elements in mushrooms-24415-en 2011.pdf - JRC Publications ...

Chemical elements in Ascomycetes and Basidiomycetes 

The reference mushrooms as instruments for investigating 

bioindication and biodiversity 

Roberto Cenci, Luigi Cocchi, Orlando Petrini, 

Fabrizio Sena, Carmine Siniscalco, Luciano Vescovi 

Editors: R. M. Cenci and F. Sena 

EUR 24415 EN 2011 

1

The mission of the JRC-IES is to provide scientific-technical support to the European Union’s policies 

for the protection and sustainable development of the European and global environment. 

European Commission 

Joint Research Centre 

Institute for Environment and Sustainability 

Via E.Fermi, 2749 I-21027 Ispra (VA) Italy 

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Commission is responsible for the use which might be made of this publication. 

2 

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A great deal of additional information on the European Union is available on the Internet. 

It can be accessed through the Europa server http://europa.eu/ 

JRC Catalogue number: LB-NA-24415-EN-C 

Editors: R. M. Cenci and F. Sena 

JRC65050 

EUR 24415 EN 

ISBN 978-92-79-20395-4 

ISSN 1018-5593 

doi:10.2788/22228 

Luxembourg: Publications Office of the European Union 

Translation: Dr. Luca Umidi 

© European Union, 2011 

Reproduction is authorised provided the source is acknowledged 

Printed in Italy

Attached to this document is a CD containing: 

• A PDF copy of this document 

• Information regarding the soil and mushroom sampling site 

locations 

• Analytical data (ca, 300,000) on total samples of soils and 

mushrooms analysed (ca, 10,000) 

• The descriptive statistics for all genera and species analysed 

• Maps showing the distribution of concentrations of inorganic 

elements in mushrooms 

• Maps showing the distribution of concentrations of inorganic 

elements in soils 

3

4 

Contact information: 

Address: Roberto M. Cenci - European Commission - DG JRC Institute for Environment and 

Sustainability, Land Management and Natural Hazards Unit, T.P. 280, I-21027 Ispra (VA), 

Italy 

E-mail: roberto.cenci@jrc.ec.europa.eu 

Tel: +39 0332 789771 

Fax: +39 0332 786394 

http://eusoils.jrc.it/index.html 

Address: Luigi Cocchi – Member of the National Steering Group, the Organisational Office of the 

National Scientific Committee and of the Commission of Mycotoxicology belonging to the 

Bresadola mycological Association. Vice President of the “Renzo Franchi” mycologicalnaturalist 

group in Reggio Emilia, Italy. 

E-mail: luigi.cocchi@libero.it 

Address: Orlando Petrini – Istituto Cantonale di Microbiologia, via Mirasole, 22A, CH-6500 

Bellinzona, Republic and Canton of Ticino, Switzerland 

E-mail: orlando.petrini@ti.ch 

Tel: +41 91 814 6031 

Fax: +41 91 814 6019 

http://www.ti.ch/dss/DSP/ISTCM/ 

Address: Fabrizio Sena - European Commission - DG JRC Institute for Environment and 

Sustainability, Rural, Water and Ecosystem Resources Unit, T.P. 270, I-21027 Ispra (VA), 

Italy 

E-mail: fabrizio.sena@jrc.ec.europa.eu 

Tel: +39 0332 785399 

Fax: +39 0332 786645 

http://ies.jrc.ec.europa.eu/rural-water-and-ecosystem-resources-unit 

http://www.jrc.ec.europa.eu/ 

http://ies.jrc.ec.europa.eu/ 

Address: Carmine Siniscalco - Istituto Superiore per la Protezione e la Ricerca Ambientale (ISPRA), 

Dipartimento Difesa della Natura, “Progetto Speciale Funghi”, via Curtatone, 3, I-00185 

Roma, Italy 

E-mail: carmine.siniscalco@isprambiente.it 

Tel: +39 06 5007 4302 

Fax: +39 06 5007 4013 

http://www.isprambiente.it 

Address: Luciano Vescovi – Technician at Enia S.p.A. Laboratories, Reggio Emilia, Italy 

E-mail: luciano.vescovi@eniaspa.it

Chemical elements in Ascomycetes and Basidiomycetes 

The reference mushrooms as an instrument 

for investigating bioindication and biodiversity 

Fungi in the wild are among the principal 

agents in biogeochemical cycles; those 

cycles of matter and energy that enable 

ecosystems to work. 

By investigating the biodiversity of Italian 

fungal species and concentration levels of 

chemical elements in them, it may be 

possible to use these fungi as biological 

indicators for the quality of forest, woodland 

and semi-natural environments. 

The database of this EUR Report record the 

dry-material concentrations of 35 chemical 

elements, including heavy metals, in over 

9,000 samples of higher mushrooms 

(Ascomycetes and Basidiomycetes). These 

samples represent approximately 200 

genera and a thousand species. As the 

database has attained statistical stability it 

has been possible to define the concept of a 

“reference mushroom”. The use of a 

“reference mushroom” may benefit – 

perhaps only as a methodological approach 

– various fields of mycological and 

R. M. Cenci, L. Cocchi, O. Petrini, 

F. Sena, C. Siniscalco e L. Vescovi 

environmental research; from biodiversity 

and bioindication, through taxonomy right up 

to health and sanitation issues. 

The sheer volume of the collected data may 

prove to be useful as a comparison for data 

collected in the future; such results would 

also allow a better and more exhaustive 

interpretation of the effects of environmental 

protection laws that have been in place over 

the years to reduce or remedy current 

climate change phenomena and the 

environmental damage caused by human 

activity. 

Studies pertaining to the frequency of 

occurrence and the ecology of the various 

fungal species found on Italian soil have 

tended to link the reference habitats used to 

European classification guidelines (Natura 

2000, CORINE Land Cover, CORINE 

Biotopes and EUNIS). Thereby the 

foundations have been laid for the use of 

mushrooms as biological indicators for the 

measurement of soil and ecosystem quality. 

5

6 

Thanks 

The following people contributed to the creation of this document: 

DR. ANNA BENEDETTI (Paragraph 2.4) 

Consiglio per la Ricerca e la sperimentazione in Agricoltura - Centro per lo Studio delle 

Relazioni tra Pianta e Suolo [Advisory board for Research and Experimentation in Agriculture – 

Centre for the study of plant-soil relationships] 

(anna.benedetti@entecra.it) 

DR. PIETRO MASSIMILIANO BIANCO (Paragraph 2.3) 

Istituto Superiore per la Protezione e la Ricerca Ambientale - Dipartimento Difesa della Natura 

[Superior Institute for Environmental Protection and Research – Nature Protection Department] 

(pietro.bianco@isprambiente.it) 

PROF. MANUELA GIOVANNETTI (Paragraph 2.2) 

Dipartimento di Biologia delle Piante Agrarie - Università di Pisa 

[Department of Agrarian Plant Biology – University of Pisa] 

(mgiova@agr.unipi.it) 

DR. CARLO JACOMINI (Paragraphs 2.3 and 2.4) 

Istituto Superiore per la Protezione e la Ricerca Ambientale - Dipartimento Difesa della Natura 

[Superior Institute for Environmental Protection and Research – Nature Protection Department] 

(carlo.jacomini@isprambiente.it) 

DR. STEFANO MOCALI (Paragraph 2.4) 

Consiglio per la Ricerca e la sperimentazione in Agricoltura - Centro per lo Studio delle 

Relazioni tra Pianta e Suolo [Advisory board for Research and Experimentation in Agriculture – 

Centre for the study of plant-soil relationships] 

(stefano.mocali@entecra.it) 

DR. LILIANE PETRINI (Paragraphs 2.1, 2.5 and 3.1) 

Lugano (CH) 

The creation of such broad and complex books always requires specific, detailed and thorough 

information that can only come from experts. We would like to thank everyone who helped us with 

their valuable contributions, without which this work would not have been possible, and in particular: 

DR. FAYÇAL BOURAOUI 

Comunità Europea - Centro Comune di Ricerca di Ispra - Istituto dell’Ambiente e Sostenibilità 

[European Community – Ispra Communal Research Centre – Environment and Sustainability 

Institute] 

(faycal.bouraoui@jrc.ec.europa.eu)

DR. NAZARIA MACCHI 

Servizio Geologico, Sismico e dei Suoli della Regione Emilia Romagna 

[Geological, seismic and soil-information service of the Emilia Romagna region] 

(nmarchi@regione.emila-romagna.it) 

PROF. GIUSEPPE RASPA 

Dipartimento di Ingegneria Chimica e dei Materiali e Ambiente - Università “Sapienza” di Roma 

[Department of Chemical Engineering and Materials and Environments – Sapienza University, 

Rome] 

(giuseppe.raspa @ uniroma1.it). 

We also wish to thank: 

Franco BERSAN, Enrico BIZIO, Giorgio BUIZZA, Luca CAMPANA, Emanuele CAMPO, Maurella 

CASTOLDI, Maurizio CHIARI, Paolo FRANCHI, Luca GORRERI, Pier Giovanni JAMONI, Angela 

LANTIERI, Giorgio MARASCA, Mauro MARCHETTI, Giovanni MONTI, Carlo PAPETTI, Giovanni 

ROBICH, Mauro SARNARI, Cesare TUGLIOZZI, Gianfranco VISENTIN, the Gruppo Micologico 

dell’Etruria meridionale – AMB, the AMB Archives – CSM, the Società Veneziana di Micologia – 

AMB, all the mycologists and mycological group participants at the AMB who have submitted their 

findings to the herbarium at the Natural History Museum of Venice. 

A heartfelt thank you goes to members and friends who have worked for more than twenty years, in 

various ways and with different skills, on the work presented here.. It is therefore fitting and correct to 

offer collective thanks to the Associazione Micologica Bresadola and special thanks also to all 

members of the Comitato Scientifico Nazionale and the Gruppo Micologico e Naturalistico “R. 

Franchi” in Reggio Emilia. 

We thank the Province of Reggio Emilia for the sponsorship and support bestowed in 2004 to 

researching the “Presence of Chemical elements in higher mushrooms”. 

We also wish to thank the following people for their help in collecting mushroom samples: 

Maria Luisa BORRETTINI, President of the GGEV group in the Province of Reggio Emilia; Gioacchino 

PEDRAZZOLI, ex-President of the Emilia-Romagna WWF and member of the Consiglio Direttivo del 

Parco nazionale dell’Appennino Tosco-Emiliano; Fabio SIMONAZZI, from the Centro di Informazione 

ed Educazione Ambientale dei Territori Canossani della Val d'Enza; Roberto BARBANTINI, Davide 

BOTTAZZI, Tito FABBIANI, Willy REGGIONI and Stefania ZANNINI, wardens of the Regional Park of 

Alto Appennino Reggiano - Parco del Gigante (the Giants’ Park), an entity closed down in 2005 

following the creation of the Parco nazionale dell'Appennino Tosco-Emiliano. We also thank Dr. 

Bruno CAVALCHI, ex-Chairman of ARPA (ex PMP) in Reggio Emilia and Dr. Roberto SOGNI of 

ARPA (ex PMP) in Piacenza. 

A special thank you goes to Tjakko STIJVE (St. Légier La Chiesàz, CH) for critical and constructive 

discussions, evaluations and data supplied. 

We also wish to thank Mr. Pietro SPAGNI. 

7

Index 

INDEX ............................................................................................................................................. 9 

1 PRESENTATIONS............................................................................................................. 11 

1.1 ICM ............................................................................................................................. 11 

1.2 AMB............................................................................................................................ 12 

1.3 ISPRA.......................................................................................................................... 13 

1.4 EU - CCR - IES............................................................................................................ 14 

1.5 ENIA ........................................................................................................................... 15 

2 INTRODUCTION............................................................................................................... 17 

2.1 MUSHROOM TAXONOMY .............................................................................................. 17 

2.2 BIOLOGICAL NOTES ON MUSHROOMS ........................................................................... 21 

2.3 MUSHROOMS AND ENVIRONMENTS FOR GROWTH......................................................... 25 

2.4 MUSHROOMS AS A SOIL-QUALITY BIOINDICATOR......................................................... 35 

2.5 THE REFERENCE MUSHROOM........................................................................................ 46 

2.6 BIODIVERSITY AND BIOINDICATION: EC AND INTERNATIONAL LEGISLATION............... 53 

3 DATA SYNTHESIS............................................................................................................ 55 

3.1 CONSIDERATION OF STATISTICS AND THE STATISTICAL METHODS EMPLOYED.............. 55 

3.2 APPLIED GEOSTATISTICAL ANALYSIS ........................................................................... 59 

4 MATERIALS AND METHODS........................................................................................ 63 

4.1 METHODS FOR CHEMICAL ANALYSIS OF SOIL AND MACROMYCETES ............................. 63 

4.2 DISTRIBUTION MAP OF ELEMENTS IN SOIL .................................................................... 64 

4.3 DISTRIBUTION MAP OF ELEMENTS IN THE MUSHROOMS................................................ 70 

4.4 SAMPLING: A DATA SHEET EXAMPLE .......................................................................... 107 

5 CONCLUSIONS ............................................................................................................... 111 

6 BIBLIOGRAPHY ............................................................................................................. 113 

7 APPENDIX........................................................................................................................ 121 

9

1.1 ICM 

Even if it is not always apparent, mushrooms affect 

our daily lives in many important ways. They are 

indispensible elements for food production, and yet 

many of them, as plant pathogens, cause great 

agricultural damage. Others, fortunately only a few 

– although their numbers are growing – are 

pathogenic agents for animals; including man. 

Mycorrhizae are symbiotic associations between 

fungi and plants that are important in agriculture 

and, last but not least, several species of 

basidiomycetes are valuable edible mushrooms. 

Therefore, it is important to know and understand 

mushrooms, and even more important to classify 

them accurately. 

For years the taxonomy of basidiomycetes and 

ascomycetes was based almost entirely on 

morphological characters; we can see this today in 

the identification keys still used by amateur and 

professional mycologists alike. However, 

mycologists were quick to realise that morphology 

was not enough to build exhaustive and trustworthy 

classifications, especially when the organisms to be 

classified were of simple shape or when the 

morphological features were limited or varied little. 

Therefore attempts were made to classify these 

organisms not only by their morphology, but also 

by their physiological and biochemical 

characteristics. Even at the beginning of the 20 th 

Century, for example, staining and biochemical 

reactions were being studied in bacteriology: these 

same properties are now used in mycology. 

Orlando Petrini 

Chapter I 

Presentations 

Director of the Istituto Cantonale di Microbiologia, Bellinzona, Switzerland 

Thus arose the problem of reconciling 

morphological classification with the type of 

classification established by genetic and 

biochemical methods. In fact, genetic analysis often 

leads us into creating taxonomic schemata which 

are not, certainly at first glance, entirely compatible 

with existing classifications. A trustworthy 

classification must take into consideration not just 

phylogenetic properties (being connected to the 

evolution of organisms over time) and phenotype 

(the observable morphology and physiology of an 

organism), but also ecological peculiarities. Such 

an approach is commonly referred to as “polyphase 

taxonomy”. 

The work contained in this book provides a new 

and important piece of the puzzle which is the 

taxonomy of mushrooms. I hope that these data 

will be of help to taxonomists in completing their 

research and that they might become part of a 

taxonomic scheme which will go towards resolving 

the difficult issue of the definition of taxa. By their 

very nature, these data are also relevant from 

physiological and ecological viewpoints. I hope 

therefore that this book will also be of service to 

physiologists who aim to better understand the 

biochemical aspects connected to absorption and 

retention of chemical elements in mushrooms, and 

that it will serve the ecologists who are trying to 

achieve a complete picture of the biodiversity of 

these fascinating organisms. 

11

1.2 AMB 

Research into the presence of chemical elements, 

including heavy metals and radioactive isotopes, in 

higher mushrooms began in 1986 at the “Renzo 

Franchi” mycological-naturalist group in Reggio 

Emilia, Italy and was then taken up by the National 

Scientific Committees of the Bresadola 

Mycological Association (AMB). This is not a 

mere “historical fact”, but instead attests to an 

essential aspect of the scenario: given the size of 

the project and the level of skills it involved, this 

research could not have been achieved outside of 

the AMB and of its network of associated groups. 

One of the most delicate issues here, as in all 

analogous studies into higher mushrooms, is that of 

having a positive identification of the fungal 

exemplars being studied and analysed: herein lies 

the skill and know-how that the AMB provides at a 

level which is widely recognised as the gold 

standard. We believe this characteristic is 

something over which we should be highly 

protective, in a spirit of militant volunteerism, 

fuelled by passion, respect and love for nature and 

its equilibrium. 

Such equilibrium is increasingly necessary for the 

future of humankind; understanding it and its 

mosaic of different pieces, possibly thanks to 

collaboration and communication between various 

scientific disciplines, is becoming an essential task 

for all those who hold the future of the planet to 

heart. 

The piece that we bring to this mosaic is our 

knowledge of mycological classification and 

taxonomy, a science created by the great naturalists 

such as Linnaeus, and which, in Italy, culminated 

with the lofty heights of Giacomo Bresadola. 

This knowledge is born of the intense work carried 

Luigi Villa 

President of the Associazione Micologica Bresadola 

12 

out by our association, which consists of 130 

groups from around the country and over 12,000 

members. Our members are involved in capillary 

networks of activities, which include research, 

information and education programmes regarding 

mycology and the environment through hundreds 

of local initiatives, ranging from mycology courses 

and mycological exhibitions to mycological studies 

and investigations. 

The AMB is also an important national and 

international mycological publisher, with two 

magazines to its name Rivista di Micologia 

(Mycology Magazine) and Pagine di Micologia 

(Mycology Pages) and various books that deal with 

basic mycology all the way up to highly-specialised 

tomes. 

Two National Scientific Committee meetings are 

held each year: one in spring, the other in autumn, 

and these involve hundreds of mycologists. 

The work of the National Scientific Committee 

meetings is the main driver of the National 

Herbarium, which has now surpassed 10,000 dry 

specimens and has recently been admitted to the 

Index Herbariorum. 

The recognition that the publication of this EU 

report constitutes for a research project that has 

lasted over 20 years is not only thanks to the 

authors and their patience and dedication, but 

should be extended to the whole AMB. 

The fact that this research has led to fruitful 

collaboration with European Community 

institutions, the Italian state, with private 

companies and non-EC research institutes, such as 

the Istituto cantonale di Microbiologia, in 

Bellinzona, Switzerland has brought the statutory 

purposes of the AMB to the highest levels.

1.3 ISPRA 

The “Special Mushrooms Project” by the ISPRA’s 

“Natural Protection” Department promotes and 

carries out studies on fungal species and is thereby 

well placed in the project using them as indicators 

of environmental quality. 

Mushrooms are important diversity indicators in 

terms of population richness and abundance at a 

genetic level and can therefore be used in the study 

and monitoring of biodiversity in a particular 

environment or ecosystem. 

One of the 16 research themes in the “Special 

Mushrooms Project” was aimed at developing an 

IT system for micotoxicological aspects, including 

phenomena of bioaccumulation and 

bioconcentration of heavy metals and xenobiotic 

substances in the mushrooms. The idea behind this 

was to facilitate bioremediation plans for degraded 

environments and also to promote studies regarding 

the health and hygiene aspects related to the human 

consumption of mushrooms. 

The work described in this volume, edited by JRC- 

IES, is the fruit of collaborations between five 

different institutions that have together been able to 

create one of the first ever applications for 

bioindication using mushrooms. 

The work springs from the results of an intense 

sampling campaign carried out in Italy by the 

Andrea Todisco 

Direttore Dipartimento Difesa della Natura di ISPRA, Roma, Italia 

Associazione Micologica Bresadola which was 

started over 20 years ago and has conducted 

chemical analysis on over 9,000 mushroom 

samples and which has led through statistical 

inference to the definition of a “reference 

mushroom”. These last two aspects will go on to 

play a fundamental role when the fungal diversity 

indicators for richness and abundance at the genetic 

level are defined. By linking fungal species to 

habitats it will, in fact, be possible to use 

mushrooms to study and monitor the biodiversity 

of an ecosystem or environment with numerous 

elements of evaluation. Those species occurring 

more frequently will act as the first sample of 

ecological value and as environmental quality 

indicators. 

In the near future then, we will have a deeper 

understanding of both those mechanisms which 

maintain and regulate the evolution of ecosystems 

and a new knowledge of those mycotoxicological 

aspects which will be used to inform protective 

legislation regarding human mushroom 

consumption. 

13

1.4 CCR - IES 

It was a great pleasure to be asked to write a small 

preface to the volume edited by Roberto Cenci, 

Fabrizio Sena and colleagues. Through this work 

there flows a clear love for the environment and a 

strongly-motivated scientific interest in a littleunderstood 

or studied field: mushrooms. 

The study of soil ecology and, more specifically, 

the use of mushrooms to evaluate the health of the 

soils in which they grow would, on the face of it, 

be a complex affair; both due to the limited 

knowledge we have about the field today and also 

to the objective difficulties presented by the 

formulation of a model by which we might “read” 

the soil. The book is a complete and exhaustive 

illustration of the characteristics of mushrooms and 

their role in the soil compartment. At the same time 

it lays out a solid base for the use of biodiversity 

and bioindication as diagnostic measures to better 

understand the health and quality of soil. 

The impressive amount of detail given to trace 

elements in over 9,000 analyzed mushroom 

samples makes this book as a guide that covers 

many other fields, such as food and nutrition. We 

should bear in mind that fungi are used in many 

different foodstuffs, and so knowledge of heavy- 

Leen Hordijk 

Direttore dell’Istituto dell’Ambiente e della Sostenibilità, Ispra, Italia 

14 

metal presence in certain types would allow the 

creation of healthier products for the consumer. 

The data collected here may aid the future 

formulation of EC food guidelines. The database 

has already been bought by the Italian Health 

Ministry (its acquisition taking place in 2008) for 

use in debating and defining Community 

Regulation n. 628/2008. Another important aspect 

of this work is the vast sampling area covered and 

the high number of fungal species identified. These 

today are of fundamental importance but will be 

even more important in the future when further 

sampling campaigns will be carried out: at that 

stage we shall have a clear view of whether, and to 

what extent, the activities of man and climatic 

change influence the biodiversity of mushrooms. 

One final, but by no means unimportant effect of 

this book relates to the use of the information it 

contains as a database for the numerous experts in 

the field who will now be able to better describe, 

and so improve, their investigations. 

I am sure that this book will be recognised for its 

merits and that it will be a great aid and step 

forward to both international and national experts 

and researchers in the field of mycology.

1.5 ENIA 

We are honoured to be among the list of authors of 

this report, which stemmed from the historical 

connection between the “Renzo Franchi” 

mycological-naturalist group in Reggio Emilia, 

Italy and the Bresadola Mycological Association – 

Mycological Study Centre, which has previously 

seen so many fruitful collaborations. 

It may, on the face of it, seem rather odd that a 

multi-utility company such as ours should be 

working with groups so far away from our daily 

activity. In reality, however, the logic that moved 

and continues to move us is the goal of widening 

our vision and forging bonds with the most 

interesting partners in the territory so as to favour 

an exchange of knowledge to our mutual 

advantage. These choices have enabled Enia to 

establish a new concept of "territorial proximity” 

by going beyond the usual partners one would 

expect an energy and environmental services 

company to have and instead seeking out different, 

but like-minded companions to explore and travel 

the road ahead. 

As always, in this relationship we have given our 

best; above all we have made our professionality 

and know-how available, in this case those 

belonging to our Reggio Emilia Laboratories. This 

project represents a very positive exchange of 

knowledge and also a serious means of supporting 

Andrea Allodi 

Chairman of Enìa Spa, Parma, Italia 

research initiatives in Italy and it placed Enia 

among the authors of an instrument which extends 

far beyond Italy in its uniqueness. 

Allow me then one last thought, not just as 

Chairman of a company, but also as a Mycologist – 

a passion which allows me to appreciate the 

findings of this book even more. Often, when one 

thinks of the instruments of innovation and 

research, habit leads us to think of modernized 

gadgets and sharp, complicated machines, as cold 

as only machinery can be. However, sometimes we 

only have to look around us and nature comes to 

the rescue. In this case it is mushrooms that we 

have to thank; “simple” mushrooms – and I ask 

mycologists to forgive me this definition. 

These mushrooms lend us a help that should make 

us reflect upon and rethink our relationship with the 

world that surrounds us and of which we are part. It 

is a world which we ever more frequently know 

little about and often mistreat. We waste, dirty and 

even ruin a bounty which should be there for 

everybody. 

If the research in this book sets us to thinking about 

this, we will have reached another goal and 

provided another significant result which can be 

added to the extraordinary value of this Report. 

15

2.1 Mushroom taxonomy 

2.1.1 Introduction 

Mushrooms, as all living beings, are scientifically 

named according to their “binomial nomenclature”, 

which indicates their “genus” (capitalised) and their 

“species” (not capitalised); both names are usually 

italicised. The binomial denomination of living 

creatures, which had previously been partially used 

by Theophrastus and Pliny, was given a scientific 

basis by Linnaeus, who established “taxonomy” 

(based on a binomial denomination using Latin or 

Latinised names) as a real and proper scientific 

discipline. Linnaeus (Linnaeus, 1753) described 

fungi for the first time, grouping them into just ten 

genera. Since then the taxonomy of mushrooms and 

fungi has made great leaps forward. 

Later on, (1816 - 1817) Nees broadened and 

refined Linnaeus’s work by enhancing taxonomy. 

First considered as vegetables and then being 

assigned to the realm of plants, mushrooms were 

regarded from early on as "abnormal" bodies, the 

taxonomic position of which remained very 

unclear. These organisms attracted the attention of 

biologists, and, after some initial important work in 

Italy, Saccardo’s monumental efforts (1882-1931) 

culminated in a veritable encyclopaedia of 

mycology. 

Initially, fungi were classified into four divisions. 

The Basidiomycetes were distinguished from the 

Ascomycetes and from the Deuteromycetes largely 

based on morphological characters such as the way 

their spores formed, the colour and shape of those 

spores and the appearance of their bodies. Clearly, 

the most studied fungi were Basidiomycetes; 

visible to the naked eye and of interest for their 

gastronomic properties and economic value. 

Chapter II 

Introduction 

2.1.2 Classification of fungi into 

kingdoms and phyla 

Fungi are eukaryotes (therefore having a nucleus 

delimited by a nuclear membrane). Their cell walls 

contain chitin and glucans (and rarely cellulose) 

and their nucleus may be haploid or diploid, 

dikaryon, homokaryon or heterokaryon. The 

fructifications can be microscopic or macroscopic, 

differentiated or undifferentiated. Their ecological 

role is very varied and among themselves they may 

be symbionts, saprobes, parasites, commensals or 

hyperparasites. 

Fungi are fairly cosmopolitan: around 70,000 

species have been described, but Hawksworth 

(1991) estimates that there may be as many as 1.5 

million. Only around 300 are known pathogens for 

humans, while many cause plant diseases and a 

considerable number play a fundamental role in the 

ecosystem, both as destroyers of vegetable detritus 

and as symbionts (Mycorrhizae and lichens). 

Until 1980, taxonomists considered fungi a 

compact, though not necessarily homogenous, 

group. Müller and Löffler (1976), in their book on 

mycology (still considered a classic reference for 

mycologists) included not only the Ascomycetes, 

the Basidiomycetes, the Deuteromycetes and 

Zygomycetes, but also some groups of organisms 

that have since been transferred into the kingdoms 

of Protista and Chromista. In 1981, Cavalier-Smith 

(1981) proposed a separate kingdom for the “higher 

mushrooms” (the Ascomycetes, Basidiomycetes, 

Zygomycetes and their asexual forms, grouped 

together as Deuteromycetes) and transferred most 

of the “lower fungi” (single-celled organisms or 

hyphae, often with flagella spores) into the Protista 

and Chromista. Cavalier-Smith’s work, which 

showed how mushrooms are more similar to 

animals than to plants (fig. 1) opened the door to a 

more detailed re-elaboration of the taxonomy and 

phylogeny of fungi. 

Kendrick (1992) described the ideas of Cavalier- 

Smith very well in his book, “The Fifth Kingdom”. 

17

18 

Fig. 1. The “Tree of Life”, the position of fungi in the phylogenic tree . Source: “Tree of Life Project” (Maddison and Schulz, 2007) 

It was the same Cavalier-Smith who, over the 

following years, proposed ever more complex and 

detailed models (Cavalier-Smith, 1993; 1998; 

2004; 2006), which were then taken and 

formalised, at least as regards the kingdom of 

fungi, by Hibbett and Binder (2007) (fig. 2). 

Currently, living organisms are divided into seven 

kingdoms (Eubacteria, Archaebacteria, Archaezoa, 

Protozoa, Plantae, Animalia, Fungi). To this list 

should be added the kingdom of Chromista, itself 

an area of considerable controversy. 

Fig. 2. The phylogeny and classification of fungi. Source: Hibbett (Hibbett and Binder, 2007), amended. The length of the branches of the 

cladogram is not proportional to the genetic distance between taxa. 

Based on the work by Cavalier-Smith (1993) and 

Hibbett and Binder (2007) fungi, that until 1980 

were grouped together in one kingdom, are now 

subdivided as follows: 

• the kingdom of fungi includes the 

Ascomycota and Basidiomycota 

(Dikarya), the Glomeromycota (which 

includes the endotrophic mycorrhizae), 

the Chytridiomycota, the Neocallimastigomycota, 

the "Mucormycotina"

(traditionally Zygomycota), the 

Blastocladiomycota, the Entomophthoromycotina, 

the Zoopagomycotina, the 

Kickxellomycotina and a group of 

unicellular human parasites Microsporidia; 

• the myxomycetes have been transferred 

into the kingdom of Protozoa 

(Amoebozoa, Eumycetozoa); 

• the Oomycetidae and Thraustochytridae 

are now assigned to the kingdom of 

Chromista. 

This new classification is the result of intensive 

phylogenic work, based in particular on genetic 

analysis, but it is interesting to note that the results 

are in perfect concordance with previously stated 

hypothesis based on observations of morphology 

and physiology (Müller et al., 1976). 

2.1.3 Problems with taxonomy in 

mycology: fungal variability 

Within any wide taxon, not only molecular 

genetics, but also morphology and physiology are 

useful for reaching taxonomic conclusions. 

Morphologically speaking, fungi are incredibly 

variable. Beyond the morphological variations 

present both at a macroscopic (just think of the 

difference between porcini mushrooms and an 

amanita muscaria, for example) and at a 

microscopic level, fungi create further difficulties 

by their ability to express two distinct phenotypes 

in their reproductive and vegetative forms. 

Furthermore, often a fungus will produce just one 

form (sexual or vegetative) through having reduced 

or lost the capacity to reproduce either sexually or 

vegetatively (asexually). 

Fig 3. Example of a holomorph: Eurotium and its anamorph, Aspergillus. 

From a phylogenetic point of view, morphological 

analysis can lead to organisms being erroneously 

identified as belonging to two distinct taxa when, in 

reality, there exists only one, either in its sexual or 

asexual form. Only in cases where both forms are 

found together can a morphological analysis give a 

trustworthy classification; in other cases genetic 

analysis becomes indispensible. 

Mushrooms are pleomorphic and therefore assume 

a different shape depending not only on the type of 

reproductive organ they develop, but also on the 

environmental and physiological conditions 

influencing their growth. 

Regarding pleomorphic growth in different 

physiological and ecological conditions, see the 

dimorphism of certain bodies (especially animal 

pathogens) in the form of yeast that grow in 

particular conditions (e.g. Paracoccidioides 

brasiliensis: yeast at temperatures above 37°C, 

hyphae at temperatures below 37°C). 

Also important is the pleomorphism of the sexual 

and vegetative forms. Over the last years this led 

to a reconsideration of nomenclature in fungi. The 

fungus considered as a complete entity, or “the 

whole fungus” was named the “holomorph”: this is 

the sum of the “anamorph” (from “anatomic 

morphology”: the asexual form), and the 

“teleomorph”. A holomorph is thus normally 

composed of a teleomorph and its anamorphic 

form. However, in some cases one of the forms will 

be unknown and so the holomorph will correspond 

either to the teleomorph or the anamorph. Not 

infrequently, a single holomorph may have several 

diverse anamorphic forms. One typical example of 

this is Aspergillus (an anamorphic form) and its 

teleomorphic form Eurotium (fig. 3). 

19

2.1.4 The phyla of fungi relevant to this 

work: Basidiomycota and Ascomycota 

During our work we were able to examine a large 

number of samples belonging to the Phylum 

Basidiomycota and also many specimens belonging 

to the Ascomycota although fewer than those 

belonging to the former group. 

We have followed the classification proposed by 

Hibbett and Binder (2007) for higher-level 

taxonomy, while inside each family we have kept 

to the classification used by CAB International 

(www.indexfungorum.org). This was not an 

“ideological choice” (in this period of continuous 

systematic and taxonomic upheaval that is only 

partly mitigated by the International Code of 

Botanical Nomenclature, "marrying" systematics to 

taxonomy is, indeed, often arbitrary), but rather an 

operative choice – made simply to render the work 

more easily accessible to all. 

In any case, the systematics and taxonomy are 

proposed while fully taking into account the results 

that continuously arise from phylogenetic analysis. 

A greater difficulty from our point of view was the 

lack of a universally-accepted definition of 

“species” in mycology. This not only creates 

problems when one is attempting to attribute a 

name to a species, but also means that it is not 

20 

always easy to be sure that carpophores identified 

by different mycologists (or even by the same 

mycologist, but during a different period) belong to 

the same species. 

The problem also arises, obviously, when we want 

to compare our data with those of other researchers, 

although fortunately for us this was only an issue in 

very few cases and has no bearing on data after the 

level of genus. More challenging (and surely 

requiring further study) is the attempt to define a 

species. For consolidated species the difficulties are 

only quantitative in nature (having data on a great 

number of samples) but species which are still 

being debated are far more complex and, beyond 

quantitative issues, present systematic and 

taxonomical questions to the resolution of wich our 

research may perhaps be able to contribute. 

2.1.5 Glossary 

Many terms used in mycology are interpreted 

differently by different researchers. For definitions 

of some of these and in particular those concepts 

such as classification, phylogeny, systematics and 

taxonomy, where there are several schools of 

thought we list here their simplified definitions. 

Haploid Cells that contain just one set of chromosomes (one Ascomycetes Fungi that exogenously produce sexual spores (i.e. those 

chromosome of each type) 

formed after meiosis) in containers called asci. 

Basidiomycetes Fungi that produce sexual spores (i.e. those formed after Chromista The kingdom that includes unicellular or multicellular living 

meiosis) exogenously on their basidia. 

organisms called eukaryotes. These are mostly photosynthetic, 

but Chromista also encompasses organisms previously 

classified among the "lower fungi". 

Classification Very broad term that denotes an organisational scheme. Commensal Organisms that live on or in other organisms, usually 

It is often the result of a taxonomic scheme, 

receiving some advantage from the relationship (food or 

notwithstanding that classification does not necessarily 

give names to the organisms classified. 

protection) but without damaging the host organism. 

Deuteromycetes (a non-taxonomic term) fungi that develop asexual 

spores (i.e. formed after mitosis). 

Dikaryon A hypha or cell with two nuclei. 

Diploid Cells that contain two copies of each chromosome. Heterokaryons A cell containing several genetically different nuclei. They are 

artificially generated by the fusion of two or more cells and in 

nature are only found in fungi. 

Phylogeny The hierarchical structure that orders living organisms 

according to their reciprocal evolution. 

Hyperparasites A parasite that lives on another parasite. 

Lichens Forms of life resulting from a combination of Mycorrhiza A symbiotic association between a fungus and a higher plant, 

autotrophic organisms (algae or cyanobacteria, mostly 

chlorophyta) and a fungus, usually an ascomycete or 

basidiomycete mushroom. They are classified according 

to the taxonomic position of the fungus. 

found in the root wall of the vegetable symbiont. 

Homokaryon Cells that contain two or more identical nuclei, usually Parasite An organism that obtains nutrition and/or shelter from another 

produced by the fusion of one or more cells of the same 

species. 

organism without giving anything in return. 

Pleomorphism A characteristic of mushrooms that develop different Protist The kingdom including unicellular eukaryote organisms, 

forms at different stages of their lifecycles. 

including protozoa, algae and fungi. 

Saprobic An organism reeceiving nutrition from non-living Symbiosis A close relationship between diverse organisms whereby each 

organic material. 

organism receives a reciprocal benefit. Lichens and 

Systematics The process of classifying living creatures based on their Taxonomy 

Mycorrhizae are examples. 

The assegnation of names to organisms. A term frequently 

phylogenetic position. 

considered a synonym of systematics.

