 

phylogenetics 

next-generation sequencing 

hydrogenosome 


Introduction 

Since their discovery by Colin Orpin in the 1970s, the 

anaerobic fungi, now members of the recently erected 

phylum Neocallimastigomycota, have aroused the curiosity 

of mycologists, not only due to their distinctive physiology 

but also because of their biotechnological potential, for more 

efficient animal nutrition and also biomass conversion/biofuel 

production. This contribution, based on a Special Interest 

Group session held during IMC9, brought together mycologists 

interested in anaerobic fungi to share recent discoveries and 

to discuss how best to move forward research in this area. Six 

speakers kindly agreed to speak at the session, with a further 

seven non-speakers in attendance. Given the small numbers 

of mycologists now engaged in research on anaerobic fungi, 

this was a pleasing quorum, with expertise in genomic, 

phylogenetic, morphological, physiological and ecological 

aspects of the biology of these fungi being represented. 

This synopsis of the session amounts to a mini-review of the 

current state of research on these fascinating fungi. 

Contributions 

The session began with a presentation from Scott Baker on 

the status of two genome sequencing projects (Piromyces E2 

and Orpinomyces SR2). The AT richness of these genomes 

(approaching 80 % in non-coding regions) has caused 

significant technical problems for genomic sequencing and 

assembly. The Joint Genome Initiative (JGI) is working through 

technical issues with assembling the genome, but does plan 

to release the sequence in the near future. In contrast, the 

Expressed Sequence Tag (EST) library sequencing project 

did not encounter any technical issues and will be made 

available concurrent with the eventual release of the genome 

sequence. Until then, contact Scott for information on 

accessing ESTs (scott.baker@pnl.gov). Much of the interest 

in these fungi relates to the genes/enzymes important for 

biorefining and biofuel production, notably xylose isomerases 

and glycosyl hydrolases (xylanases, cellulases). 

Kate Fliegerova, who also presented a poster (4.015) at 

the main IMC9 congress, further explored the biotechnological 



Attribution: 






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ARTICLE 

potential of these fungi. Some of the cellulases of anaerobic 

fungi originated via horizontal gene transfer from bacteria, 

so these are the only fungi known to possess cellulosomes, 

cell-wall associated multienzyme complexes (Garcia-Vallve 

et al. 2000, Steenbakkers et al. 2001). Combined with 

their anaerobic metabolism and ability grow at elevated 

temperatures (39 ºC), they have great biotechnological 

potential. In Prague, Kate and her colleagues have explored 

the use of anaerobic fungi to improve the hydrolytic phase of 

biogas production. They have also investigated which fungi 

are present in the cow manure used to prime the biogas 

fermentations (Fliegerova et al. 2010), finding members of 

the genus Cyllamyces, the sixth and most recently discovered 

group of anaerobic fungi (Ozkose et al. 2001) to be dominant, 

with the ‘most famous’ (and type) genus Neocallimastix 

comprising only a small proportion of the population. This 

imbalance is also reflected in the literature, possibly a result 

of the widespread use of wheat straw for the culture-based 

isolation of these organisms (Griffith et al. 2009). 

The application of culture-independent approaches to 

assessing the diversity of anaerobic fungi was the subject of 

Audra Liggenstoffer’s presentation. Her PhD project at the 

Oklahoma State University used barcoded 454 sequencing 

to determine the fungal symbionts present in the faeces of 30 

species of larger herbivores, many from Oklahoma City Zoo. 

Her findings have recently been published (Liggenstoffer 

et al. 2010) and demonstrated not only confirmation of the 

occurrence of anaerobic fungi in a non-mammal host (green 

iguana) but also the existence of eight novel groups of 

fungi, with these new taxa (likely to represent new genera) 

comprising almost 40 % of the >250,000 ITS sequences 

obtained. Whilst it can be difficult to be certain that zoo animals 

have not acquired new symbionts whilst in captivity, some of 

these novel groups (NG) did show some host specificity, for 

example with NG6 comprising nearly all the fungi in kudu and 

NG8 being found only in Somali wild ass (Fig. 1). 

With such a high rate of taxon discovery, the 

Neocallimastigomycota, the newest of the fungal phyla 

Fig. 1. Network graph highlighting shared OTUs between different anaerobic fungal communities in different animal hosts. The graph is colourcoded 

by animal host phylogeny (family). Circular nodes indicate animal data sets, whereas smaller square, grey nodes represent individual 

OTUs. Data sets with a higher proportion of shared OTUs are pulled to the middle, whereas data sets with a high proportion of unique OTUs 

remain on the periphery. The distance between any two data sets is a function of the number of shared OTUs between the two. Figure supplied 

by Audra Liggenstoffer. 

182 

 

i m a f U N G U S


(Hibbett et al. 2007) clearly has more taxonomic gems 

awaiting discovery, notably in habitats beyond the digestive 

tracts of vertebrates. Anaerobic fungal sequences do appear 

in environmental clone libraries (Lockhart et al. 2006) but this 

attests more to the resilience of their resting spores (Ozkose 

2001) than to their active metabolism in these habitats. 

However, it is already clear that a robust taxonomic scheme 

based on gene sequence data rather than the meagre 

morphological traits is needed. Kerstin Voigt specialises in the 

phylogenetics of the lower fungi and questions the acceptability 

of the phylum Neocallimastigomycota (Ebersberger et 

al. 2010). In collaboration with Kate Fliegerova and Ingo 

Ebersberger from Vienna, Kerstin has taken a phylogenomic 

approach (concatenated supermatrices of data) to generate 

more robust phylogenies. A key element of this approach is 

the use of orthologous genes for comparisons, requiring first 

the identification of the original member of any gene family 

within an organism, prior to any interspecific comparisons. 

Application of this more robust approach for the anaerobic 

fungi requires sequence data from genes other than the 

rRNA locus, however, Kerstin’s initial analyses, like Audra’s 

454 data, revealed the presence of four novel genera (Fig. 

2). It was also clear that some GenBank accessions are 

mis-labelled, highlighting the difficulty in morphological 

classification of these fungi. 

If there is one feature of the Neocallimastigomycota that 

intrigues microbiologists more generally, it is their obligately 

anaerobic metabolism. Prior to their “official” discovery by Colin 

ARTICLE 

Fig. 2. Phylogenetic tree based on a maximum likelihood analysis using RAxML v. 7.2.6 (Stamatakis 2006) with the aligned ITS1-5.8SITS2 

region out of a total of 186 chytrids. 

v o l u m e 1 · n o . 2 

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Griffith et al. 