2.2 Biological notes on 

mushrooms 

The word mushroom immediately gives one the 

idea of edible mushrooms, those we gather from 

the woods or we buy straight from the market, 

however, mushrooms are just one type of fungi 

along with many others. Other fungi include the 

moulds that sometimes attack cultivated plants or 

the walls of our houses, those that contaminate our 

food, produce damaging toxins; or those that are 

used industrially in the production of food, drinks 

and pharmaceuticals. Fungi are employed 

commercially in the production of antibiotics, 

steroids, cyclosporin and enzymes for use in 

cooking and in the production of food and drink. 

Just try to imagine a world without fungi: we 

would have to say goodbye to wine, beer, bread, 

several types of fine cheese, antibiotics and other 

therapeutic chemical compounds. 

Fungi, as all eukaryotic organisms, possess cells 

that contain a nucleus wrapped up in a membrane, 

more than one chromosome and organelles such as 

mitochondria. 

They have many unique features in terms of their 

structure, cellular components and organisation. 

They are in fact filamentous, multicellular 

organisms made up of long, branching tubular cells 

called hyphae which are of varying lengths, but 

uniform diameter, of 2-30 μm, and are together 

known as mycelium. 

The hyphal walls are composed of polysaccharides 

(80-90%), proteins, lipids, polyphosphates and 

organic ions, but their main constituent is chitin, a 

polymer of N-acetylglucosamine, which is derived 

from glucose (Carlile et al., 2001). 

The hyphae grow from the tip; the extension zone 

may be between 30 to 400 micrometres and the 

hyphal walls stiffen rapidly. The part of the hypha 

immediately below the extension zone ages 

progressively and its oldest parts can either be 

lysed by the organism’s own enzymes (autolysis) 

or by other organisms (heterolysis). Protoplasm 

moves continuously from the old parts of the hypha 

towards the tip and so the hyphae continuously 

grow from one end and continuously age from the 

other; all the while the protoplasm shifts along 

from the aging part to the new. The hyphae of most 

fungi are divided at regular intervals by transversal 

septa, but these are not present, for example, in the 

hyphae of the Glomeromycota, except where they 

are used to isolate the hyphae’s own dead or 

decaying regions. In any case, the functional 

subdivision of fungi into those with septa and those 

without is not so clear since fungal septa contain 

pores through which cytoplasm, and in some cases 

even nuclei, may pass. Therefore septate hyphae 

are composed of interconnecting compartments and 

function as integrated units. Inside the hyphae the 

cytoplasm and nuclei move and there may be 

varying numbers of them in each compartment 

separated by septa, from one to dozens of them, 

right up to tens of thousands of them in coenocytic 

fungi (without septa) (Gregory, 1984) (fig. 4). 

Fig. 4. Epifluorescent photo of the many nuclei present in the fungal hyphae of a coenocytic fungus, detected by staining with DAPI. 

A distinctive aspect of fungi is the ability of their 

hyphae to fuse to form a closely inter-connected 

mycelium where the identities of the individual 

hyphae are lost in favour of the sharing of nutrients 

and genetic heritage: a fundamental characteristic 

allowing fungal colonies to resist environmental 

stresses (Brasier, 1992; Glass et al., 2004) (fig. 5). 

21

22 

Fig. 5. View of hyphal fusions that give rise to a closely interconnected fungal mycelium. 

Fungi can reproduce both sexually and asexually. 

Asexual reproduction takes place by mitosis, with 

the production of spores that are dispersed in the air 

and then fall on an appropriate substrate where they 

germinate and the cycle may repeat. Sexual 

reproduction instead occurs after the fusion of two 

haploid cells, with the creation of one diploid cell. 

After meiosis and subsequent mitosis this cell 

develops single spores. In the Ascomycetes, these 

are formed inside closed receptacles called asci and 

are known as ascospores. In the Basidiomycetes 

they are found externally on structures called 

basidia and are called basidiospores (Carlile et al., 

2001). 

Fungi are chemoheterotrophic organisms and 

therefore need a source of organic nutrients from 

which to draw energy for their cellular metabolism. 

Given a simple energy source, such as glucose, 

many fungi can then absorb all the other cellular 

components they need from inorganic sources 

(ammonia or nitrate, phosphate and other minerals 

such as calcium, potassium, magnesium and iron). 

Their cell walls are composed of complex 

polysaccharides, such as chitin, and fungi can 

absorb simple soluble nutrients through their cell 

walls and membranes. 

Thanks to their ability to produce extracellular 

enzymes, they can break down complex polymers 

such as cellulose and lignin so as to then reabsorb 

the simple sugars remaining. Fungi produce a wide 

range of enzymes that can degrade the most varied 

and recalcitrant polymers such as lignin (Carlile et 

al., 2001). 

Yeasts also belong to the kingdom of fungi; these 

are; mostly-unicellular organisms with globose or 

oval cells that measure 6-12 micrometres in 

diameter. 

Yeasts may reproduce through budding, division or 

sexually by formation of asci or basidia and live in 

sugar-rich environments. The species most 

commonly used in the industrial fermentation 

process is Saccharomyces cerevisiae, the “brewer’s 

yeast”. Each industry will have its own selected 

strains, which are treated as genuine industrial 

secrets in the production of wine, beer, cider and 

bread. 

When yeasts reproduce asexually through the 

production of buds on the cell surface, each bud 

will grow until it reaches the same dimensions as 

the mother cell. At this stage the new cell will 

detach itself, leaving a scar that will be covered 

with chitin. Since chitin (the principal constituent 

of insects’ and crustaceans’ exoskeletons) is a 

polysaccharide with a rigid consistency. new buds 

cannot form at these chitin-covered scars and as a 

consequence, the mother cell, upon having 

produced as many budsto cover her whole surface 

with chitin, dies. 

Yeasts have a special metabolism that allows them 

to exist in the presence of oxygen, by breathing, 

and also in oxygen’s absence, by fermenting 

(Carlile et al., 2001).

Fungi are an important component of the 

ecosystem, instrumental in the continuation of 

biogeochemical cycles and represent the main 

agents in the decomposition of organic matter 

containing carbon, nitrogen, sulphur and 

phosphorus into mineral compounds that can be 

used again by plants. 

In terms of the carbon cycle, the main components 

of organic matter to be decomposed are cellulose, 

hemicellulose and lignin, which make up around 

70% of all material in plant cell walls. Fungi can 

completely break down cellulose by producing 

three principal enzymes: Endocellulase, which acts 

randomly inside the chain of cellulose, breaking up 

the molecules into smaller fragments; Exocellulase, 

which only acts at the ends of the cellulose chains, 

releasing cellobiose units; and the third enzyme; 

cellobiase, which breaks the disaccharide 

cellobiose into two molecules of glucose that can 

be absorbed by the fungus. 

The three enzymes act synergistically and are 

carefully regulated to ensure that a fungus that 

degrades cellulose does not release sugars at higher 

rates than it can absorb them. 

The regulation of cellulose degradation is achieved 

through a feedback system called catabolyte 

repression, in which genes that encode enzymes are 

repressed when readily usable substrates (including 

glucose) are available in the environment. 

The most notable species of cellulolytic fungi are 

Chaetomium cellulolyticum, Humicola grisea and 

Trichoderma reesei. Fungi are also the only 

organisms capable of completely degrading lignin, 

a recalcitrant compound consisting of units of 

phenylpropane linked together by chemical bonds 

of different types. Among these, the most studied is 

the basidiomycete Phanerochaete chrysosporium, 

which is also capable of degrading other molecules 

which have a nature similar to lignin (Bosco et al., 

2008a). 

The mushrooms and truffles that we gather in the 

woods are none other than the sexual fructification 

of filamentous fungi that grow in soil, most of 

which live in close symbiosis with the roots of 

various forest plants such as chestnut, oak, beech, 

fir, larch, pine, hazel and linden. 

To date we are unable to cultivate boletus, amanita, 

chantrelle, milkcap or russula mushrooms, or 

Caesar’s mushrooms or white or black truffles: for 

these we must await until completion of their 

lifecycles inside the plant roots, which terminates 

with the production of their fruiting bodies and 

frequently depends on the season and 

environmental conditions. 

A series of studies have been using techniques to 

inoculate sterile plants with truffle spores, create 

symbiosis in the laboratory and then transplant the 

mycorrhizal plants into the field (Bosco et al., 

2008b). 

There are, of course, other mushrooms that do not 

live in symbiosis with plants and as such can be 

cultivated on an industrial scale and are available 

all year round, such as Agaricus bisporus (J. E. 

Lange) Imbach. These mushrooms are usually 

grown on inexpensive material such as straw and 

wood residues to which manure is added, and 

under well-controlled light conditions, temperature 

and humidity. After inoculation of the mycelium of 

the fungus obtained in pure culture invading 

hyphae start growing all over the substrate and after 

3 weeks from sowing begin to produce fruiting 

bodies that can be collected, packaged and 

distributed to retailers. A mushroom mycelium 

obtained from an axenic culture is used to inoculate 

the substrate and the hyphae begin to grow. Three 

weeks after seeding, the fruiting bodies begin to 

grow and can be gathered, packaged and sent to 

resellers. 

Another mushroom produced on a large scale and 

particularly favoured in Japan and the Far East is 

Lentinula edodes (Berk.) Pegler (“Shetake”), which 

is able to degrade the cellulose content in trees. 

Small logs are hydrated by soaking in water and the 

fungal mycelium is put into pre-drilled holes in 

each log. After about a year the first batch of 

fruiting bodies appear (Carlile et al., 2001). 

Fungi living in symbiosis with the roots of plants 

form associations called mycorrhizae (table 1), 

which can be found in around 90% of terrestrial 

plants: these symbioses involve over 6,000 species 

of fungi and 240,000 vegetable species. The two 

symbiotic organisms, the fungus and the plant, 

initiate a very close physiological, ecological and 

reproductive relationship that works to their mutual 

advantage. Fungi colonise the root without causing 

damage and get sugars, which they are unable to 

synthesise, and the plants receive mineral nutrients 

and water absorbed and translocated through the 

large hyphal network (known as the Wood Wide 

Web) that extends from the mycorrhizal roots to the 

surrounding ground and acts as a true auxiliary 

absorption system. Many types of mycorrhizae 

exist, with diverse morphological and physiological 

features, and they have colonised very diverse 

environments (Bosco et al., 2008b). 

23

In our woodlands a large number of ectomycorrhiza 

can be found in forest plants such as the fir, 

larch, pine, birch, chestnut, beech, hazel and oak, 

and often their fruiting bodies (e.g. the well-known 

amanita, pinaroli, russula mushrooms and truffles) 

are visible to the naked eye. 

In total ectomycorrhizal symbionts include over 

500 species of mushrooms, among which we find 

members of the genera Boletus, Lactarius, Russula, 

Suillus, Amanita, Paxillus, Morchella and Tuber. 

24 

Table 1. Types of mycorrhiza, host plants and symbiont fungi. 

Previously, the number of fruiting bodies found in 

association with various plant species was 

considered indicative of the total number of fungal 

species in an ecosystem: in fact, molecular studies 

have shown that some species found in the roots 

produce few fruiting bodies, while other species 

producing many carpophores in the forest were 

rarely found among the roots of plants. 

Type of mycorrhiza Host plant Symbiont fungi 

ectomycorrhiza Evergreen forest plants and trees such as fir 

and pine and other forest trees such as beech, 

chestnut and oak 

Ecto- endomycorrhiza Ectomycorrhizal plants (pine and larch), 

Ericales (Arbutus, Arctostaphylos), and 

Monotropa and Pyrola 

Ericoid mycorrhiza Some Ericales species such myrtle, heather, 

Calluna, Rhododendron 

About 5,000 species of fungi belonging to 

Basidiomycota (Amanita, Boletus, Laccaria, 

Ascomycetes (Tuber) and Glomeromycota 

(Endogone) 

Ascomycota, Basidiomycota, some 

ectomycorrhizae (Boletus, Laccaria) 

Two species of Ascomycota, Hymenoscyphus 

ericae and Oiodendron maius 

Orchid mycorrhiza All species of orchids Eight genera of Basidiomycota, belonging to the 

genus Rhizoctonia 

Arbuscular mycorrhiza Bryophytes, Pteridophytes, Gymnosperms Around 150 species of the phylum 

and Angiosperms (around 80% of plant 

species) 

Glomeromycota 

It seems then that just a few fungal species (fiveten) 

are alone able of colonizing around 50-70% of 

the roots of the plants on this Earth. Furthermore, 

some species form symbiosis with many species of 

forest plants (e.g. Cenococcum geophylum 

colonises approximately 150 host plant species), 

while others tend to associate with very few or only 

one type of host (for example, Suillus luteus is 

found only in black pine and Suillus grevillei in the 

larch tree) (Bosco et al., 2008b). 

In the same forests and woodlands ericoid 

mycorrhizae associate with such plants as myrtle, 

heather, Calluna and Rhododendron. Arbuscular 

mycorrhizae are the most commonly found 

mycorrhizal symbiosis in nature, associating with 

around 80% of plant species and most cultivated 

foodstuffs such as wheat, corn, barley, potatoes, 

tomatoes, vegetables, citrus fruits, grapes, olives, 

fruit trees, cotton, sugarcane, rubber tree and 

meadow flowers. 

In this type of mutual relationship, the symbiont 

fungus forms characteristic branch-like structures 

called “arbuscules” inside the root cells of the host 

plant and it is through these structures that the 

exchange of nutrients between fungus and plant 

occurs. 

Irrespective of the type of mycorrhiza, the plants 

that host the fungal symbionts in their roots 

demonstrate not only better growth, due to the 

improved absorption of minerals effected through 

the fungal hyphae that stretch between the root and 

surrounding ground, but also a higher tolerance of 

biotic and abiotic stresses, and therefore a general 

fitness far larger than plants devoid of these fungal 

symbionts (Giovannetti and Avio, 2002). 

Recently, scientists have demonstrated that the 

sugars synthesised by a plant can be transported to 

other plants, even belonging to other species, if 

both plants share the same type of symbiotic fungus 

and if the two symbionts are joined by the same 

network of mycorrhizal fungal hyphae. 

This demonstrates that mycorrhizal fungal 

symbionts, beyond absorbing and carrying mineral 

nutrients to the host plant, also have an important 

role to play in the redistribution of energy resources 

within plant communities: in fact, adult plants can 

transmit nutrients via the fungal network to 

younger plants, thus increasing their chances of 

survival (Simard et al., 1997). 

The existence of networks of hyphae that explore 

the environment and act as a vehicle for nutrients 

seems to be of fundamental importance for plants,

that must grow, develop and reproduce while 

anchored to the same spot. The importance of these 

subterranean fungal networks which link several 

plants together can be fully appreciated if we 

consider fungi’s characteristic capacity for 

indefinite growth. Fungi can extend for hundreds of 

metres in every direction and in the most 

extraordinary case, reported by several North 

American researchers in 1992, one unique fungal 

individual was shown to colonise 15 hectares in a 

forest (Smith et al., 1992). 

The mechanisms underlying the formation of these 

fungal networks are still little understood, despite 

recent data showing that hyphae originating from 

one individual are able to recognise and form 

anastomoses with the hyphae from another 

compatible individual and thus create networks of 

indefinite length (Giovannetti et al. 1999; 2001; 

2004; 2006) (fig. 6). 

In conclusion, studies on fungi have helped us to 

understand their importance in natural ecosystems 

and in agroecosystems: they are able to modify the 

availability, capture and use of soil resources, such 

as water and mineral nutrients, and to directly 

intervene in the trophic relationships of plant 

communities and in the regeneration of soil 

fertility. 

Fig 6. Graphic representation of the networks of fungal mycorrhizae connecting diverse plants. 

2.3 Mushrooms and environments 

for growth 


The Italian Istituto Superiore per la Protezione e la 

Ricerca Ambientale (ISPRA) (Superior Institute for 

Environmental Protection and Research) inherited 

the role and responsibilities of the APAT (Agenzia 

per la Protezione dell’Ambiente e per i Servizi 

Tecnici) (Environmental Protection and Technical 

Services Agency), the ’ICRAM (Istituto Centrale 

per la Ricerca Applicata al Mare) (Central Institute 

for Applied Maritimes Studies) and the INFS 

(Istituto Nazionale per la Fauna Selvatica) 

(National Wild Fauna Institute). 

As part of the institutional activities of ISPRA’s 

Dipartimento Difesa della Natura (Nature Defence 

Dept.) the “Special Mushrooms Project” was 

established to develop understanding and 

awareness of these ecosystem components, 

regarding which very few national-level studies had 

been previously carried out. 

To date, one of the main studies carried out by 

ISPRA, in collaboration with, above all, the 

25

Associazione Micologica Bresadola – Centro Studi 

Micologici (AMB-CSM) and other partners, 

constituted a census of Italian myxomycetes and 

macromycetes so as to compile a checklist of the 

national mycological flora which could then be 

used in developing mycological cartography. 

A critical step in the acquisition of field data on 

comparable and acceptable national mycoflora is 

the classification of the habitats where fungi are 

found according to standardised systems 

recognised at a European level. 

Thanks to their diffusion, different trophic forms 

and specific ecological characteristics, fungi can be 

used as indicators of biodiversity and 

environmental quality (a series of monthly 

seminars on this subject has been taking place at 

ISPRA since 2007). Therefore it would be 

advantageous for fungi to be mentioned in 

environmental protection laws as soon as possible. 

As things currently stand, no fungal species are 

contained in the attachments to the Bern 

Convention (European Commission, 1982) or the 

Habitats Directive (European Commission, 1992) 

which detail the main acts of European legislation 

aimed at protecting wild species and their habitats. 

Furthermore, in Italy, unlike in the case of vascular 

flora, information regarding the links between 

fungal species and their habitat is sporadic and 

localised. Other difficulties have also arisen from 

attempts to link mycoflora to their Italian habitats: 

• Great numbers of species 

• Difficulties in understanding the 

taxonomy of many taxa. 

• Carpophore phenology, the emergence 

cycles of which (from seasonal to 

multiannual) display an apparent absence 

of the species for years. 

Such problems, given the overall importance this 

kingdom has within ecosystems can easily be 

overcome by increasing understanding of the role 

that mycoflora play in these same ecosystems. The 

acquisition of such understanding, especially in the 

light of the dearth of institutional studies on the 

matter, should be considered a national priority 

and, consequently, supported by adequate financial 

resources to allow further research in this field. 

In recent years the role that fungi play as natural 

ecosystem regulators has been recognised at a 

European level and as such ever more attention is 

being paid to mycoflora. We have, in fact, seen Red 

Lists of mushrooms gaining greater diffusion in at 

least 35 EU member states. In August 2003, for 

26 

example, a report was presented by the Swedish 

Environmental Protection Agency (Naturvårdsverket) 

and by the European Council for the 

Conservation of Fungi (ECCF) to the Environment 

Directorate-General of the European Commission 

(Dahlberg et al., 2003). That document proposes 

the inclusion of 33 European fungal species into 

Appendix 1 of the Bern Convention and into the 

Habitats Directive. The species recommended for 

inclusion are rare in Europe and are already 

contained in some countries’ Red Lists. These 33 

species are only a fraction of the threatened 

varieties throughout Europe, but the document 

represents a first step towards official recognition 

of the importance of mycoflora and of its 

conservation by the European Commission. 

In countries where research in this field is more 

developed it has been recognised that 

macromycetes are threatened to a far greater degree 

than vascular flora. In Switzerland, 32% of the 

macromycete species recorded in the country have 

been placed on the Red List (Senn-Irlet et al., 

2007). The endangered species are concentrated 

mainly in dry grasslands and swamps. 

These data, extrapolated to cover the Italian 

ecological situation, underline the necessity of 

taking rapid action to build awareness of mycoflora 

and to make focused interventions to protect it. 

With the aim of extending awareness and 

establishing monitoring systems across the national 

territory as part of an international network, the 

Special Mushrooms Project involved a systematic 

data-collection drive which specifically sought to 

associate the environments where mycoflora was 

found in Italy with European classification systems 

governing soil use [CORINE Land Cover (APAT, 

2005)] and biotopes [CORINE Biotopes (AAVV, 

1991), EUNIS (Davies et al., 2004), NATURA 

2000 (European Commission, 2007)]. Beyond 

enabling us to learn more about the ecology of 

various environments of national and European 

Community interest, this laid the foundations for 

the use of fungal species as possible ecological 

indicators in varying thematic cartography projects. 

These in turn make possible global biodiversityevaluation 

and other nature conservation initiatives. 

2.3.2 Materials and methods 

Based on the mycological lists currently available 

from the Special Mushrooms Project and from 

those records which were comparable to our 

classification systems, it has been possible to

construct a database which correlates fungal 

species to their habitats. 

The lists used at this juncture were: 

• Database of heavy metals in 

macromycete, edited by L. Cocchi and L. 

Vescovi, from the Gruppo Micologico e 

Naturalistico “R. Franchi” di Reggio 

Emilia - AMB (4.956 records). 

• Database of species deposited in the 

Herbarium mycologicum at the Natural 

History Civic Museum of Venice, edited 

by G. Robich and M. Castoldi, Società 

Veneziana di Micologia - AMB, 

21.823 records). 

• ISPRA Database (edited by C. 

Siniscalco, ISPRA - Gruppo Micologico 

dell’Etruria Meridionale - AMB, 

5.334 records). 

• For the genus Russula, Sarnari’s 

monograph (2000). 

• For alpine fungi, works by Bizio, Campo, 

(1999) and Jamoni (2008) regarding 

alpine and sub-alpine flora. 

• For dune environments, works by Monti 

et al. (2000) (Tuscany) and Lantieri 

(2003) (Sicily). 

The fields currently in the database, always 

expandible, are: 

• Nomenclature: Genus, Species, Variety. 

• Ecology: trophic features, host plants, 

habitats. 

• Geography: area, municipality, province, 

altitude, latitude, longitude. 

• Habitat: CORINE Land Cover third and 

fourth level, CORINE Biotopes third 

level, fourth level and fifth level, Natura 

2000. 

• Sample data: collector, identifier, date. 

Statistical analyses regarding the percentage of 

occurrence for each type of habitat were carried 

out. 

First, data regarding the occurrence of each species 

in different habitats of the same area were analysed 

while excluding those data pertaining to recordings 

of the same species in the same type of habitats in 

that area. 

Cluster analysis was used to group the CORINE 

Biotopes – fourth level data – with regards to the 

distribution and frequency of mycological species’ 

occurrence. To this end, the Minitab software was 

used. We applied single-linkage, complete-linkage 

and Ward methods to the Manhattan and Euclidean 

distances calculated on standardised values. Bonds 

that continued to repeat themselves during analysis 

were identified. Finally, based on the ordered 

tables, “characteristic species” and “differential 

species” were derived for each category of habitat. 

2.3.3 Results 

We employed a process of upscaling that better 

enables mycological components to be classified at 

the various levels into which the Italian ecological 

environment may be separated. We present here the 

comparative and statistical analyses, following their 

hierarchical classification. The species have been 

ordered according to the frequency they occurred in 

the records regarding the diverse habitats analysed. 

2.3.3.1 CORINE Land Cover, Third level 

Using the CORINE Land Cover (Third level) 

categories, the species-habitat association provides 

macro-level information. 

This type of classification made it possible to 

obtain useful information on the distribution of 

macromycetes even across densely anthropic areas, 

and has furthermore enabled a broad spectrum 

analysis of species common to the coniferous and 

deciduous forests. The species were ranked 

according to the frequency they occurred in the 

records regarding the diverse habitats analysed. 

Land Cover code, third level: 1.4.1. Green 

urban areas 

(508 records, 158 species) 

Agaricus bitorquis (Quél.) Sacc.; Agaricus 

campestris L.; Agaricus bresadolanus Bohus; 

Agaricus xanthodermus Genev.; Coprinus comatus 

(O.F. Müll.) Pers.; Inocybe rimosa (Bull.) P. 

Kumm.; Lepista sordida (Schumach.) Singer; 

Lepiota subincarnata J. E. Lange; Leucoagaricus 

leucothites (Vittad.) Wasser; Lyophyllum decastes 

(Fr.) Singer; Mitrophora semilibera (DC.) Lév.; 

Morchella hortensis Boud.; Psathyrella 

candolleana (Fr.) Maire; Russula ochrospora 

(Nicolaj ex Quadr. & W. Rossi) Quadr.; Boletus 

rubellus Kromb. 

27

Lignicolous species: 

Agrocybe cylindracea (DC.) Gillet; Armillaria 

mellea (Vahl) P. Kumm.; Flammulina velutipes 

(Curtis) Singer; Pleurotus ostreatus (Jacq.) P. 

Kumm.; Polyporus squamosus (Huds.) Fr. 

Land Cover code, third level: 3.2.1. Natural 

grasslands 


Marasmius oreades (Bolton) Fr.; Lycoperdon 

utriforme Bull.; Agaricus macrocarpus (F. H. 

Møller) F. H. Møller; Agaricus campestris L.; 

Amanita vittadini (Moretti) Sacc.; Leucoagaricus 

leucothites (Vittad.) Wasser; Volvariella 

gloiocephala (DC.) Boekhout & Enderle; Agaricus 

arvensis Schaeff.; Helvella crispa (Scop.) Fr.; 

Agaricus xanthodermus Genev.; Hygrocybe ingrata 

J. P. Jensen & F. H. Møller; Hygrocybe conica 

(Schaeff.) P. Kumm.; Hygrocybe quieta (Kühner) 

Singer; Hygrocybe calyptriformis (Berk.) Fayod; 

Hygrocybe pratensis (Fr.) Murrill; Macrolepiota 

procera (Scop.) Singer; Hygrocybe nitrata (Pers.) 

Wünsche; Hygrocybe psittacina (Schaeff.) P. 

Kumm.; Panaeolina foenisecii (Pers.) Maire; 

Hygrocybe ceracea (Wulfen) P. Kumm.; 

Hygrocybe punicea (Fr.) P. Kumm.; Hygrocybe 

irrigata (Pers.) Bon; Inocybe fraudans (Britzelm.) 

Sacc.; Calvatia gigantea (Batsch) Lloyd; Laccaria 

laccata (Scop.) Cooke; Coprinus comatus (O. F. 

Müll.) Pers.; Entoloma mougeotii (Fr.) Hesler; 

Hygrocybe citrinovirens (J. E. Lange) Jul. Schäff.; 

Hygrocybe coccinea (Schaeff.) P. Kumm.; 

Chlorophyllum rhacodes (Vittad.) Vellinga; 

Lycoperdon pratense Pers. 

Land Cover code, third level: 3.1.1. Broadleaved 

forest 

(2,653 records, 590 species) 

Boletus subtomentosus L.; Cantharellus cibarius 

Fr.; Russula vesca Fr.; Russula cyanoxantha 

(Schaeff.) Fr.; Boletus rhodopurpureus Smotl.; 

Amanita caesarea (Scop.) Pers.; Boletus reticulatus 

Schaeff.; Boletus calopus Pers.; Boletus luridus 

Schaeff.; Boletus edulis Bull.; Boletus pinophilus 

Pilát & Dermek; Amanita rubescens Pers.; 

Cortinarius caperatus (Pers.) Fr.; Amanita 

muscaria (L.) Lam; Mitrophora semilibera (DC.) 

Lév.; Verpa bohemica (Krombh.) J. Schröt.; 

Russula nigricans (Bull.) Fr.; Amanita phalloides 

(Vaill. ex Fr.) Link; Infundibulicybe geotropa 

(Bull.) Harmaja; Clitocybe nebularis (Batsch) P. 

Kumm.; Lactarius piperatus (L.) Pers.; Boletus 

28 

appendiculatus Schaeff.; Russula delica Fr.; 

Morchella esculenta (L.) Pers.; Russula acrifolia 

Romagn.; Amanita vaginata (Bull.) Lam.; 

Calocybe gambosa (Fr.) Donk; Russula chloroides 

(Krombh.) Bres.; Amanita pantherina (DC.) 

Krombh.; Boletus aereus Bull.; Boletus 

pulchrotinctus Alessio; Fistulina hepatica 

(Schaeff.) With.; Russula albonigra (Krombh.) Fr.; 

Agaricus silvicola var. silvicola (Vittad.) Peck; 

Armillaria tabescens (Scop.) Emel; Boletus queletii 

Schulzer; Boletus satanas Lenz; Gymnopus fusipes 

(Bull.) Gray; Entoloma sinuatum (Bull.) P. Kumm.; 

Boletus rubellus Krombh. 

Land Cover code, third level: 3.1.2. Coniferous 

forest (including aforestation) 


Agaricus silvicola var. silvicola (Vittad.) Peck; 

Amanita muscaria (L.) Lam; Amanita rubescens 

Pers.; Boletus calopus Pers.; Boletus edulis Bull.; 

Boletus erythropus Pers.; Boletus pinophilus Pilát 

& Dermek; Craterellus lutescens (Fr.) Fr.; 

Chalciporus piperatus (Bull.) Bataille; 

Chroogomphus rutilus (Schaeff.) O. K. Mill.; 

Clitocybe gibba (Pers.) P. Kumm.; Clitocybe 

nebularis (Batsch) P. Kumm.; Clitopilus prunulus 

(Scop.) P. Kumm.; Rhodocollybia butyracea (Bull.) 

Lennox; Entoloma hirtipes (Schumach.) M. M. 

Moser; Geopora arenosa (Fuckel) S. Ahmad; 

Hebeloma laterinum (Batsch) Vesterh.; 

Hygrophorus agathosmus (Fr.) Fr.; Hygrophorus 

latitabundus Britzelm.; Hygrophorus marzuolus 

(Fr.) Bres.; Inocybe arenicola (R. Heim) Bon; 

Inocybe bongardii (Weinm.) Quél.; Inocybe 

dunensis P. D. Orton; Catathelasma imperiale (Fr.) 

Singer; Inocybe geophylla (Fr.) P. Kumm.; Inocybe 

mixtilis (Britzelm.) Sacc.; Inocybe nitidiuscula 

(Britzelm.) Lapl.; Inocybe cincinnata var. major (S. 

Petersen) Kuyper; Inocybe piceae Stangl & 

Schwöbel; Inocybe fraudans (Britzelm.) Sacc.; 

Inocybe rimosa (Bull.) P. Kumm.; Inocybe 

splendens R. Heim; Lactarius chrysorrheus Fr.; 

Lactarius deliciosus (L.) Gray; Lactarius 

deterrimus Gröger; Lactarius salmonicolor R. 

Heim & Leclair; Lactarius sanguifluus (Paulet) Fr.; 

Lactarius scrobiculatus (Scop.) Fr.; Lycoperdon 

perlatum Pers.; Mycena pura (Pers.) P. Kumm.; 

Rhizopogon roseolus (Corda) Th. Fr.; Cortinarius 

caperatus (Pers.) Fr.; Russula torulosa Bres.; 

Sarcosphaera coronaria (Jacq.) J. Schröt.; Suillus 

bellinii (Inzenga) Watling; Suillus collinitus (Fr.) 

Kuntze; Suillus granulatus (L.) Roussel; Suillus

luteus (L.) Roussel; Suillus mediterraneensis 

(Jacquet. & J. Blum) Redeuilh; Suillus variegatus 

(Sw.) Kuntze; Tricholoma myomyces (Pers.) J. E. 

Lange; Tricholoma scalpturatum (Fr.) Quél.; 

Tricholomopsis rutilans (Schaeff.) Singer; Boletus 

badius (Fr.) Fr. 

The comparison between the lists at this level has 

allowed us to extrapolate a great deal of ubiquitous 

species listed below. Knowledge of these species 

has facilitated the interpretation of lists derived 

from the application of the Nature 2000, EUNIS 

and CORINE Biotopes classification systems, 

allowing a "cleaning up" of the species 

characteristics and differentials identification 

tables. 

Strongly ubiquitous species (found both in 

forests and meadows) 


Lycoperdon utriforme Bull.; Inocybe fraudans 

(Britzelm.) Sacc.; Agaricus arvensis Schaeff.; 

Helvella crispa (Scop.) Fr.; Cantharellus cibarius 

Fr.; Macrolepiota procera (Scop.) Singer; Russula 

cyanoxantha (Schaeff.) Fr.; Hygrocybe conica 

(Schaeff.) P. Kumm.; Calocybe gambosa (Fr.) 

Donk; Hygrocybe quieta (Kühner) Singer; 

Clitocybe nebularis (Batsch) P. Kumm.; Boletus 

luridus Schaeff.; Lactarius deliciosus (L.) Gray; 

Infundibulicybe geotropa (Bull.) Harmaja; Boletus 

reticulatus Schaeff.; Agaricus silvicola var. 

silvicola (Vittad.) Peck; Lepista nuda (Bull.) 

Cooke; Laccaria laccata (Scop.) Cooke; Mycena 

pura (Pers.) P. Kumm.; Clitocybe gibba (Pers.) P. 

Kumm.; Lycoperdon perlatum Pers.; Amanita 

vaginata (Bull.) Lam.; Lepista flaccida (Sowerby) 

Pat.; Lepista sordida (Schumach.) Singer; Boletus 

ferrugineus Schaeff.; Inocybe splendens R. Heim; 

Phallus impudicus L.; Russula virescens (Schaeff.) 

Fr.; Bovista aestivalis (Bonord.) Demoulin; 

Paxillus involutus (Batsch) Fr.; Lyophyllum 

decastes (Fr.) Singer; Amanita crocea (Quél.) 

Singe; Morchella elata Fr.; Clitocybe phyllophila 

(Pers.) P. Kumm.; Clavulina coralloides (L.) J. 

Schröt.; Inocybe inodora Velen.; Schizophyllum 

commune Fr.; Hygrocybe persisitens (Britzelm.) 

Singer; Marasmius oreades (Bolton) Fr.; Agaricus 

macrocarpus (F. H. Møller) F. H. Møller; Agaricus 

campestris; Suillus collinitus (Fr.) Kuntze; 

Leucoagaricus leucothites (Vittad.) Wasser; 

Lactarius deterrimus Gröger; Agaricus 

xanthodermus Genev.; Inocybe mixtilis (Britzelm.) 

Sacc.; Cortinarius praestans Cordier; Hygrocybe 

nitrata (Pers.) Wünsche; Boletus rubellus Krombh.; 

Morchella esculenta (L.) Pers.; Chlorophyllum 

rhacodes (Vittad.) Vellinga; Coprinus comatus (O. 

F. Müll.) Pers.; Calvatia gigantea (Batsch) Lloyd; 

Inocybe piceae Stangl & Schwöbel; Clavulina 

rugosa (Bull.) J. Schröt.; Russula delica Fr.; 

Entoloma mougeotii (Fr.) Hesler; Chroogomphus 

rutilus (Schaeff.) O. K. Mill.; Russula torulosa 

Bres.; Clitocybe rivulosa (Pers.) P. Kumm.; 

Leccinum duriusculum (Schulzer ex Kalchbr.) 

Singer; Leucoagaricus barssii (Zeller) Vellinga; 

Agaricus osecanus Pilát; Gymnopus fusipes (Bull.) 

Gray; Lycoperdon excipuliforme (Scop.) Pers.; 

Boletus dryophilus Thiers; Entoloma incanum (Fr.) 

Hesler; Agaricus augustus Fr.; Hygrophorus 

hypothejus (Fr.) Fr. 

Species ubiquitous to forests 


Amanita muscaria (L.) Lam; Boletus pinophilus 

Pilát & Dermek; Boletus calopus Pers.; Amanita 

rubescens Pers.; Cortinarius caperatus (Pers.) Fr.; 

Russula vesca Fr.; Boletus erythropus; Chalciporus 

piperatus (Bull.) Bataille; Inocybe rimosa (Bull.) P. 

Kumm.; Amanita phalloides (Vaill. ex Fr.) Link;; 

Clitopilus prunulus (Scop.) P. Kumm.; Tricholoma 

scalpturatum (Fr.) Quél.; Russula foetens (Pers.) 

Pers.; Gyromitra gigas (Krombh.) Cooke; 

Lactarius piperatus (L.) Pers.; Inocybe geophylla 

(Fr.) P. Kumm.; Sarcosphaera coronaria (Jacq.) J. 

Schröt.; Tricholoma myomyces (Pers.) J.E. Lange; 

Pluteus cervinus (Schaeff.) P. Kumm.; Lactarius 

chrysorrheus Fr.; Amanita pantherina (DC.) 

Krombh.; Russula albonigra (Krombh.) Fr.; 

Hydnum repandum L.; Russula acrifolia Romagn.; 

Rhodocollybia butyracea (Bull.) Lennox; 

Tricholoma saponaceum (Fr.) P. Kumm.; Xerula 

radicata (Relhan) Dörfelt; Gyroporus castaneus 

(Bull.) Quél.; Rhodocybe gemina (Fr.) Kuyper & 

Noordel.; Amanita citrina (Pers.) Pers.; Amanita 

excelsa (Fr.) P. Kumm.; Clitocybe odora (Bull.) P. 

Kumm.; Hebeloma laterinum (Batsch) Vesterh.; 

Rhizopogon roseolus (Corda) Th. Fr.; Russula 

fragilis Fr.; Russula romellii Maire; Gymnopus 

dryophilus (Bull.) Murrill; Lactarius volemus (Fr.) 

Fr.; Tricholoma columbetta (Fr.) P. Kumm.; 

Boletus chrysenteron Bull.; Boletus pruinatus Fr. & 

Hök; Kuehneromyces mutabilis (Schaeff.) Singer & 

A. H. Sm.; Tricholoma sulphureum (Bull.) P. 

Kumm.; Hypholoma fasciculare (Huds.) P. 

Kumm.; Trametes versicolor (L.) Lloyd; Inocybe 

leucoblema Kühner; Suillus lakei (Murrill) A. H. 

29

Sm. & Thiers; Tricholoma imbricatum (Fr.) P. 

Kumm.; Ramaria pallida (Schaeff.) Ricken; 

Boletus armeniacus Quél.; Tapinella atrotomentosa 

(Batsch) Šutara; Auricularia auricula-judae (Bull.) 

Berk.; Hebeloma sinapizans (Fr.) Sacc.; Russula 

risigallina (Batsch) Sacc.; Craterellus tubaeformis 

(Schaeff.) Quél.; Helvella acetabulum (L.) Quél.; 

Amanita submembranacea (Bon) Gröger; 

Cortinarius laniger Fr.; Lactarius pallidus Pers.; 

Lepiota ignivolvata Bousset & Joss. ex Joss.; 

Stropharia aeruginosa (Curtis) Quél.; Trametes 

pubescens (Schumach.) Pilát; Tricholoma 

orirubens Quél.; Hydnum rufescens Pers.; 

Lactarius vellereus (Fr.) Fr.; Russula heterophylla 

(Fr.) Fr.; Hydnum albidum Peck; Inocybe 

oblectabilis (Britzelm.) Sacc.; Lepiota clypeolaria 

(Bull.) P. Kumm.; Tricholoma portentosum (Fr.) 

Quél.; Amanita gemmata (Fr.) Bertill.; Coprinopsis 

picacea (Bull.) Redhead, Vilgalys & Moncalvo; 

Lyophyllum rhopalopodium Clémençon; 

Gymnopilus penetrans (Fr.) Murrill; Inocybe 

erubescens A. Blitt; Lactarius uvidus (Fr.) Fr.; 

Ramaria botrytis (Pers.) Ricken; Ramaria gracilis 

(Pers.) Quél.; Russula amoena Quél.; Russula 

parazurea Jul. Schäff.; Tricholoma batschii 

Gulden; Agaricus urinascens (Jul. Schäff. & F. H. 

Møller) Singer; Russula luteotacta Rea; 

Auriscalpium vulgare Gray; Hypholoma capnoides 

(Fr.) P. Kumm.; Limacella guttata (Pers.) Konrad 

& Maubl.; Paxillus filamentosus Fr.; Pholiota 

squarrosa (Bull.) P. Kumm.; Polyporus lepideus 

30 

Fr.; Ramaria stricta (Pers.) Quél.; Russula 

persicina Krombh. 

2.3.3.2 CORINE Biotopes Third level 

Use of the third level of the CORRINE Biotopes 

classification system characterises several habitats 

of particular importance and interest for the 

European Community. 

45.2 Cork (Habitat Natura 2000: 9330 Forests of 

Quercus suber) 


Characteristic and differential species: 

Amanita ponderosa Malençon & R. Heim; Boletus 

pseudoregius (Heinr. Huber) Estadès; Gymnopilus 

suberis (Maire) Singer; Plectania platensis (Speg.) 

Rifai; Russula albonigra (Krombh.) Fr.; 

Trichaptum biforme (Fr.) Ryvarden. 

45.3 Ilex (Habitat Natura 2000: 9340 Forests of 

Quercus ilex and Quercus rotundifolia) 



Boletus aemilii Barbier; Boletus pulchrotinctus 

Alessio; Boletus rhodoxanthus (Krombh.) Kallenb.; 

Russula ilicis Romagn., Chevassut & Privat (Fig. 7 

(Chiari et al., 2008)); Leccinellum lepidum 

(Bouchet ex Essette) Bresinsky & Manfr. Binder. 

Fig. 7. Russula ilicis Romagnesi, Chevassut & Privat (Photo: Maurizio Chiari).

Fig. 8. Real distribution of Russula ilicis Romagnesi, Chevassut & Privat in the Province of Venice obtained by associating location data from 

mycological records and the polygons from the Carta della Natura (Nature Map) (ISPRA; 2008) of the relative habitats (ilex: CORINE 

Biotopes 45.3; Natura 2000). 

41.9 Chestnut (Habitat Natura 2000: 9260 

Forests of Castanea sativa) 

Frequent species: Cantharellus cibarius Fr.; 

Russula vesca Fr.; Amanita caesarea (Scop.) Pers.; 

Boletus reticulatus Schaeff.; Boletus subtomentosus 

L.; Fistulina hepatica (Schaeff.) With.; Russula 

cyanoxantha (Schaeff.) Fr.; Lactarius piperatus 

(L.) Pers.; Gymnopus fusite (Bull.) Gray; Amanita 

phalloides (Vaill. ex Fr.) Link; Amanita rubescens 

Pers.; Boletus edulis Bull.; Lactarius volemus (Fr.) 

Fr.; Ramaria formosa (Pers.) Quél. 

42.8 Mediterranean Pine forests (Habitat 

Natura 2000: 9540 Mediterranean pine forests 

with endemic Mesogean pines) 



Buchwaldoboletus lignicola (Kallenb.) Pilát; 

Suillus mediterraneensis (Jacquet. & J. Blum) 

Redeuilh; Mycena seynesii Quél.; Rhizopogon 

roseolus (Corda) Th. Fr.; Boletus obscuratus 

(Singer) J. Blum. 

Frequent species: 

Lactarius sanguifluus (Paulet) Fr.; Cantharellus 

cibarius Fr.; Lactarius deliciosus (L.) Gray; Suillus 

collinitus (Fr.) Kuntze; Tricholoma saponaceum 

(Fr.) P. Kumm. 

At this level by superimposing the Habitats Map 

and the Nature Map (Various Authors, 2004; 

AAVV, 2009th; AAVV, 2009b) with the maps of 

the stations where the mycological recordings were 

made, one obtains the real and potential distribution 

maps (fig. 8 and fig. 10). 

2.3.3.3 CORINE Biotopes Fourth level 

At the fourth level of the CORINE Biotopes system 

it is possible to identify other leading species, 

especially in those environments considered as 

being of particular ecological importance at 

European level (Dir. 92/43 CEE). 

16.27 Dune juniper thickets and woods (Habitat 

Natura 2000: 2250 Coastal dunes with Juniperus 

spp. – Priority) 



Geastrum minimum Schwein.; Geastrum schmidelii 

Vittad.; Helvella juniperi M. Filippa & Baiano; 

Marcelleina atroviolacea (Delile ex De Seynes) 

Brumm.; Melanoleuca rasilis (Fr.) Singer; Pithya 

cupressi (Batsch) Fuckel. 


Geopora arenicola (Lév.) Kers; Greletia 

planchonis (Dunal ex Boud.) Donadini; Inocybe 

dulcamara (Alb. & Schwein.) P. Kumm.; 

Pustularia patavina (Cooke & Sacc.) Boud.; 

Octospora convexula (Pers.) L. R. Batra; Xerula 

mediterranea (Pacioni & Lalli) Quadr. & Lunghini. 

31

16.29 Wooded dunes (Habitat Natura 2000: 

2270 Dunes with forests of Pinus pinea and/or 

Pinus pinaster – Priority) 


32 

Fig. 9. Inocybe psammobrunnea Bon (Photo: M. Marchetti). 


Inocybe pseudodestricta Stangl & J. Veselský; 

Inocybe psammobrunnea Bon (Fig. 9); Melanoleuca 

microcephala (P. Karst.) Singer; Rhizopogon 

luteolus Fr. & Nordholm. 

Fig. 10: Real distribution of Inocybe psammobrunnea Bon in the Province of Rovigo obtained by associating location data from mycological 

records and the polygons from the Carta della Natura (Nature Map) (ISPRA; 2008) of the relative habitats (wooded dunes: CORINE 

Biotopes 16.29, Habitat Natura 2000: 2270 - Priority). 

Very common species: 

Geopora arenosa (Fuckel) S. Ahmad; Inocybe 

arenicola (R. Heim) Bon; Inocybe dunensis P. D. 

Orton; Inocybe heimii Bon; Inocybe inodora 

Velen.; Inocybe dulcamara (Alb. & Schwein.) P. 

Kumm.; Melanoleuca rasilis (Fr.) Singer; Inocybe 

rufuloides Bon.

36.11 Boreo-Alpic acid snow-patch communities 

(Habitat Natura 2000: 6150 Siliceous alpine and 

boreal grasslands) 



Hebeloma bruchetii Bon; Octospora humosa (Fr.) 

Dennis; Naucoria tantilla J. Favre; Cortinarius 

cinnamomeoluteus P. D. Orton; Cortinarius 

anomalus (Pers.) Fr.; Entoloma papillatum (Bres.) 

Dennis; Galerina pseudotundrae Kühner; Helvella 

queletii Schulzer; Inocybe bulbosissima (Kühner) 

Bon; Inocybe giacomi O. K. Mill.; Laccaria 

montana Singer; Lactarius dryadophilus Kühner; 

Peziza alaskana E. K. Cash; Russula laccata 

Huijsman; Russula saliceti cola (Singer) Kühner ex 

Knudsen & T. Borgen; Scutellinia kerguelensis 

(Berk.) Kuntze; Scutellinia superba (Velen.) Le 

Gal. 


Cortinarius favrei D. M. Hend.; Helvella corium 

(O. Weberb.) Massee; Inocybe salicis-herbaceae 

Kühner; Mycena pura (Pers.) P. Kumm.; Helvella 

lacunosa Afzel. 

36.12 Boreo-Alpic calcareous snow-patch 

communities (Habitat Natura 2000: 6170 Alpine 

and subalpine calcareous grasslands) 


Helvella alpestris Boud.; Cortinarius 

chamaesalicis Bon; Cortinarius phaeochrous J. 

Favre; Hebeloma alpinum (J. Favre) Bruchet; 

Helvella capucina Quél.; Helvella solitaria (P. 

Karst.) P. Karst.; Inocybe albovelutipes Stangl; 

Inocybe canescens J. Favre; Inocybe favrei Bon; 

Inocybe geraniodora J. Favre; Inocybe lacera (Fr.) 

P. Kumm.; Inocybe splendens var. phaeoleuca 

(Kühner) Kuyper; Inocybe subbrunnea Kühner; 

Inocybe taxocystis (J. Favre & E. Horak) Senn- 

Irlet; Inocybe umbrinodisca Kühner; Lactarius 

salicis-reticulatae Kühner; Peziza saniosa Schrad.; 

Russula subrubens (J. E. Lange) Bon; Russula 

nana Killerm. 

Other Frequent species: 

Inocybe fraudans (Britzelm.) Sacc.; Inocybe 

godfrinioides Kühner; Inocybe calamistrata (Fr.) 

Gillet; Inocybe nitidiuscula (Britzelm.) Lapl.; 

Tricholoma scalpturatum (Fr.) Quél. 

44.61 Mediterranean riparian poplar forests. 

(Habitat Natura 2000: 9240 Quercus faginea 

and Quercus canariensis Iberian woods) 


Helvella spadicea Schaeff.; Inocybe leucoblema 

Kühner; Lactarius controversus (Pers.) Pers.; 

Leccinum nigellum Redeuilh; Pholiota populnea 

(Pers.) Kuyper & Tjall.-Beuk.; Tricholoma 

populinum J. E. Lange. 


Leccinum duriusculum (Schulzer ex Kalchbr.) 

Singer; Morchella esculenta (L.) Pers.; Pluteus 

cervinus; Mitrophora semilibera (DC.) Lév. 

Even at this level it is possible to match the 

mycological data with the Natura Map (AAVV, 

2004; AAVV, 2009a; AAVV, 2009b) so as to obtain 

distribution data for each species in their relative 

habitats. 

Fig. 11: Cluster analysis over 33 CORINE Biotopes categories, Fourth level and 975 species of macromycetes. 

33

2.3.3.4 CORINE Biotopes Fifth level 

Using the fifth level where possible, particularly 

relating to catenal ecological series, it was possible 

to achieve a greater level of detail, especially 

regarding the interpretation of dune environments, 

which, as also shown in figure 11, are significantly 

distinct from other habitats in terms of 

macromycete populations: 

16.211 Embryonic dunes (Habitat Natura 2000: 

2110 Embryonic shifting dunes) 


Cyathus stercoreus (Schwein.) De Toni; Diderma 

spumarioides (Fr.) Fr.; Geopora arenosa (Fuckel) 

S. Ahmad; Pustularia patavina (Cooke & Sacc.) 

Boud.; Psathyrella ammophila (Durieu & Lév.) P. 

D. Orton; Rhodocybe malençonii Pacioni & Lalli. 

16.212 Biscay grey dunes (Habitat Natura 2000: 

2120 Shifting dunes along the shoreline with 

Ammophila arenaria “white dunes”) 


Agaricus menieri Bon; Agrocybe pediades (Fr.) 

Fayod; Gyrodon lividus (Bull.) Fr.; Agaricus 

aridicola Geml, Geiser & Royse; Gyrophragmium 

delilei Mont.; Hygrocybe persistens (Britzelm.) 

Singer; Lepiota brunneolilacea Bon & Boiffard; 

Marasmius oreades (Bolton) Fr.; Melanoleuca 

cinereifolia (Bon) Bon; Montagnea arenaria (DC.) 

Zeller; Panaeolus cinctulus (Bolton) Sacc. 

Other frequent species: 

Agrocybe pediades (Fr.) Fayod; Psathyrella 

ammophila (Durieu & Lév.) P. D. Orton; Peziza 

pseudoammophila Bon & Donadini. 

16.221 Northern Atlantic grey dunes (Habitat 

Natura 2000: 2130 Fixed coastal dunes with 

herbaceous vegetation “grey dunes”, Priority) 


Arrhenia spathulata (Fr.) Redhead; Xerula 

mediterranea (Pacioni & Lalli) Quadr. & Lunghini; 

Clitocybe barbularum (Romagn.) P. D. Orton; 

Rhizopogon roseolus (Corda) Th. Fr.; Tulostoma 

brumale Pers. 


Geopora arenosa (Fuckel) S. Ahmad; Hebeloma 

ammophilum Bohus. 

34 

16.223 Ibero-Mediterranean grey dunes 

(Habitat Natura 2000: 2210 Crucianellion 

maritimae fixed beach dunes) 


Marasmius anomalus Peck; Peziza boltonii Quél.; 

Galerina laevis (Pers.) Singer; Arrhenia retiruga 

(Bull.) Redhead; Clitocybe barbularum (Romagn.) 

P. D. Orton; Gymnopus aquosus (Bull.) Antonín & 

Noordel.; Conocybe blattaria (Fr.) Kühner; 

Conocybe leucopus Kühner ex Kühner & Watling; 

Conocybe rickeniana P. D. Orton; Coprinus 

xanthothrix Romagn.; Crinipellis scabella (Alb. & 

Schwein.) Murrill; Hydnocystis piligera Tul.; 

Hygrocybe conica (Schaeff.) P. Kumm.; Mucilago 

crustacea P. Micheli ex F. H. Wigg.; Octospora 

leucoloma Hedw.; Pachyella celtica (Boud.) 

Häffner; Peziza varia (Hedw.) Fr. 


Agrocybe pediades (Fr.) Fayod; Cyathus olla 

(Batsch) Pers.; Peziza pseudoammophila Bon & 

Donadini; Volvariella gloiocephala (DC.) 

Boekhout & Enderle. 

2.3.4 Discussion 

Mycological communities, as can be seen in fig. 11 

are far more differentiated than the natural habitats 

in which they reside; this can also be seen in the 

specificity of the relationships between plants and 

fungi. 

The CORINE Land Cover system, by using wider 

units to classify vegetation, simplifies linking 

species to habitats and enables the use of a greater 

volume of information; This renders ecological and 

biogeographical analyses less detailed, but it also 

allows us to create a map on a scale of 1:100,000 

for the entire Italian territory which is comparable 

to analogous territorial analyses at a European 

level. 

The CORINE Biotopes system, instead, makes 

possible a connection to the national project called 

“Carta della Natura” (Nature Map) (AAVV, 2004; 

AAVV, 2009a; AAVV, 2009b), which is itself useful 

for determining the real distribution of 

macromycetes but which could also be used to map 

out the potential distribution of fungal species. 

The availability of data regarding mycological 

diversity in the CORINE Biotopes and EUNIS 

categories also allows us to gather more useful

information for a full evaluation of habitat diversity 

and the vulnerability of these habitats. 

During the creation of the mycological datacards, 

available from ISPRA, several fundamental 

guidelines for their use in high-level CORINE and 

EUNIS classification schemes were established. 

Beyond the data normally contained in mycological 

datacards, it was found to be essential to: 

• For mixed consortia, besides the 

dominant plant species, state the codominant 

ones also, both in the canopy, 

and the shrub layer and ground cover. 

Usually, three or four species are 

sufficient to identify the habitat. 

• Increase as far as possible the site 

information with geological substrata 

information, height and geographic 

location (WGS 84 for the Carta della 

Natura and CORINE Land Cover). 

• For garigue and mediterranean 

undergrowth formations, indicate the 

height of natural surface levels, as the 

CORINE Biotopes and EUNIS categories 

distinguish between high undergrowth 

(>3m) low undergrowth (1-3 m) and 

garigue (< 1 m). 

• For conifers, it is essential to establish 

whether they are in a natural forest or a 

reforestation (in the latter case, check 

whether the installation was done with 

native or alien species). For this type of 

analysis prior knowledge of the natural 

formations in the analysed area is 

necessary. 

2.4 Mushrooms as a soil quality 

bioindicator 


2.4.1.1 The quality of the soil-system 

The environmental quality of an area or territory 

can be estimated by the use of effective indicators 

whereby these indicators are designated as 

instruments capable of representing particular 

environmental conditions. 

The quality of a selected environmental system 

cannot be described by one indicator alone, but 

usually needs a combination of different indicators 

which work over different scales and which 

therefore each have a different weight in the overall 

analysis (Benedetti et al., 2006). 

A good indicator needs to have certain characteristics 

that guarantee representativity, accessibility, 

trustworthiness and operability. Each indicator has 

to furthermore guarantee a certain level of political 

relevance and utility, analytical validity and 

measurability (OECD, 1999). 

The need of turning to synthetic indicators to 

establish soil quality stems from the fact that, being 

a very complex system, often, through objective 

difficulties in its measurement, its importance tends 

to be overlooked. This has led to a decrease of over 

10% in the productive capacity of cultivated land 

worldwide since the early ‘80s as soil erosion, 

pollution, aggressive modern farming methods, 

grazing, salification and, above all, desertification 

due to the loss of organic substances and 

biodiversity all take their toll. 

Among the various definitions of soil quality, one 

of the most widely-accepted is that of Doran and 

Parkin (1994) who defined it as “the ability of soil 

to interact with the ecosystem to maintain 

biological productivity, environmental quality and 

to promote plant and animal health”. In reality, 

many scientists link soil quality with a fundamental 

conceptual place in regards to territorial planning 

and company management, adding the vocational 

concept of “fit for” to it and then focusing 

primarily on what the soil will be used for. 

From inspection of the scientific literature on the 

subject one quickly notes that there is no single 

parameter or universal indicator capable of defining 

every single situation or environmental pressure; 

instead, each time a situation arises it is necessary 

to determine the most appropriate parameters to 

measure that particular environment, which, 

afterwards may be used to monitor the state of the 

soil under its new use (Doran et al., 1994). 

In the Mediterranean area, which is at risk of 

desertification owing to the marked loss of organic 

substances and biodiversity of its soil, bioindicators 

– those living organisms that can be used as environmental 

indicators – are of particular importance. 

Beyond the traditional existence of “index species”, 

which underline the necessity of having systems of 

indicators at diverse trophic levels to obtain precise 

answers from such complex objects as the 

35

Mediterranean soil, there is now the need to replace 

this with a concept of ecosystem functionality and 

to employ those organisms or indicators that can 

tell us something about how regularly an ecological 

system is carrying out its role or how much an 

ecological function is slowing down or accelerating 

in response to environmental or anthropic 

pressures. 

Of particular usefulness to environmental scientists 

are biochemical indicators that describe the 

metabolic processes at work in the soil and, 

therefore, provide a summary of how well-working 

nutritional cycles are. We can discover this 

information by looking at the dosage of molecules 

or process-marking elements in them. 

This is the case, for example, for the determination 

of soil respiration by measuring the CO 2 flows, 

which define all aerobic processes in the soil and 

therefore govern the organic-matter mineralization 

processes as well. Or, the determination of 

enzymatic levels, which in addition to metabolic 

functions can impact genetic diversity by 

representing the phenotypic expression of one 

producing organism rather than another. 

2.4.1.2 Soil ecology 

Soil is in a close relationship with the plants it 

supports and thereby forms a unique ecosystem 

with these and other microorganisms. Its fertility is 

in fact defined as the capacity of the soil to render 

crops productive. Normally we speak about 

chemical fertility (the sum of nutritional elements 

capable of being absorbed by crops), physical 

fertility (structure, terrain consistency, etc.) and 

biological fertility (Bloem et al., 2006). 

The concept of biological fertility, however, has 

only really been established over the last 20 years; 

its aim has usually been to characterise soil 

metabolism and microbic turnover. 

The function of microorganisms in the soil is multifaceted. 

It is expressed in both pedogenic processes 

and in the regulation of nutrient cycles, and thus in 

plant nutrition. Microorganisms are involved in the 

mineralisation of organic matter, in the synthesis of 

nitrogen, in the formation of humus and also 

impact the mobilisation of mineral elements 

(Lavelle et al., 2001). 

The soil, however, is also an extremely vital entity 

and as such parallel studies have been carried out 

aiming to understand the synergistic and 

competitive relationships between microorganisms 

36 

and mycorrhizae/carpophores in diverse 

pedological situations (Steinaker et al., 2008). 

Recently soil fauna has been the object of careful 

research and correlations between their presence 

and the development of fungal fruiting bodies has 

started to emerge. For example, to guarantee the 

normal development of a Tuber ascocarp, that 

organism needs to absorb the nutritional substances 

– in particular, small organic molecules and 

mineral salts – from a land mass which is fairly rich 

in humus and which is equal to around double the 

average range of the truffle (Granetti et al., 2005). 

This volume of earth, unfortunately, cannot be 

explored by the strands of hyphae that are formed 

by the ascocarp because they are of insufficient 

length. Instead, to facilitate nutrition, truffles take 

advantage of the soil microfauna, the biological 

activity of which ensures a continuous supply of 

nutrients to the immediate vicinity of the 

carpophore. Even the relationships between 

ectomycorrhizal hyphae and soil fauna, including 

the relation between the various biotic components 

and their potential function as indicators, have been 

examined in detail during research into the most 

precious Tuber species (Callot et al., 1999). These 

studies analysed the roles and relationships 

between the various components and established 

new foundations for future experimentation on soil 

bioindication. 

The term microfauna of the soil, in its widest 

sense, includes a complex series of animal species 

of varying shape, dimension and role. Species with 

individuals between 0.01 and 0.2 mm fall into this 

category: protozoa that mineralize nitrogen, 

phosphorus and sulphur, making easily absorbed 

nutritional substances available to hyphae; and 

nematodes that feed on bacteria, protozoa and 

fungal hyphae and are involved in the 

decomposition of underground carpophores 

(Lavelle et al., 2001; Callot et al., 1999). 

The mesofauna consists of animal species measuring 

from 0.2 to 2 mm, mostly mites and ticks, 

other microarthropods (protura, diplura, symphyla, 

pauropoda, pseudoscorpions, etc.), and 

enchytraeidic annelids (oligochaetes), which 

display some of the most diverse and specialised 

eating habits around (Lavelle et al., 2001; Siepel, 

1994); many microfitofagous and detritivorous 

species feed on mycorrhizal hyphae and tufts of 

mycelium from carpophores. Their catabolites (rich 

in undigested fungal hyphae) are a food source for 

earthworms (annellids) and nutritional hyphae of 

fungal species (Callot et al., 1999), the spores of

wich are inoculated in mesofauna fecal pellets, 

sometimes in organic and organo-mineral 

compounds (Lavelle et al., 2001; Siepel, 1994), 

which have an enormous positive effect on the 

growth rates of fungal hyphae. 

Macrofauna includes all individuals larger than 

2 mm; among others earthworms, snails, isopods, 

spiders, opiliones, chilopoda (centipedes), 

diplopoda (millipedes) and many insects, mostly 

larvae and adult beetles, flies, termites and ants. 

These organisms contribute to varying degrees 

towards decomposition and vertical and horizontal 

remixing of organic matter, which can be of 

vegetable, animal or fungal origin (Lavelle et al., 

2001; Menta, 2008). 

The function fulfilled by earthworms takes on 

particular relevancy, where, by feeding on organic 

fragments, the excretions of mites and ticks and 

mineral particles, they produce catabolites of 

around 1 mm in size, surrounded by a mucus-like 

intestinal material (Lavelle et al., 2001; Lavelle, 

1997), which are easily and quickly colonised by 

the nutritional hyphae of the fungal species. The 

fungi also derive great benefits from the aeration 

produced by the tunnels these earthworms dig, 

which can be up to 2.5 metres deep. Even ants 

provide soil aeration with their tunnels, which can 

be at depths of 70-80 cm, but their most important 

function is the moving of material accumulated 

during the excavation of tunnels from lower strata 

towards the surface and thus eliminating the 

negative impact of rainwater runoff on nutrients. 

(Granetti et al., 2005). 

2.4.2 Mushrooms as indicators 

2.4.2.1 mushrooms as indicators of particular 

soil characteristics 

Mushrooms may assume the role of bioindicators 

in a given ecosystem. In particular we should 

underline the enormous contribution to this 

situation made by the studies carried out over the 

last 40 years on valuable truffles (ectomycorrhizal 

species) to achieve a proper artificial cultivation 

and production of them. 

Analysis of soil characteristics (profile, particle 

size, pH, content of minerals, including trace 

elements, organic component, macro and micro 

porosity) have provided a database that is helping 

to shed light on the complex relationships between 

soil and fungi. Investigations based on ecological 

study in artificial ecosystems have shown that some 

fungi can be identified as indicators of undisturbed 

natural forests and can demonstrate the degree of 

trunk decomposition (Holmer, 1997). According to 

these authors, however, studies of fungal 

propagation should consider all the mycelium and 

not only the fruiting bodies, which constitute a 

minor part in the vegetative body of a fungus. In 

general, epigean and hypogean fungi and other 

microorganisms that colonise the humic layer of 

soil tend to prefer acid or sub-alkaline reactant 

substrates; alternatively they tend to be resistant to 

thermal stress and stresses deriving from water. It 

is not by chance that microbial taxonomy splits 

organisms into groups that demonstrate such 

characteristics as being thermophilic, cryophilic or 

alkaphilic, according to whether they need to exist 

at very high temperatures or in places with near 

eternal snowfall or in ground with sub-alkaline pH. 

Mushrooms tend to grow in soils with varying pH 

levels, from sub-alkaline to subacid, although many 

species prefer soil of around pH 7. Studies on the 

pH of natural soils conducted in different wooded 

areas showed a range of levels from 4.8 to 8 with 

an optimal value of 7.2 found in woods with green 

cover in excellent health with no loss of branches 

and/or leaves. The pH of these forest floors is 

affected by the number of trees per hectare and the 

type of vegetation cover of the soil (Bersan, com. 

pers., 2002). 

Research carried out to increase knowledge and 

improve cultivation of the ecological characteristics 

of hypogeous fungi, especially the precious tuber 

varieties, showed soil pH levels ranging from 7 to 

8.3 depending on the species studied (Granetti, 

1994). 

In general, Tuber ascocarps have improved growth 

when their hyphae live in micro-environments with 

a pH 6.0, while other ectomycorrhiza develop best 

in substrates that range from subalkaline to alkaline 

(from pH 7 to over 8) (Granetti et al., 2005). 

Various studies carried out on naturally occurring 

Tuber sites have demonstrated the following 

characteristics for the different species: 

1. T. melanosporum Vittad. (Fig. 12) usually 

prefers richly skeletal soil with the remaining 

parts made up of fine soil (sandy-loam 

texture). The pH is very uniform and has an 

average of 8.0 ± 0.4 pH (having extremes of 

5.7 and 8.25). Most of the soils studied in 

three regions of central Italy have a pH of 

around 8, and an average skeleton of 52% in 

37

38 

Abruzzo, 54% in Lazio and 52% in Umbria (Bencivenga et al., 1990 ). 

Fig. 12. Tuber melanosporum Vitt. (Prized black truffle from Norcia) (Photo: AMB – CSM Archives). 

2. For T. aestivum Vittad. (Fig. 13) the soil is 

19 cm deep on average, with a soil skeleton 

made of limestone (20%) and fine earth 

(80%); of which 16% is sand, 56% silt and 

28% clay. The pH is 7.7 on average 

(Bencivenga et al., 1996).2. 

Fig. 13. Tuber aestivum Vitt. (Summer truffle or scorzone) (Photo: AMB – CSM Archives), 

3. T. aestivum Vittad. f. uncinatum (Chatin) 

Montecchi & Borelli 1 ; prefers soil on average 

28 cm deep, with a skeleton consisting of 10% 

limestone and the remaining 90% fine earth 

4. T. mesentericum Vittad., on average, is 

found in soil deeper than 30 cm, soft or 

compacted, as in road cuttings where 

macadam limestone with a neutral or subalkaline 

pH accumulates (Palenzona et al., 

1976). Other studies in Irpinia (Campania) 

found it in sandy-loam soil with low levels of 

(28% sand, 56% silt and 47% clay). The pH 

varies from 7.0 to 7.8 with diverse amounts of 

organic matter (Bencivenga et al., 1996). 

soil skeleton and greater percentages of 

limestone [which almost always ensures a pH 

close to neutral (7.7)] and in some cases, the 

pH drops to subacid levels (Bencivenga et al., 

1996). 

1 This taxonomic entity is recorded as a variety (described in 1998 by Ian R. Hall, Peter Buchanan, Wang Yun and Anthony L.J. Cole), referred to 

and recorded in the CABI (Commonwealth Agricultural Bureaux International) “Index Fungorum” database (RecordID = 318194) 

(http://www.indexfungorum.org). After appropriate evaluation, we prefer to classify this entity as a form, as suggested by Montague & Borelli in 

1995 and in (Montecchi et al., 2000).

5. T. magnatum Pico (Fig. 14) can be found in 

deep soil with a poor skeleton but rich in silt 

and clay, which make up a skeleton of 68.4%. 

pH levels are close to 8 and vary little 

(Bencivenga et al., 1988). 

Fig. 14. Tuber magnatum Pico (Prized white truffle from Alba) (Photo: AMB – CSM Archives). 

6. T. borchii Vittad.; is found in soil with an 

average skeleton of 31.7%, the remainder 

being on average 66.3% sand, 23.2% silt, and 

13.2% clay. The pH varies from 7.5 to 8.0 

with average values of 7.6 (Giovagnotti et al., 

1999). 

2.4.2.2 Mushrooms as indicators of ongoing 

degradation 

Some fungal species, by their mere presence and 

quantity, indicate a current imbalance in the 

ecosystem and can predict other detectable forms 

of degradation in advance. A fungal species present 

on the woody remains and an indicator of 

significant amounts of nitrogen in the substrate is 

Megacollybia platyphylla (Pers.) Kotl. & Pouzar, 

which, by its intrinsic characteristic of acting on 

large surfaces with its mycelial cords and 

producing basidiomycetes directly on these 

surfaces (Fig. 15), is considered a good indicator of 

ongoing forest degradation. 

In these cases, the genetic and functional ranges of 

fungi provide a large number of species, which 

indicate large ecosystem suffering due to excessive 

dead biomass (necromass). We note, among others, 

Cerrena unicolor (Bull. : Fr.) Murr., Coriolopsis 

gallica Fr. and Trametes trogii Berk., the spores of 

which precede the hyphal entrance of Megacollybia 

platyphylla (Pers. : Fr.) Kotlaba & Pouzar (Bersan, 

com. pers., 2002). Even Clitocybe phaeophthalma 

(Pers.) Kuyper is an indicator species, suggesting 

excessive amounts of nitrogen in the substrate, but 

unlike Megacollybia platyphylla it has manifold 

functions and so its presence needs to be evaluated 

on a case-by-case basis. For example, an 

accumulation of substrate in a place with stagnant 

water and low ventilation, related to a number of 

C. phaeophtalma fruiting bodies, may indicate the 

decay of plants in the area circumscribed by the 

basidiomycetes as the excess necromass inhibits the 

recycling processes related to other species of fungi 

(Bersan, com. pers., 2002). 

39

40 

Fig. 15. Rhizomorphs of Megacollybia platyphylla (Pers.: Fr.) Kotl. & Pouzar, with fruiting bodies highlighted (Photo: C. Siniscalco). 

Other species of gasteroid fungi (Sarasini, 2005), 

belonging to diverse families (Phallaceae Corda, 

Lycoperdaceae Corda, Clathraceae E. Fisch.) are 

also indicators of ongoing decay processes: for 

2.4.2.3 Mushrooms as indicators of future decay 

activites 

There are species which, with their fruiting bodies, 

by feeding on the by-products of other fungal 

species (primary degraders) indicate ecosystem 

changes that will only be perceivable to us in the 

distant future (when these primary degraders, with 

their long cycles, fruit). 

Owing to characteristics connected to their 

biological cycle, several species of the genus 

Mycena (Pers.) Roussel have this predictive 

Fig. 16. Clathrus ruber P. Micheli ex Pers. (Photo: C. Siniscalco). 

example, Mutinus caninus (Huds. : Pers.) Fr., 

Lycoperdon pyriforme Schaeff. : Pers., Clathrus 

ruber Micheli : Pers. (Fig. 16) (Bersan, com. pers., 

2002). 

capacity (Robich, 2003): Mycena rosea (Schumac.) 

Gramberg; Mycena pura (Pers. : Fr.) P. Kumm.; 

Mycena pelianthina (Fr.) Quél.; Mycena 

galericulata (Scop. : Fr.) Gray; Prunulus niveipes 

Murril [Sin. Mycena niveipes (Murrill) Murrill]; 

Mycena polygramma (Bull. : Fr.) Gray; Mycena 

amicta (Fr.) Quél.; Mycena flavoalba (Fr.) Quél. 

Mycena rosea has established itself as an excellent 

indicator of biodegradation caused by primary 

fungal agents (Fig. 17) (Bersan, com. pers., 2002).

2.4.2.3 Mushrooms as indicators of habitat 

diversity 

Organisms capable of indicating biological 

diversity in terms of richness and population 

abundance are very important for the understanding 

and conservation of ecosystems. Even mushrooms, 

then, can be used in the study and monitoring of 

biodiversity of an ecosystem or an environment 

(Benedetti et al., 2006). 

Regarding mycological elements, several studies 

were conducted by APAT (today the ISPRA) as of 

2003 to build databases in collaboration with the 

Associazione Micologica Bresadola, Centro Studi 

Micologici (AMB. - CSM), who have the mandate 

to conduct the first stages of habitat-linking 

between the nationally-recognised habitats to their 

CORINE Biotopes and Natura 2000 equivalents 

(Siniscalco, 2008; 2009). 

As an example of a similar activity, we can cite the 

preliminary analysis conducted on some habitats of 

European importance that has allowed for certain 

guide species to be identified for dune 

environments (Bianco et al., 2009). 

Species lists for each habitat have been created. 

These are based on available national data and are 

made according to the frequency of occurrence of 

each species. 

Characteristic and differential species have 

emerged when comparing their presence and 

frequency with other habitats; frequent species are 

those with high levels of occurrence, but also 

present in other habitats. These species (n. =177) 

represent an initial sampling of precious ecological 

elements and environmental quality indicators 

(Bianco et al., 2009). 

Fig. 17. Mycena rosea (Schumac.) Gramberg (Photo: AMB – CSM Archives). 

2.4.2.4 The trophic qualities of fungi as a key 

function of processes linked to soil fertility 

Mushrooms are increasingly becoming a principal 

tool in land-quality monitoring thanks to their 

specialised trophic activities which guarantee them 

a spot in all the national habitats. 