ARTICLE 

Orpin (1974), these organism had been reported by several 

rumen microbiologists, and even named by Liebetanz (1910) 

as the flagellate protozoan Callimastix. During the 1960's, 

Hungate and Prins had also noted these organisms but had 

dismissed them as contaminants. Thus the dogma that there 

are no anaerobic fungi was not easily overturned. Despite 

the near-absence of any useful fossil record, increasingly 

accurate molecular clock approaches consistently show 

that the fungi diverged from the more primitive metazoans 

(animals) some 1000 million years ago. At that time, between 

the two great oxygenation events (at ca. 2400 and 600 

Myr), primitive eukaryotes (including the earliest fungi) were 

exposed to low atmospheric oxygen levels (


Liebetanz E (1910) Die parasitischen Protozoen des 

Wiederkäuermagens. Archiv für Protistenkunde 19: 19–80. 

Liggenstoffer AS, Youssef NH, Couger MB, Elshahed MS (2010) 

Phylogenetic diversity and community structure of anaerobic 

gut fungi (phylum Neocallimastigomycota) in ruminant and nonruminant 

herbivores. ISME Journal 4: 1225–1235. 

Lockhart RJ, Dyke MI van, Beadle IR, Humphreys P, McCarthy 

AJ (2006) Molecular biological detection of anaerobic gut 

fungi (Neocallimastigales) from landfill sites. Applied and 

Environmental Microbiology 72: 5659–5661. 

Orpin CG (1974) Rumen flagellates Callimastix frontalis and Monas 

communis - Zoospores of phycomycete fungi. Journal of Applied 

Bacteriology 37: R9–R10. 

Ozkose E (2001) Morphology and molecular ecology of anaerobic 

fungi. PhD thesis, University of Wales, Aberyswyth. 

Ozkose E, Thomas BJ, Davies DR, Griffith GW, Theodorou MK 

(2001) Cyllamyces aberensis gen.nov sp.nov., a new anaerobic 

gut fungus with branched sporangiophores isolated from cattle. 

Canadian Journal of Botany 79: 666–673. 

Rezaeian M, Beakes GW, Parker DS (2004) Distribution and 

estimation of anaerobic zoosporic fungi along the digestive tracts 

of sheep. Mycological Research 108: 1227–1233. 

Stamatakis A (2006) RaxML-VI-HPC: maximum likelihood-based 

phylogenetic analyses with thousands of taxa and mixed models. 

Bioinformatics 22: 2688–2690. 

Steenbakkers PJM, Li XL, Ximenes EA, Arts JG, Chen HZ, et 

al. (2001) Noncatalytic docking domains of cellulosomes of 

anaerobic fungi. Journal of Bacteriology 183: 5325–5333. 

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Polyphasic taxonomy of Aspergillus section Sparsi 

János Varga 1,2 , Jens C. Frisvad 3 and Robert A. Samson 1 

1 

CBS-KNAW Fungal Biodiversity Centre, Uppsalalaan 8, NL-3584 CT Utrecht, The Netherlands 

2 

Department of Microbiology, Faculty of Science and Informatics, University of Szeged, H-6726 Szeged, Közép fasor 52, Hungary; corresponding 

author e-mail: jvarga@bio.u-szeged.hu 

3 

Centre for Microbial Biotechnology, Department of Systems Biology, Building 221, Technical University of Denmark, DK-2800 Kgs. Lyngby, 

Denmark 

ARTICLE 

Abstract: Aspergillus section Sparsi includes species which have large globose conidial heads with colours 

ranging from light grey to olive-buff. In this study, we examined isolates of species tentatively assigned to section 

Sparsi using a polyphasic approach. The characters examined include sequence analysis of partial b-tubulin, 

calmodulin and ITS sequences of the isolates, morphological and physiological tests, and examination of the 

extrolite profiles. Our data indicate that the revised section Sparsi includes 10 species: A. anthodesmis, A. 

biplanus, A. conjunctus, A. diversus, A. funiculosus, A. implicatus, A. panamensis, A. quitensis, A. sparsus, 

and the new taxon A. haitiensis. The recently described A. quitensis and A. ecuadorensis are synonyms of 

A. amazonicus based on both molecular and physiological data. The white-spored species A. implicatus has 

also been found to belong to this section. Aspergillus haitiensis sp. nov. is characterised by whitish colonies 

becoming reddish brown due to the production of conidial heads, and dark coloured smooth stipes. The taxon 

produces gregatins, siderin and several unknown but characteristic metabolites. 

Key words: 

Aspergillus section Sparsi 

b-tubulin 

calmodulin 

Eurotiales 

extrolites 

ITS 

polyphasic taxonomy 


Introduction 

The Aspergillus sparsus species group (Aspergillus section 

Sparsi; Gams et al. 1985) was established by Raper & Fennell 

(1965) to accommodate four species isolated from tropical or 

subtropical soils. Species assigned to this group have large 

globose conidial heads, which irregularly split with age, with 

colours ranging from light grey to olive-buff. Samson (1979) 

suggested that A. gorakhpurensis should also be placed 

to this section. However, phylogenetic analysis of parts of 

the ribosomal RNA gene cluster indicated that this species 

belongs to Aspergillus section Cremei (Peterson 1995, 2000). 

According to the recent data of Peterson et al. (2008) and 

Peterson (2008), the monophyletic section Sparsi belongs 

to subgenus Nidulantes, and in addition to A. sparsus, A. 

biplanus, A. diversus and A. funiculosus, originally placed to 

this section by Raper & Fennell (1965), it also includes A. 

panamensis and A. conjunctus previously assigned to section 

Usti, and A. anthodesmis which was previously placed in the 

A. wentii group (Raper & Fennell 1965). 

In this study, we examined available isolates of the species 

proposed to belong to section Sparsi to clarify the taxonomic 

status of this section. The methods used include sequence 

analysis of the ITS region (including internal transcribed 

spacer regions 1 and 2, and the 5.8 S rRNA gene of the 

rRNA gene cluster), and parts of the b-tubulin and calmodulin 

genes, analysis of macro- and micromorphological characters 

and extrolite profiles. 


Morphological examinations 

The strains examined are listed in Table 1. The strains were 

grown for 7 d as three-point inoculations on Czapek agar, 

Czapek yeast autolysate agar (CYA), malt extract agar 

(MEA), and oatmeal agar (OA) at 25 °C and 37 °C (medium 

compositions in Samson et al. 2010). 

Analysis for Extrolites 

The cultures were analysed according to the HPLC-diode 

array detection method of Frisvad & Thrane (1987, 1993) as 

modified by Smedsgaard (1997). The isolates were analysed 

on CYA and YES agar using three agar plugs (Smedsgaard 

1997). The secondary metabolite production was confirmed 

by identical UV spectra with those of standards and by 

comparison to retention indices and retention times for pure 

compound standards (Frisvad & Thrane 1993, Rahbaek et 

al. 2000). 



Attribution: 






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Table 1. The Aspergillus section Sparsi isolates examined in this study. 