Mushrooms, along with bacteria and other 

microorganisms, ensure the catabolic degradation 

of organic substances so as to obtain simple 

molecules in the form of water, carbon dioxide and 

mineral salts and also ensure the metabolic 

synthesis of complex organic and organomineral 

molecules that are involved in the formation of 

humus (Zanella et al., 2001). 

Therefore mushrooms and microorganisms play a 

fundamental role in guaranteeing soil fertility and 

in the absence of which the soil would be simply an 

inert mechanical support. 

Recent observation (Papetti, pers. comm.) seem to 

lend weight to the hypothesis that in meadows and 

mountains, the presence of fruiting bodies of 

Hygrophoraceae (grass symbionts) is limited by 

excess nitrogen of mineral and organic origin. The 

reduction of anthropic nutrition seems then to allow 

a return to the soil’s original condition, probably 

because the mycelium causing mycorrhizal activity 

is not, in this case, permanently affected by 

nitrogen pollution of the soil. 

2.4.2.5 Mycorrhizal fungi as indicators of plant 

health 

Ectomycorrhizae, besides being a physical barrier 

to the penetration of parasites and being able to 

qualitatively and quantitatively change the 

metabolites released in the plant rhizosphere, also 

41

tend to produce antibiotic compounds that represent 

a barrier against many toxic soil microorganisms 

(Montecchio, 2008). Understanding the mycorrhizal 

metabolism and its working mechanisms 

provides a number of bioindication keys, given that 

the roots of an adult forest plant, normally, can 

simultaneously bear mycorrhizal fungi from 30 to 

50 different species, each able to best develop only 

under certain environmental, phenological and soil 

microclimate conditions (Koide et al. 2000). 

These new bioindication resources allow us to say 

that mycorrhizae producing their fruiting bodies, 

enable us to monitor their beneficial effects on host 

42 

plants. For example, Rhizopogon vinicolor A. H 

Sm. confers greater resistance to drought to 

seedlings of Pseudotsuga menziesii (Mirbel.) 

Franco, while Laccaria lacquer (Scop.) Cooke 

shows greater resilience than Hebeloma 

crustuliniforme (Bull.) Quélet (Fig. 18), staying 

alive far longer than the plant itself after cutting 

(Parke et al., 1983). 

H. crustuliniforme shows better efficiency in the 

nitrogen mobilization from proteinous substances 

in Betula pendula Roth than both Amanita 

muscaria (L.) Lam. (Fig. 19) and Paxillus involutus 

(Batsch) Fr (Abuzinadah et al., 1989). 

Fig. 18. Hebeloma crustuliniforme (Bull.) Quél. (Photo: AMB – CSM Archives). 

Mycorrhizal communities are complex and many 

different factors influence their dynamics in various 

ways, so that is not possible to speak of a single 

“mycorrhizal effect”, but many associated effects. 

For example, there is a very complex syndrome 

that is usually designated with the generic term 

"deterioration". In recent years, “deterioration” has 

Fig. 19. Amanita muscaria (L. : Fr.) Hooker. (Photo: AMB – CSM Archives). 

been interpreted and evaluated according to 

different criteria owing to various studies on the 

mycorrhizal community. Research has demonstrated 

that the absorbing roots of deteriorating trees 

often show significant variations in the make-up of 

mycorrhizal communities (Blaschke, 1994; Causin 

et al. 1996, Mosca et al., 2007). Often, the

symptoms of deterioration are observably more 

markedly in conditions when there is little water, or 

where the water is heavily saline, which means that 

these environmental factors can play a more 

relevant role than others in determining the 

deterioration of the lesser resistant plant genotypes 

(Schütt et al., 1985; Shi et al., 2002; Manion et al., 

1992). In these cases, it has been observed that the 

frequency of some ectomycorrhizae is associated 

with plant health, thus meaning that it may be 

possible to identify the level and intensity of plant 

deterioration, even in a preventative manner, 

through objective underground parameters. It has 

been observed, in fact, that trees deteriorate 

gradually, progressively losing their ability to 

select the most efficient mycorrhizal symbionts and 

allowing them to be replaced with those better 

adapted to the changing environmental conditions. 

The relative frequency of ectomycorrhizal 

communities typically varies significantly between 

healthy and slightly/heavily deterioration points, 

allowing these communities to be used as 

bioindicators of the presence and degree of 

deterioration (Lilleskov et al. 2001; Loreau et al., 

2001). 

2.4.2.6 The presence of heavy metals in 

mushrooms: a possible new instrument for soil 

bioindication 

The ability of living beings to exchange elements 

and substances creates an ecological cycle that will 

continue until the sun's energy is no longer 

available, provided that no external factors with an 

intensity of disturbance exceeding the homeostatic 

capacity of the interested ecosystem are introduced. 

The presence of man on Earth has had a powerful 

impact on these natural cycles by artificially 

manipulating the chemical elements of life and 

dispersing in the environment synthetic and foreign 

(xenobiotic) substances that have already entered 

the metabolic cycle of organisms (Ravera, 1981). A 

survey conducted by the American Chemical 

Registry found that over fourteen million different 

chemicals are available on the world market and 

ten thousand new ones become available every 

week. The vast majority of these chemicals have 

been (and are still being) released into the 

environment and are interfering with the balance of 

terrestrial ecosystems (Sequi, 1981; Siniscalco et 

al., 2002). 

For a long time it was considered that soil was able 

to retain pollutants and very quickly dampen their 

harmful effects. Therefore more attention was paid 

to those environmental compartments such as air or 

surface water resources where the effects of 

anthropogenic pollution are more immediately 

obvious. The capacity of soil to accumulate 

pollutants can in fact prevent immediate 

contamination of neighbouring environmental 

assets, but it may also lead to a sudden release of 

pollutants once retention limits are exceeded 

(Gallini, 2002). 

Mycelium in the soil is in direct contact with the 

external environment and can absorbe and 

accumulate heavy ions. These, in turn, may be 

transferred within the cell. This phenomenon 

occurs in different ways across various fungal 

families and species. There have been numerous 

studies over the past twenty years, particularly in 

Europe, examining the presence of heavy metals in 

mushrooms and the results show heterogeneous 

behaviour between species (Siniscalco et al., 2001). 

Many metals that are present in trace amounts on 

earth are essential for growth and reproduction of 

microorganisms. Different concentrations of heavy 

metals in soil affect the composition of fungal 

communities present in the litter and soil (Onofri et 

al., 1999). In the last decade there has been a 

growing conviction that the results of these studies 

require a refinement of analysis so as to achieve the 

creation of a comparative tool, to be called the 

“reference mushroom” (Cocchi et al., 2006). When 

such a reference has been developed for each 

species, this could help to shed light on the 

physiological function of chemical elements in 

fungi; it will give us more information on 

bioindication and taxonomic assessment and no 

less importantly, it will provide an estimate of the 

volume of heavy metals that are ingested through 

diet by man and other living beings (Cocchi 2009). 

Reference mushrooms are probably a very valid 

instruments for evaluating soil biodiversity and the 

ecosystems which are connected to the soil (Petrini 

et al., 2009). 

From a functional point of view the complex 

formed by hyphal emanations from ectotrophic 

mycorrhizae and its connected mycoclena 

mobilises various minerals, starting with proteins, 

in order to protect the apex from those toxic 

pollutants (including heavy metals) that are in the 

soil at mycotoxic concentration levels (Rousseau et 

al., 1994). 

43

Absorption of heavy metal not only inhibits fungal 

growth but also causes physiological and 

morphological changes. Their toxic effect seems to 

be mainly exerted on enzymes. Growth inhibition 

may be caused by catalytically active groups being 

masked; protein denaturation; steric conformation 

being modified; or by the activation of other sites 

involved in the formation of enzyme-substrate 

complexes competing with those normally present. 

These toxic actions vary from species to species 

and depend on metal concentrations and exposure 

time (Onofri et al., 1999; Tyler et al., 1989). 

2.4.2.7 mycorrhizal fungi as indicators of the 

quality and health of the plant-soil complex 

In mycorrhizal symbiosis, the positive effects of 

nutritional exchanges can be observed in the 

metabolisms of both partners. The efficiency of 

these associations varies according to a series of 

dynamic interactions which involve not only the 

plant and the fungus, but other environmental and 

pedological factors and the relationships that are 

established between these variables (Montecchio, 

2008). 

Mycorrhizal fungi can be used as soil-quality 

indicators because they carry out key functions 

within the soil. The identification of easily 

monitorable metabolic markers makes the 

observation and evaluation of changes which can 

affect soil environment functionality possible. A 

good example for this is glomalin, a hydrophobic 

glycoprotein produced by arbuscular mycorrhizal 

(AM) fungi (Wright et al., 1996). Glomalin 

accumulates in soil in the form of a proteinous 

substance known as Glomalin Related Soil Protein 

(GRSP). GRSP is an easily measurable marker that 

can determine the medium- to long-term activity of 

AM fungi. It has been demonstrated that this 

marker is not only sensitive to environmental 

changes such as the increase of atmospheric CO 2 

(Rillig et al., 2000) and to various soil use and 

management systems (Bedini et al., 2007), but also 

bears a high correlation to the stability of particle 

aggregations within the soil (Bedini et al., 2009), 

an important parameter of the soil’s own 

functionality. 

2.4.2.8 Mushrooms and the health of the plantsoil 

complex 

It is rather difficult to make use of the values 

supplied by mycological and microbiological 

44 

parameters because soil microorganisms and fungi 

in the litter react very quickly to seasonal changes 

and rapidly adapt to different environmental needs. 

Thus it becomes difficult to distinguish natural 

fluctuations from changes caused by human 

activity, especially when the data are recorded 

without a control group, as is the case in natural 

systems. 

Several authors have proposed solutions to this 

issue. Brookes (1994), for instance, states that no 

parameter can be used in isolation but rather a 

whole series of correlated parameters should be 

jointly considered so as to create a kind of “internal 

control group”, as is the case when considering 

carbon in microbial biomass and the total organic 

carbon in soil. When a soil sample presents a 

significant variation from “normal” values 

(Cbiomass/C total organic in soil) in a particular soil 

management system under particular climatic and 

soil type conditions, this value then becomes an 

indicator of the soil’s changing ecosystem 

functionality. 

There is actually an almost linear relationship 

between these two variables, even if large 

discrepancies may exist between soils with 

differing physical characteristics or which are 

managed differently (Bloem et al., 2006). 

Many studies have been conducted on the 

possibility of using microbiological and biochemical 

parameters to characterize soil microbial 

diversity in both genetic and functional terms. 

Firstly, one must determine the presence of 

microbial life in soil and its order of magnitude; 

next it is essential to understand which functions 

the living population has and how active it is; then 

finally, it is important to determine the structure of 

the microbial and mycological communities therein 

and the relationships they establish with plants 

(ISPRA, 2009). 

Microbiological and biochemical methods are now 

able to provide necessary information on soils. 

Recently Bloem et al. (2006) have divided these 

methods into four groups depending on the type of 

information they can provide: 

• I. Measurement of biomass and 

microbial load: includes all methods that 

define the weight of soil and the number 

of microorganisms it contains, both in 

terms of total load and as nutritional or 

physiological groups, such as plate 

counts, colorimetric microscopy, and 

biochemical methods able to provide

information on active populations; to this 

group one should add the study of 

mycorrhizae linked to the mapping and 

inventory of fruiting bodies of 

macromycetes. 

• II. Measurement of microbial activity: 

includes all biochemical methods that 

provide information about the metabolic 

processes of microbial communities, both 

in their entirety and divided into 

functional groups. The biochemical 

methods can be divided into two 

subgroups: the first includes methods that 

count the active population in its entirety 

and that, depending on the outcome and 

the type of information it provides, 

should be included in the first group of 

methods, regarding weight and number, 

mentioned above. The second subgroup 

contains methods that define the current 

activity and the activity potential of 

individual organisms or metabolic 

groups, such as respirometric tests, 

nitrogen mineralisation, etc. Other 

methods can determine the maximum 

activity potential that can be reached in 

specific substrates (Benedetti, 2004). 

• III. Microbial and structural diversity 

of a community: this includes the most 

up-to-date methods for acquiring 

ecological and molecular data. 

Traditionally, analysis of microbial 

communities in soil was carried out using 

cultivation techniques, but only a small 

fraction (

2.5 The reference mushroom: a 

useful instrument for defining the 

capacity for bioindication of 

superior fungi 


The first hurdle to be overcome, in order to be able 

to interpret and evaluate the significance of the 

presence of chemical elements, especially heavy 

metals in higher fungi, was to have internal and 

external standards, previously absent in scientific 

literature. The kingdom of fungi is very complex 

and holds more species than the animal and plant 

kingdoms, and it is estimated that the still 

undescribed fungal species are still hundreds of 

thousands (Hawksworth, 1991). Fungi are thus one 

of the most significant components of biodiversity 

and the scientific potential for the development of 

studies of these living beings is great. These 

considerations led us to further our work in trying 

to identify an object of comparison for fungi; the 

“reference mushroom”, (Cocchi et al., 2006). The 

fungal metabolism, however, is still far from being 

fully understood and it is therefore hard to find an 

example in nature which, with reference to the 

concentrations of chemical elements it contains, 

can be described as "pure". The variables involved 

are many, as is shown by the high number of 

deviations from standard concentrations measures 

(as found in data from all other authors) for almost 

all chemical elements (the few exceptions taking on 

especial importance) and not all the variables are 

known. In this situation statistical analysis becomes 

essential. 

The idea for the definition of a “reference 

mushroom” came from an article by Markert 

(1992), who stated that “Two thirds of naturally 

occurring chemical elements in eco-systems are not 

investigated since they are viewed as nonessential 

or nontoxic to biota. In view of the important role 

plants play in most ecosystems, their inorganic 

chemical characterization, according to modern 

instrumental multi-elements techniques, the 

establishment of a “Reference plant”, comparable 

to the “Reference man” by the International 

Commission on Radiological Protection (ICRP), 

can be a useful tool for this type of chemical 

“fingerprinting” […] In the future, more attention 

should focus on establishing baseline values for 

46 

“normal” elemental concentrations in ecosystem 

components […]” 

The idea of extending the same concept to 

macromycetes, based on the statistical stability 

reached by our database, was thus evident (Cocchi 

et al., 2006). As our database contains information 

only on selected ascomycetes and basidiomycetes, 

it only allows us to establish a “reference 

mushroom” of initial approximation; the following 

objective will be to refine analyses (and therefore 

gather more data for those species for which it is 

necessary) to arrive at the definition of “reference 

mushrooms” for different taxa, up to the level of 

species. 

In summary, the concept of a “reference mushroom” 

aids us in understanding whether the 

concentrations of chemical elements in higher 

mushrooms might play a role in: 

• Bioindication. 

• Taxonomic evaluation 

• Estimation of the consumption of heavy 

metals by different species of edible 

mushrooms 

It should be noted that when speaking of taxonomic 

assessments, we refer to a species paradigm 

conceptually different from the morphological 

species concept defined by both macroscopical and 

microscopical characters used so far almost 

exclusively in basidiomycete taxonomy, but widely 

and historically considered obsolete. Indeed in the 

animal and plant kingdoms, the concept of species 

is essentially biological and historically based on 

both classical and molecular phylogenetics. 

Molecular methods are now increasingly being 

applied to fungal taxonomy and the concept of 

species to which we refer in our analyses is 

analogous to that of a “biological species”, because 

the concentrations of the various chemical elements 

depend heavily on the different metabolisms of 

each species. It is obvious, however, that it would 

be absurd to completely disregard the systematics 

and taxonomy currently used in mycology; these 

being the sole criteria available for the taxonomic 

determination of the fungal samples analysed. For 

this reason, identifications were all controlled by 

expert mycologists from the Associazione 

Micologica Bresadola (many of our samples came 

from the National Scientific Committee of the 

AMB) and from the Gruppo Micologico e 

Naturalistico “Renzo Franchi” di Reggio Emilia 

(AMB). In fact, the comparison and exchange of

information with other researchers must be 

underpinned by the "certainty" that the samples 

each researcher is studying come from the same 

species – because systematics and morphological 

taxonomy, even under the rules of the International 

Code for Botanical Nomenclature, still leave ample 

room for manoeuvre in naming species and in the 

use of synonyms. 

Clearly, the "reference mushroom" is strongly 

influenced by the size and composition of the 

global sample used. Despite this, but by employing 

a large sample, the use of this concept allows us to 

determine to a good degree the average values of 

the variables under consideration. 

Our work has shown that macromycetes can 

accumulate large quantities of various chemical 

elements in their mycelia. Some authors have 

suggested that this accumulation may be specific to 

species and genera, but certainly the composition of 

substrate may affect concentrations in the 

mycelium and then in the fruiting bodies. During 

this study, which lasted more than 20 years, we 

analysed the distribution of over 30 chemical 

elements in the fruiting bodies of more than 9,000 

samples of ascomycetes and basidiomycetes 

collected in Italy and, to a lesser extent, in other 

European regions. The data are presented in this 

work, both in extended form in the Appendix and 

in summarised form in Table 2 in paragraph 3.1.7. 

This type of presentation should enable interested 

parties to analyse the information contained in the 

database in detail, potentially even starting from the 

raw data included in the accompanying CD. 

2.5.2 Elaboration of the “reference 

mushroom”: an example 

We herein aim to clarify the procedure we used to 

define our “reference mushroom”. We identified 

the species for which we had data from at least 20 

samples, and from these (about 60 species), we 

selected some at random. 

This is a list of the selected species. We have 

indicated, for each of the species, the symbol that 

represents it on the graph in Fig. 20 and the number 

of samples analysed: 

• Agaricus arvensis Schaeff. : (AA). Nr. 58 

• Agaricus bisporus (J. E. Lange) Imbach: 

(AB). Nr. 43 

• Agaricus bitorquis (Quél.) Sacc. : (AC). 

Nr. 37 

• Agaricus urinascens (Jul. Schäff. & F. H. 

Møller) Singer [sin. A. alberti Bon; A. 

macrosporus (F.H. Møller & Jul. Schäff.) 

Pilát]: (AM). Nr. 51 

• Amanita caesarea (Scop.) Pers. : (AI). 

Nr. 27 

• Amanita muscaria (L.) Lam. : (AF). Nr. 

197 

• Amanita phalloides (Vaill. ex Fr.) Link: 

(AP). Nr. 24 

• Armillaria mellea (Vahl) P. Kumm. : 

(AR). Nr. 26 

• Boletus edulis Bull. : (BE). Nr. 115 

• Boletus luridus Schaeff. : (BL). Nr. 37 

• Boletus pinophilus Pilát & Dermek: (BP). 

Nr. 78 

• Calocybe gambosa (Fr.) Donk: (CG). Nr. 

20 

• Lycoperdon utriforme Bull. [sin. Calvatia 

utriformis (Bull.) Jaap]: (CU). Nr. 49 

• Cantharellus cibarius Fr. : (CC). Nr. 42 

• Craterellus lutescens (Fr.) Fr. [sin. 

Cantharellus lutescens Fr.]: (CL). Nr. 39 

• Infundibulicybe geotropa (Bull.) Harmaja 

[sin. Clitocybe geotropa (Bull.) Quél.]: 

(CA). Nr. 23 

• Entoloma saundersii (Fr.) Sacc. : (ES). 

Nr. 41 

• Helvella crispa (Scop.) Fr. : (HC). Nr. 29 

• Hydnum repandum L.: (HR). Nr. 37 

• Marasmius oreades (Bolton) Fr. : (MO). 

Nr. 66 

• Mitrophora semilibera (DC.) Lév. [sin. 

Morchella semilibera DC.]: (MS). Nr. 27 

• Cortinarius caperatus (Pers.) Fr. [sin. 

Rozites caperatus (Pers.) P. Karst.]: 

• (RC). Nr. 52 

• Russula cyanoxantha (Schaeff.) Fr. : 

(RA). Nr. 33 

47

48 

• Russula vesca Fr : (RV). Nr. 39 

• Boletus rubellus Krombh. [sin. 

Xerocomus rubellus (Krombh.) Quél.]: 

(XR). Nr. 77 

• Boletus subtomentosus L. [sin. 

Xerocomus subtomentosus (L.) Quél.]: 

(XS). Nr. 31 

Fig. 20. Results of a “multidimensional scaling” analysis applied to the species listed in the preceding text. Abbreviations: see text. RM: 

calculated values of the reference mushroom (stress coefficient = 0.129). 

Statistical analysis and multi-dimensional scaling 

(MDS; a powerful multivariate data analysis 

technique that helps to identify key dimensions 

among a large set of variables) were applied to the 

concentration data of the selected species. MDS, 

unlike others, does not imply particular 

mathematical assumptions about the data 

distribution type (e.g. data linearity or distribution 

normality) and is therefore a general procedure 

which is not, in practice, subject to significant 

mathematical distortions. The stress coefficient of 

the final configuration is an index that measures the 

quality of reduction and indicates if the model is 

applicable to the sampled data: data range from 0 

(the "ideal value" in a practical sense) to 1 (a poor 

value). Two coordinates were calculated for each 

species and these helped to identify one point, a 

depiction of the species, in the two-dimensional 

graph in Fig. 20. 

The quality of reduction is normally considered 

good when the stress coefficient is less than 0.2. 

The point that represents the "reference mushroom 

(RM) was measured on all samples using the mean 

values. From Fig. 20 one can note that the RM 

point, as expected, is very near to zero. From the 

graph one can also see that points BE (Boletus

edulis), BP (Boletus pinophilus), AP (Amanita 

phalloides) differ significantly from RM. This 

difference is characteristic of the species. What we 

must do then, is to check to see what the causes of 

these respective positions are. 

To this end we may refer to the tables containing 

the specific average data and their confidence 

intervals as determined in our research. From these 

tables it is clear that factors influencing distance 

from the "reference mushroom" for A. phalloides 

include high chlorine concentrations (Cl); for 

B. edulis and B. pinophilus (and to a lesser extent, 

for Agaricus bitorquis) high concentrations of Se 

are responsible. 

In summary we indicate below the key features of 

some species on which we have carried out the 

same statistical analysis: 

• Mitrophora semilibera (DC.) Lév. [sin. 

Morchella semilibera DC.] (MS): high 

levels of aluminium (Al), barium (Ba), 

calcium (Ca), cobalt (Co), iron (Fe), 

nickel (Ni), phosphorus (P), strontium 

(Sr). 

• Lycoperdon utriforme Bull. [sin. 

Calvatia utriformis (Bull.) Jaap] (CU): 

high levels of copper (Cu), potassium 

(K), lead (Pb) (the fact that this species, 

always collected in high-altitude 

grasslands, presents relatively high 

concentrations of Pb as compared to 

other species, requires further 

investigation), sulphur (S), zinc (Zn). 

• Agaricus arvensis Schaeff. (AA): high 

levels of silver (Ag), cadmium (Cd), 

cobalt (Co), copper (Cu), phosphorus (P); 

• Agaricus urinascens (Jul. Schäff. & F. 

H. Møller) Singer [sin. A. alberti Bon; 

A. macrosporus (F. H. Møller & Jul. 

Schäff.) Pilát] (AM): high levels of silver 

(Ag), cadmium (Cd), cobalt (Co), copper 

(Cu), phosphorus (P). 

• Amanita muscaria (L.) Lam. (AM): high 

levels of vanadium (V), zirconium (Zr). 

• Boletus edulis Bull. (BE): high levels of 

mercury (Hg), selenium (Se), sulphur (S) 

and low potassium (K). 

• Boletus pinophilus Pilát & Dermek (BP): 

high levels of mercury (Hg), selenium 

(Se), sulphur (S). 

2.5.2.1 Procedures to follow 

The methodology described above can be applied 

to varying types of both ecological and taxonomic 

data so long as care is taken to establish control 

structures each time it is used. In general the 

procedure to follow will be: 

• Checking that the data represent a 

“statistically stable” set. 

• Using in a first step descriptive/ 

exploratory statistics methods (mean, 

median, confidence intervals, maximum, 

minimum, and standard deviation). 

• If the descriptive methods indicate 

possible differences, moving on to 

multivariate analysis. Here one can either 

use summary data or the raw data, 

depending on the purpose of each 

analysis. 

Below, we present a concrete example of how the 

data gathered can be analysed. The graphs in Figs. 

21, 22, and 24 are largely similar to those given in 

Petrini et al. (2009). 

2.5.2.2 Univariate analysis 

49

Control of sample group homogeneity: 

50 

• By geographic region. 

• By matrix or support. 

• By elevation 

When confidence intervals overlap 

one cannot postulate significant 

statistical differences between the 

samples. 

Second step: 

Choice of reference mushroom 

(Reference) 

Fig. 21. Univariate analysis: the first step. 

For example, if one would aim to study Amanitales, Boletales and 

Russulales): 

• Reference calculated for all samples 

• Reference for the Amanitales order 

• Reference for the Boletales order 

• Reference for the Russulales order

Explorative analysis – comparison 

of groups: 

The P content in samples of 

Lycoperdon utriforme Bull. [sin. 

Calvatia utriformis (Bull.) Jaap] 

and Calvatia gigantea (Batsch) 

Lloyd [sin. Langermannia 

gigantea (Batsch) Rostk.] is 

much higher than in other 

studied samples. 

The Pb content in samples of 

Lycoperdon utriforme Bull. [sin. 

Calvatia utriformis (Bull.) Jaap] 

is much higher than in other 

studied samples. 

Applicability to other samples: 

The Se content in samples of the Boletus 

edulis group gathered in Calabria and in the 

Province of Massa are different from those 

found in other Italian, European and 

worldwide regions. 

Fig. 22. Univariate analysis: the third step. 

Fig. 23. Univariate analysis: the fourth step. 

51

2.5.2.3 Multivariate analysis 

Example: classification of several Boletus species. 

The use of MDS (Figure 24) to classify certain 

species of Boletus – also investigated by Vizzini et 

52 

al. (2008) – by using their content of certain 

chemical elements has produced results that are in 

complete agreement with those produced by 

molecular biology (details in Petrini et al., 2009). 

Fig. 24. Results of analysis on several species from the genus Boletus using MDS (Petrini et al., 2009). Final configuration stress:

of biomass as well as the storing and 

transformation of energy and mineral/organic 

elements; the soil also acts as a filter protecting 

subterranean waters and exchanges gas with the 

atmosphere; it constitutes a support for life and 

whole ecosystems. Furthermore, soil is a reserve of 

genetic resources and raw materials, the custodian 

of lost civilisations, and a pillar of the landscape. 

To allow the soil to carry out these functions, it 

must be defended from degradation and from other 

threats to its well-functioning. The Communication 

lists the main threats such as erosion, organic 

matter decline, local and diffuse contamination, 

sealing, compaction, salinisation, landslides and 

flooding as well as adding the loss of biodiversity 

as a full problem in its own right. This last point 

assumes strategic importance as it was one of the 

first times the word “biodiversity” featured clearly 

in any official EC documents. 

Two other directives are believed to be 

fundamental to safeguarding the environment and 

its biodiversity: Directive 79/409/EEC, better 

known as the "Birds Directive" and Directive 

92/43/EEC, the "Habitats Directive". With these 

two, signatory countries were asked to make efforts 

to conserve biodiversity through the conservation 

of natural habitats and of wild fauna and flora, 

through the establishment and maintenance of a 

coherent ecological network of special areas of 

conservation. The message sent out was to preserve 

and restore plant and animal biodiversity. 

So as to better implement the strategy, in 

September 2006, the European Commission 

adopted a series of tools. These were: the SFD Soil 

Framework Directive, COM 232 (2006), 

Commission Communication, COM 231 (2006) 

and the Impact Assessment SEC 620 (2006). These 

tools established soil as having a central role to 

play and consequently saw biodiversity as a key 

feature in its preservation and restoration. 

The United Nations Framework Convention on 

Climate Change (UNFCCC), and the subsequent 

Kyoto Protocol defined strategies for containing 

emissions of greenhouse gases. They also 

recognised that the terrestrial biosphere plays a 

fundamental role in the conservation of 

ecosystems, plants and the creation of new forests 

which are all important steps for combating the 

greenhouse effect and restoring biodiversity. 

The documents required signatory countries to 

quantify the spatial distribution of six different 

categories of land use (forests, wetlands, meadows, 

farmland, urban areas and other). Furthermore, for 

each land category, information regarding the type 

of management it requires, the biomass associated 

with it, the changes that occur there over time and 

an evaluation of the type of transformation there 

should be provided. In relation to these aspects the 

concepts of biodiversity and bioindication take on 

an ever-greater relevance. 

Regarding forests, the Forest Principles, adopted 

during the Earth Summit on Sustainable 

Development, called on States to maintain or 

increase the extent of forest cover; an essential 

strategy to protect and increase biodiversity. 

The European Landscape Convention, signed in 

Florence in 2000, acknowledges that, "the quality 

and diversity of European landscapes constitute a 

common resource, and that it is important to cooperate 

towards their protection, management and 

planning". Actions aimed at guiding and 

harmonising the transformation of the area, such 

transformation being caused by processes linked to 

social, economic or environmental development, 

are a valid means for the sustainable management 

of the “landscape resource”. To provide an idea of 

the importance attached to maintaining the 

extension of natural and semi-natural areas for 

sustainable development, it should be mentioned 

that the "Land Use Change" indicator is part of a 

number of indicators proposed by the United 

Commission on Sustainable Development. More 

recently, the European Environment Agency, 

through the IRENA project (Indicator Reporting on 

the Integration of Environmental Concerns into 

Agriculture Policy) has selected Land Use Change 

as one of the 35 agri-environmental indicators for 

monitoring the integration of environmental needs 

with the Common Agricultural Policy. 

Throughout the Italian national territory, the 

Sustainable Use of Natural Resources Sector of the 

Natural Resources and Parks Service – APAT, 

(ISPRA, today), launched a study of changes in 

land use and vegetation cover that took place in 

Italy between 1990 and 2000 using the CORINE 

Land Cover Database for each year. It is evident 

how biodiversity, its maintenance and its 

restoration play an important role in this work, 

often with aspects related to the concept of bioindication 

going hand in hand with biodiversity and 

almost assuming a single, interchangeable identity. 

Unfortunately at present not enough is known 

about soil biodiversity. This will be addressed in 

the Seventh Framework Programme, aimed at 

gathering a better understanding of biodiversity as 

an environmental function. This process of 

53

knowledge building will also be supported by 

ongoing initiatives connected to the Convention on 

Biological Diversity and the "Forest Focus" 

programme. 

In conclusion, the data relating to mushrooms 

presented in this book augment and complete the 

information requested by the various international 

agencies. 

The biodiversity of the Italian fungal species we 

have studied and the use of the chemical 

concentration levels in them, single out fungi as 

54 

potential biological indicators for the quality of 

forest, woodland and semi-natural habitats. 

In addition, the very broad range of data included 

here can be used over the next few decades to 

enable a comparison with future data, which could 

then make possible a better and more 

comprehensive interpretation of the effectiveness 

of current environmental protection legislation in 

minimising or negating the effects of climate 

change.

3.1 Consideration on statistics 

and statistical methods employed 

One of the fundamental problems in taxonomy in 

general is the selection of internal and external 

control groups for the samples analysed. 

Phylogenetics researchers have found a solution 

with external samples (outgroups), but classical 

taxonomy does not yet have trustworthy models in 

this field. 

Therefore, we have recently suggested making use 

of centroids to establish internal and external 

references in cases where significant amounts of 

data are available that can be summarised with 

parametric and nonparametric descriptive statistics 

(Petrini et al. 2009). 

3.1.1 Sample choice 

During our work we collected data for about 9,000 

samples (carpophores) of basidiomycetes and 

ascomycetes. Our work can be described as a nonrandom 

semi-quantitative census, because the 

samples tested were provided in most cases by 

colleagues and friends belonging to the AMB. In 

general, the exact origin of each carpophore was 

recorded for all those mushrooms provided by 

Italian and foreign mycologists. However, we also 

examined samples from exhibitions of mycology in 

Italy, and the origin of these was not always 

traceable. The majority of samples were collected 

in the province of Reggio Emilia (50% of 

mushrooms were from exhibitions). Therefore, the 

representativeness of our sample is somewhat 

reduced, because the data collected are especially 

representative of Reggio Emilia and we cannot 

exclude the possibility that analysis of a different 

set of sample data may provide a different set of 

Chapter III 

Data Synthesis 

results. Descriptive analysis of subsamples, 

however, showed that at least for Italian 

mushrooms, our results are representative. 

Furthermore, our sample is particularly 

representative of basidiomycetes, because only 

relatively few ascomycetes were examined. We 

note that our tables, which include the mean values 

and 95% confidence intervals for the various 

chemical elements studied, summarise all the data 

we collected and most of these tables are included 

in the CD attached to this document. Please note 

that only samples of taxa for which at least 20-30 

carpophores (originating from different sites) were 

examined provide reliable values. When only a few 

carpophores of a given species were studied, the 

values are to be regarded as indicative only. 

3.1.2 Statistical measurements 

In our work we mainly used descriptive statistics, 

particularly mean, median and confidence intervals. 

Of the different types of mean (such as arithmetic, 

geometric, and harmonic), we used the arithmetic 

mean, which together with the median proved best 

suited to describe the datasets we collected. 

The 95% confidence interval (95% CI) is used to 

estimate "real" values. By definition, the actual 

value of the entire population can only be 

estimated, because one never has access to the 

entire population. The CI therefore describes a set 

of values inside which most likely lies the "real" 

value, calculated using observed results from the 

sample, given a certain margin of error. 

The calculated data can then be plotted using 

different techniques. An example is shown in Fig. 

25. 

55

Fig. 25. Arsenic content in specimens of Boletus edulis collected in different geographical regions. The circle represents the average, while the 

vertical bars enclose all values within the CI of 95%. 

3.1.3 Statistical stability 

To perform reliable statistical analyses, it is 

important that the variation of data within 

taxonomic (or ecological) groups to be examined 

be stable and remain constant over random samples 

56 

Ag (mg/Kg) 

during resampling procedures and when new 

elements are introduced (Cocchi et al., 2006). Our 

database has reached statistical stability, as shown 

in Figure 26. 

Fig. 26. Content (average and 95% CI) of Ag in the genus Boletus. Results of nine resamplings with the introduction of new elements in the 

samples. The final sample contains 504 elements. 

3.1.4 The reference mushroom 

A great aid in statistical analysis is the use of a 

“reference mushroom”. A propos this we hereby 

12 

10 

8 

6 

4 

2 

0 

Boletus (N=504) 

1 2 3 4 5 6 7 8 9 

reproduce a part of a document we published in 

2006 (Cocchi et al. 2006).

Markert (1992), has proposed the adoption of a 

"reference plant", an ideal plant that would have the 

anatomical and physiological characteristics of the 

average plant in the sample. This idea is intimately 

linked to that of the "reference man" proposed by 

the ICRP and which corresponds to a person with 

the anatomical and physiological characteristics of 

the average individual. Similarly, the "reference 

mushroom" is a fungus that has the anatomical and 

physiological characteristics of the average fungus 

in the sample group studied (Cocchi et al., 2006). 

In summary, the concept of the "reference 

mushroom" is used to determine whether the 

concentrations of chemical elements in higher 

mushrooms may have a role in determining the 

possibility of their use in bioindication, taxonomic 

evaluation and estimating the dietary intake of 

heavy metals through the consumption of edible 

mushroom species. 

Obviously the "reference mushroom" is strongly 

influenced by the size and composition of the 

global sample group used. Nevertheless, with a 

large sample group, the use of this concept allows 

us to gain a good approximation of the average 

value of the variables under consideration. 

Therefore one must not forget that any average will 

depend greatly on the size and composition of the 

sample group and that consequently a “reference 

mushroom” needs to be identified on a case-bycase 

basis. 

3.1.5 Data analysis 

During this study, which lasted more than 20 years, 

we analysed the distribution of over 30 chemical 

elements in the fruiting bodies of more than 9,000 

samples of ascomycetes and basidiomycetes 

collected in Italy and, to a lesser extent, in other 

European regions. 

3.1.5.1 Procedures followed 

The above-described methodology can be applied 

to various data-types, including both ecological and 

taxonomic data. 

3.1.5.2 Multivariate analysis 

The purpose of multivariate analysis is often to 

detect clusters (taxonomic, ecological, etc..) that 

can lead to a classification which can be used for 

identification purposes. In this work, we have not 

used any ordering or classification techniques, 

since the purpose of this document is purely 

descriptive, and instead we aim to very briefly 

describe some methods that could be applied to the 

data presented in this work. 

There are various methods available for grouping 

("ordering") data, and include, to mention only the 

most popular, cluster analysis, factor analysis, 

multidimensional scaling (MDS) and multiple 

correspondence analysis. Each of them has 

advantages and disadvantages, but all result in a 

reduction of a model from “n” to just a few values, 

this being mostly accomplished by optimising/ 

reducing system variance. A deeper but simple 

description of the process, written for non 

statisticians and with more-detailed bibliographical 

references can be found in Petrini and Sieber 

(2000) and Sieber et al. (1998). 

As regards the identification of samples, canonical 

discriminant analysis is perhaps the best known 

technique. Here we refer to more specialised texts, 

also cited in Petrini and Sieber (2000). 

3.1.6 Software used 

The data were gathered using a Microsoft® 

Access® database and analysed with SPSS, version 

17 (SPSS Inc., Chicago, IL, USA). 

3.1.7 Results 

The CD that accompanies this report contains 

descriptive statistics describing the reference 

mushrooms for each of the families, genera and 

species of fungi studied. It also contains the 

number of samples, the average values and the 

relative 95% CI for the taxa studied. As a 

representative example of the analyzed data we 

present here the levels of all chemical elements 

considered in determining the universal "reference 

mushroom” (the centroid of the total sample of 

about 9,000 carpophores). 

57

Table 2. Total samples, average values and 95% CI for all samples (N= 9328) (in mg/kg dry weight or bq/kg dry material by Cs 134 , Cs 137 and K 40 ). 

58 

Element N Average 95% CI 

Al 9074 346 333 - 360 

Ag 9326 3.44 3.27 - 3.61 

As 9327 15.4 11.6 - 19.2 

B 8881 9.64 9.06 - 10.2 

Ba 9279 3.84 3.59 - 4.08 

Be 7222 0.014 0.01 - 0.01 

Ca 9326 914 848 - 980 

Cd 9328 4.20 3.92 - 4.49 

Cl 845 3670 3290 - 4040 

Co 9240 0.40 0.38 - 0.42 

Cr 9327 1.49 1.40 - 1.58 

Cs 7852 2.28 2.04 - 2.52 

Cs 134 328 91.8 44.7 - 139 

Cs 137 328 2590 1740 - 3435 

Cu 9327 58.8 56.4 - 61.3 

Fe 9323 330 318 - 343 

Ge 1182 0.033 0.03 - 0.04 

Hg 9296 1.19 1.11 - 1.28 

K 9327 39630 39310 - 39950 

K 40 328 1350 1290 - 1410 

La 6534 0.34 0.29 - 0.39 

Li 9248 0.36 0.35 - 0.38 

Mg 9327 1310 1300 - 1330 

Mn 9327 34.7 33.0 - 36.4 

Mo 9216 0.20 0.19 - 0.21 

Na 9327 328 314 - 342 

Ni 9327 1.87 1.79 - 1.96 

P 9300 7195 710 – 7286 

Pb 9320 1.61 1.51 - 1.72 

Rb 9327 138 133 - 144 

S 9317 3364 3314 - 3415 

Sc 5623 0.27 0.25 - 0.3 

Se 9327 4.13 3.87 - 4.39 

Sr 9307 3.22 2.97 - 3.48 

Ti 8102 10.2 9.8 - 10.6 

V 9327 3.22 2.83 - 3.61 

Y 5620 0.20 0.18 - 0.22 

Zn 9327 117 115 - 119 

Zr 6633 0.42 0.37 - 0.47

3.2 Applied geostatistical analysis 


Geostatistics is the branch of statistics that deals 

with the analysis of spatial data derived from 

sampling. In environmental analysis and modelling 

it is an essential tool for the management, 

understanding, and correct use of data from 

environmental surveys and measurements (such as 

meteorological data, pollutant concentrations, 

piezometers, etc.). 

Geostatistical analysis consists in modelling the 

phenomenon one wishes to investigate with a 

random variable characterized by a spatial, 

temporal or spatiotemporal law. This approach 

allows one to highlight and describe the regional or 

temporal variability (qualitative and quantitative) 

of the data analysed and to map out the results. It 

measures the effect of the position of the measuring 

point on the variability of the data observed. 

Geostatistical methods are valid for all fields of 

applied science in which the phenomena to be 

studied are spatial in nature. Over the past three 

decades, it has been used, for instance, in soil 

science, hydrology, hydrogeology, geochemistry, 

meteorology, oceanography, environmental health, 

agronomy, and imaging analysis. 

Let us take a spatial phenomenon, for example the 

heavy metal pollution of a site. In general, by 

indicating the concentration of the pollutant with z, 

and the generic coordinate point of the field (xlat, 

xlong) i with x i, Z(x) is a variable that represents the 

concentration of pollution in certain points of the 

site. 

If one property varies in a more or less continuous 

way through space, it can be taken as a regionalised 

variable (Goovaerts, 1997) and analysed with 

geostatistical instruments. 

Estimating by kriging interpolation enables a more 

detailed local spatial variation of the properties 

under study to be achieved. This type of 

interpolation is appropriate only where the property 

varies in a continuous manner and the data are 

spatially dependent or correlated. The model of 

spatial variation for geostatistical estimation is as 

follows: 

Z(x) = μ�v + ε(x) 

where Z(x) is the random variable in location x, μ v 

is the local average of Z in the predefined limits of 

location x, and ε(x) is a random term with an 

expectation of zero, and a variance equal to: 

var[ε�(x) −ε�(x + h)]= E[ε�(x) −ε�(x + h) 2 ]= 2γ(h) 

variance is calculated for all couples of locations 

x + x+h, where h is a distance vector (lag) for all 

distances and directions. γ is the semivariance 

between two locations, which, where this is 

stationary (μ v is locally constant) will be equivalent 

to: 

γ(h) = 1 /2 var [Z(x) – Z(x+h)] = 1 /2 E [Z(x) – Z(x+h) 2 ] 

and defines the variogram of Z. The variogram 

provides an unbiased description of the scale and 

pattern of spatial variation, the spatial model 

needed for kriging, and a basis for designing 

optimal sampling schemes (McBratney et al., 

1981). Theoretical variogram modelling, starting 

from experimental variogram modelling, can 

indicate which approach to take in predictive 

investigation. 

3.2.2 Ordinary kriging 

At this point values can be estimated in points or in 

blocks through kriging, a shifting mean weighted to 

observed values based on the variogram inside 

determined limits defined by an area N (inside 

which the stationary values of the variable are 

assumed to hold). For a regionalised variable Z, 

with values measured as being z(xi) at site (x i), 

I = 1, 2, …, n, the ordinary kriging algorithm will 

be: 

ZB ( ) 

N 

i = 1 

λ . zx 

i 

where Z(B) represents the value estimated for 

block B and λi the weights assigned to internal 

points at N. The kriging estimator can be defined as 

non-distorted (the weights add up to one) and 

optimal (the weights are selected so as to reduce 

variance to a minimum). Unlike with classical 

interpolation methods (inverse square of the 

distance, triangulations, variable mean ...), this 

interpolation allows not only obtaining an 

estimation map of the parameter but also a map of 

the estimation variance (the kriging error), allowing 

evaluation of the reliability of prediction. 

59

One of the most common tasks in the processing of 

spatial data is the construction of thematic maps, 

i.e. geo-referenced maps relating to selected 

geographic areas, in which, by an appropriate 

method of representation, the trend of a variable 

under study – in our case the concentration of 

metals – is given. 

Iso-value contour maps, which in cartographic 

jargon are also known as vector maps, are just one 

of many ways of representing a geographic 

variable. They are not obtained directly, but 

through creating a regular grid to represent the 

variable, itself obtained by an estimation 

60 

Fig. 27. Construction of a map, starting from measurement points. 

These maps are normally made by starting from the 

values of the variable as measured within the area. 

For example, from the initial situation shown in 

Fig. 27a, which represents the location of the 

samples and the measured values of a variable, we 

aim to build an iso-value contour map such as that 

shown in Fig. 27b. Note the non-uniform 

distribution of the samples in the example. 

calculation (Fig. 28a). The contour lines are created 

by interpolating the values on the mesh axes (Fig. 

28b). It is thus clear that the quality of the map is 

wholly dependent on the equation that produced the 

grid values. 

Fig. 28. Construction of a vector map: a) reconstruction of the variable to a regular grid; 

b) interpolation tracing of the iso-value lines.

3.2.3 Geostatistical analysis applied to 

the distribution of inorganic elements in 

soil and in fungi 

For this book, our geostatistical analysis was 

focused on the following elements or physiochemical 

parameters found in the surface soil and 

in mushrooms gathered in the Province of Reggio 

Emilia: pH, aluminium, arsenic, cadmium, 

chromium, mercury, nickel, lead, copper, selenium, 

vanadium, zinc and zirconium. Irregular gridpattern 

sampling was carried out over a large area 

in the southern part of the province of Reggio 

Emilia. 

An exploratory analysis of experimental data using 

basic statistics was carried out, along with data 

frequency distributions and experimental 

variograms. All elements tested showed a tendency 

to lognormal distribution, characterised by the 

presence of a small number of sampling points with 

values of between one to two orders of magnitude 

greater than the database. Analysis of the spatial 

correlation of the data was conducted by 

In some cases, a logarithmic transformation of data 

was applied to normalise distributions and to 

clarify the spatial correlation, by inserting a new 

transformation in the original scale of the estimate 

data. 

The figures, relating to the spatial distribution of 

concentration parameters, were generated with the 

program Surfer 8.0, while the data grid was 

Table 3. Classification of variograms used. 

calculating the experimental variogram, which was 

then used to develop a model of interpolation 

according to the Kriging method (Isaaks and 

Srivastava, 1989) for the methodological 

framework and Carlon et al. (2000), for specific 

application to contaminated soils. 

The spatial correlation of the data were generally 

good and proved sufficient to furnish an 

interpolation function. In a few cases however, the 

correlation was very weak, returning a spatial 

representation of the data which was too low to 

allow an estimate of the distribution of the chemical 

elements by interpolation. In these cases, the 

distribution of elements cannot be represented by a 

vector map, but instead by maps of sampling points 

divided by different classes of concentration. 

The experimental variograms were modelled with 

spherical functions characterised by the range, sill 

and nugget values shown in Table 3, a value of zero 

intercept was imposed at the outset (nugget = 0) in 

order to honour the measured values. 

obtained by Kriging (Isaaks and Srivastava, 1989; 

Clark and Harper, 2004). The variographic model 

was tested by cross-validation. The data provided 

relate to the sampling and chemical analysis 

procedures as described in previous chapters, and 

they are all listed in the attachments to this 

document. 

61

4.1 Methods for chemical 

analysis: soil and macromycete 

carpophores 

4.1.1 Carpophores 

For a meaningful comparison of analytical results it 

is important to consider the degree of maturity at 

harvest of the carpophores that were used to 

determine the concentration of inorganic elements. 

Based on experience and the standard practice 

followed by many mycologists, maturity can only 

be estimated empirically as there are no analytical 

methods to measure this parameter. Nevertheless, 

the empirical method that assigns "numbers" to the 

various stages of carpophore maturity combined 

with the experience of the mycologist works quite 

well and certainly contributes to reducing the errors 

that could arise through variations of chemical 

concentrations due to the varying age of 

carpophores. The scale used is as follows: 

1. Primordial (carpophore is still in embryonic 

form) 

2. Young (carpophore already formed, with 

hymenium not exposed to air in carpophores 

having a secondary veil like a cloak or a ring) 

3. Almost mature (spores begin to mature and 

the hymenium is completely exposed to the 

air) 

4. Mature (the spores are ripe and in most 

species the hymenium takes on the colour of a 

spore) 

5. Old (the first signs of deterioration or 

putrefaction have set in) 

It is clear that a more-refined evaluation (by use of 

half numbers) is the highest degree of sensitivity 

that this empirical scale of carpophore maturity 

could attain. 

Most of the carpophores analysed during this study 

had a maturity-level of 4, therefore mature 

carpophores formed the basis of the study. 

Chapter IV 

Materials and Methods 

After being gathered, the carpophores were very 

carefully cleaned with special, soft brushes to 

remove any soil or sand particles, plant debris, 

insects, larvae or other material. After cleaning, the 

carpophores were roughly cut into slices to verify 

that there was no foreign material on the inside 

either. 

At this stage the amount of fungal material to be 

dried was weighed out: 20 grams can usually be 

considered sufficient. During this operation it was 

important to maintain the real weight proportions 

between the different parts of each carpophore 

(stem, cap, hymenium). The cut material was then 

set in a crystallizer and placed in a ventilated oven 

at a temperature of 45°C for 48 hours. At the same 

time, for a small number of samples, drying was 

carried out at 105°C to measure water loss. 

Mushrooms were not kept for longer than 

necessary in contact with metal objects to avoid 

contaminating the samples, especially when damp. 

Instead we made use of other tools and treatment 

methods that involved plastic or glass tools. 

After being dried at 45 o C the samples were ground 

with an agate pestle and mortar and the ground part 

was then put into a pre-washed polyethylene 

container with two stoppers. Into this receptacle we 

put a 10 mm-diameter sphere of Teflon or glass to 

homogenise each sample before weighing and 

before following treatments. 

The acid mineralisation of all carpophores was 

performed by placing a quantity of sample 

weighing 0.5-0.7 grams in a microwave oven with 

aqua regia. After mineralisation, the sample was 

increased to 50 ml in a volumetric flask with 

ultrapure H2O (Cenci et al., 2008). 

The quantitative determination of all elements 

(except Hg) was performed with a ICP-AES Perkin 

Elmer Optima 3000 XL spectrophotometer. The 

determination of mercury was performed using an 

Perkin Elmer FIMS 100 atomic absorption 

spectrophotometer designed for cold-vapour 

mercury determination. 

The preparation of calibration standards was 

carried out starting from certified ultrapure monoelement 

solutions of 1000 mg/l by ICP instruments. 

63

Certified Reference Materials (NBS and SRM) 

with a matrix similar to that of fungi were 

mineralised and subsequently analysed. The 

concentration levels obtained were within tolerance 

intervals. 

4.1.2 Soil 

At the base of the carpophore stems 4-5 cubes of 

soil, each measuring 5x5x5 cm, were collected and 

all grass, stones, leaves and other matter were 

immediately removed. 

The soil samples were subsequently placed in 

plastic bags and mixed manually to form a single 

sample. In the laboratory each sample was dried 

and then passed through a 2 mm sieve. The 

particles of the sample which were equal to or 

smaller than 2mm were then ground with a mortar 

and pestle and then placed in polyethylene bags, as 

was the case for the mushroom samples; likewise, 

64 

Table 4. detection limits (d.l.) expressed in mg/kg. 

the mineralisation process was also carried out in 

the same way as for the mushrooms. 

Determination of the concentrations of the various 

elements under investigation was carried out using 

the same analytical instruments as for the 

mushroom carpophores. 

The Certified Reference Materials for soil and 

sediment (CRM 141 R, Calcareous Loam Soil e 

CRM 277, Estuarine Sediment) were mineralised 

and subsequently analysed. The concentration 

levels obtained were within tolerance intervals. 

4.1.3 Criteria for data collection 

Table four indicates the element-by-element 

detection limits (d.l.) obtained by measuring each 

sample ten times . The d.l. represents the triple of 

the standard deviation. This criterion is widely used 

in analytical laboratories. 

d.l. d.l. d.l. d.l. d.l. d.l. 

Al 1 Cd 0.05 Cr 0.1 Hg 0.05 Cu 0.2 Ti 0.05 

Ag 0.05 Ca 2 Fe 0.1 Mo 0.2 Rb 0.5 V 0.1 

As 1 Cs 0.1 P 5 Ni 0.2 Se 2 Zn 0.2 

Ba 0.1 Cl 5 Mg 0.2 Pb 0.5 Na 3 Zr 0.05 

B 0.2 Co 0.1 Mn 0.05 K 500 Sr 0.3 S 10 

When a measurement was less than the detection 

limit, a value equal to one tenth of the d.l. was 

entered in the archive: this is because otherwise, in 

our statistical analysis, it would not have been 

possible to distinguish a cell with no value (where a 

measurement for that element in that sample was 

not made) from a cell in which the measurement 

was less than the d.l.. 

Although the tables show values for samples and/or 

elements for which also very few measurements 

were taken, only samples and/or elements having at 

least 30 measurements are to be considered reliable 

in this statistical analysis. 

4.2 Distribution map of elements 

in soil 

An exploration of the thematic maps resulting from 

our studies will now follow, aiming to interpret the 

spatial distribution of several inorganic trace 

elements, also known as persistent inorganic 

contaminants. Furthermore, both aluminium and 

the pH levels of surface soil will be taken into 

consideration. 

After appropriate treatment, around 180 samples of 

surface soil, taken from depths of between 0 to 

5cm, were used to assess concentration levels of 

aluminium, arsenic, cadmium, chromium, copper, 

mercury, nickel, lead, vanadium, selenium, zinc, 

zirconium, and pH levels. 

The area of investigation is represented by the 

province of Reggio Emilia (Fig. 29), more 

precisely, starting from State Road 9, (known as

Via Aemilia) to the Apennine border with Tuscany, 

including a small area of the Tuscan-Emilian 

National Park. 

This area can be divided into five sections starting 

from the city of Reggio Emilia. In this segment of 

lowlands the towns Bibbiano, Cavriago and 

Montecchio are situated. Farther to the south is a 

narrow strip of land with hills spread out between 

the plains and the northern footpath. The towns 

which in part make up this area are: San Polo, 

Before commencing with a description of the 

thematic maps pertaining to the elements under 

study, it is important to emphasise that what is 

proposed and illustrated in the figures represents 

concentration distributions. This aim to 

demonstrate the real concentration base levels. 

These are the sum of geochemical contents, 

understood as purely natural phenomena, and 

concentration values due to anthropogenic activity. 

Fig. 29. Geographical map of the Province of Reggio Emilia. 

Quattro Castella, Albinea, Scandiano and 

Casalgrande. 

Continuing south, one comes across a wide hilly 

area. The towns it encompasses are Vezzano, 

Baiso, Viano and Canossa. The towns of Carpineti, 

Toano, Vetto and Castelnovo nei Monti lie between 

the Enza and Dolo rivers. Finally in the area with a 

purely mountainous morphology are the resort 

towns of Villa Minozzo, Ligonchio Besana, 

Ramiseto and Collagna. 

For all twelve elements under study, general 

remarks can be made stating that the concentration 

distribution throughout Reggio Emilia is mostly 

monotonic and qualitatively comparable. 

In general, the many human activities that have 

acted and are acting on the Reggio Emilia area, 

have not left significant signs of contamination, 

furthermore, the data we have been obtained are 

very similar to findings reported by Marks et al. 

(2009) in their study of the soil of Emilia Romagna. 

65

Further confirmation of this was provided by the 

chemical data obtained by analysing the surface 

soils collected in Scandiano, Carpineti, and their 

large surrounding areas. For most of the elements, 

the concentration levels found during this 

66 

monitoring study yielded very similar results again 

(Cenci et al., 2005). 

Aluminium (Fig. 30) can be seen to have a rather 

homogeneous concentration distribution. Lower 

levels can be found in the hills. 

Fig. 30. Spatial distribution of the concentration levels of aluminium (mg/kg dry weight) in surface soils. 

In the plains and mountains concentrations are 

higher; this is mainly due to the geology of the 

area. Arsenic (Fig. 31) mirrors the findings for 

aluminium, while for cadmium (Fig. 32), there are 

two areas, San Polo d'Enza and Reggio Emilia, 

where one could reasonably expect to see 

significant man-made effects. For the remaining 

part of the area, levels remain rather uniform. 

Fig. 31. Spatial distribution of the concentration levels of arsenic (mg/kg dry weight) in surface soils.

Fig. 32. Spatial distribution of the concentration levels of cadmium (mg/kg dry weight) in surface soils. 

Chromium and nickel (Fig. 33) have a very similar 

spatial distribution of concentration levels, as one 

would expect, and both are present in the western 

zone, in two areas lying congruent to ophiolitic 

ultramafic outcrops of the Enza basin: the levels 

found here were quite high. The small area near 

State Road 9, which is already known for its high 

cadmium levels, should also be considered a point 

of contamination for chromium. Often chromium is 

found to be enriched with many elements, but not 

nickel, which, in cases where there has been a 

geogenic malfunction, grows in proportion to the 

chromium. 

Fig. 33. Spatial distribution of the concentration levels of nickel (mg/kg dry weight) in surface soils. 

The elements copper and mercury (Figs. 34 and 35) 

show a fairly uniform distribution of concentration 

levels which, as a whole, are fairly low. Only two 

localized areas in the mountains showed higherthan-expected 

levels, presumably due to local 

contamination. 

67

68 

Fig. 34. Spatial distribution of the concentration levels of copper (mg/kg dry weight) in surface soils. 

Fig. 35. Spatial distribution of the concentration levels of mercury (mg/kg dry weight) in surface soils. 

The spatial distribution of the concentration levels 

of lead were also fairly homogeneous throughout 

the territory (Fig. 36); such values suggest a 

general enrichment due to contribution by man. 

There are two areas that have certainly been 

“contaminated”: one near State Road 9 and the 

other in the Baiso area. 

For vanadium and selenium, the Apennine area 

appears to have lower concentration levels, the 

latter also being confined to the mountainous areas. 

The largest concentration levels are diametrically 

opposed: high values for selenium were found in 

the plains, while for vanadium, high levels were 

observed in the mountains. Regarding the latter, 

there appears to be also a localised contamination 

in this area (near Reggio Emilia and State Road 9) 

of cadmium, chromium and lead.

Fig. 36. Spatial distribution of the concentration levels of lead (mg/kg dry weight) in surface soils. 

For zinc, as with cadmium, chromium, lead and 

vanadium; soil contamination is found near State 

Road 9. The remaining area contains practically 

uniform concentration levels, except for the higher 

levels found in the mountainous ranges to the east. 

Low concentration levels were found consistently 

for zirconium and it was only in the Apennines that 

higher concentrations of it were registered. The 

zirconium levels observed in this study are on 

Fig. 37. Spatial distribution of pH levels in surface soils. 

average one order of magnitude lower than those 

obtained by Marchi et al. (2009). 

Lastly, the acidity levels of Reggio Emilia soils 

measured spatially (Fig. 37) are considered. 

Throughout most of the territory pH levels of 

around 6.5 are the norm. In two areas, one located 

in the mountains and the other in the south, there 

are more acidic soils, while those in the Apennines 

tend toward neutral-alkaline pH. 

69

4.3 Distribution map of elements 

in mushrooms 

Heavy metals can be regarded as one of the main 

sources of environmental pollution. For several 

decades the output of heavy metals into the 

environment due to human activity exceeded those 

of natural origin such as volcanic eruptions and 

large forest fires. A large part of these persistent 

inorganic contaminants ends up in the soil, 

increasing chemical element concentration levels. 

The soil, thus enriched with heavy metals, may, as 

a result of leaching processes, allow some of these 

metals to leak into groundwater tables. The fact 

that mushrooms are bioaccumulators means that 

their fruiting bodies may take on these 

contaminants and pass them on to animal biota – 

with potentially hazardous results for humans. 

Regarding the heavy-metal and macro-element 

concentration levels which accumulate in 

mushrooms, they can depend on and be influenced 

by factors such as soil type and the concentration 

levels of the metals. It should be noted that 

concentrations of heavy metals in soils, in this 

study area of the province of Reggio Emilia, are 

completely consistent with the geology of the area 

(Marchi, pers. comm., 2010). 

Other factors that may affect this accumulation are 

the chemical forms of the heavy metals involved, 

the content of organic matter and the concentration 

of hydrogen ions in the soil, and many other factors 

(Garcia et al., 2009) that, together, may play a 

decisive role in the process of bioaccumulation. 

Other aspects relate to the genera and species of 

mushrooms, and which are able to bioaccumulate 

heavy metals, and to what degree. It should also be 

remembered that aspects such as density, depth and 

age of the fungal mycelia; their lifecycles that can 

last for months to years – all these factors affect the 

growth of fungal fruiting bodies (carpophores), 

affecting and conditioning the processes of 

bioaccumulation. Knowledge about the 

mechanisms which transport heavy metals from the 

soil-substrate to the mycelium and from thence to 

the carpophores, however, are as yet poorly 

understood (Svoboda and Kalač, 2000). 

The data collection in this book is really quite vast, 

with more than 9,000 carpophore samples collected 

from all over Italy, corresponding to a universe of 

about 1,000 species of fungi. For each sample the 

concentrations of 32 elements were quantified. In 

70 

addition, approximately 350 surface soil samples 

were examined so as to obtain a better 

interpretation of bioaccumulation processes. For 

the surface soil samples, in addition to the 32 

elements, their respective concentrations of 

hydrogen ions were evaluated. 

The graphic representation of over 300,000 sample 

data cannot be shown and discussed in a 

comprehensive way here; therefore it was decided 

to present, evaluate and interpret data only for areas 

in the province of Reggio Emilia where carpophore 

collection was carried out with greatest intensity. 

These zones extend from State Road 9 up to the 

southern ridge of the Tosco Emiliano Apennines 

and include areas of the Tuscan mountainside (Alta 

Garfagnana and Lunigiana). 

We selected for examination the elements which, 

based on the scientific data and literature to hand, 

were the most significant and highly-studied: 

aluminium (Al), arsenic (As), cadmium (Cd), 

chromium (Cr), copper (Cu), mercury (Hg), nickel 

(Ni), lead (Pb) and zinc (Zn). Selenium (Se) was 

only examined for Boletus edulis, as it is not found 

to a significant degree in other taxa; Vanadium (V) 

and zirconium (Zr) that were investigated only for 

Amanita muscaria (L.) Lam., as they were not 

found to a significant degree in any other taxa. 

Also included in the study were those taxa that had 

an "abundance" of sufficiently large sample groups 

(a total of 24,000 concentration level values) in 

order to create distribution maps for the average, 

minimum and maximum concentration values, 

thereby forming a complete overview, which is 

summarised in Table 5. 

� Subdivision Basidiomycotina - Subclass 

Agaricomycetideae 

• Order Tricholomatales and its Genus 

Clitocybe. 

• Genus Amanita and the species Am. 

Muscaria. 

• Genus Russula. 

• Genus Agaricus and the sections 

Bitorques [Subgenus Agaricus (L. : Fr.) 

Heinem.] and Arvenses [SottoGenus 

Flavoagaricus Wasser]. 

• Group of Boletus edulis (B. aereus, B. 

reticulatus, B. edulis, B. Pinophilus).

� Subdivision Basidiomycotina - Subclass 

Aphyllophoromycetideae 

• Genus Cantharellus. 

• Genus Ramaria. 

� Subdivision Ascomycotina - Subclass 

Pezizomycetideae 

• Genus Morchella. 

Table 5. average, minimum and maximum values for the elements analysed relative to taxa included in the study. 

Order Tricholomatales 

Order Tricholomatales 

Genus Clitocybe 

Genus Amanita 

Amanita muscaria 

Genus Russula 

Genus Agaricus 


Section Bitorques 


Section Arvenses 

Genus Boletus 

Gruppo B. edulis 

Genus Cantharellus 

Genus Ramaria 

Genus Morchella 

The discussion deals separately with each of the 

various taxa considered. Below, there are 

distribution maps of certain elements in conjunction 

with evaluations regarding different 

enrichment factors and with data obtained from 

analysis of the principal components of this study. 

Al As Cd Cr Cu Hg Ni Pb Zn Se V Zr 

Average 217 1.0 2.37 1.94 36.4 0.35 1.37 0.94 157 - - - 

min Value 16 0.1 0.09 0.10 4.0 0.01 0.14 0.05 50.0 - - - 

Max Value 2400 59 14.5 63.5 93.0 4.0 24.5 18.8 340 - - - 

Average 178 0.4 1.08 1.26 93.7 2.24 1.44 1.97 98.2 - - - 


Max Value 1520 9.0 3.88 5.30 319 6.00 5.00 21.5 165 - - - 

Average 334 0.2 5.02 2.03 63.6 1.19 1.37 1.32 131 - - - 


Max Value 3410 7.0 33.4 29.9 740 24.3 10.1 64.3 328 - - - 

Average 225 0.4 12.2 1.01 31.7 0.73 0.84 0.91 138 - 99.0 4.66 

min Value 33 0.1 2.74 0.10 6.0 0.20 0.02 0.05 58.0 - 12.5 0.05 

Max value 1500 2.0 33.4 4.90 69.0 2.55 3.30 19.1 280 - 195 19.4 

Average 288 0.2 3.33 1.06 53.7 0.52 1.69 1.83 91.0 - - - 


Max Value 2940 7.0 24.6 6.10 189 4.06 13.5 53.6 864 - - - 

Average 488 0.1 1.58 6.07 161 2.29 3.15 1.09 110 - - - 


Max Value 1290 1.0 5.83 125 934 7.26 27.0 3.70 203 - - - 

Average 539 0.3 1.99 1.90 132 5.25 2.89 3.94 100 - - - 


Max Value 2770 4.0 9.20 7.40 812 25.3 14.1 29.5 273 - - - 

Average 119 1.5 40.9 0.74 188 4.99 2.39 2.52 156 - - - 


Max Value 797 21 391 10.9 1410 19.3 14.3 22.7 361 - - - 

Average 252 0.1 3.52 1.52 37.3 3.52 2.49 0.80 138 51.2 - - 

min Value 3 0.1 0.34 0.10 5.0 0.13 0.75 0.05 28.0 5.0 - - 

Max Value 2490 3.0 15.7 22.1 98.0 28.9 16.6 9.10 447 223 - - 

Average 270 0.1 0.49 2.91 42.9 0.23 2.33 1.97 71.3 - - - 


Max Value 1360 0.1 2.87 57.3 111 1.44 28.6 5.50 143 - - - 

Average 327 8.7 5.74 3.54 51.4 0.99 10.19 0.98 71.1 - - - 


Max Value 970 40 32.3 28.6 244 8.23 48.3 3.20 167 - - - 

Average 709 0.1 0.94 3.27 53.6 0.09 2.40 1.19 144 - - - 


Max Value 6100 3.0 4.12 30.7 123 0.25 12.2 11.0 281 - - - 

4.3.1 Order Tricholomatales 

(Subdivision Basidiomycotina – Subclass 

Agaricomycetideae) 

The distribution maps for the elements arsenic, 

cadmium, chromium, copper, mercury, nickel, lead 

and zinc, were far from uniform. Aluminium, 

71

chromium and nickel tend to have a similar 

concentration distribution and are generally 

consistent throughout, but it is interesting to note 

that for chromium and nickel there is a strong 

overlap with the distribution maps of soil. Figs. 38 

and 39, that show copper and zinc, highlight 

Cadmium, mercury and lead (figs. 40, 41 and 42) 

have higher levels in hilly areas and, only for 

cadmium, even in the mountainous border with 

Tuscany. 

72 

Fig. 38. Spatial distribution of copper for the order Tricholomatales. 

Fig. 39. Spatial distribution of zinc for the order Tricholomatales. 

common areas where concentrations are similar in 

proportion. In the Albinea and Quattro Castella 

areas, maximum concentration levels for arsenic 

were observed. Regarding pH, areas with more 

highly-acidic soils display correspondingly high 

levels of copper and zinc. 

Cadmium: the average concentration level was 2.37 

mg/kg. Slightly lower levels (1.67 mg/kg) were 

found east of the Black Sea (Demirbaş, 2001), 

while also in Turkey (Soylak et al., 2005) in Ma.

oreades and T. argyraceum 0.63 and 0.91 mg/kg 

were found. In the same area Yamaç et al. (2007) 

repeatedly found 0.58 and 1.99 mg/kg in I. 

geotropa and T. equestre. In the area of Epirus and 

Mercury: the average level of mercury was 

0.35mg/kg; this is far lower than in the Czech 

Republic in the proximity of silver mines, where 

Svoboda et al. (2006) found levels of 10.5 and 8.9 

Lead: the average concentration level was 0.94 

mg/kg. In an area east of the Black Sea, a level of 

0.06 mg/kg was observed in the species T. terreum 

Fig. 40. Spatial distribution of cadmium for the order Tricholomatales. 

Fig. 41. Spatial distribution of mercury for the order Tricholomatales. 

Macedonia, Ouzoun et al. (2009) found, 

respectively, concentrations of 1.8, 1.67 and 0.25 

mg/kg in three species, Ar. tabescens, Ar. mellea 

and T. rutilans. 

mg/kg in Le. nuda and Ar. mellea respectively. On 

the other hand, in Turkey, levels of 0.07 and 0.09 

mg/kg (Demirbaş, 2000) were recorded in 

La. laccata and T. terreum. 

(Demirbaş, 2001). In a recent study, significantly 

higher values were observed in central Spain by 

Campos et al. (2009). 

73

During analysis of I. geotropa, T. ustaloides and 

T. rutilans, the authors found levels of 4.73; 3.33 

and 3.23 mg/kg respectively (although the method 

of measurement used should be considered here). 

In the Black Sea area, Sesli et al. (2008) found a 

concentration of 2.6 mg/kg in Le. nuda. Demirbaş 

(2001) in the same area, found levels of 2.43 mg/kg 

in the species T. terreum, while a level of 0.86 

mg/kg was found in La. laccata (Demirbaş, 2000). 

In Turkey, Yamaç et al. (2007) found 1.22 and 

1.59 mg/kg in I. geotropa and T. equestre 

respectively. 

The enrichment-factor levels show that the 

elements tend to accumulate in carpophores and 

their accumulation usually varies with fungal 

species. In general it was observed that arsenic, 

chromium, nickel and lead do not tend to 

accumulate in the fungi of this order, despite the 

soil being rich in them (Kalač and Svobova, 2000; 

García et al., 2009). In contrast, cadmium, copper, 

mercury and zinc can accumulate in carpophores, 

with above average concentration factors, even 

when the soil only holds low concentrations. This 

has been further confirmed for mercury (Falandysz 

et al., 2002). 

In addition, Kalač and Svobova (2000), found 

enrichment factors equal to 50-300 and 30-500 for 

74 

Fig. 42. Spatial distribution of lead for the order Tricholomatales. 

In accordance with our own findings, in Macedonia 

and in the area of Epirus, Ouzouni et al. (2009), 

found levels of 0.79; 0.49 and 1.16 mg/kg in 

Ar. tabescens, Ar. mellea and Le. nuda 

respectively, while in Turkey (Soylak et al., 2005) 

levels of 1.05 and 1.89 mg/kg were recorded in Ma. 

oreades and T. argyraceum. In France; in the area 

surrounding Paris, C. nebularis, Ma. oreades and 

T. terreum were found to have levels equal to 42.5; 

33.6 and 24.3 mg/kg. 

cadmium and mercury and 0.1-0.2 for lead, as 

previously observed by Kalač et al. (1989b). 

The results obtained from principal component 

analysis are presented in Table 6. The first three 

components describe 63% of the total variance. In 

the first component, soil acidity does not seem to 

affect the increase in cadmium, chromium, copper 

and zinc; this is in contrast to the behaviour of lead 

as described by Kalač and Svobova (2000). Main 

component 2 describes the relationship between 

nickel, cadmium, mercury and lead that tend to 

build up under moderately acidic conditions. 

Cadmium, mercury and lead are all typically 

affected by human activities. The third component 

describes the relationship between alkalinity of the 

soil and high levels of arsenic bioaccumulation.

4.3.2 Genus Clitocybe (Subdivision 

Basidiomycotina - Subclass Agaricomycetideae 

– Order Tricholomatales) 

The genus Clitocybe, as a whole and also for 

selected species, will be examined here in 

comparison with species from the order 

Tricholomatales. In the genus Clitocybe, spatial 

distribution of element concentrations displays two 

types of associations. The first is represented by 

chromium, nickel and zinc with maximum levels of 

5.3, 6 and 165 mg/kg, respectively which were 

observed in the areas of Toano and Reggio Emilia. 

The total remaining area, with the exception of zinc 

concentrations in Collagna bore uniform 

concentration values. A second association is 

represented by aluminium and copper and it is the 

area of Reggio Emilia which provided the highest 

levels. 

The maps representing arsenic, mercury, cadmium 

and lead are very different both among themselves 

and compared to other distribution maps. They 

confirm that the only overlap with the soil acidity 

map is the mercury content in mushrooms. The 

Spatial distribution of element concentrations 

shows a few types of associations. The first is 

represented by chromium and nickel with the 

highest levels, 125 and 27 mg/kg, respectively, 

found in Montecchio; other areas displaying 

uniform concentration levels. A second association 

is represented by copper and lead with the area of 

Carpineti which registered the highest values for 

those elements. 

Aluminium, zinc and mercury partly overlap, while 

arsenic and cadmium are quite different both 

Table 6. Results of principal component analysis . 