Species Strain No. Origin 

A. amazonicus CBS 124228 T = E19D Soil, Makas, Ecuador 

A. anthodesmis CBS 552.77 T = NRRL 22884 Soil, Ivory Coast 

A. biplanus CBS 468.65 T = NRRL 5071 Soil, Tilaran, Costa Rica 

A. biplanus CBS 469.65 = NRRL 5073 Soil, Tilaran, Costa Rica 

A. biplanus NRRL 5072 Soil, Tilaran, Costa Rica 

A. conjunctus CBS 476.65 T = NRRL 5080 Forest soil, Palmar, Province of Punteras, Costa Rica 

A. diversus CBS 480.65 T = NRRL 5074 Soil, Esparta, Costa Rica 

A. ecuadorensis CBS 124229 T =E19F Soil, Makas, Ecuador 

A. funiculosus CBS 116.56 T = NRRL 4744 Soil, Ibadan, Nigeria 

A. haitiensis CBS 464.91 T = NRRL 4569 Soil under sage and cactus, Haiti 

A. haitiensis CBS 468.91 = NRRL 4568 Desert soil, Haiti 

A. implicatus CBS 484.95 T Soil, Ivory Coast 

A. panamensis CBS 120.45 T = NRRL 1785 Soil, Panama 

A. panamensis NRRL 1786 Soil, Panama 

A. quitensis CBS 124227 T = E19C Soil, Makas, Ecuador 

A. sparsus CBS 139.61 T = NRRL 1933 Soil, Costa Rica 

A. sparsus NRRL 1937 Soil, San Antonio, Texas, USA 

Isolation and analysis of nucleic acids 

The cultures used for the molecular studies were grown on 

malt peptone (MP) broth using 1 % (w/v) of malt extract (Oxoid) 

and 0.1 % (w/v) bacto peptone (Difco), 2 mL of medium in 15 

mL tubes. The cultures were incubated at 25 °C for 7 d. DNA 

was extracted from the cells using the Masterpure yeast 

DNA purification kit (Epicentre Biotechnologies) according to 

the instructions of the manufacturer. Fragments containing 

the ITS region were amplified using primers ITS1 and ITS4 as 

described previously (White et al. 1990). Amplification of part 

of the b-tubulin gene was performed using the primers Bt2a 

and Bt2b (Glass & Donaldson 1995). Amplifications of the 

partial calmodulin gene were set up as described previously 

(Hong et al. 2005). Sequence analysis was performed with 

the Big Dye Terminator Cycle Sequencing Ready Reaction 

Kit for both strands, and the sequences were aligned with 

the MT Navigator software (Applied Biosystems). All the 

sequencing reactions were purified by gel filtration through 

Sephadex G-50 (Amersham Pharmacia Biotech, Piscataway, 

NJ) equilibrated in double-distilled water and analyzed on 

the ABI PRISM 310 Genetic Analyzer (Applied Biosystems). 

The unique ITS, b-tubulin, and calmodulin sequences were 

deposited at the GenBank nucleotide sequence database 

under accession numbers FJ491645–FJ491675, and 

FJ943936–FJ943941. 

Data analysis 

The sequence data was optimised using the software package 

Seqman from DNAStar Inc. Sequence alignments were 

performed by MEGA v. 4.0 (Tamura et al. 2007) and improved 

manually. For parsimony analysis, the PAUP v. 4.0 software 

was used (Swofford 2002). Alignment gaps were treated as a 

fifth character state and all characters were unordered and of 

equal weight. Maximum parsimony analysis was performed 

for all data sets using the heuristic search option with 100 

random taxa additions and tree bisection and reconstruction 

(TBR) as the branch-swapping algorithm. Branches of zero 

length were collapsed and all multiple, equally parsimonious 

trees were saved. The robustness of the trees obtained 

was evaluated by 1000 bootstrap replications (Hillis & Bull 

1993). An A. ochraceoroseus isolate belonging to section 

Ochraceorosei of subgenus Nidulantes (Peterson et al. 2008) 

was used as outgroup in these experiments. The alignments 

were deposited in TreeBASE () 

under accession number S11028. 

Results and discussion 

Phylogeny 

We examined the genetic relatedness of section Sparsi 

isolates using sequence analysis of the ITS region of the 

ribosomal RNA gene cluster, and parts of the calmodulin 

and b-tubulin genes. The calmodulin data set included 566 

characters, with 288 parsimony informative characters. 

One of the 56 MP trees is shown in Fig. 1 (tree length: 741, 

consistency index: 0.7247, retention index: 0.8903). During 

analysis of a part of the b-tubulin gene, 494 characters were 

analysed, among which 196 were found to be parsimony 

informative. The single MP tree based on partial b-tubulin 

genes sequences is shown in Fig. 2 (length: 507 steps, 

consistency index: 0.7179, retention index: 0.8938). The 

ITS data set included 559 characters with 58 parsimony 

informative characters. One of the 702 MP trees is presented 

in Fig. 3 (tree length: 180, consistency index: 0.8056, 

retention index: 0.8763). 

Phylogenetic analysis of b-tubulin, calmodulin and ITS 

sequence data indicated that Aspergillus section Sparsi 

includes 10 species. Aspergillus biplanus and A. diversus 

are closely related to each other on all trees, while another 

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A. conjunctus NRRL 5080 

A. conjunctus CBS 476.65 

A. anthodesmis CBS 552.77 

A. panamensis NRRL 1785 

A. panamensis CBS 120.45 

A. amazonicus CBS 124228 

A. ecuadorensis CBS 124229 

A. quitensis CBS 124227 

A. diversus NRRL 5074 

A. diversus CBS 480.65 

A. biplanus NRRL 5072 



A. biplanus CBS 468.65 


A. implicatus CBS 484.95 

A. funiculosus NRRL 4744 

A. sparsus NRRL 1933 

A. sparsus CBS 139.61 


A. hai6ensis NRRL 4569 

A. hai6ensis CBS 464.91 

Fig. 1. One of the MP trees obtained based on 


phylogenetic analysis of calmodulin sequence data of 


Aspergillus section Sparsi. Numbers above branches 

A. ochraceoroseus CBS 550.77 

are bootstrap values. Only values above 70 % are 

indicated. 

ARTICLE 


























Fig. 2. The single MP tree obtained based on 

phylogenetic analysis of b-tubulin sequence data of 

Aspergillus section Sparsi. Numbers above branches 

are bootstrap values. Only values above 70 % are 

indicated. 

v o l u m e 1 · n o . 2 

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A. anthodesmis NRRL 22884 











Fig. 3. One of the MP trees obtained based on 

phylogenetic analysis of ITS sequence data 

of Aspergillus section Sparsi. Numbers above 

branches are bootstrap values. Only values 

above 70 % are indicated. 

clade includes A. panamensis, A. anthodesmis, A. conjunctus, 

and the recently described A. amazonicus, A. quitensis and 

A. ecuadorensis isolates on the trees based on b-tubulin and 

ITS sequence data (Figs 2, 3; Mares et al. 2008). Although 

Mares et al. (2008) found that these three isolates have 

identical ITS sequences, they were suggested to represent 

distinct species based on morphological data (length of talks, 

diameter of vesicles, morphology of conidia and number of 

phialides), and were placed in Aspergillus section Wentii. 

However, these three isolates could not be distinguished from 

each other based on molecular, morphological or extrolite data 

in our study, and clearly belong to section Sparsi (Figs 1–4). 

Aspergillus amazonicus is chosen as the correct name for the 

taxon and A. quitensis and A. ecuadorensis are considered 

synonyms. Aspergillus implicatus, a white-spored species 

originally assigned to Aspergillus section Candidi (Maggi & 

Persiani 1994), also belongs to this section. This species was 

described to produce conidiophores surrounded by sterile 

hyphae, not yet seen in any other species of the Aspergillus 

genus. Unfortunately the ex-type culture showed only poor 

sporulation and only a few conidiophores with sterile outgrowth 

could be observed (Fig. 5). 

Phylogenetic analysis of sequence data indicated that 

the four examined A. sparsus isolates fall into two closely 

related clades. The three phylogenies were concordant, 

with no conflict between the topologies of the gene trees, 

in accordance with the phylogenetic species recognition 

concept detailed by Taylor et al. (2000). The ex-type strain 

of A. sparsus (CBS 139.61 T ) together with an isolate from 

Texas, USA form one clade, while two isolates came from soil 

from Haiti form another clade on all trees (Figs 1–3). Both 

of the latter isolates were found by Raper & Fennell (1965) 

to differ from the ex-type strain of A. sparsus in producing 

more restrictedly growing colonies in shades of reddish 

brown on MEA plates, while one of the isolates (CBS 464.91 

= NRRL 4569) also produced “small fragmentary sporulating 

structures adjacent to the agar surface that bear conidia 

similar to those of normal heads” (Raper & Fennell 1965). 

Here we describe this new species as Aspergillus haitiensis. 

Regarding the value of the different loci for species 

delimitation in section Sparsi, all species could be 

distinguished using either ITS, b-tubulin or calmodulin 

sequence data. However, the resolving power was much 

higher for the protein coding genes than for the ITS region. 

The situation is more difficult in other sections of Aspergilli, 

including for example sections Nigri (Samson et al. 2007), 

Clavati (Varga et al. 2007), and Cervini (J. Varga, unpubl. 

observ.), where the ITS region cannot be used reliably to 

distinguish all species assigned to the given section. 

Extrolite profiles 

Among the species assigned to Aspergillus section Sparsi, 

A. panamensis produces cyclogregatin and gregatins (also 

called graminins or aspertetronins; Anke et al. 1980a, b, 

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Fig. 4. Aspergillus amazonicus (CBS 124228). A–C. Colonies of 7 d grown at 25 °C; A on CYA, B on MEA, C on CREA. D–I. Conidiophores and 

conidia. Bars = 10 µm. 

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Table 2. Extrolites produced by species of Aspergillus section Sparsi. The structures of the extrolites in brackets have not yet been elucidated. 

Species 

Extrolites 

A. amazonicus an aszonalenin, (dob-indol, fot, Vurs1, vurs2, stan) 

A. anthodesmis gregatins, siderin (alk-769gl; AMF1, AMF2, AMF3, ANTW, kota, met k, tidmyco1, tidmyco2, senmyco1, senmyco2, 

senmyco3, UNTW) 

A. biplanus auroglaucin, (BLØDO, CUR-678, KONI, OKSI-1121, RAI-701, RAI-843, SKOT, VERN-652, VERN-655, VERN- 

661, VERN-673, vers-965, vers-979, vers-1049, vers-1107) 

A. conjunctus auroglaucin, siderin?, (alk-1538, alk-1756, blæam, CONJ1, CONJ2, CONJ3, DUTS, INSUX,JON1, JON2, JON3, 

JON4, kola, kola2, SVIF1, SVIF2, UT, verruc1, verruc2, vers-1049, vers-1107), a falconensin (? by A. conjunctus 

SRRC 423) 

A. diversus auroglaucin, mycophenolic acid?, (alka-704, CONJ1, kola2, OKSI-1, OKSI-2, vers-965, vers-979, vers-1049, 

vers-1107; OKSI-3, OKSI-4, OKSI-5, OKSI-6 by NRRL 5075) 

A. funiculosus arugosin E, ethericin A, funicin = ethericin B, terrein?, (AQ-798, AQ-1456, bianthron-1396, DERH, DRI, emon, 

hæms, NOL, RAI-921, RAI-972, storå, SULTI-1, SULTI-2, vers-818, vers-856) 

A. haitiensis NRRL 4568 (ATROV, GYLA, NIDU, tidmyco1, tidmyco2, tidmyco3, spar1, spar2, spar3) 

A. haitiensis NRRL 4569 gregatins, siderin, (AMF1, AMF2, AMF3, senmyco1, senmyco2, senmyco3, tidmyco1, tidmyco2, tidmyco3) 

A. implicatus a versicolorin, an austalide derivative (?) 

A. panamensis gregatins, siderin, (AQ-1456, OTTO), 

A. sparsus (NIDU, senmyco1, senmyco2, senmyco3, spar1, spar2) 

Fig. 5. Aspergillus implicatus (CBS 484.95). A–B. Conidal heads showing sterile outgrowths. Bars = 100 µm. 

1988), while A. funiculosus has been found to produce 

ethericin A (also called violaceol I or aspermutarubrol), and 

ethericin B (or funicin; König et al. 1978, 1980, Nakamura et 

al. 1983) (Table 2). Ethericin A was first isolated and called 

aspermutarubrol from A. sydowii, causing the red colouration 

of the medium, as this unstable compound will turn into a 

red dye by oxidation (Shibata et al. 1978). The ethericins (or 

violaceols) are also produced by A. versicolor and several 

Emericella species (Fremlin et al. 2009). Gregatins are also 

produced by A. anthodesmis and one of the A. haitiensis 

isolates (Table 2). Siderin is related to kotanins produced by 

some black Aspergilli and A. clavatus (Samson et al. 2007, 

Varga et al. 2007), and is also produced by A. panamensis, 

A. anthodesmis, A. conjunctus and by an A. haitiensis isolate 

(NRRL 4569). Auraglaucin production is shared by A. biplanus, 

A. conjunctus and A. diversus, and is also produced by some 

Eurotium species (Gould & Raistrick 1934, Quilico et al. 1949). 

Aspergillus implicatus (Fig. 5) has been found to produce a 

versicolorin derivative. The two A. haitiensis isolates produced 

quite distinct extrolite profiles, but shared the production of 

several unknown compounds including those tentatively 

named tidmyco1-3. Several of the other extrolites produced by 

species assigned to Aspergillus section Sparsi have also been 

detected in other species assigned to sections Nidulantes, Usti 

and Versicolores, justifying the assignment of section Sparsi to 

Aspergillus subgenus Nidulantes (Peterson et al. 2008). 