Component 

1 2 3 

pH_s -.042 -.060 .803 

As_FA .006 .093 .721 

Cd_FA .437 .304 .003 

Cr_FA .839 .004 -.037 

Cu_FA .897 .191 -.055 

Hg_FA .209 .788 .054 

Ni_FA .242 .629 -.350 

Pb_FA -.102 .692 .131 

Zn_FA .932 -.013 -.019 

between themselves and with other distribution 

maps. The map showing soil acidity has a 

resemblance to that displaying the zinc and 

mercury content in mushrooms and also a 

quantitative overlap between the aluminium in the 

soil and the concentrations of aluminium in 

mushrooms has been observed. 

The maps in Figs. 43, 44 and 45 illustrate the 

concentration distribution of arsenic, copper and 

mercury, the average levels of which are, 

respectively: 0.43, 93.7 and 2.24 mg/kg. 

Arsenic: the areas of highest concentration were 

found between Casina and Canossa, where the 

maximum is 9 mg/kg, while for the rest of the 

territory, the distribution is mostly uniform. Other 

authors found higher values (1.76 mg/kg) in La. 

laccata (Demirbaş, 2001) at the Black Sea. In 

samples from the various European nations and 

Brazil Slejkovec et al. (1977) found a value of 0.66 

mg/kg in La. laccata, while their highest 

concentration was found in La. fraterna, having 30 

mg/kg. Konuk et al. (2007) reported a level of 0.44 

mg/kg in a sample of Ar. mellea collected in 

Turkey. 

Copper: Fig. 44 shows the distribution of 

copper. High values are present in the plains 

around the city of Reggio and area to the north of 

the Pedemontana road with a maximum level of 

319 mg/kg. In a previous study, different values 

were reported in La. laccata; respectively 12.9 and 

92.5 mg/kg (Demirbaş, 2000, 2001). 

75

In Macedonia and the area of Epirus, Ouzoun et 

al. (2009) have found levels of 17.38 and 17.47 

mg/kg in samples of Ar.mellea and Ar. tabescens 

respectively. The same authors, Ouzoun et al 

(2007), found concentrations of 16 and 4.65 mg/kg 

in Hy. eburneus and Hy. chrysodon. In Turkey 

Yamachiche et al. (2007) found 144.2 and 82.4 

Mercury: the Mercury map is shown in Fig. 45. 

The Quattro Castella site displays the highest 

76 

Fig. 43. Spatial distribution of arsenic for the genus Clitocybe. 

Fig. 44. Spatial distribution of copper for the genus Clitocybe. 

mg/kg in Le. nuda and I. geotropa respectively. In 

an area of the Black Sea, Sesli et al (2008) found 

concentrations of 32.8, 52.4 and 20.1 mg/kg in the 

species La. amethystina, Cl. gibba and Le. nuda. In 

France there was a concentration of 56.9 mg/kg in 

samples of Cl. nebularis (Michelot et al., 1998). 

levels, equal to 6 mg/kg. Moderately high values 

were also observed on the Apennine border. In

literature there are relatively few data on mercury. 

Demirbaş (2000, 2001), in two studies carried out 

east of the Black Sea, obtained levels which were 

different (0.39 and 0.072 mg/kg) and much lower 

than those observed in Reggio Emilia, by using and 

analyzing samples of La. laccata. Cocchi et 

al. (2006), in the province of Reggio Emilia have 

reported levels of 6.25, 0.12, 0.22, 0.77, 1.74 and 

1.78 mg/kg in Le. nuda, La. laccata, Hy. penarius, 

Hy. russula, Ly. decastes and Ma. oreades 

As has already been observed for other genera and 

species of fungi, arsenic, chromium, nickel and 

lead do not tend to accumulate in carpophores. This 

is evidenced by the enrichment factors for lead and 

confirmed by the studies of Garcia et al. (2009). In 

contrast the elements mercury (especially in Le. 

nuda) (Kalač et al., 1989b), cadmium, copper and 

Fig. 45. Spatial distribution of mercury for the genus Clitocybe. 

Table 7. Result of principal component analysis. 


1 2 3 

pH_s -.128 -.096 .961 

As_FA .975 .175 -.090 

Cd_FA .948 .256 -.123 

Cr_FA .701 .304 .267 

Cu_FA .980 .097 -.128 

Hg_FA .215 .964 -.105 

Ni_FA .988 .040 -.132 

Pb_FA .911 .312 -.085 

Zn_FA .977 .159 -.113 

respectively. In Cl. nebularis there was a 

significantly higher concentration, equivalent to 

62.9 mg/kg (Michelot et al., 1998). In the Czech 

Republic, in an area adjoining silver mines, 

Svoboda et al. (2006) found levels of 12.9 and 4.2 

mg/kg in Ar. mellea and Le. nuda. These values are 

considerably higher than both the average 

concentration level and the maximum value 

obtained in the province of Reggio Emilia. 

zinc tend to accumulate, but not in such a marked 

manner in Clitocybe carpophores, even though the 

soil is low in concentration levels of these. All this 

is confirmed for mercury by Falandysz et al. 

(2002). 


analysis are presented in Table 7. 

77

The first three components describe and explain 

94% of the total variance. It is noted that mercury 

is in contrast with other elements and is dissociated 

by the acidity of the soil. The second component 

reinforces what has been described, mercury being 

separated from the other elements, and the third 

constituent describes the "weak" link between 

acidity and chrome. The remaining elements are 

not related to pH. 

4.3.3 Genus Amanita (Subdivision 


- Order Amanitales) 

The chart of the concentration levels for arsenic, 

cadmium, chromium, copper, mercury, nickel, lead 

and zinc, was quite uniform. The exceptions are the 

pairs of aluminium and nickel and also chromium 

and cadmium that are qualitatively similar. As for 

the comparison between the concentration 

distributions in the mushrooms with those of the 

soil, the map does not appear related to the pH 

distribution, while the content of chromium, nickel 

and aluminium in the soil displays a 

correspondence with the mushrooms. 

Copper: the distribution of copper is shown in Fig. 

47. The average level is 63.59 mg/kg. The highest 

values are found in the plains of the province while 

78 

Fig. 46. Spatial distribution of cadmium for the genus Amanita. 

Cadmium: Fig 46 shows the distribution of 

cadmium, for which the average value is 2.5 mg/kg 

with a maximum of 33.4 mg/kg located in the 

Apennines. High levels are also found in the air 

between Vezzano and Viano. In Turkey, Tüzen 

(2003) found a similar level in Am. solitaria, equal 

to 7.5 mg/kg, while in Macedonia and in the area of 

Epirus, Ouzoun et al. (2009) found a rather modest 

level of 1.3 mg/kg in Am. caesarea. Similar values 

were found in the Black Sea in M. muscaria, Am. 

rubescens and Am. vaginata, with values of 1.6, 

0.79 and 0:56 mg/kg, respectively (Demirbaş, 

2001). In Turkey, Yamaç et al. (2007) measured a 

level of 2.46 mg/kg in Am. Caesarea, which is in 

the middle between what is stated in our 

investigation and that of the other authors cited. A 

significant number of samples from species in the 

genus of Amanita was analysed by Michellot et al. 

(1998) in the Paris region. These included Am. 

Excelsa var. excelsa, Am. gemmata, Am. muscaria, 

Am. ovoidea, Am. pantherina, Am. phalloides, Am. 

rubescens, Am. solitaria, Am. excelsa var. spissa 

and Am. vaginata. The values obtained were 

similar to those observed by us, being, respectively: 

6; 14.9; 13.9; 2.9; 10.3; 1.5; 2.4; 2.6; 2.5 and 7.7 

mg/kg. 

other areas are very uniform and quite close to the 

average level. A similar value of 50.8 mg/kg was 

found in Turkey in Am. Caesarea (Yamachiche et

al., 2007), while Tüzen (2003) found 96.2 mg/kg in 

Am. solitaria. Demirbaş (2001) recorded slightly 

erratic values of 23.5, 51.2 and 5.1 mg/kg in 

Am. muscaria, Am. rubescens and Am. vaginata in 

the Black Sea area. In Macedonia and Epirus, 

Ouzoun et al.(2009) recorded 19.3 mg/kg in 

samples of Am.Caesarea. Also regarding copper, 

Michellot et al. (1998) observed similar levels in 

Mercury: The highest values of mercury were 

found in the area of Casalgrande (24.3 mg/kg being 

the maximum level) (Fig. 48). 

The average value of 1.19 mg/kg is representative 

of a substantial part of the Reggio Emilia area. In 

the Czech Republic, in an area close to silver 

mines, Svoboda et al. (2006) found 1.55 mg/kg in 

the Am. rubescens. Values one order of magnitude 

lower were found in the Black Sea in the Am. 

muscaria (0.18 mg/kg), Am. rubescens (0.23 

mg/kg) and Am. vaginata (0.32 mg/kg) (Demirbaş, 

2001). In Poland, Falandysz et al. (2002) found 

Fig. 47. Spatial distribution of copper for the genus Amanita. 

Am. excelsa var. excelsa (75.6 mg/kg), Am. 

gemmata (44.4 mg/kg), Am. muscaria (28.4 

mg/kg), Am. ovoidea (21.7 mg/kg), Am. pantherina 

(38.5 mg/kg), Am. phalloides (29.7 mg/kg), Am. 

rubescens (41.9 mg/kg), Am. solitaria (24 mg/kg), 

Am. excelsa var. spissa (29.2 mg/kg) and Am. 

vaginata (65.9 mg/kg). 

levels between 0.07 and 1.5 mg/kg in the caps and 

between 0.021 and 1.3 mg/kg in the stalks of Am. 

muscaria. The mercury levels found by Michellot 

et al. (1998) appear excessively high in: Am. 

excelsa var. excelsa (61.3 mg/kg), Am. Gemmata 

(37.4 mg/kg), Am. muscaria (61.3 mg/kg), Am. 

ovoidea (61.4 mg/kg), Am. pantherina (64.9 

mg/kg), Am. phalloides (40.3 mg/kg), Am. 

rubescens (57 mg/kg), Am. solitaria (48.8 mg/kg), 

Am. excelsa var. spissa (58.4 mg/kg) and Am. 

vaginata (54.3 mg/kg). 

79

4.3.4 Amanita muscaria (L.) Lam. 

(Subdivision Basidiomycotina - Subclass 

Agaricomycetideae - Order Amanitales - 

Genus Amanita) 

For Am. muscaria, a medium-high to high 

frequency species, distribution maps of vanadium 

and zirconium (Figs. 49 and 50) have been drawn 

because the concentrations of these elements in this 

species are by all means exceptional. The areas 

80 

Fig. 48. Spatial distribution of mercury for the genus Amanita. 

Fig. 49. Spatial distribution of vanadium for Am. muscaria. 

with the greatest concentrations are located in the 

Apennines, where vanadium and zirconium occur 

in their highest levels of 195 and 19.4 mg/kg 

respectively. From the concentration levels of the 

two elements and the soil pH levels, no direct 

relationship is evident, as was the case with 

samples of Am. muscaria. The two elements are 

bioaccumulated regardless of the concentration 

present in soils and their acidity.

It has been noted that arsenic, chromium, nickel 

and lead do not tend to accumulate in the genus 

Amanita, even if the soil is rich in these elements 

(Kalac and Svobova, 2000, García et al., 2009). 

The elements copper and zinc tend to accumulate 

but to a lesser extent, while cadmium and mercury 

accumulate abundantly (Sova et al., 1991; Vetter, 

Fig. 50. Spatial distribution of zirconium for Am. muscaria. 

Table 8. Results of principal components analysis. 


1 2 3 4 5 

Al_a -.096 .017 .832 -.035 .054 

As_a -.038 .139 -.221 -.104 .099 

Cd_a -.084 .611 -.138 -.004 .077 

Cr_a .091 -.026 .721 -.058 -.054 

Cu_a -.065 .025 .069 .802 .137 

Hg_a .129 -.043 .057 .612 .065 

Ni_a .006 -.104 .884 -.031 .009 

Pb_a -.091 .186 .272 .020 -.052 

Zn_a -.104 .432 -.048 .664 .026 

Al_s .009 .776 .040 -.093 .457 

As_s .293 .627 .011 .171 -.049 

Cd_s .730 -.083 -.056 .106 .010 

Cr_s .872 .227 .027 .075 .086 

Cu_s .026 .056 -.056 .259 .917 

Hg_s .086 -.599 -.032 .077 -.098 

Ni_s .913 -.163 .004 -.031 -.003 

Pb_s .211 -.263 -.181 .613 -.036 

Zn_s .070 .277 -.088 .002 .913 

1994). This has been confirmed by Falandysz et al. 

(2002) for the element mercury. Kalač and 

Svobova (2000) examined enrichment factors equal 

to 50-300 and 30-500 for cadmium and mercury. 

The results obtained from principal components 

analysis are presented in Table 8. The first five 

components explained 59% of the total variance. 

81

In component one, nickel and chromium in 

particular and also cadmium are linked together 

and fall marginally below the other elements in 

soils and fungi. In component two, cadmium and 

zinc tend to increase in mushrooms with increasing 

aluminium and arsenic in soils, while the mercury 

in the soil decreases. The third component 

describes the link between aluminium, chromium 

and nickel in mushrooms of terrigenous origin, in 

contrast again with falling arsenic levels in 

82 

mushrooms. The fourth component shows an 

increase of copper and mercury in mushrooms with 

lead, while the remaining elements do not change. 

The fifth component concerns aluminium, copper 

and zinc in soils. 

As for the enrichment factor, the results obtained 

from the principal components analysis are 

presented in Table 9. These are related to 

enrichment factors, and the first three components 

describe and explain 93% of the total variance. 

Table 9. Results of principal components analysis in Amanita muscaria (L.) Lam.. 

Base levels in soils are directly correlated with the 

bioaccumulation of almost all elements except for 

copper and lead, while nickel is predominantly 

bioaccumulated in acidic soils. Component two 

explains how acidic soils are in particular 

opposition to vanadium, cadmium, copper, nickel 

and zinc. The third component describes a strong 

relationship between pH and lead, an increase in 

one also increases the other and vice versa. 

4.3.5 Genus Russula (Subdivision 


- Order Russulales) 

An overview and comparison of the maps showing 

arsenic, cadmium, chromium, copper, mercury, 

nickel, lead and zinc reveal a rather heterogeneous 

distribution of concentrations. 

Arsenic: for this element (Fig. 51), the distribution 

is constant over the entire investigated area. The 

1 


2 3 

As_FA .996 -.013 -.050 

Cd_FA .867 .451 -.044 

Cr_FA .926 .154 .170 

Cu_FA .112 .973 -.113 

Hg_FA .941 .063 .304 

Ni_FA -.265 .922 .018 

Pb_FA .070 -.088 .954 

Zn_FA .279 .923 -.187 

V_FA .587 .759 -.126 

Zr_FA .984 -.123 .110 

pH_s .416 -.460 .549 

higher values of 7 and 5 mg/kg were found near 

Ligonchio and Viano. The average level is 0.2 

mg/kg. 

Higher concentration values in four species of 

Russula were found by Demirbaş (2001) to the east 

of the Black Sea. In central Finland concentrations 

ranging from 0.1 to 0.5 mg/kg were found 

(Nikkarinen and Mertanen, 2004). 

Lead and Zinc: Figs. 52 and 53 illustrate the 

spatial distribution of lead and zinc concentrations 

that appear to be qualitatively similar, except for a 

small area represented by the area of the city of 

Reggio Emilia, where the levels of lead appear to 

be significantly higher. These levels are due to 

motor vehicle fuel containing tetraethyl lead as an 

antiknock additive (Cenci et al., 2008). The 

average level is 1.8 mg/kg, in agreement with that 

found in Finland (1.4 ÷ 2 mg/kg; Demirbaş, 2001) 

and Turkey (2.3 mg/kg; Tüzen, 2003).

Still focusing on Turkey, the lead concentration in 

Ru. delica was 3.89 mg/kg (Demirbaş, 2000), while 

Konuk et al. (2007) found the rather small level of 

0.03 mg/kg. This value, judging by observations of 

other elements in different fungi, seems somewhat 

underestimated. For zinc the average concentration 

Fig. 51. Spatial distribution of arsenic for the genus Russula. 

Fig. 52. Spatial distribution of lead for the genus Russula. 

of 91mg/kg was higher than average levels, that 

ranged between 19 and 32 mg/kg (Demirbaş, 

2001), whereas the value of 78 mg/kg repored from 

Turkey (Tüzen, 2003) looks quite similar to the one 

observed in our study. 

83

The values of enrichment factors showed that 

arsenic, chromium, nickel and lead do not tend to 

accumulate in mushrooms even if the soil has high 

concentrations of these elements. This phenomenon 

has been further confirmed by Garcia et al. (2009). 

By contrast, cadmium, copper, zinc and mercury 

tend to accumulate in the fruiting bodies of fungi 

even when these elements are at low concentrations 

84 

Fig. 53. Spatial distribution of zinc for the genus Russula. 

Table 10. Results of principal component analysis. 

Components 

1 2 3 4 5 

pH_s .325 .123 .282 -.662 -.403 

Al_r -.083 .850 .022 .059 .136 

As_r .075 .139 -.096 -.144 .365 

Cd_r -.011 -.162 -.332 -.127 .018 

Cr_r .058 .904 .101 .017 .101 

Cu_r .110 -.204 -.472 .165 .129 

Hg_r .188 .090 -.148 -.031 -.258 

Ni_r -.030 .767 .073 -.143 -.235 

Pb_r .203 .383 -.358 -.005 .476 

Zn_r .239 .181 -.071 .046 .289 

Al_s .065 .146 -.004 .719 -.491 

As_s .196 -.068 .114 .893 -.083 

Cd_s .196 -.211 .574 .051 .471 

Cr_s .267 -.033 .851 .064 -.183 

Cu_s .939 -.018 .047 -.072 .116 

Hg_s .027 -.080 .055 -.043 .681 

Ni_s .094 -.061 .884 -.102 .166 

Pb_s .834 -.086 .213 .116 -.060 

Zn_s .910 .009 .060 .067 .124 

in the soil (Kalac and Svoboda, 2000). This has 

been confirmed for mercury in Ru. emetica 

(Falandysz et al., 2002) and for cadmium in Ru. 

cyanoxantha (Vetter, 1994). 


analysis are presented in Table 10. The first five 

components describe and explain 60% of the total 

variance.

In component 1, only copper, lead and zinc are 

associated with soil pH. With increasing pH the 

three elements increase, therefore lower soil acidity 

is directly related to an increase in the 

concentration levels of those three elements. The 

association of chromium and nickel in the genus 

Russula is affected by aluminium. The same 

association, along with cadmium, can be found in 

soil and it represents a countertrend to copper and 

lead in mushrooms. Increases in the aluminium 

concentration in soil are associated with increases 

in arsenic and run counter to pH levels. In the fifth 

component we can see a divergence between the 

pH level and aluminium against arsenic and lead in 

soil and cadmium and mercury in mushrooms. For 

these last two items the increase in concentration 

inside carpophores is independent of concentration 

levels of the substrate (Kalač et al., 1989b; Jorhem 

and Sundström, 1995; Falandysz et al., 2002). 

4.3.6 Genus Agaricus (Subdivision 


- Order Agaricales) 

Spatial distribution of the element concentrations 

highlights a few types of "associations". The first is 

represented by chromium and nickel with the 

highest values, 125 and 27 mg/kg respectively. 

They are characteristic of the area of Montecchio, 

while the remaining area displays mainly uniform 

Fig. 54. Spatial distribution of cadmium for the genus Agaricus. 

concentration values. A second association is 

represented by copper and lead: we registered the 

maximum levels in the area of Carpineti. The levels 

of aluminium, zinc and mercury partly overlap, 

while arsenic and cadmium are quite different both 

from each other and from other concentration 

distribution maps. The map showing soil acidity 

has a resemblance to that displaying zinc and 

mercury in mushrooms: one even notes a 

quantitative overlap between the aluminium in the 

soil; apparently, aluminium accumulates in fungi. 

Maps 54, 55 and 56 illustrate the concentration 

distribution of cadmium, mercury and lead. The 

average levels are respectively 1.58, 2.29 and 1.09 

mg/kg. 

Cadmium: the areas of highest concentration were 

found near Carpineti, Vezzano and Reggio Emilia. 

Demirbaş (2001) reported similar values, of 3.84 

and 1.04 mg/kg in Ag. bisporus and Ag. Silvicola 

near the Black Sea. The same author in Turkey 

(Demirbaş, 2000) reported 2.17 mg/kg by 

analysing the species Ag. bitorquis. In Greece 

Ouzoun et al. (2007) found 0.15 mg/kg of cadmium 

in Ag. cupreobruneus. 

Mendil et al. (2004) reported a value of 0.1 mg/kg 

in Ag. bisporus in an area with high vehicular 

traffic in Turkey. 

85

Mercury: the distribution of mercury 

concentrations over the territory was not uniform 

(Fig. 55). The highest values were seen in the area 

of Carpineti. Also, those areas comprising the 

towns of Castelnovo Monti, Vetto and Ramiseto 

have comparatively high values. 

in the Black Sea Demirbaş (2001) recorded lower 

levels, 0.6 and 0.15 mg/kg, in Ag. bisporus and Ag. 

Lead: Fig. 56 shows that the highest levels are 

those in the areas of Carpineti and the town of 

86 

Fig. 55. Spatial distribution of mercury for the genus Agaricus. 

Fig. 56. Spatial distribution of lead for the genus Agaricus. 

silvicola respectively. The same author (Demirbaş, 

2000) reported 0.14 mg/kg in Ag. bitorquis in 

Turkey. These levels are significantly lower than 

those observed in this study. In the Czech Republic, 

in an area close to silver mines the average 

concentration levels reported for Ch. rhacodes 

were 2.59 mg/kg (Svoboda et al., 2006). 

Reggio Emilia. The latter probably due to its high 

volume of vehicular traffic.

Different levels were reported by Demirbaş (2000, 

2001): in samples of Ag. bitorquis, Ag. bisporus 

and Ag. Silvicola they found 1.34, 2.41 and 0.92 

mg/kg respectively. Mendil et al. (2004), reported a 

level of 6.9 mg/kg in Ag. bisporus in an area with 

high traffic levels in Turkey. Campos et al. (2009) 

found a 2.53 mg/kg in an Ag. campestris sample in 

central Spain. García et al. (2009), in the province 

of Lugo (Spain) analyzed Ag. bisporus, Ag. 

campestris, Ag. urinascens and Ag. silvicola, and 

found concentrations in the mushroom caps of 0.35, 

3, 1.4 and 1.4 mg/kg. The same authors evaluated 

enrichment factors, concluding that lead does not 

tend to accumulate in fungi, even if it is present in 

high concentrations in the soil. 

Arsenic, chromium, nickel and lead do not tend to 

accumulate in fungi. This aspect has been shown by 

enrichment factors, and convincingly confirmed for 

lead by García et al. (2009). In contrast the 

elements mercury (in particular), cadmium, copper, 



1 2 3 4 

pH_s .486 .462 .621 .086 

Al_s .203 -.562 .284 .727 

As_s .551 -.592 .187 .538 

Cd_s .961 -.180 .078 -.062 

Cr_s .898 .056 .286 .274 

Cu_s .944 .242 .186 -.028 

Hg_s .102 .684 -.102 -.266 

Ni_s .979 .120 .144 -.032 

Pb_s .918 -.019 .256 .269 

Zn_s .944 -.264 .123 .083 

AL_a .101 -.756 .305 -.074 

As_a .014 -.440 .159 -.755 

Cd_a -.003 .835 .253 .055 

Cr_a .420 -.219 .810 .017 

Cu_a -.190 .710 -.133 -.017 

Hg_a -.495 .359 -.068 -.486 

Ni_a .301 .214 .828 -.107 

Pb_a .177 .811 .186 .032 

Zn_a .002 .256 -.838 -.144 

and zinc tend to accumulate in the genus Agaricus, 

even where the soil has low concentrations, 

however this accumulation is not particularly strong 

or marked. This was confirmed for the element 

mercury by Falandysz et al. (2002). 


analysis are presented in Table 11. The first four 

components describe and explain 75% of the total 

variance. 

The first component shows that the mercury in 

mushrooms goes against the majority of elements 

in the soil and is not affected by the mercury 

content of the soil itself. The second component 

supports this idea and links pH levels to cadmium, 

lead and copper in mushrooms in contrast to 

aluminium and arsenic in the soil. The third 

component describes the link between nickel and 

chromium in fungi with the soil pH level. The 

fourth component explains how the concentration 

of arsenic in mushrooms is independent and 

detached from soil. 

87

4.3.7 Section Bitorques (Subdivision 


- Order Agaricales – Genus 

Agaricus – Subgenus Agaricus) 

Only one pattern adequately summarises the spatial 

distribution of the concentration levels of arsenic, 

chromium, mercury and nickel. For arsenic and 

other more-strongly “associated” elements, the 

areas of highest concentration are between Canossa 

and the northern border of the Apennines, for 

chromium and mercury we can also include the 

area of Reggio Emilia and Castellarano. As regards 

the other elements, aluminium reaches its highest 

levels (2,771 mg/kg) in the Apennine zone of 

Ramiseto; cadmium (9.2 mg/kg) in the area of Villa 

Minozzo, copper (812 mg/kg) in Casalgrande and 

Castellarano, lead (29.5 mg/kg) in certain areas 

88 

near the city of Reggio Emilia and zinc (273 

mg/kg) in Montecchio. 

Overlap with the values in soils is possible with 

nickel and to a lesser extent for chromium and lead. 

The remaining elements and pH levels have 

apparently no direct relationship to the 

concentration levels in mushrooms. 

Arsenic: the distribution of the concentration levels 

of arsenic is shown in Fig. 57. An average level of 

0.3 mg/kg was found over most of the territory. 

The maximum levels were found in a vast area 

ranging from Canossa to Ligonchio. Slejkovec et 

al. (1977), analysed dozens of mushroom samples 

from European countries and Brazil and found a 

level of 1 mg/kg in the species Ag. bisporus, which 

is three times higher than that we found. Always in 

the Black Sea, Demirbaş (2001) reported a 

concentration level of 0.76 mg/kg in Ag. bisporus. 

Fig. 57. Spatial distribution of arsenic for the section Bitorques of the genus Agaricus. 

Cadmium: Fig 58 shows the distribution of 

cadmium. The highest levels were recorded in the 

Apennine area and the whole southern zone that 

runs from Casalgrande to Villa Minozzo. The 

average level, equal to 1.99 mg/kg, is characteristic 

of a good part of the territory. Values higher than 

those we encountered were reported by Demirbaş 

(2000, 2001) to the east of the Black Sea and in 

Turkey respectively in Ag. bisporus (3.48 mg/kg) 

and Ag. bitorquis (2.17 mg/kg). Mendil et al. 

(2004) in an area of high traffic in Turkey reported 

an extremely low level of 0.1 mg/kg in the species 

Ag. bisporus, a fact not easily explained, because 

from the environmental context in which it was 

found one would expect far higher concentration 

levels. 

Lead: lead is shown in Fig. 59. The average level 

of 3.94 mg/kg was found in almost every part of 

the territory investigated, the highest values for this 

species were in the vicinity of the city of Reggio 

Emilia. Mendil et al. (2004) reported a level of 6.9 

mg/kg in Ag. bisporus in an area with high 

vehicular traffic.

Fig. 58. Spatial distribution of cadmium for the section Bitorques of the genus Agaricus. 

Demirbaş (2000, 2001) found a level slightly below 

the average level of 2.41 mg/kg in Ag. bisporus 

east of the Black Sea, while in Turkey on Ag. 

bitorquis the concentrations were significantly 

lower (1.34 mg/kg). García et al. (2009), in the 

province of Lugo (Spain), analysed Ag. bisporus 

and observed levels of 0.35 mg/kg in the caps and 

0.54 mg/kg in the stems which represent fairly low 

concentration levels. 

In the Paris region, Michellot et al. (1998) observed 

31 mg/kg in Ag. maleolens and 32 mg/kg in Ag. 

silvaticus. 

Fig. 59. Spatial distribution of lead for the section Bitorques of the genus Agaricus. 

Arsenic, chromium, nickel and lead do not tend to 

accumulate in the fruiting bodies, while mercury 

(Vetter, 1994), cadmium, copper, and zinc, in 

descending order tend to accumulate in the fruiting 

bodies of section Bitorques, but not in a 

particularly marked manner. 

89

The results of the principal component analysis are 

presented in Table 12. The first four components 

Component one describes how a great deal of the 

elements in soils are themselves linked and present 

a countertrend to arsenic and, to a lesser extent with 

cadmium, in mushrooms. Component two in 

particular describes the conflict that exists between 

the concentrations of mercury, zinc, cadmium and 

copper in soils in relation to those found in fungi. It 

also describes the lack of any link between 

chromium, nickel, and to a lesser extent for arsenic 

and lead in mushrooms with the elemental contents 

of the soil. Component three groups several 

elements in soil which have trends contrary to 

mercury, while the fourth constituent explains how 

cadmium and zinc in fungi are not related to the 

concentration levels in the soil for the same 

elements and are both opposed to arsenic in soil. 

4.3.8 Section Arvenses (Subdivision 


- Order Agaricales – Genus 

Agaricus - Subgenus Flavoagaricus) 

Detailed observation rules out significant 

similarities between the concentration distributions 

90 

Table 12. Results of the principal component analysis. 


1 2 3 4 

Al_b -.013 .667 -.165 .210 

As_b -.425 .375 .051 -.369 

Cd_b -.125 -.108 -.027 .825 

Cr_b .002 .939 .069 -.012 

Cu_b .037 .197 .101 .812 

Hg_b .470 .607 -.071 .035 

Ni_b .013 .858 .138 -.191 

Pb_b .516 .195 .353 .080 

Zn_b .370 .522 .202 .395 

Al_s .315 -.007 .761 .215 

As_s .234 .220 .856 .041 

Cd_s .849 .202 .033 .113 

Cr_s .728 .015 .525 -.192 

Cu_s .050 -.232 .568 -.084 

Hg_s .464 -.417 -.650 -.111 

Ni_s .709 .040 .529 -.208 

Pb_s .930 .030 -.034 -.061 

Zn_s .702 -.164 .431 .047 


of the elements investigated. A kind of association 

can only be seen for aluminium and lead. The 

remaining elements generally show far from 

uniform distributions. For cadmium the highest 

levels (390 mg/kg) were found between Ligonchio 

and Collagna, while in two areas (Casalgrande and 

Ligonchio-Collagna) the highest concentration 

level (19.2 mg/kg) was recorded for mercury.. For 

chromium, the distribution was fairly uniform 

throughout the area: the highest levels (10.9 mg/kg) 

were seen in the central Apennines. The maximum 

concentration level for copper (1.41 mg/kg), was 

recorded in the city of Reggio Emilia, where 

anthropogenic activity is of considerable 

importance, and to a lesser extent, on the border of 

the Apennines. Zinc and nickel were recorded as 

having their maximum levels, respectively, in 

Montecchio-Cavriago (361 mg/kg) and Quattro 

Castella (14.3 mg/kg). 

Arsenic: Fig 60 shows the distribution of arsenic in 

the Reggio Emilia area. The maximum 

concentration of 21 mg/kg was reported in Villa 

Minozzo. Other areas of high concentration were

observed, while the average level of 1.49 mg/kg 

was recorded over the majority of the territory. 

Cocchi et al. (2006) reported levels of 1.06 and 

1.52 mg/kg in Ag. arvensis and Ag. silvicola 

respectively. In samples from some European 

countries and Brazil, Slejkovec et al. (1977) found 

6.24 and 3.32 mg/kg in Ag. silvicola and Ag. 

Macro-carpus, levels significantly higher than 

those observed in our study. 

Fig. 60. Spatial distribution of cadmium for section Arvenses of genus Agaricus. 

Lead: Fig. 61 shows that the maximum level (22.7 

mg/kg), can be found in the vicinity of the city of 

In the region of Paris (France), Michelot et al. 

(1998) analysed 92 species of mushroom including 

Fig. 61. Spatial distribution of lead for section Arvenses of genus Agaricus. 

Reggio Emilia. The average concentration level of 

2.52 mg/kg was found across most of the territory. 

Ag. arvensis, Ag. silvicola, Ag. altipes, finding 

levels of 22, 31 and 33.4 mg/kg respectively. 

91

Demirbaş (2001) reported an average level of 0.92 

mg/kg in Ag. silvicola, to the east of the Black Sea. 

In the province of Lugo (Spain), Garcia et al. 

(2009) found the same level (1.4 mg/kg) in both the 

cap and the stem of Ag. silvicola. Cocchi et al. 

(2006) reported 1.78 and 3.08 mg/kg in Ag. 

arvensis and Ag. silvicola respectively in Reggio 

Emilia. 

For the section Arvenses, as well as for the other 

taxa presented here, the elements arsenic, 

chromium, nickel and lead tend not to accumulate 

in the fruiting bodies, while cadmium, mercury, 

copper and zinc, in descending order, do, despite 

low concentrations of these elements in the soil. 

4.3.9 Group Boletus edulis (Subdivision 

Basidiomycotina – Subclass 

Agaricomycetideae - Order Boletales) 

For the group Boletus edulis (B. aereus, B. 

reticulatus, B. edulis, B. pinophilus), in addition to 

arsenic, cadmium, chromium, copper, mercury, 

nickel, lead and zinc, selenium was also taken into 

consideration. An overview and a comparison of 

the spatial maps showing the distribution of the 

92 

Table 13. Rresults of the principal component analysis. 


presented in Table 13. The first three components 


The first component comprehensively describes the 

aggregation of cadmium, mercury, copper and zinc 

which run in countertendency to the other elements 

in the soil and soil acidity. The second component 

associates with arsenic and lead, while the third is 

influenced by chromium and nickel, and is opposed 

to soil acidity and other elements. It seems clear 

how these two groups of elements, having a totally 

different behaviour in bioaccumulation, were 

separated during the analysis of the main 

constituents. 


1 2 3 

As_FA .168 .703 .022 

Cd_FA .978 .048 .026 

Cr_FA .271 .033 .427 

Cu_FA .937 .225 .160 

Hg_FA .928 .093 .074 

Ni_FA -.121 .204 .827 

Pb_FA .137 .891 -.047 

Zn_FA .784 .512 .107 

pH_s -.119 .314 -.741 

element concentrations, highlights four types of 

"associations." The first is represented by 

aluminium, chromium, nickel and lead (figs. 62 → 

65). The maximum concentration levels are located 

in the hilly areas of Casina and Canossa. As for the 

similarities with the soil, there are only affinities 

for chromium and nickel, as also evinced by other 

species of fungi.

Average concentration levels are respectively 252 

mg/kg for aluminium; 1.52 for chromium, 2.49 for 

nickel and 0.8 mg/kg for lead. A second group is 

composed of copper, mercury and zinc, the average 

levels of which are respectively 37, 3.52 and 138 

mg/kg. The third group is composed of cadmium 

and selenium, with maximum levels (15.7 and 223 

mg/kg, respectively) found at the border with 

Tuscany. The fourth group is formed by arsenic 

Fig. 62. Spatial distribution of aluminium for the group Boletus edulis. 

Fig. 63. Spatial distribution of chromium for the group Boletus edulis. 

only: its maximum level of 3 mg/kg was observed 

near Quattro Castella. 

Similar concentration levels of almost all elements 

analysed were found in samples of B. edulis in 

Finland (Nikkarinen and Mertanen, 2004). We 

found lead, selenium and zinc to have lower levels, 

however at: 0.18, 18.5 and 91 mg/kg. Falandysz et 

al. (2008) found selenium at lower amounts still, 

ranging between 8.7 and 32 mg/kg in the 

mountainous regions of Poland. 

93

In Greece Ouzoun et al. (2009) observed lower 

concentration levels for cadmium (0.23 mg/kg), 

chromium (0.86 mg/kg), nickel (1.61 mg/kg), lead 

0.09 mg/kg) and zinc (89 mg/kg), and an almost 

identical level to copper (41 mg/kg) in B. aereus. 

Higher concentration values in the Boletaceae were 

found by Demirbaş (2001) in an area east of the 

Black Sea for arsenic (1.41 mg/kg), nickel (65 

mg/kg) and lead (6.9 mg/kg). Lower levels were 

found by the same author for cadmium (1.36 

94 

Fig. 64. Spatial distribution of nickel for the group Boletus edulis. 

Fig. 65. Spatial distribution of lead for the group Boletus edulis. 

mg/kg), chromium (0.86 mg/kg), copper (11.5 

mg/kg), mercury (0.48 mg/kg) and zinc (19.6 

mg/kg). 

In the region of Paris (France), Michelot et al. 