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Fig. 6. Aspergillus haitiensis (CBS 464.91). A–C. Colonies of 7 d grown at 25 °C; A on CYA, B on MEA, C on CREA. D–I. Conidiophores and 

conidia. Bars = 10 µm. 

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Taxonomy 

Aspergillus haitiensis Varga, Frisvad & Samson, sp. 

nov. 


(Fig. 6) 

Speciebus Aspergilli sect. Sparsi similes, sed coloniis porphyreis et 

stipitibus fuscatis, laevibus distinguitur. 

Typus: Haiti: isolated from soil under sage and cactus, W. 

Scott (as 113a) (CBS H-20503 -- holotypus, cultures exholotype 

CBS 464.91 = NRRL 4569). 

Colonies on MEA 50–60 mm, on CYA 30–35 mm, after 14 

d at 25 °C, moderate growth on MEA after 7 d at 37 °C. 

Conidial heads produced sparsely on CYA, colony colour first 

white then reddish brown, colony texture floccose, reverse 

creamish to light brown. Conidial heads radiate; stipes 

5–9 µm, thick-walled, dark brown in colour; vesicles 10–25 

µm wide, biseriate; metulae covering the whole vesicle, 

measuring 2.5–4 × 5–7 µm. Conidiogenous cells (phialides) 

2–2.5 × 7–8 µm. Conidia globose to ellipsoidal 4–5.6 × 5–6 

µm, smooth. Fragmentary sporulating structures in addition 

to the normal conidial heads are also present. 

Additional isolate studied: Haiti: Port de Paix, from desert 

soil, W. Scott (as 103b) (CBS 468.91 = NRRL 4568). 

Diagnostic features: Thin whitish colonies turning to reddish 

brown colour on CYA, brown-coloured smooth stipes, 

and production of unknown extrolites tentatively called 

tidmyco1-3. 


We are grateful to Tineke van Doorn who helped with the 

morphological data, Uwe Braun with the Latin diagnosis, and our 

referees. 

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Aspergillus sect. Aeni sect. nov., a new section of the genus for A. 

karnatakaensis sp. nov. and some allied fungi 

János Varga 1,2 , Jens C. Frisvad 3 and Robert A. Samson 1 

1 

CBS-KNAW Fungal Biodiversity Centre, Uppsalalaan 8, NL-3584 CT Utrecht, The Netherlands 

2 

Department of Microbiology, Faculty of Science and Informatics, University of Szeged, Közép fasor 52, H-6726 Szeged, Hungary; 

corresponding author e-mail: jvarga@bio.u-szeged.hu 

3 

Department of Systems Biology, Building 221, Søltofts Plads, Technical University of Denmark, DK-2800 Kgs. Lyngby, Denmark 

ARTICLE 

Abstract: The new species Aspergillus karnatakaensis sp. nov. is described and illustrated. All three 

isolates of this species were isolated from Indian soil; two from soil under a coconut palm in a coffee 

plantation in Karnataka, and one from soil in the Machrar river bed in Bansa district. This species is 

closely related to, but clearly distinct, from A. aeneus based on b-tubulin or calmodulin sequence data. 

Sequences of the ITS region of these two species are identical. Aspergillus karnatakaensis produced 

terrein, gregatins, asteltoxin, karnatakafurans A and B and the unknown metabolite, provisionally 

named NIDU. Aspergillus karnatakaensis belongs to a well-defined clade within Aspergillus subgenus 

Nidulantes together with eight other species including A. aeneus, A. crustosus, A. eburneocremeus, A. 

heyangensis, and the teleomorph producing-species Emericella bicolor, E. discophora, E. spectabilis, 

and E. foeniculicola. This clade is placed in a new section, Aspergillus sect. Aenei sect. nov. All 

teleomorph species assigned to this section are able to produce sterigmatocystin. 

Key words: 

Aspergillus subgen. Nidulantes 

b-tubulin 

calmodulin 

Eurotiales 

extrolites 

ITS 

polyphasic taxonomy 


Introduction 

Aspergillus subgenus Nidulantes is one of the largest 

subgenera of the genus Aspergillus, including about 80 

species (Peterson 2008, Peterson et al. 2008). Several 

species of this subgenus have a teleomorph assigned to 

Emericella (Pitt et al. 2000, Samson 2000, Frisvad & Samson 

2004). Species of subgenus Nidulantes are important as 

opportunistic human pathogens (Verweij et al. 2008, Varga 

et al. 2008), as producers of various secondary metabolites 

which are useful for the pharmaceutical industry (e.g. 

penicillin, echinocandins, ophiobolins), and mycotoxins 

which are harmful to animals and humans (e.g. aflatoxins, 

sterigmatocystin; Frisvad et al. 2004, 2005, Frisvad & 

Samson 2004, Zalar et al. 2008). 

During surveys of Aspergillus isolates from soil samples 

from subtropical regions, two interesting isolates were 

recovered which did not match any known species of the 

genus. We used the polyphasic approach, including sequence 

analysis of parts of the b-tubulin and calmodulin genes and the 

ITS nrDNA region, macro- and micromorphological analyses, 

and examination of the extrolite profiles of the isolates to 

differentiate the new species Aspergillus karnatakaensis sp. 

nov. We also analysed strains of species which appeared 

to be closely related to the new species for the production 

of extrolites and found sterigmatocystin in all species 

with a teleomorphic state studied and also in Aspergillus 

ebureocremeus. 


Isolates 

The strains used in this study are listed in Table 1. 

Morphological analysis 

For macromorphological observations, Czapek yeast 

autolysate (CYA), malt extract autolysate (MEA) agar, Yeast 

Extract Sucrose agar (YES), creatine sucrose agar (CREA), 

and oatmeal agar (OA) were used (Samson et al. 2010). The 

isolates were inoculated at three points on each plate of each 

medium and incubated at 25 °C and 37 °C in the dark for 7 

d. For micromorphological observations, microscopic mounts 

were made in lactic acid from MEA and OA colonies and a 

drop of alcohol was added to remove air bubbles and excess 

conidia. 

Extrolite analysis 

The isolates were grown on CYA and YES at 25 °C for 7 d. 

Extrolites were extracted after incubation. Five 6 mm plugs 



Attribution: 






v o l u m e 1 · n o . 2 

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Varga et al. 

ARTICLE 

Table 1. Isolates of Aspergillus and Emericella spp. examined in this study. 

Species Strain No. a Substratum, country, location GenBank No. 

β-tubulin calmodulin ITS 

A. aeneus CBS 128.54 T = NRRL 4769 Forest soil, Modilen, Somalia EF652298 EF652386 EF652474 

A. crustosus CBS 478.65 T = NRRL 4988 Skin scrapings, man, Kankakee, Illinois, USA EF652313 EF652401 EF652489 

A. eburneocremeus CBS 130.54 T = NRRL 4773 Forest soil, Modien Forest, Somalia EF652300 EF652388 EF652476 

A. heyangensis CBS 101751 T Placentae of Gossypium sp., Heyang, Shaanxi FJ491520 FJ491521 FJ491522 

Province, China 

A. karnatakaensis CBS 102800 T = IBT 22153 Soil under coconut palm in coffee plantation, EU482438 EU482431 EU482441 

India, Karnataka 

A. karnatakaensis CBS 102799 = IBT 22154 Soil under coconut palm in coffee plantation, EU482436 EU482430 EU482443 

India, Karnataka 

A. karnatakaensis NRRL 4649 Soil in the Machrar river bed located in district EF652292 EF652380 EF652468 

Bansa, state Madhya Pradesh, India 

E. bicolor CBS 425.77 T Soil from Artemisia grassland, USA, Wyoming, EF652335 EF652423 EF652511 

Teton Basin 

E. discophora CBS 469.88 T = IBT 21910 Soil, Spain AY339999 EU443970 EU448272 

E. discophora CBS 470.88 = IBT 21911 Forest soil, Spain AY340000 EU443969 EU448266 

E. foeniculicola CBS 156.80 T Foeniculum vulgare seed, China EU443990 EU443968 EU448274 

E. heterothallica CBS 489.65 T Soil, Costa Rica EU076369 EU076361 AB248987 

E. spectabilis CBS 429.77 T Coal mine spoil material, Wyoming, USA, 

Seminole no. 1 mine 

EU482437 EU482429 EU482442 

a 

Cultures are deposited in/were obtained from the following collections: CBS, CBS-KNAW Fungal Biodiversity Centre, Utrecht, the Netherlands; 

IBT, Culture Collection of Fungi, Mycology Group, BioCentrum-DTU, Technical University of Denmark, Lyngby, Denmark; NRRL, Agricultural 

Research Service Culture Collection, Peoria, IL, USA. 

of each agar medium were taken and pooled together into 

the same vial for extraction with 0.75 mL of a mixture of ethyl 

acetate/dichloromethane/methanol (3:2:1) (v/v/v) with 1 % 

(v/v) formic acid. The extracts were filtered and analyzed by 

HPLC using alkylphenone retention indices and diode array 

UV-VIS detection as described by Frisvad & Thrane (1987, 

1993), with minor modifications as described by Smedsgaard 

(1997). The column used was a 50 × 2 mm Luna C-18 (II) 

reversed phase column (Phenomenex, CA, USA) fitted with 

a 2 × 2 mm guard column. 

Genotypic analysis 

The cultures used for the molecular studies were grown on 

malt peptone (MP) broth using 1 % (w/v) of malt extract (Brix 

10) and 0.1 % (w/v) bacto peptone (Difco), 2 mL of medium 

in 15 mL tubes. The cultures were incubated at 25 °C for 7 

d. DNA was extracted from the cells using the Masterpure 

yeast DNA purification kit (Epicentre Biotechnology.) 

according to the instructions of the manufacturer. The ITS 

region and parts of the b-tubulin and calmodulin genes were 

amplified and sequenced as described previously (Varga et 

al. 2007a–c). 

Data analysis 

The sequence data was optimised using the software package 

Seqman from DNAStar Inc. Sequence alignments were 

performed by MEGA v. 4.0 (Tamura et al. 2007) and improved 

manually. For parsimony analysis, PAUP v. 4.0b10 software 

was used (Swofford 2003). Alignment gaps were treated as a 

fifth character state and all characters were unordered and of 

equal weight. Maximum parsimony analysis was performed 

for all data sets individually using the heuristic search option 

with 100 random taxa additions and tree bisection and 

reconstruction (TBR) as the branch-swapping algorithm. 

Branches of zero length were collapsed and all multiple, 

equally parsimonious trees were saved. The robustness 

of the trees obtained was evaluated by 1000 bootstrap 

replications (Hillis & Bull 1993). Eurotium heterothallica was 

used as outgroup in these analyses (Houbraken et al. 2007). 

The alignments were deposited in TreeBASE () under accession number 

S11027. 

RESULTS AND DISCUSSION 

Phylogeny 

Of the aligned β-tubulin sequences, a portion with 438 

positions, including 107 parsimony informative characters, 

was selected for the analysis; MP analysis of the sequence 

data resulted in two similar, equally most parsimonious 

trees (tree length = 289 steps, consistency index = 0.7855, 

retention index = 0.7919), one of which is shown in Fig. 1. The 

calmodulin data set consisted of 492 characters, including 188 

198 

 

i m a f U N G U S

Aspergillus karnatakaensis sp. nov. 

“A. aeneus” NRRL 4649 

A. karnatakaensis CBS 102799 


A. aeneus NRRL 4769 

A. aeneus CBS 128.54 

ARTICLE 

E. discophora CBS 469.88 


A. eburneocremeus CBS 130.54 

A. eburneocremeus NRRL 4773 

E. foeniculicola CBS 156.80 

E. bicolor CBS 425.77 

E. spectabilis CBS 429.77 

A. heyangensis CBS 101751 

A. crustosus CBS 478.65 

A. crustosus NRRL 4988 

E. heterothallica CBS 489.65 

Fig. 1. One of the two equally MP trees obtained based on phylogenetic analysis of b-tubulin sequence data of Aspergillus sect. Aenei. Numbers 

above branches are bootstrap support values. Only values above 70 % are indicated. 















A. crustosus NRRL 4988 

Fig. 2. The single MP tree obtained based on phylogenetic analysis of calmodulin sequence data of Aspergillus sect. Aenei. Numbers above 

branches are bootstrap support values. Only values above 70 % are indicated. 

v o l u m e 1 · n o . 2 

199

Varga et al. 

ARTICLE 







A. eburneocremeus NRRL 4773 









Fig. 3. One of four equally MP trees obtained based on phylogenetic analysis of ITS sequence data of Aspergillus sect. Aenei. Numbers above 

branches are bootstrap support values. Only values above 70 % are indicated. 

parsimony informative sites; MP analysis resulted in a single 

most parsimonious tree (length = 485, consistency index = 

0.7402, retention index = 0.8040), which is presented in Fig. 

2. The ITS data set consisted of 451 characters, including 

43 parsimony informative sites; MP analysis resulted in four 

equally most parsimonious trees (length = 105, consistency 

index = 0.8190, retention index = 0.8541), one of which is 

presented in Fig. 3. 

The two isolates from Karnataka, India were found to be 

closely related to Aspergillus aeneus based on phylogenetic 

analysis of protein coding sequences (Figs 1, 2), and had 

identical ITS sequences to A. aeneus (Fig. 3). One additional 

isolate also from India, “A. aeneus” NRRL 4649 (= IMI 086833) 

was found to be conspecific with these two isolates. This 

isolate was obtained from soil of the Machrar river bed in 

the district of Bansa, Madhya Pradesh (Rai et al. 1964), and 

is morphologically similar to the other two Indian isolates. 