(1998) analysed 92 species of mushroom including 

B. edulis and found 5.35 mg/kg for nickel and 21.2 

mg/kg for lead. Smaller amounts were found for 

cadmium (1.39 mg/kg), chromium (1.34 mg/kg), 

copper (14.9 mg/kg), mercury (40.6 mg/kg) and 

zinc (55.4 mg/kg). In a recent paper by Frankowski

et al. (2010), the concentration levels of heavy 

metals in specimens of B. edulis collected in 

Poland are reported. The concentrations in the 

mushroom caps are significantly higher (Cd 5.5; Cu 

47; Hg 4.9 and Zn 190 mg/kg). Similar levels were 

observed in the mountains in Poland (Falandysz et 

al., 2008); the following concentration intervals 

have been proposed: Cd (4-18 mg/kg), Cu (26-57 

mg/kg), Hg (0.95-2.39 mg/kg) and Zn (150-210 

mg/kg). 

Enrichment factors confirm the tendency of 

arsenic, chromium, nickel and lead to not 

bioaccumulate, even in soils with high 

concentrations of the same elements. This is in 

strong contrast to copper, zinc and especially 

cadmium, mercury and selenium, which have a 

pronounced ability to bioaccumulate in the fruiting 

The results of the principal component analysis, 

which relate the concentrations of heavy metals in 

mushrooms and the soil as well as pH levels, are 

presented in Table 14. The first four components 




1 2 3 4 

Al_b .918 .020 -.042 -.202 

Cd_b -.153 -.086 .394 .436 

Cr_b .952 .064 .054 -.222 

Cu_b -.037 -.048 .731 .160 

Hg_b .195 .007 .418 .141 

Ni_b .936 -.052 -.005 -.199 

Pb_b .939 -.001 -.069 -.171 

Se_b -.022 -.251 .157 .483 

Zn_b -.321 .113 .692 -.030 

pH_s .130 .398 -.401 -.621 

Al_s -.353 -.056 -.027 .682 

As_s -.200 .211 -.281 .751 

Cd_s .140 .476 -.689 .169 

Cr_s .790 .322 -.029 .204 

Cu_s .070 .876 -.037 .041 

Hg_s .229 -.095 .808 -.237 

Ni_s .825 .339 .037 -.132 

Pb_s .080 .667 -.157 -.369 

Se_s .076 .576 -.423 .640 

Zn_s .166 .917 .062 -.078 

bodies of fungi even at low concentrations of 

metals in the soil. This has been confirmed by 

García et al. (2009); Falandysz et al. (2002) Jorhem 

and Sundström (1995), Kalač et al. (1989b) and 

Cocchi et al. (2006). In particular, the last group of 

researchers has shown that the species of the 

Boletus edulis are able to accumulate large 

quantities of selenium. A further confirmation 

comes from a study by Frankowski et al. (2010) 

who also analysed the soil collected in the vicinity 

of the fungi. The enrichment factors were higher 

for Zn, Cd, Cu and Hg, in ascending order, 

demonstrating the ability of Boletus edulis to 

bioaccumulate these four elements, even if the soil 

has low concentrations of them (Zn 22; Cd 0.35, 

Cu 2.8 and Hg 0.04 mg/kg). 

In component one, it is clear that the pH does not 

affect and is not linked to any elements either in the 

fungi or in the soil, while chromium and nickel in 

soil have a similar qualitative behaviour as they do 

in fungi in addition to lead and aluminium. Zinc in 

95

species of Boletus edulis and aluminium in soils 

exist in countertendency. 

Component two shows that, as Cocchi et al. (2006) 

observed, selenium in the species of the Boletus 

edulis is not related to the concentration of the 

same element in the soil. Copper, lead and zinc are 

linked and tend to increase regardless of the levels 

in the soil. The third constituent demonstrates that 

soil pH, selenium and cadmium in the soil are 

linked together and in countertendency to copper, 

zinc and mercury in fungi, and to the latter element 

in the soil. This highlights the bioaccumulating 

properties of mercury (Falandysz et al., 2002). The 

fourth constituent binds cadmium and selenium in 

4.3.10 Genus Cantharellus (Subdivision 

Basidiomycotina – Subclass 

Aphyllophoromycetideae – Order 

Cantharellales) 

The maps show a similarity between the couples of 

cadmium/mercury and chromium/nickel (Figs. 66 

→ 69). The remaining elements (arsenic, copper, 

lead and zinc) have very heterogeneous 

concentration distributions. 

Cadmium and mercury: these have average levels 

of 0.49 and 12.23 mg/kg respectively, while their 

highest levels are found in hilly areas of Casina and 

96 

Table 15. Rresults of principal component analysis. 

the species of the Boletus edulis to arsenic and 

selenium in the soils and in countertendency to lead 

in soils, thereby confirming the strong capacity for 

bioaccumulation of selenium (Cocchi et al., 2006). 

Enrichment factors (Table 15) show that the pH is 

unconnected to most elements. Component two 

clarifies that high acidity levels are inversely 

proportional to high concentration levels of 

chromium, mercury, and selenium, themselves 

running in countertendency to lead. This is 

illustrated by the values of main component three. 


1 2 3 

pH_s .055 -.428 -.769 

As_FA .915 .361 -.042 

Cd_FA .686 .003 .212 

Cr_FA .419 .869 -.047 

Cu_FA .922 .317 -.012 

Hg_FA .179 .946 .019 

Ni_FA .833 .306 .236 

Pb_FA .301 -.244 .677 

Se_FA .436 .835 .183 

Zn_FA .857 .435 .099 

Baiso and in the mountain range between Busana 

and Villa Minozzo. In the area of Epirus and 

Macedonia, Ouzoun et al. (2009) recently reported 

cadmium concentrations of 0.38 mg/kg, in perfect 

agreement with those observed in this study. Also 

in Greece (Ouzoun et al. (2007) 0.41 mg/kg of 

cadmium was found in Ca. cibarius. For mercury, 

in the Czech Republic, in an area with many silver 

mines, the average level was 0.25 mg/kg (Svoboda 

et al., 2006). In Svoboda et al. (2000), reported 

mercury concentrations were similar to those 

observed by us in the Reggio Emilia Apennines.

Chromium and nickel: these elements (Figs. 68 

and 69) have average levels of 2.9 and 2.3 mg/kg 

respectively, while their maximum levels are 57 

and 29 mg/kg. These levels were found in the 

mountains between Vetto and Ramiseto. There 

seems to be no link between these and soil acidity, 

although concentration levels in the soil do 

correlate to mushroom concentration levels. In 

Greece, lower concentration levels of chromium 

Fig. 66. Spatial distribution of cadmium for the genus Cantharellus. 

Fig. 67. Spatial distribution of mercury for the genus Cantharellus. 

(1.6 mg/kg) and nickel (1.1 mg/kg) were observed 

(Ouzoun et al., 2009). 

Enrichment Factor values showed, like the genus 

Russula, little tendency for arsenic, chromium, 

nickel and lead to bioaccumulate in these 

mushrooms despite the soil being rich in the same 

elements. On the other hand, cadmium, copper, 

mercury and zinc were able to bioaccumulate in 

carpophores even where lower concentrations of 

these metals were found in the soil. 

97



describe and explain 79% of the total variance. In 

component one the acidity of the soil is 

disconnected from all the elements. While amounts 

of arsenic, copper, mercury, lead, zinc and, to a 

lesser extent, cadmium increase, those of nickel 

and chromium tend to decrease. In component two, 

as nickel and chromium increase, a decrease in 

98 

Fig. 68. Spatial distribution of chromium for the genus Cantharellus. 

Fig. 69. Spatial distribution of nickel for the genus Cantharellus. 

mercury and lead can be seen; this fact could be 

linked to the origin of these two elements, both 

notoriously influenced by human activities. The 

third constituent shows that basic soil corresponds 

to a reduction in cadmium concentrations, while 

other elements remain unchanged. This aspect of 

reduced bioaccumulation for cadmium has been 

shown in Ca. cibarius (Jorhem and Sundström, 

1995).

4.3.11 Genus Ramaria (Subdivision 

Basidiomycotina - Subclass Aphyllophoromycetideae 

– Order Clavariales) 

Comparison of the maps showing the spatial 

distribution of element concentrations reveals three 

Table 16. Results of principal components analysis. 

Constituents 

1 2 3 

pH_s .076 .072 .860 

As_FA .963 .040 -.037 

Cd_FA .421 .149 -.708 

Cr_FA .227 .811 .034 

Cu_FA .935 .081 -.244 

Hg_FA .499 -.652 .177 

Ni_FA -.119 .760 .020 

Pb_FA .796 -.468 .215 

Zn_FA .919 -.093 -.153 

Fig. 70. Spatial distribution of aluminium for the genus Ramaria. 

types of "associations." The first is represented by 

four elements, namely aluminium, lead, zinc and, 

to a lesser extent, copper (Figs. 70 → 72). The 

maximum concentration levels are observed in flat 

areas and those in the Apennine border. 

99

The second group includes arsenic, nickel and 

chromium (Fig. 73.) The areas with the highest 

concentrations are found in Ramiseto, Vetto and in 

Castelnovo nei Monti. The third group is composed 

of cadmium and mercury: the areas with the 

greatest concentrations were located between San 

100 

Fig. 71. Spatial distribution of zinc for the genus Ramaria. 

Fig. 72. Spatial distribution of copper for the genus Ramaria. 

Polo, Bibbiano and Montecchio. The 

concentrations of aluminium and chromium in the 

soil overlap with their distribution maps in 

mushrooms, while the remaining elements only 

display a superposition of levels in some areas.

Figs. 74, 75 and 76 illustrate the distribution of the 

concentration levels of arsenic, cadmium and lead. 

The average levels were 8.72, 5.74 and 0.98 mg/kg 

respectively. In Turkey, in Ra. Flava, arsenic 

concentrations of 0.02 mg/kg were observed 

(Konuk et al., 2007); this level is quite different to 

Cadmium: as for cadmium, very low levels (1.13 

mg/kg) were reported in the area of Epirus and 

Macedonia in samples of Ra. Largentii by Ouzoun 

Fig. 73. Spatial distribution of chromium for the genus Ramaria. 

Fig. 74. Spatial distribution of arsenic for the genus Ramaria. 

that seen in the province of Reggio Emilia. A more 

comparable level (3.7 mg/kg) was reported by 

Slejkovec et al. (1977) who analysed samples of 

mushrooms from several European countries and 

Brazil. 

et al. (2009). The value observed by Konuk et al. 

(2007) in samples of Ra. flava collected in Turkey 

(0.01 mg/kg) was 50 times lower than the 

101

minimum we recorded (0.52 mg/kg). In the region 

of Paris (France), Michelot et al. (1998) analyzed 

Lead: the average concentration levels we 

observed were significantly higher than those 

reported in previous literature [0.12 mg/kg 

Enrichment Factor values showed that cadmium, 

mercury, copper, zinc, arsenic and nickel, in 

descending order, accumulate in Ramaria 

102 

Fig. 75. Spatial distribution of cadmium for the genus Ramaria. 

Fig. 76. Spatial distribution of lead for the genus Ramaria. 

92 species of fungi, including Ramaria sp., and 

found levels of 4.32 mg/kg. 

(Ouzouni et al., 2009) and 0.018 mg/kg (Konuk et 

al., 2007)]. 

carpophores in the presence of small concentrations 

of these metals in the soil, while lead and, to a 

lesser degree, chromium, do not tend to accumulate

in mushrooms even if the soil is rich in these 

elements. 




The first component explains how the acidity of the 

soil is not related to bioaccumulation of the 

elements we tested: all elements except lead tend to 

4.3.12 Genus Morchella (Subdivision 

Ascomycotina - Subclass Pezizomycetideae 

– Order Pezizales) 

One type of distribution is able to represent the 

concentration levels of aluminium, mercury, nickel, 

zinc and to a lesser extent chromium and lead. The 

remaining elements arsenic, cadmium and copper 

show different types of "associations" for each of 

the six metals listed above. For aluminium and 

other, more “associated”, elements, the areas with 

the greatest concentrations were Casalgrande and 

San Polo, and excluding chromium and lead, also 

the hilly areas. Arsenic values reach their 

maximum near Reggio Emilia, cadmium in the 

areas of Toano, Villa Minozzo and Quattro 

Castella, while copper is found in high 

concentrations near Castellarano and Castelnovo 

Monti. 

Considering a quantitative overlap with soil 

concentrations, lead showed the greatest affinity, 

Tabella17. results of the principal component analysis. 

1 


2 3 

pH_s .129 .011 .985 

As_FA .909 .187 .090 

Cd_FA .960 .040 .060 

Cr_FA .757 .549 .216 

Cu_FA .927 .047 .061 

Hg_FA .819 .380 .308 

Ni_FA .952 .280 .073 

Pb_FA .063 .956 -.020 

Zn_FA .950 -.079 .141 

increase their concentration in the fruiting bodies of 

fungi. 

The second component confirms no relation to 

other elements or pH levels for the 

bioaccumulation of lead and chromium. The third 

confirms the independence of the acidity of the soil 

from all elements considered. 

but to a lesser extent also chromium and copper. 

Cadmium: the distribution of cadmium 

concentrations is shown in Fig. 77, the average 

level is 0.94 mg/kg and covers much of the 

territory. The highest levels (4.12 mg/kg) were 

observed between Toano and Villa Minozzo and in 

the area of Canossa. In the same area Cocchi et al. 

(2006) observed 12.55 mg/kg in Mo. esculenta. In 

Turkey (Tüzen, 2003) Mo. esculenta contained 1.43 

mg/kg. Also in Turkey, in Mo. esculenta, Mo. 

esculenta var. umbrina, Mo. vulgaris, Mo. costata, 

Mo. deliciosa, and Mo. rigida cadmium 

concentrations were 0.031, 0.002, 0.036, 0.024, 

0.029, 0.007 mg/kg respectively (Konuk et al., 

2007). These levels are 20-200 times lower than 

those reported in this paper and given in literature. 

In France, in samples of Mo. esculenta, a 

concentration of 3.6 mg/kg was observed (Michelot 

et al., 1998). 

103

Nickel: Fig. 78 represents the concentration of 

nickel in the Reggio Emilia area. The maximum 

level of 12.2 mg/kg was observed in the lowlands 

near Scandiano. The average level of 2.4 mg/kg is 

commonly found all over the area. In France a 

concentration of 15.4 mg/kg, higher than the 

maximum level we measured, was found in a 

sample of Mo. esculenta (Michelot et al., 1998). 

Tüzen (2003) in Turkey recorded 1.18 mg/kg in 

104 

Fig. 77. Spatial distribution of cadmium for the genus Morchella. 

Fig. 78. Spatial distribution of nickel for the genus Morchella. 

Mo. esculenta; a value in line with the data 

presented here. In Turkey Mo. esculenta, Mo. 

esculenta var. umbrina, Mo. vulgaris, Mo costata, 

Mo. deliciosa, and Mo. rigida were found to 

contain nickel concentrations of 0.07; 0.68; 0.04; 

0.4; 0.23; 0.41 mg/kg, respecitvely (Konuk et al., 

2007); these levels are at least an order of 

magnitude lower than those we recorded.

For completeness, we provide here several data for 

two species of the genus Helvella (order Pezizales): 

Cadmium: 1.97 mg/kg in Helvella crispa (Scop.) 

Fr. (Cocchi et al., 2006) and 0.033 mg/kg in 

Helvella leucopus Pers. (Konuk et al., 2007); 

Nickel: 0.3 mg/kg in Helvella leucopus Pers. 

(Konuk et al., 2007) 

For the genus Morchella as it was for other taxa, 

arsenic, chromium, nickel and lead do not tend to 

In the first component soil acidity and copper are 

linked but at the same time independent of other 

elements, instead, in the second copper and zinc 

move in the same direction and are differentiated 

from both the pH and from almost all other 

elements. The third reinforces the neutrality 

towards soil acidity for all elements, except, in part, 

for mercury that tends to increase with increasing 

pH, while the lead does the opposite and tends to 

decrease as soil pH rises. 

4.3.13 Conclusions 

The analytical results obtained by analysing 

thousands of different fungi belonging to hundreds 

of species have yielded distribution maps of heavy 

metals in an area with diverse geomorphological 

features ranging from lowland areas to the 

Apennine peaks. The land use includes residential 

zones such as the city of Reggio Emilia and other 

major urban centres, busy main roads, industrial 

areas, agricultural areas and other intensively 

farmed pasture, and woods and forests. There is a 

rich diversity of flora, fauna and countryside where 

many daily activities are carried out, leaving 

"footprints" that combine to change the very nature 

of the environment. 

Table 18. Analysis of main constituents. 

1 

Constituents 

2 3 

pH_s -.043 .120 .964 

As_FA .934 .307 -.029 

Cd_FA .976 .142 -.014 

Cr_FA .966 -.154 .032 

Cu_FA .140 .954 .174 

Hg_FA .816 .161 .329 

Ni_FA .957 .140 -.131 

Pb_FA .802 .377 -.206 

Zn_FA .797 .502 -.091 

accumulate in the fruiting bodies. In contrast, and 

in descending order, the elements zinc, cadmium, 

mercury and copper (Falandysz et al., 2002) tend to 

accumulate in the fruiting bodies (though not in a 

marked manner) even if the soil has low 

concentrations. 




The results obtained from analysing mushrooms 

cannot tell us how we should behave as regards the 

chemical elements contained inside them, in 

particular heavy metals. The results of the spatial 

concentration distributions are quite different from 

each other and "dominated" by species-specific 

factors for each fungus such as the length, depth 

and age of the mycelium, the type of element under 

investigation, its chemical form and its availability 

in the soil substrate, the type of soil-substrate, the 

organic matter in the soil substrate, vegetation, the 

degree of humidity, and many other unknown 

factors, some of which are still unknown. 

One important aspect that should perhaps be 

investigated further is the ability of fungi to 

accumulate heavy metals, even if the soil has no 

high concentrations of these. In this study we have 

seen that some elements, such as cadmium, copper, 

mercury and zinc tend to accumulate in different 

fungal species regardless of the nature and 

concentrations of elements in the soil-substrate. 

Such has been confirmed for mercury (Falandysz et 

al., 2002) and for mercury, and cadmium (Vetter, 

1994, Kalac and Svobova, 2000). It has also been 

observed that, in general, arsenic, chromium, nickel 

and lead do not tend to accumulate in fungi, even if 

the soil is rich in these elements, a fact confirmed 

105

for lead (Kalac and Svobova, 2000, García et al., 

2009). 

Giving "guidelines" at present is rather difficult and 

premature. In the light of current knowledge, the 

use of mushrooms as bioindicators of soil quality 

and the environment, is still in the embryonic 

stages and not yet viable, even if some fungal 

species could be used as “warning mushrooms”. 

Future investigations in the same area could expand 

the information available for one or more species of 

mushroom, allowing some to be defined and used 

as environmental bioindicators. 

Based on the wealth of data and considerations 

presented here, we think it is feasible to claim that, 

given the concentration levels of chemical elements 

in mushrooms in conjunction with the 

concentrations of these metals in soils, this study 

could facilitate the identification of "threshold 

limits" for these heavy metal concentrations, 

bearing in mind their impact on human health, 

which would apply to the sale of certain species of 

edible mushrooms. 

The names of the fungal species considered in this 

chapter have been written in an abbreviated form to 

make the text more readable,. The species are now 

listed here alphabetically, alongside with their full 

names, in accordance with the taxonomy specified 

in paragraph 2.1.4. 

• Ag. altipes = Agaricus altipes (F. H. Møller) 

F. H. Møller 

• Ag. arvensis = Agaricus arvensis Schaeff. 

• Ag. bernardii = Agaricus bernardii Quél. 

• Ag. bisporus = Agaricus bisporus (J.E. Lange) 

Imbach 

• Ag. bitorquis = Agaricus bitorquis (Quél.) 

Sacc. 

• Ag. campestris = Agaricus campestris L. 

• Ag. cupreobrunneus = Agaricus cupreobruneus 

(Jul. Schäff. & Steer) Pilát. 

• Ag. macrocarpus = Agaricus macrocarpus 

(F.H. Møller) F.H. Møller 

• Ag. silvaticus = Agaricus silvaticus Schaeff. 

• Ag. silvicola = Agaricus silvicola (Vittad.) 

Peck. 

• Ag. urinascens = Agaricus urinascens (Jul. 

Schäff. & F.H. Møller) Singer 

• Am. caesarea = Amanita caesarea (Scop.) 

Pers. 

106 

• Am. excelsa var. excelsa = Amanita excelsa 

var. excelsa (Fr.) P. Kumm. 

• Am. excelsa var. spissa = Amanita excelsa var. 

spissa (Fr.) Neville & Poumarat 

• Am. gemmata = Amanita gemmata (Fr.) 

Bertill. 

• Am. muscaria = Amanita muscaria (L.) Lam. 

• Am. ovoidea = Amanita ovoidea (Bull.) Link 

• Am. pantherina = Amanita pantherina (DC.) 

Krombh. 

• Am. phalloides = Amanita phalloides (Vaill. 

ex Fr.) Link 

• Am. rubescens = Amanita rubescens var. 

rubescens Pers. 

• Am. solitaria = Amanita solitaria (Bull.) Fr. 

• Am. vaginata = Amanita vaginata (Bull.) 

Lam. 

• Ar. mellea = Armillaria mellea (Vahl) P. 

Kumm. 

• Ar. tabescens = Armillaria tabescens (Scop.) 

Emel 

• B. aereus = Boletus aereus Bull. 

• B. reticulatus = Boletus reticulatus Schaeff. 

• B. edulis = Boletus edulis Bull. 

• B. pinophilus = Boletus pinophilus Pilát & 

Dermek 

• Ca. cibarius = Cantharellus cibarius Fr. 

• Cl. gibba = Clitocybe gibba (Pers.) P. Kumm. 

• Cl. nebularis = Clitocybe nebularis (Batsch) 

P. Kumm. 

• He. crispa = Helvella crispa (Scop.) Fr. 

• He. leucopus = Helvella leucopus Pers. 

• Hy. chrysodon = Hygrophorus chrysodon 

(Batsch) Fr. 

• Hy. eburneus = Hygrophorus eburneus (Bull.) 

Fr. 

• Hy. penarius = Hygrophorus penarius Fr. 

• Hy. russula = Hygrophorus russula (Schaeff.) 

Kauffman 

• geotropa = Infundibulicybe geotropa (Bull.) 

Harmaja 

• La. amethystina = Laccaria amethystina 

Cooke 

• La. fraterna = Laccaria fraterna (Sacc.) 

Pegler 

• La. laccata = Laccaria laccata (Scop.) Cooke 

• Le. nuda = Lepista nuda (Bull.) Cooke 

• Ly. decastes = Lyophyllum decastes (Fr.) 

Singer 

• Ma. oreades = Marasmius oreades (Bolton) 

Fr.

• Mo. costata = Morchella costata (Vent.) Pers. 

• Mo. deliciosa = Morchella deliciosa Fr. 

• Mo. esculenta = Morchella esculenta (L.) 

Pers. 

• Mo. esculenta var. umbrina = Morchella 

esculenta var. umbrina (Boud.) S. Imai 

• Mo. rigida = Morchella rigida (Krombh.) 

Boud. 

• Mo. vulgaris = Morchella vulgaris (Pers.) 

Boud. 

• Ru. cyanoxantha = Russula cyanoxantha 

(Schaeff.) Fr. 

• Ru. delica = Russula delica Fr. 

• Ru. emetica = Russula emetica (Schaeff.) 

Pers. 

4.4 Sampling: a data sheet 

example 

Each sampling site, where both soil and fungal 

samples were collected, is described in detail and 

• Ra. flava = Ramaria flava (Schaeff.) 

• Ra. largentii = Ramaria largentii Marr & D.E. 

Stuntz 

• T. argyraceum = Tricholoma argyraceum 

(Bull.) Gillet 

• T. equestre = Tricholoma equestre (L.) P. 

Kumm. 

• T. terreum = Tricholoma terreum var terreum 

(Schaeff.) P. Kumm. 

• T. rutilans = Tricholomopsis rutilans 

(Schaeff.) Singer 

• T. ustaloides = Tricholoma ustaloides 

Romagn. 

correctly georeferenced, so that the analytical data 

of the samples can then be used to create 

geostatistical maps. 

Here follows an example of one sample data sheet 

used in this study to describe the sample sites. 

107

Area and sample description 

108 

Sheet 1 

Map 1 Map 2 

Photo nr.1 Photo nr.2 

ID Toponym - Location: Fonte dell’Anatella 

Municipality: Rocca di Mezzo 

Geographic area: Abruzzo, Provincia dell’Aquila, Parco del Sirente, Velino 

Sampling date: 7 Settembre 2005 

Coordinates: Geographic coordinate system: UTM/UPS Map Datum: WGS 84 

33T Long. 0379918 Lat. 4671322 

Altitude and inclination: 1400 m; 5% 

Area description: Specimen found in the city of Fonte Anatella, on the basal portion of a beech 

(Fagus sylvatica L.) in the municipality of Rocca di Mezzo, in a beech forest on 

limestone matrix on the slopes of Mount Sirente. Wooded slopes, dense 

vegetation, steep terrain

Habitat: natural woodlands 

Substrate: Woody matrix 

Phylum: Basidiomycota 

Class: Basidiomycetes 

Order: Polyporales 

Family: Meripilaceae 

Fungus 

Specimen name: basidiocarp of Meripilus giganteus (Pers. : Fr.) P. Karsten 

leg. Fabio Siniscalco, det. Carmine Siniscalco 

Height: 94.5 cm (highest point of basidiocarp complex) 

Width: 139.5 cm (widest point of basidiocarp complex) 

Weight: 142.68 kg (total basidiocarp weight) 

Note: basidioma growing at the base of a 

beech tree between the collar and large 

emerging roots. 

element 

concentration 

mg/kg 

Analytical results 



mg/kg 

Meripilus giganteus (Pers.) P. Karst., 



mg/kg 

Ag 1.11 Ge 0.001 Se 0.1 

Al 115 Hg 0.05 Sr 0.62 

As 0.2 K 27900 Ti 9.37 

Ba 1.18 La n.d. V 0.11 

Be 0.01 Li 0.13 Y n.d. 

B 10.4 Mg 1910 Zn 50.8 

Cd 2.25 Mn 4.67 Zr 0.18 

Ca 185 Mo 0.11 Cl n.d. 

Cs 0.06 Na 80.0 P 6270 

Cr 3.34 Ni 2.02 S 1310 

Co n.d. Pb 9.13 

Cu 107 Rb 23.4 

Fe 216 Sc n.d. 

134 Cs n.d. 

137 Cs n.d. 

40 K n.d. 

109

110 

Location of sample area 

Starting from the macroarea, 

the three figures on 

the left describe the exact 

spot where the discovery 

and the collection of the 

fungus took place.

Bioindication allows evaluating the effects of 

anthropic activity on the environment through the 

observation of living organisms. The widespread 

use of macromycetes for bioindication is often 

limited by taxonomic difficulties, by incomplete 

knowledge of fungal metabolism and physiology, 

and by the lack of data on the quality and 

characteristics of the substrate. 

Further obstacles to the correct interpretation of 

environmental data through fungi are the lack of 

precision and accuracy in describing their habitat. 

This EUR report aims to help develop a working 

methodology and to show applicable concrete 

examples of the paths to take in the future. 

The first aspect which was studied herein was the 

measure of chemical substances contained in the 

carpophores of spontaneous macromycetes. 

The second aspect considered was the “reference 

mushroom”, a tool already in use for other 

organisms, and which could prove to be a key 

element in unravelling the issues surrounding 

chemical elements in macromycete carpophores. 

We arrived at this by attempting to overcome the 

initial difficulties of interpreting, with no available 

parameters, the significance of the presence of 

different chemical elements (heavy metals in 

particular) in fungi. The presence of these elements 

was often surprising due to unexpected 

concentration levels and large differences between 

very closely-related taxonomic species. 

The third aspect dealt with in this work was linking 

fungal species with their habitats. This allowed us 

to observe important biodiversities which might be 

further investigated and studied in future work, 

with the aid of new technologies and with the help 

of environmental coding carried out by the 

CORINE programme and the European system of 

natural information, EUNIS. Following this it will 

Chapter V 

Conclusions 

be possible to be guided by objective criteria when 

describing habitats with specific macromycetes 

growth. 

We hope that our work may sparkle new ideas for 

study and research that could help support the use 

of these organisms in environmental assessments. 

Such a feature will be important when considering 

the mycological components of terrestrial 

ecosystems, which is becoming an increasingly 

important and relevant factor in the evaluation of 

global ecological balances. 

This work aims not to close but to open a new 

vision with new opportunities for scientific 

research on the world of higher mushrooms, which 

is little understood and too often undervalued. By 

making available to everyone this volume of data, 

that, in itself, constitutes an important contribution 

to the documentation of fungal biodiversity, we 

have above all aimed to describe a particular 

working methodology and some practical 

applications. We are convinced that it can be used 

to produce new research in this field. 

For example, we are convinced that taxonomic 

research should be based on a polyphasic approach 

that includes the collection and analysis of macro 

and microscopic data, including morphological, 

physiological, and biochemical features; further, 

the most accurate possible definition of each 

macromycete’s habitat; we believe that in this 

context the measurement of concentrations of 

different chemicals in and molecular characteristics 

of that habitat should be analysed. 

Last but not least we hope that our work will prove 

useful to all those who have different levels of 

political and administrative responsibility of the 

territories, especially regarding the use of 

mushrooms as food. 

. 

111

112

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from phylogenetic relationships to specific 

primers”. Pagine di Micologia, (30), pp. 49- 

52. 

Whittaker R. H. (1969): “New Concepts of Kingdoms of 

Organisms”. Science 163, pp. 150-160. 

Woese C. R., Kandler O., Wheelis M. L. (1990): 

“Towards a natural system of organisms: 

Proposal for the domains Archaea, Bacteria, 

and Eucarya”. Proceedings of the National 

Academy of Science USA, 87, pp. 4576-4579. 

Wright S. F., Upadhyaya A. (1996): “Extraction of an 

abundant and unusual protein from soil and 

comparison with hyphal protein from 

arbuscular mycorrhizal fungi”. Soil Science, 

n. 161, pp. 575-586. 

Yamaç M., Yıldız D., Sarıkürkcü C., Çelikkollu M., 

Solak M. H. (2007): “Heavy metals in some 

edible mushrooms from the Central Anatolia, 

Turkey”. Food Chemistry, Volume 103, Issue 

2, pp. 263-267. 

Zanella A., Tomasi M., De Siena C., Frizzera L., 

Jabiol B., Nicolini G., Sartori G., Calabrese 

M. S., Mancabelli A., Nardi S., Pizzeghello 

D., Odasso M. (2001): “Humus Forestali”. 

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Zimmermannová K., Kalač P. (2000): “Concentrations 

of mercury, cadmium, lead and copper in 

fruiting bodies of edible mushrooms in an 

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mercury smelter”. The Science of The Total 

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119

120

For a better understanding of the book, we have 

decided to include descriptions and pictures of 

some fungal species studied, as well as some of the 

most interesting and commonly known fungal 

species. Pictures and descriptions are taken from 

Volumes I (1999), II (2001) and III (2009) of the 

series “Atlante Photografico dei Funghi d’Italia” 

("Photographic Atlas of Italian Mushrooms"), 

published by the Fondazione Centro Studi 

Micologici of the Associazione Micologica 

Bresadola, edited by Giovanni Consiglio, Carlo 

Papetti and, only for Vol. 1, Giampaolo Simonini. 

In some casees the names of the species used by 

these authors are different from those used here, for 

the reasons described in paragraph 2.1.4(page 20). 

As already explained, the nomenclature follows 

www.indexfungorum.org; synonyms are given in 

square brackets. 

VII Appendix 

The descriptions also provide information on the 

edibility of each species (please note: “edible” is 

appropriate only if the mushroom is thoroughly 

cooked!).. 

The number of images presented here is limited, 

compared to the number of fungal species 

occurring in nature; we believe, however, that it 

represents appropriately the considerable potential 

for biodiversity possible in the fungal kingdom. 

Species are listed in alphabetical order for purely 

practical reasons and not according to rigorous 

systematic criteria. 

The photographers are credited in “Atlante 

Photografico dei Funghi d’Italia”, with the sole 

exception of the image showing Agaricus 

urinascens, which was taken by Ennio Carassai. 

121

122

CAP 80-140 mm, hemispheric, then convex-flat, hairless, 

white, bordering on a silky shine, at maturity yellowish on 

circumference, finally, from yellow-citrina to vivid ochre 

all over; edges hanging with residues of veil. 

GILLS at the extremes from whitish to pale grey-pink, 

then pink-greying and only with age from brown-purple 

to brown-black, crowded, free, width of 6-8 mm, with 

pale gill edge. 

STIPE 80-120 × 12-25 mm, cylindrical, widening towards 

base but not bulbous, white, to the touch yellow-citrina, 

annulus white, persistent, high, broad, flaring, yellow 

when handled, in two layers, lower layer has coarse 

white-ochre-ish scales , with appearance of a gear wheel. 

FLESH white, with age light hint of ochre, thick in the 

centre, strong smell of aniseed, flavour of hazelnut. 

MICROSCOPY: ellipsoidal spores, smooth, under 

microscope dark brown, dimensions 6.5-8 × 4.2-5.4 µm; 

tetrasporophyte basidia; marginal cells from clavate to 

vescicle. Brown-black spores. 

Agaricus arvensis Schaeff. : Fr. 

HABITAT: often gregarious, rarely isolated, from the 

spring to autumn, in meadows, in pastures, around the 

edges of woods. Infrequent. 

EDIBILITY: edible 

NOTE – Good to eat. The species of the subsection 

Flavescentes are however, natural concentrators of silver, 

cadmium and mercury. The Flavescentes produce 

carpophores with an anise or hazelnut odour and are 

weakly yellow at the extremities. The caps tend to be 

quite large, the gills at the extremes are a very pale fleshy 

grey, there will usually be evident yellowing, the base of 

the stipe is large but not bulbous, all factors which 

facilitate the identification of this lovely species. It could 

be confused with Agaricus nivescens, which, however 

differs by having a more modest yellowing, and smaller 

and rounder spores. Another similar species is Agaricus 

xanthodermus, which has a slight odour of phenol, 

marginally bulbous stipe and on scratching shows a 

golden yellow colour which is particularly evident at the 

base of the stipe and to a lesser degree in the flesh. 

123

CAP 60-160 mm, fleshy, initially globular, then 

hemispheric or hemispheric-truncated, then flat-convex 

with slight central depression; edges convoluted even in 

mature examples, striated on lower parts; cap covering 

white, white-greying, ochre-ish or even brownish, in the 

wide central part, often dissociated in large irregular 

scales on a light background. 

GILLS free, crowded, wide, initially whitish, then 

progressively dirty pink, brown, dark brown, blackishbrown; 

sterile surface, light, minutely denticulate. 

STIPE 40-80 × 20-35 mm, cylindrical or ventricular with 

pointed base, robust, has a small annulus, simple and fine 

towards the middle; above the annulus white and smooth, 

under the annulus whitish or grey-ochre-ish, smooth or 

with transversal bands concolour with the cap. 

FLESH firm white, reddish to gill edge. Odour normally 

strong and unpleasant of fish or of fish-eating birds. Cross 

reaction with Schäffer’s reagent, negative. 

124 

Agaricus bernardii Quél. in Cooke & Quél. 

MICROSCOPY: spore mostly ellipsoidal to subglobose, 

mono- or biguttulate, more rarely multiguttulate, 6-8 × 

4.5-6.5 µm. basidia clavate, tetrasporic. Cheilocystidia 

very numerous, multiform, usually cylindrical, fusiform 

or clavate. Spores blackish-brown. 

HABITAT: gregarious, in small groupings or in “witches’ 

circles”, both in dune or in coastal areas, also in urban 

areas (for example under Cedrus) or in mountainous 

zones, but never inside woods; from the end of spring to 

autumn. 

EDIBILITY: of no value 

NOTE - Is often confused, due to its similar appearance, 

with A. litoralis (Wakef. & A. Pearson) Pilát, which, 

however has a flaring annulus, or with A. bitorquis 

(Quél.) Sacc., which, however has two lower annuli.

CAP 50-130 mm, fleshy, initially hemispheric, then flatconvex, 

finally flat; edges curving at the extremes; cap 

covering variable from white to ochre, up until brownish, 

outside the disc a little dissociated in triangulated 

fibrillated scales, adpressed, on a lighter background. 

GILLS free, fairly crowded, wide, initially whitish, but 

soon pink then brown-reddish, finally brown-blackish, 

with light surface. 