The three isolates are described here as a new taxon, A. 

karnatakaensis sp. nov. A typical characteristic is the formation 

of a crust of Hülle cells. The strains were incubated on various 

media for ascoma production, but in none of the strains were 

ascomata or ascospores found. Also, a mating experiment 

with the three strains did not induce ascoma production. 

Aspergillus karnatakaensis formed a well-supported 

clade together with four Emericella species, E. foeniculicola, 

E. bicolor, E. spectabilis and E. discophora, and four species 

known to reproduce only asexually, including A. aeneus, 

A. eburneocremeus, A. crustosus and A. heyangensis 

on the trees based on calmodulin (Fig. 4), β-tubulin, and 

ITS sequence data (data not shown). Based on these 

observations, we describe Aspergillus sect. Aenei sect. nov. 

to accommodate these species within subgenus Nidulantes. 

This group of species was originally assigned to section 

Nidulantes (Raper & Fennell 1965, Christensen et al. 1978, 

Samson 1979, Udagawa & Muroi 1979, Sun & Qi 1994). 

Extrolites 

Aspergillus karnatakaensis isolates were found to produce 

karnatakafurans A and B (Manniche et al. 2004), terrein, 

gregatins, asteltoxin (until now only detected in CBS 102799) 

and the partially characterised metabolite NIDU. Both 

gregatins and NIDU are also produced by A. granulosus, 

while karnatakafurans are produced in common with A. 

aeneus and A. multicolor. However, phylogenetic analysis 

of sequence data of A. multicolor (Peterson 2008) and A. 

granulosus (Houbraken et al. 2007) indicated that they are 

not closely related to A. karnatakaensis, while A. aeneus is. 

Among the other species found to belong to the same 

clade as A. karnatakaensis, Emericella bicolor produces 

sterigmatocystin, versicolorins, some anthraquinones, and a 

polar extrolite with end-absorption; E. foeniculicola produces 

sterigmatocystin (and many other sterigmatocystin and 

versicolorin-related compounds), xanthocillin derivatives, 

and the partially characterized (but common) metabolite DRI; 

E. spectabilis produces two members of the shamixanthone 

biosynthetic family (both more polar than shamixanthone itself) 

and a member of the sterigmatocystin biosynthetic family; 

A. heyangensis produces a decaturin in common with A. 

aeneus and A. karnatakaensis and NIDU, while E. discophora 

produces sterigmatocystin and versicolorins (Zalar et al. 

200 

 

i m a f U N G U S


E. gemmata CBS 853.96 

E. quadrilineata CBS 591.65 

E. nidulans CBS 589.65 

E. corrugata CBS 191.77 

E. falconensis CBS 271.91 

E. striata NRRL 4699 

ARTICLE 

E. cleistominuta CBS 200.75 

E. rugulosa CBS 171.77 

E. echinulata CBS 120.55 

E. violacea NRRL 2240 

E. violacea NRRL 4178 

E. foveolata CBS 279.81 

E. desertorum CBS 654.73 

E. navahoensis CBS 351.81 

E. fruticulosa CBS 989.72 

A. recurvatus NRRL 4902 

E. unguis CBS 132.55 

E. variecolor CBS 598.65 

E. astellata CBS 135.55 

A. caespitosus NRRL 1929 

E. undulata CBS 261.88 

A. aureolatus NRRL 5126 

A. asperescens CBS 110.51 

E. purpurea CBS 754.74 

A. sydowii CBS 593.65 

Sec%on 

Nidulantes 

A. versicolor NRRL 4642 

A. versicolor NRRL 227 

A. tabacinus NRRL 4791 

A. versicolor CBS 583.65 

A. protuberus CBS 602.74 

A. protuberus CBS 110382 









A. karnatakaensis NRRL 4649 



Fig. 4. Phylogenetic affinities of Aspergillus section Aenei to section Nidulantes based on neighbor-joining analysis of calmodulin sequence data 

of selected species assigned to these sections. Numbers above branches are bootstrap values. Only values above 70 % are indicated. 

2008). Decaturins are antiinsectan metabolites which have 

previously been identified in Penicillium species including P. 

thiersii and P. decaturense (Zhang et al. 2003, Li et al. 2005). 

Aspergillus eburneocremeus has both sterigmatocystin and 

mer NF-8054X in common with E. heterothallica. Aspergillus 

crustosus is different from all these species in producing only 

PR-toxin and related mycotoxins, and has no extrolites in 

common with the other species in sect. Aenei. All Emericella 

species in sect. Aenei produce sterigmatocystin, while the 

Aspergillus species without a known teleomorph apparently 

cannot produce it, with the exception of A. eburneocremeus. 

However, sterigmatocystin is common throughout the different 

sections of subgenus Nidulantes, and has even been found 

in sections Ochraceorosei and Flavi (Frisvad et al. 2005). 

Other extrolites such as shamixanthones, mer NF-8054X and 

the related emesterones, and terrein have also been found in 

other species in section Nidulantes. Aspergillus heyangensis 

is only known from ex-type cultures and re-examination of 

the cultures showed that the taxon has great similarities with 

the species mentioned above, including its inability to grow 

at 37 °C, and the shape of the conidial heads and vesicles, 

although this species does not produce Hülle cells (Fig. 5). 

That species also produces the unknown metabolite NIDU, 

as do A. karnatakaensis and E. discophora (Table 2). 

v o l u m e 1 · n o . 2 

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Varga et al. 

ARTICLE 

Fig. 5. Aspergillus heyangensis (CBS 101751). A–C. Colonies incubated at 25 °C for 7 d; A on CYA, B on MEA, C on CREA. D–I. Conidiophores 

and conidia. Bars = 10 µm. 

202 

 

i m a f U N G U S


Table 2. Extrolites produced by members of Aspergillus section Aenei. 

Species Culture collection number Extrolites 

A. aeneus IMI 069855ii = CBS 128.54 asteltoxin, fumitremorgin B, karnatakafurans, a decaturin, GUUM* 

A. crustosus IMI 135819 = CBS 478.65 PR-toxin 

A. eburneocremeus IMI 069856 = CBS 130.54 mer-NF 8054X, sterigmatocystin 

A. heyangensis CBS 101751 = IBT 29634 a decaturin, NIDU* 

A. karnatakaensis IBT 22154 = CBS 102799 asteltoxin, gregatins, karnatakafuran A and B, quinolactacin, terrein, NIDU*, 

GUUM* 

A. karnatakaensis IBT 22153 = CBS 102800 asteltoxin, gregatins, karnatakafuran A and B, physcion, quinolactacin, terrein, 

NIDU*, GUUM* 

A. karnatakaensis IMI 086833ii = WB 4649 a decaturin, karnatakafuran A and B, terrein, GUUM* 

E. bicolor CBS 425.77 = IBT 22833 sterigmatocystin 

E. discophora CBS 469.88 = IBT 21910 sterigmatocystin 

E. discophora CBS 470.88 = IBT 21911 sterigmatocystin, NIDU* 

E. foeniculicola CBS 156.80 = IBT 22831 DRI, sterigmatocystin, xanthocillin FA 

E. heterothallica WB 5097 = IBT 22604 DRI, emeheteron, sterigmatocystin, mer-NF 8054X, stellatin 

E. heterothallica CBS 489.65 = WB 5096 = IBT 22607 DRI*, NIDU*, versicolorins, mer-NF 8054X 

E. heterothallica WB 4981 = IBT 22605 DRI*, sterigmatocystin, mer-NF 8054X 

E. heterothallica WB 4983 = IBT 22606 DRI*, sterigmatocystin, mer-NF 8054X 

E. spectabilis CBS 429.77 = IBT 22891 extrolites with shamixanthone chromophore, trace of sterigmatocystin 

ARTICLE 

*NIDU, GUUM, and DRI are common extrolites with a characteristic UV chromophore. Their structure has not been elucidated yet. 

Aspergillus karnatakaensis Varga, Frisvad & 

Samson, sp. nov. 


(Fig. 6) 

Coloniis Emericellae similibus. Conidiophoris cum stipitibus laevibus, 

conidiis subglobosis vel late ellipsoideis. Aggregationibus insignibus 

cum tegumento ex cellulis globosis efferentibus. 

Typus: India: Karnataka, near Chickmagalur, Netraconda 

Estate, isolated from soil under coconut palm (Cocos nucifera) 

in coffee plantation, 20 Dec. 1996, J.C. Frisvad (CBS H-20502 

-- holotypus, culture ex-holotype CBS 102800). 

Colonies on CYA, at 25 °C: 31–37 mm diam after 7 d, reverse 

orange; on MEA, at 25 °C: 12–19 mm, reverse yellow; on 

YES, at 25 °C: 33–45 mm, reverse pink to raspberry-red 

reverse; on OAT, at 25 °C: 16–23 mm, Hülle cells present; on 

CYA, at 37 °C: no growth to micro-colony (

Varga et al. 

ARTICLE 

Fig. 6. Aspergillus karnatakaensis (CBS 102800). A, B. Colonies incubated at 25 °C for 7 d, A on CYA, B on MEA. C, Crusts of Hülle cells,. D, 

E, and G–I. Conidiophores and conidia. F. Hülle cells. Bars = 10 µm, except F = 100 µm. 

204 

 

i m a f U N G U S



We are grateful to Dr R Naidu for permission to sample soil for 

mycological examinations in the Coffee Research Station and 

associated estates near Chickmagalur, Karnataka, India. Tineke van 

Doorn helped with the morphological data and Uwe Braun kindly 

provided the Latin diagnosis. We are also indebted to our referees. 

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text article 

INSTRUCTIONS TO AUTHORS 

EDITORIAL BOARD

Editorial (1) 

News 

The home stretch for fungal barcoding (2) 

Fungi and the Convention on Biological Diversity (3) 

Mycologists go political, with success (4) 

Mushrooms – the new plastic? (4) 

White truffles still demand a high price (4) 

Fungi on German TV: focus on black fungi (5) 

Reports 

IMC9: The Biology of Fungi. A personal reflection (8) 

International Commission on the Taxonomy of Fungi (ICTF) (11) 

Awards and Personalia 

IMA Medals awarded at IMC9 (15) 

Order of Canada: Stanley J Hughes (17) 

Obituary: C Terence Ingold (1905–2010) (17) 

IMA Young Mycologist Awards (18) 

Correspondence 

A vision for the future of the ICTF (20) 

Research News 

Carbonaceous spherules exposed as fungal sclerotia, are not evidence of the impact of a comet (22) 

Physiological differences between wet- and dry-distributed conidia (23) 

Mobile chromosomes: the clue to pathogenicity in Fusarium species (23) 

Ectomycorrhizal symbiosis in basidio- and ascomycete fungi have different origins (24) 

A basal bryophilous fungus associates with cyanobacteria (24) 

Molecular clocks and evolutionary rates (25) 

Numbers of fungi in China (26) 

Society and Association News 

International Society for Fungal Conservation (27) 

Mycological Society of America (30) 

Iranian Mycological Society (31) 

Book News (32) 

Forthcoming Meetings (36) 

Articles 

“The enigma of Calonectria species occurring on leaves of Ilex aquifolium in Europe” by Christian Lechat, Pedro W. Crous and Johannes 101 

Z. Groenewald 

“How to describe a new fungal species” by Keith A. Seifert and Amy Y. Rossman 109 

“What is Johansonia?” by Pedro W. Crous, Robert W. Barreto, Acelino C. Alfenas, Rafael F. Alfenas and Johannes Z. Groenewald 117 

“The history, fungal biodiversity, conservation, and future perspectives for mycology in Egypt” by Ahmed M. Abdel-Azeem 123 

“IMC9 Edinburgh Nomenclature Sessions” by Lorelei L. Norvell, David L. Hawksworth, Ronald H. Petersen and Scott A. Redhead 143 

“Fungal phoenix rising from the ashes?” by Michael J. Wingfield, Martin P.A. Coetzee, Pedro W. Crous, Diana Six and Brenda D. Wingfield 

149 

“Modelling fungal colonies and communities: challenges and opportunities” by Ruth E. Falconer, James L. Bown, Eilidh McAdam, Paco 155 

Perez-Reche, Adam T. Sampson, Jan van den Bulcke and Nia A. White 

“Colletotrichum: species, ecology and interactions” by Ulrike Damm, Riccardo Barroncelli, Lei Cai, Yasuyuki Kubo, Richard O’Connell, 161 

Bevan Weir, Kae Yoshino and Paul F. Cannon 

“Cryptic species in lichen-forming fungi” by Ana Crespo and H. Thorsten Lumbsch 167 

“Sex in Penicillium series Roqueforti” by Jos Houbraken, Jens C. Frisvad and Robert A. Samson 171 

“Anaerobic fungi: Neocallimastigomycota” by Gareth W Griffith, Scott Baker, Kate Fliegerova, Audra Liggenstoffer, Mark van der 

181 

Giezen, Kerstin Voigt and Gordon Beakes 

“Polyphasic taxonomy of Aspergillus section Sparsi” by János Varga, Jens C. Frisvad and Robert A. Samson 187 

“Aspergillus sect. Aeni sect. nov., a new section of the genus for A. karnatakaensis sp. nov. and some allied fungi” by János Varga, Jens C. 197 

Frisvad and Robert A. Samson

Volume 1 Â· No. 2 Â· December 2010 V o lu m e 1 Â· N o ... - IMA Fungus

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