STIPE 40-90 × 10-25 mm, cylindrical or progressively 

widened towards base, which is often a little bulbous, 

straight, tightly fistular, white, under the annulus finely 

fibrillated often with white mycelial cords. Annulus thick, 

normally triangular at division; due to its complex 

structure, to be considered intermediate (neither growing 

upwards nor downwards), even though it can be pendant 

(drooping or skirt-like) and more rarely, sheathing 

(opening upwards around the stipe). 

Agaricus bisporus (J.E. Lange) Imbach 

FLESH firm white, weakly pink-reddish to gill edge. 

Odour weak, fungal. Cross reaction with Schäffer’s 

reagent, negative. 

MICROSCOPY: spores mostly ellipsoidal, generally 

multiguttulate, 6.5-7.5 × 5-6 µm. basidia bisporic, but also 

monosporic (often very numerous) and more rarely 

trisporic or tetrasporic (these last most common in var. 

eurotetrasporus). Very numerous Cheilocystidia, clavate 

or a little fusiform, spores brown-blackish . 

HABITAT: gregarious, in small groupings with isolated 

or clustered examples; common to fertilised ground or 

close to livestock enclosures, rare in “wild” areas, 

generally under Cupressaceae. Spring and late autumn. 


NOTE – This is the only Agaricus with a great deal of 

bisporic basidia. It is good to eat and particularly suited to 

cultivation. It is produced industrially and sold almost all 

around the world. 

125

CAP 50-120 mm, first hemispheric, then convex-flat, 

fleshy, firm, from pure white to dirty whitish, sometimes 

pale ochre-ish, smooth or with some under developed 

fibrils often covered by earth; edges convoluted and only 

with age distended. 

GILLS from free to lightly adnate, close (4-6 mm), 

crowded, initially pale pink, soon becoming full and dirty 

flesh pink, and with age purple-black; gill edge whitish. 

STIPE 40-80 × 20-40 mm, cylindrical, attenuated at base, 

full, rigid, from whitish to dirty pink, at tip smooth or 

white and flocculent; bearing two membranous, white 

annuli, situated in the medial and basal zones of the stipe. 

FLESH thick, almost hard, gill edge, pale hazelnut, then 

taking tones of pink, red wine; sweet flavour, odour 

pronounced and pleasant. 

MICROSCOPY: spore mostly ellipsoidal, almost 

rounded, smooth, under microscope brown to yellowish 

126 

Agaricus bitorquis (Quélet) Saccardo 

shades; dimensions 4.5-6 × 4-5.5 µm; tetrasporic basidia; 

clavate cells. Brown-purple spores. 

HABITAT: often gregarious on bare, sandy or compact 

soil at roadsides, in parks, riverside areas and streams, 

even under asphalt, from late spring to late autumn. 


NOTE - Good to eat. The species is recognisable among 

other taxa from the section Bitorques by the clear set of 

double annuli, pleasant odour and the short stipe when 

compared to the diameter of the cap. The validus variety 

grows in clusters, has more compact and more heavily 

reddened flesh. It can be confused with the more fragile 

A. campestris, which has an ephemeral annulus and more 

brightly-coloured gills, and A. bisporus, which has a 

simple annulus and bisporic basidia.

CAP 50-120 mm, initially globular, soon convex-flat, 

fleshy, white to whitish, sometimes suffused with pink, 

silky-fibrillated, under developed fibrils more or less 

concentric, scales turn brown with age; at the extremes 

convoluted, rarely distended often bearing remnants of 

veil. 

GILLS free, close, with lamellule, pink while still young 

concolour with gill edge, then purple-black in maturity. 

STIPE 50-80 × 10-18 mm, more or less cylindrical, but 

often attenuated at base, full, rigid white, bearing fine 

white dirty fibrils, browning with age; annulus tight and 

fragile, flaring, of cottony consistency, white, just 

reddening especially in the stipe-cap join, thick centrally, 

with pleasant odour and sweet flavour. 

MICROSCOPY: oval spores, under microscope pale 

grey-brown, with just visible germinating pores, 

dimensions 6.5-8 × 4.5-5.5 µm; tetrasporophyte basidia. 

HABITAT: in fertilised meadows, in wheel tracks, in city 

parks, in small groupings or in circles of many 

individuals; from the start of summer to late autumn. 

Agaricus campestris L. : Fr. 


NOTE - Good to eat. It is one of most renown and 

gathered edible Agaricus species, and is well known 

everywhere. It is easily recognisable by its shape, its 

odour, which is neither anise nor of almonds, the flesh, 

which when bruised assumes a shade of red, the fugacious 

annulus and the stipe, which is often pointed at its base. It 

belongs to the genus Agaricus characterised by their 

fragile carpophores, flaring annuli and gills which tend to 

red at the extremes. There are many forms and varieties 

about which the Authors do not agree; the var. 

squamulosus, in particular, which displays brownish 

concentric scales on its cap. Confusable with the Agaricus 

species in the section Xanthodermatei, which are 

poisonous, but which have yellowing flesh and an inky 

smell. 

127

128 

Agaricus cupreobrunneus (Jul. Schäff. & Steer ex F.H. Möller) Pilát 

CAP 30-90 mm, initially globular, then hemispheric with 

truncated edges and with flattened centre, finally flatconvex 

with slight central depression; cap covering 

brownish copper, or even of varying tones of brown, in 

adulthood whitish silky-fibrillated edges; often the cap 

can be completely white with brown scales towards the 

centre. 

GILLS free, fairly crowded, wide, with lamellule, from 

light pink to vivid pink up until brown-reddish, at 

maturity brown-blackish; edges concolour with surface. 

STIPE 20-70 × 8-20 mm, clavate-ventricular often with 

pointed base, but also cylindrical, from full to tightly 

fistular, white, under the annulus smooth or finely 

fibrillated but bearing several more or less complete labra 

of the same colour as the cap. 

FLESH white, uniform to gill edge, or just pink in the top 

part of the stipe and close to the gills. Light odour, fungal. 

Cross reaction with Schäffer’s reagent, negative. 

MICROSCOPY: ellipsoidal spores, generally mono- and 

biguttulate or with granular content, 7-9 × 4.5-6 µm. 

basidia clavate, tetrasporic. Cheilocystidia isolated, 

almost absent. Spores brown-dark purple. 

HABITAT: gregarious in small groupings or in “witches’ 

circles” in large grassy areas, both in the mountains and 

near coastal and urban areas; autumn. 


NOTE - Fairly rare but quite widespread, in its place of 

growth it is prized as a delicacy, considered superior to A. 

campestris L. : Fr. for its firmer flesh. A similar species is 

A. porphyrocephalus F.H. Møller, and it is often difficult 

to distinguish them macroscopically, but the latter has 

decidedly smaller spores. Its shape, with a white cap and 

brown scales in the centre, is similar to A. campestris var. 

squamulosus (Rea) Pilát, which is today considered 

synonymous with A. cupreobrunneus.

CAP 80-150 (180) mm, hemispheric, then for a long 

while convex, finally distended, dry and silky, fairly 

regular lip, with hanging-floccules from remnants of veil; 

white, then irregularly suffused with yellowish tones, 

more ochre-ish and intense around circumference. 

GILLS highly crowded, free from stipe, intercalated with 

lamellule; initially whitish, then grey-pink, pink, and 

finally brown- dark purple-ish, keeping a (sterile) whitish 

surface. 

STIPE 80-130 (150) × 20-30 mm, robust, cylindrical, 

sometimes a little attenuated at tip, with a very squashed 

basal bulb, typically in the shape of elephant feet; white, 

stained yellow-ochre-ish at base, in some adult 

specimens, more or less flocular fibrils under the annulus. 

ANNULUS broad, thick, fairly persistent, flaring, 

coarsely decorated with the shape of a gear wheel on the 

lower surface; white. 

Agaricus macrocarpus (F.H. Møller) F.H. Møller 

FLESH white, nearly uniform in cap, tendency to turn 

yellow on stipe, compact, tender; weak odour of bitter 

almonds, mild, pleasant flavour. 

MICROSCOPY: ellipsoidal spores, smooth, 7.2-8.6 × 

4.6-5.2 µm. Cheilocystidia clavate-obese, often with 

apical appendage. 

HABITAT: grows in meadows and in pastures or in 

sparse, grassy conifer woods; first appears in late spring, 

and then fruits in autumn; infrequent. 


NOTE - A large mushroom which morphologically 

evokes the better-known and more common A. arvensis 

Schaeff. : Fr. from which, however, it differs mainly in its 

larger size, stipe base with a squashed bulb, its coarse 

irregular cog-like pattern on the surface just under the 

annulus and for its slightly smaller spores. 

129

CAP 50-100 mm, initially hemispheric or campanulate, 

then convex or a little flattened, sometimes umbonate, 

with the edges convoluted, then straight, the cap covering 

brown-ochre-ish, more or less reddish, dissociated outside 

disc in fibrillated scales on lighter background. 

GILLS free, crowded, close, greying-brownish, hint of 

pink, eventually brown dark, with the sterile surface, 

more or less pale. 

STIPE 60-150 × 7-15 mm, cylindrical, sometimes a little 

bent, with bulbous base, firm then fistular, initially white 

then greying, bearing fine flakes or scales under the 

annulus. Annulus flaring, simple, fine, the upper part 

whitish, the lower concoloured with the cap, sometimes 

evanescent. 

FLESH white, to gill edge turning to fairly intense blood 

red, with a sour odour and sweetish flavour. Cross 

reaction with Schäffer’s reagent, negative. 

130 

Agaricus silvaticus Schaeff. : Fr. 

MICROSCOPY: spore from oval to ovoid-ellipsoidal, 

monoguttulate, 5.4-6 × 3.6-4 µm. clavate tetrasporic 

basidia. Cheilocystidia very numerous, from clavate to 

mostly clavate, with brownish content. Brown-purple 

spores. 

HABITAT: isolated or gregarious, in hardwood and 

coniferous areas; summer-autumn. 


NOTE - A. silvaticus belongs to a group of species 

characterised by their slightly reddening flesh towards 

their gill edges and by their predominantly woodland 

habitat. It is distinguishable from other, similar species by 

the colour, shape and ornamentation of its cap, its flesh, 

which strongly reddens towards the gill edge and by its 

pleasant odour. According to several authors, A. silvaticus 

is a very variable, collective species,, and includes, among 

others, A. haemorrhoidarius and A. langei.

CAP 50-90 mm, ovoid or hemispheric, then convex-flat, 

with the edges convoluted, then straight and wavy, white 

cap covering, whitish cream, hairless, fibrillated, yellow 

on rubbing. 

GILLS free, fairly crowded, pale at the extremes, with a 

slight reddish shade, finally brown dark, with pale sterile 

surface. 

STIPE 60-90 × 10-15 mm, cylindrical, widened at base, 

fistular, white, silky under the annulus, often lightly pink 

then blackened above the annulus, yellowed by rubbing. 

Annulus flaring, simple, broad, white, upper part smooth, 

lower part with a flaky yellow bloom towards the edges . 

FLESH white, pink or reddish when exposed to air, with 

odour of anise and sour flavour. Cross reaction with 

Schäffer’s reagent, positive. 

MICROSCOPY: spore oval with one or two guttules, 

(5.4) 5.8-6.6 × 3.8-4.2 (4.4) µm. clavate basidia, 

tetrasporic. Cheilocystidia subglobular, obpiriform, 

Agaricus silvicola (Vittadini) Saccardo 

ellipsoidal, even in cheilocatenule, with brownish content. 

Spores purplish-brown in mass. 

HABITAT: gregarious, in large groups, in conifer woods. 

Summer to autumn, fairly frequent and common. 


NOTE - A. sylvicola belongs to a group of species which 

also includes A. essettei, A. tenuivolvatus and A. 

macrocarpus, where one can often find "family 

resemblances" between one species and another. The 

differences between them lie in characteristics such as 

size, degree of robustness, development of a universal veil 

and spore size, but in the absence of reliable studies on 

phenotypic variability such distinctions are only relatively 

valid. For example, A. syilvicola, of medium size and 

typically slender, has a stipe with a bulbous and welldefined 

base as, indeed have A. macrocarpus and A. 

essettei, but it has the smallest spore size of the group. 

131

132 

Agaricus urinascens var. urinascens 

(Jul. Schäff. & F.H. Möller) Singer 

[= A. macrosporus (Jul. Schäff. & F.H. Möller) Pilát; A. alberti Bon; 

A. stramineus (Jul. Schäff. & F.H. Möller) Singer] 

CAP 100-200 mm (sometimes up to 350-400 mm), 

fleshy, hemispheric or campanulate, then convex often 

with flattened centre, finally flat; curving edge at the 

extremes; white cap covering, alutaceous or ochre-ish, 

usually dissociated outside disc, adpressed or even clearly 

cracked scales. 

GILLS free, crowded, close, light at the extremes, then 

flesh coloured, finally brown-blackish with light surface. 

STIPE 50-100 × 25-35 mm, usually short and thick, 

frequently with pointed rooting base, but often elongated, 

tight, medullar or even a little hollow, white or whitish, 

completely flocculent-scaly under the annulus; annulus 

flaring, broad, smooth above, irregularly serrated to edges 

on the lower face and flocculent-scaly elsewhere, white. 

FLESH firm white, flesh coloured on stem to gill edge. 

Odour of bitter almonds when fresh, then of mouldy straw 

or urine. 

MICROSCOPY: spores ovoid-ellipsoidal, multiguttulate, 

8.5-12 × 5.5-6.5 µm. Tetrasporophyte basidia, clavate. 

Cheilocystidia very numerous, mostly clavate. Spores 

brown-blackish. 

HABITAT: gregarious or in “witches’ circles” in 

meadows and hilly pastures and mountains, at high 

altitude, rarely in woods; start of summer-autumn. 


NOTE - A. excellens (F.H. Møller) F.H. Møller and A. 

stramineus (Jul. Schäff. & F.H. Møller) Singer, were 

originally recorded as two distinct species, varying only 

in stipe length and the straw yellow colour of the surface 

respectively. They are now unanimously considered as 

varieties of A. urinascens.

CAP 80-150 mm, from hemispheric to convex, sometimes 

conical typically flattened at tip, finally flat; convoluted 

edges, then curving towards the bottom, distends acutely 

late on, with partial remnants of veil; white colour, 

smooth and hairless, rarely lightly scaled, sometimes 

brown in the centre, intense chrome yellow on surface 

after minimal contact or rubbing. 

GILLS free, close, quite crowded, with lamellule; initially 

whitish, then pink, pink becoming darker until chocolate 

brown; sterile gill edge. 

STIPE:50-150 × 8-25 mm, slim and cylindrical, often 

curving, with tendency to broaden at base forming a 

roundish and sometimes non-marginated bulb, up to 35 

mm in width; white, yellowing if touched or upset; clear 

and broad annulus, fairly thick, separates late from edges 

of cap, flaring, sometimes appears dissociated from teeth 

in the lower face, yellowing if touched. 

FLESH whitish, stained chrome yellow at base of stipe to 

gill edge. Odour characteristic of phenol or ink. 

Agaricus xanthodermus Genevier 

MICROSCOPY: spore ovoid, dimensions 5-6.5 × 3.5-4 

µm; spores brown with violet shades. 

HABITAT: grassy areas, parks, roadsides, meadows, 

under trees. Summer to late autumn. 

EDIBILITY: toxic 

NOTE - May cause gastrointestinal disturbances. The 

clear yellowing of the flesh and inky or phenol odour are 

important features clearly distinguishing it from other 

Agaricus species of the arvenses (e.g. A. arvensis, A. 

essettei). A. campestris can also be similar, but has 

slightly red flesh near gill edge. A. xanthodermus due to 

the variability in the appearance of its cap covering, 

should be distinguished from the following varieties: var. 

griseus, from its scaly grey ochre-ish cap, var. lepiotoides, 

from its cracked cap with large grey-brown scales. A. 

praeclaresquamosus has a cap covered with concentric 

dark grey scales, while its centre is of a blackish colour. 

133

CAP up until 150 mm, from hemispheric to convex, then 

almost flat, sometimes fairly umbonate, smooth or more 

often minutely corrugated surface, dry and tends to crack 

coarsely in dry weather or becomes slightly greasy in wet 

weather; colour white cream, chamois, breadcrust, darker 

in the centre, sometimes, uniformly dark brown especially 

in smaller specimens. 

GILLS adnate or just starting to form a tooth, curved, 

very tall (up to 13 mm) and crowded, with numerous 

lamellule; from milk white to grey-ochre-ish, finally 

tobacco colour at spore maturation. 

STIPE full, dimensions extremely variable in both length 

and diameter depending on growing conditions: often 

thick and short, more often fine and slim, longer than 

diameter of cap, supple, curving, white then ochrebrownish. 

Surface smooth or fibrillated lengthwise 

sometimes finely scaled in dry weather. Annulus high, 

broad, membranous, persistent, white and then brown 

tobacco due to deposition of spores. 

134 

Agrocybe aegerita (Briganti) Fayod 

[= Agrocybe cylindracea (De Cand. : Fr.) Maire] 

FLESH white, a little brown at base of stipe in mature 

examples, tenacious, elastic. Odour characteristic and 

indefinable, acidic, like wine. 

MICROSCOPY: ellipsoidal spores, light brown under 

microscope, dimensions 8.4-9.5 × 5.0-6.0 µm; 

tetrasporophyte basidia. Brown spores. 

HABITAT: in numerous groups, cespitose, on live or 

dead trunks of broad-leaved trees, with preference for 

Populus, Ulmus, Acer. Fruits several times a year. From 

spring to late autumn. 


NOTE - This unmistakable mushroom is common to 

plains and flatlands, is much sought after and is safe and 

very good to eat. When given the right substrate it grows 

very well and is a commonly sold species. It, however, 

rapidly kills any plant or tree which hosts it, and will 

continue to grow on the dead tree for many years 

afterwards.

CAP 80-120 (180) mm, initially hemispheric, then 

convex, eventually flat-distended, hairless, occasionally 

bearing patches of membrane from veil, finely striated at 

hem; vivid orange colour, a little lighter at edges. 

GILLS free, crowded, a little ventricular; uniformly 

chrome-yellow colour. 

STIPE 80-140 (200) × 18-25 (35) mm; subcylindrical, 

attenuated at tip, normally straight, hairless; concolour 

with gills or of lighter tones; fairly hard and fibrous, fullmedulla, 

then fistular. Base area covered with a wide 

VOLVA SACK, attenuated at base, free and laciniate at 

edge, white, soft consistency, but fairly tenacious, thick 

up to 3 mm. 

ANNULUS membranous, positioned above median zone; 

long and densely striated; yellow. 

FLESH abundant and compact on cap, more granularfibrous 

on stipe; white or lightly yellowing, more yellow 

in peripheral area. Little or no significant odour and 

pleasant taste. 

Amanita caesarea (Scop. : Fr.) Persoon 

MICROSCOPY: spore from suboval to fairly regularly 

ellipsoidal, 9.4-11 × 6.2-6.8 µm; not amyloid. 

HABITAT: loves temperate climates and fruits in 

hardwood areas not rare even though not widespread. 


NOTE - Popularly known as Ovolo Buono (the tasty 

button), this is certainly one of the most popular edible 

species; appreciated since Roman times. It has recently 

been subjected to unrestricted gathering and moreover, 

this has often included specimens which are still closed. 

Because of this, in several areas A.Caesarea is in serious 

danger of extinction. In order to ensure its continued 

ability to sporulate, the law governing the collection of 

epigeal fungi in Italy prohibits the collection of these 

mushrooms during their button stage. This restriction can 

also be considered a safety measure because it prevents 

confusion with button mushrooms of the (very similar at 

this stage) deadly Amanita (see A. phalloides and similar 

species). 

135

CAP 50-90 mm, hemispheric then flat-convex, bearing 

fairly weak pyramidal warts. cuticle completely separable, 

viscid then dry, smooth, bright, whitish then with green 

tinges; smooth edges, displaying hanging remnants of 

veil. 

GILLS sub-free, slightly crowded, whitish with green 

shades, with floccose surface. 

STIPE 90-120 × 15-30 mm, cylindrical, with bulbousrooting 

base, white, sometimes hint of green, bearing 

small sections of more or less erect warts. Annulus: 

pendant, membranous, persistent, white, striated, with the 

edge bearing small flakes. 

FLESH hard white, hints of green. Fairly strong, 

unpleasant odour and flavour. 

136 

Amanita echinocephala (Vittad.) Quél. 

MICROSCOPY: spore 10-11 × 7-8 µm, ellipsoidal, 

smooth, hyaline, amyloid. basidia 45-60 × 11-13 µm, 

tightly clavate, tetrasporic, with joints at hinge. 

HABITAT: solitary or gregarious, under wide leaved 

trees, mainly beech and oak. Summer-autumn. 


NOTE - This species is characterised by its whitish 

colour, with greenish tinges here and there, the small, 

pointed warts which confer an echinulate appearance to 

the cap (hence the name), and its membraneous annulus 

and bulbous rooting stipe.

CAP 80-120 mm, hemispheric then convex, eventually 

flat; smooth edges. Viscous cuticle in wet weather, greybrownish 

or lead-grey, bearing floury plates of greybrown 

or dirty white veil. 

GILLS free, crowded, ventricular, white, sometimes 

shaded grey. 

STIPE 100-160 × 15-30 mm, robust, cylindrical, 

sometimes attenuated at tip, very fixed in ground, straight, 

full and dense, white-greying. VOLVA whitish, 

dissociated into plates at the base of the stipe, evanescent 

on cap, which usually appears bare. 

ANNULUS broad and persistent, striated on upper part, 

white-greying. 

FLESH firm and compact on cap, fibrous on stipe, whitegreying, 

a little reddening-browning. Root-like odour, 

similar flavour. 

MICROSCOPY: spore 8.5-9.0 × 5.5-7.5 µm, ovoid or 

mostly ellipsoidal, smooth, hyaline, amyloid. basidia 30- 

Amanita excelsa var. excelsa (Fr. : Fr.) P. Kumm. 

50 × 9.5-11 µm, tetrasporic, tightly clavate, without joints 

at hinge. 

HABITAT: single or in small groups, in hardwood and 

coniferous areas. Spring-autumn. 


NOTE - the separation of A. excelsa var. excelsa from A. 

excelsa var. spissa (Fr.) Neville & Poumarat [in this 

series, A. spissa (Fr.) P. Kumm.] has caused gallons of ink 

to be spilt, since these are two extremely similar species 

whose differentiating characteristics often "cross over". A. 

spissa typically has a more robust, “boletoid” form, an 

obese-clavate stipe, a veil that breaks down into powdery 

plates and adheres to the cap, while A. excelsa is more 

slender (hence its name) has a nearly cylindrical rooting 

stipe which holds tightly to the ground and a volva that 

dissociates into fairly large flaps, which do not adhere to 

the cap. 

137

CAP 50-70 (100) mm, hemispheric-glandiform, then 

convex, only distended late on, hairless, regularly bearing 

white membranous plates, residues of veil; warm to pale 

yellow, primrose yellow, lighter at edge which is finely 

striated. 

GILLS FROM free to subfree, ventricular, crowded and 

thin; white. 

STIPE 55-85 × 10-20 mm, progressively attenuated at tip, 

bulbous-napiform at base where it is covered by a 

membranous, fine, adherent VOLVA normally clear-cut 

at edge, sometimes also dissociated in rings that persist in 

the lower part of the stipe; white, hairless, full, fistular at 

base. 

ANNULUS positioned above the medial zone, very fine, 

soon becoming dissociated-evanescent, slightly persistent, 

sometimes completely absent in adult examples. 

FLESH tender, white, just yellowish under the cuticle; 

almost odourless, pleasant taste. 

138 

Amanita junquillea Quélet 

MICROSCOPIA: spore from subglobose to short 

ellipsoidal, 9.5-11 × 7.5-8.5 µm; not amyloid. 

HABITAT: early, from April to May, in hilly woods, a 

little later in June, in mountains. Common and widespread 

both in hardwoods and coniferous areas. 

EDIBILITY: suspect 

NOTE - Thanks to its sloping shape and its decidedly 

tender consistency, A. junquillea reminds one of the 

Amanitopsis, from which it can be distinguished by its 

annulus, which is actually rather ephemeral. Its edibility is 

a controversial topic. It must always be cooked, but even 

then is not tolerated well by all individuals. Among 

related species one should be careful of the decoloured 

form of the toxic A. pantherina (see below), which is 

morphologically similar, but has a brown cap.

CAP 100-150 (250) mm initially hemispheric then 

convex, eventually flat-distended, with the centre often 

depressed, with smooth cuticle, sticky in humid weather, 

almost always covered with white pyramidal warts, 

raised, concentric, residues of the veil, whitish, finely 

striated rim, reddish-orange, red, dark red. 

GILLS free, dense, ventricular, pure white or slightly 

yellow. 

STIPE 80-140 (200) × 10-20 (30) mm; cylindrical, 

attenuated at the apex, straight, hairless or slightly 

flocculent, white, rather tough and fibrous, then fistular 

and eventually hollow. The basal zone widens into a bulb 

covered with a thin VOLVA, dissociated with concentric 

whitish warts. 

ANNULUS membranous, large, located in the upper 

middle of the stem, striated on the top, remains of the veil 

found at the edge. 

FLESH firm in the cap, more granular-fibrous on the 

stipe; white and faded on the cuticle immediately below. 

No odour and a sweet, pleasant taste. 

Amanita muscaria (L. : Fr.) Hooker 

MICROSCOPY: spore subovoid and roughly ellipsoidal, 

9.0-11.2 × 6.5-7.5 µm; not amyloid; white in mass. 

HABITAT: in mountains, acidic soil, with hardwood and 

coniferous trees. Summer-autumn. Widespread. 


NOTE - This is without doubt the mushroom which has 

most stimulated the imaginations of illustrators over the 

years. In fact there are countless representations of this 

splendid species, both realistic and less so, which are 

employed when depicting the toadstools of mythical and 

fantasy worlds. In popular belief, A. muscaria is often 

confused with the lethal A. phalloides, a fact which brings 

no shortage of consequent danger to the inexpert. From a 

taxonomic point of view, we feel it appropriate to note a 

variant with a more brittle volva and, rarely, warts on the 

cap: the form aureola. 

139

CAP 70-130 (200) mm, hemispherical, then convex, 

convex-flattened for a long time, even revolute in old age; 

edge quite often joined, non-striated, exceedingly 

appendicular; silky-shining, white, ivory, cream and ochre 

with weak shade in the centre, hairless, sometimes with a 

few dissociated strips, remnants of veil. 

GILLS free, dense and thin, white, creamy pink shade in 

adults, minutely flocculent edge, alternating with 

truncated lamellule. 

STIPE 80-170 × 15-25 (35) mm, robust, yet slender, 

gradually expanding towards the sub-bulbous base, 

rooting at the base, full, firm, white, with concolour, or 

coloured, and highly transient floccules. 

VOLVA membranous, thick, fairly high, sheathing at the 

base, free at edge; ochre-ish externally, ochre-ish-pale 

orange, light clay-coloured, whitish inside. 

ANNULUS often situated fairly very high, typically softtender 

consistency (reminiscent of whipped cream), 

dissociated-evanescent, white, striated top. 

140 

Amanita ovoidea (Bull. : Fr.) Link 

FLESH highly abundant, softened, white on cap, more 

compact on stipe, with ochre-ish shades; salty odour, 

fairly pleasant, sweet flavour. 

MICROSCOPY: spore lengthwise ellipsoidal-ovoid, 

smooth, 9.5-11 × 5.5-7 µm; amyloid. 

HABITAT: regarded as a thermophilic species, but also 

spreads to subalpine zones, although more rarely. In oak 

and coastal pine woods it is very abundant and under 

broad leaved trees in woods in the hills it is rare. 


NOTE - the size, colours and the downy-tender 

consistency of the annulus of this mushroom constitute 

excellent diagnostic elements which should hopefully 

avoid any confusion between this species and the similar 

lethal white Amanita varieties. In literature we can often 

find it compared to A. proxima, which has a more 

colourful (even tawny) volva, but which may also be 

toxic; therefore we recommend caution be taken in its 

culinary use.


convex, eventually flat-distended sometimes depressed in 

the centre, smooth, covered with minute pure white floury 

warts, often concentric, remnants of veil; finely striated to 

edge (but smooth in var. abietum, which is heavier); 

brown-ochre, brown, brown dark. 

GILLS free to just non-marginated, crowded, slightly 

ventricular; pure white. 


attenuated at tip, straight, hairless; white; firm fibrous, 

then fistular and finally hollow, with base widening into a 

broad bulb. Basal zone is covered in an adherent VOLVA 

and dissociated from annulus, white. 

ANNULUS more or less broad, typically low on stipe; 

white and striated on the upper part. 

FLESH firm on cap, more fibrous on stipe; white. Odour 

almost nonexistent or slightly earthy, sweet flavour. 

MICROSCOPY: ellipsoidal spores, 9.5-11.5 × 6.5-7.5 

µm; not amyloid; white in mass. 

Amanita pantherina (De Cand. : Fr.) Krombholz 

HABITAT: in mountains, in coniferous and hardwood 

areas, often around the edges of woods. Summer-autumn. 


NOTE – Here is another toxic Amanita, whose 

dangerousness is without a doubt greater than that of A. 

muscaria, of which, at first sight, this might be considered 

a brown variant. In reality, A. pantherina has an evident 

and sturdy volva and its annulus is distinctly lower. A. 

junquillea, whose edibility is shrouded by serious doubts, 

has a similar shape, but its colour is a strong cowslip 

yellow. A. franchetii (which is edible cooked) has 

colourful forms and these could appear similar were it not 

for its yellowish veil, while A. rubescens (also edible 

when cooked) is of a brown-reddish colour and its flesh 

can easily be seen to redden when exposed to the air or in 

the tunnels created by larvae. 

141

CAP 60-120 (150) mm, hemispheric, then convex, finally 

distended radially from crowded and fine fibrils, often 

covered with white membranous strips, residues of veil; 

greenish, green-olivish, yellow-brown-greenish, lighter at 

edge which is smooth. 

GILLS free, ventricular, crowded and thin, fairly wide; 

white. 

STIPE 75-120 (150) × 10-22 mm, progressively 

attenuated towards apex, typically decorated with yellowolivish 

zig-zag bands on a white background; full, then 

medullar. Bulbous at base where it is covered with a 

membranous, fairly fine VOLVA sack, adherent to bulb 

but free towards edge where it is usually splits into white 

petal shapes. 

ANNULUS positioned fairly high, pendant, fairly fine, 

white, persistent. 

FLESH initially firm soon becoming soft, white, just faint 

green shades under the cuticle; from odourless to slightly 

smelly; older examples smell of putrid water (water from 

old flowers); no significant flavour. 

142 

Amanita phalloides (Vaill. : Fr.) Link 

MICROSCOPY: spore from mostly ellipsoidal to 

subglobose, 9-11.3 × 7-9 µm; amyloid. 

HABITAT: found in woods of deciduous trees in summer 

(rarely in conifers?) it seems to prefer oak, chestnut and 

beech trees, is also able to grow at high altitudes where it 

is regularly associated with hazelnut trees. Very common 

and widespread. 

EDIBILITY: deadly 

NOTE - This mushroom is the primary cause of all deaths 

which occur through mushroom poisoning. It is 

(unbelievably) confused with green Russule, from which 

it differs through a series of primary characteristics: it has 

a volva at the base of its stipe, it has an annulus, free gills, 

heterogeneity between cap and stipe, changing bands of 

colour on the stipe, non-chalky flesh. In nature one 

encounters numerous colour variations, including one 

which is completely white, the fo. alba.


convex, sometimes distended, smooth, viscous in humid 

weather, with minute remnants of veil in the form of 

crowded and prominent small warts, dirty white, greying 

or sometimes also ochre; with smooth brown, reddish 

edges, paling with age or heavy rainfall with 

characteristic vinous patches. 

GILLS rounded to stipe, crowded, ventricular and large, 

strong; white, stained red wine with age or to the touch. 


attenuated at tip, normally straight, hairless or sometimes 

minutely floccose under the annulus; from whitish to 

white pinkish and finally vinous brown; more or less full; 

firm then fibrous and fragile. The base widens into a non 

marginated ovoid bulb covered with an adherent VOLVA, 

soon becoming dissociated in floccose residues the same 

colour as stipe with a tendency to disperse. 

ANNULUS positioned in the middle to upper stipe, 

membranous, pendant, whitish or pinkish, sometimes 

even a little yellow, striated on the upper part. 

Amanita rubescens Pers. : Fr. 

FLESH abundant and compact in cap more fibrous and 

fragile in stipe; white then pinkish to gill edge; typically 

vinous red in veiled area, which is almost always visible. 

Odour irrelevant, pleasant, slightly salty-sour flavour. 

MICROSCOPY: ellipsoidal spores, 7.5-9.6 × 5.6-6.5 µm; 

amyloid; white in mass 

HABITAT: ubiquitous and widespread in conifer and 

hardwood areas. 


NOTE - Good to eat; though should be cooked first, as the 

flesh contains thermolabile toxins, and are best used in 

mixes. Confusion with the toxic A. pantherina is possible 

for the inexpert; the latter, however, has a very 

individualised volva and always has uniformly white 

flesh. The species which it most resembles 

morphochromatically is A. franchetii, which has more 

uniform flesh and a yellowish veil. 

143

CAP (30) 45-70 (95) mm, initially campanulate-parabolic, 

then convex, finally distended with broad slightly 

emerging umbo; hairless, rarely with residues of veil in 

shape of large plates, finely striated to hem; of ash grey, 

pearl grey colour, sometimes with a hint of brownish 

colour (white in var. alba; lead grey in var. plumbea), 

sometimes hint of ochre in disc area. 

GILLS free, crowded and thin; white, intercalated from 

lamellule trunk. 

STIPE 65-120 × 10-22 mm, slim, attenuated at tip, white, 

smooth, or becoming flocculent adnate and coloured; 

fully-medullar, finally fistular. 

VOLVA membranous, not very fine but fragile, adherent 

in basal area, free to hem; from white to whitish. 

ANNULUS apparently absent (reduced to shreds that 

remain at the base of the stipe, inside the volva, as in 

other "Amanitopsis” specimens). 

FLESH not very abundant, tender, white or whitish; no 

particular odour, pleasant flavour. 

144 

Amanita vaginata (Bull. : Fr.) Vittadini 

MICROSCOPY: spore from subglobular to globular, 10- 

12.2 × 9.6-11.6 µm; not amyloid. 

HABITAT: ubiquitous; found especially in hardwood or 

coniferous areas, common, from summer to late autumn. 


NOTE - As is the case for its relatives, this mushroom is 

considered an edible delicacy, ideal for frying (it must be 

cooked prior to consumption). The group, A. vaginata is 

certainly the most complex of those inside the subgenus 

Amanitopsis, which includes many species without 

annulus, at least at maturity; and in fact, there are many 

entities (species, varieties or forms) that are cited in 

literature, with nearly all having a greyish colouration in 

common. A. Mairei is a little more stocky and it has no 

umbo on its cap; A. nivalis, which is paler and smaller, 

can be found on the microsilve Alps. A. lividopallescens 

has more grey tones on its cream-brownish cap. The 

varieties plumbea, argentea, and alba differ only in their 

colour.

CAP 25-60 (100) mm, hemispheric, then distended finally 

flat-umbonate with slight prediscal depression, scattered 

with fine erect brownish transient scales, with fine edge, 

initially fringed with residues of veil, then bare and 

slightly striated, becoming undulating-sinuous in adults; 

variable colour, from yellowing brown to amber, dark 

brown, sometimes with olivish hints remaining at the 

extremes of the edges, pale, whitish. 

GILLS adnate and slightly decurrent lengthwise, fairly 

crowded and fairly close; whitish, then weak beige, 

finally bearing brown reddish spots. 

STIPE 60-120 (160) × 10-22 (35) mm, cylindrical, often 

tapered at base, other times a little dilated, grows 

cespitosely, fibrous-woody, full, soon becoming filled 

with an evanescent medulla, exterior is scattered with 

flocculent dissociated membrane disorderly arranged 

under the annulus; rather transient, fibrillated at tip; from 

cream-flesh coloured to concolour with cap, clearly 

browning at base. 

ANNULUS membranous, persistent, striated on lower 

face, floccose-cottony at edge; white, sometimes yellow 

at extreme edges . 

Armillaria mellea (Vahl : Fr.) Kummer 

FLESH fairly thick on circumference, whitish, highly 

leathery in stipe, rather bitter and astringent to taste, 

fungal odour. 

MICROSCOPY: spore from ellipsoid to ovoid, smooth, 

7-8.6 × 5.4-6 µm; basidia without joints on hinge. 

HABITAT: parasitic on wood of deciduous trees, often 

late, abundant, very common. 


NOTE - Edible with care (see note to A. ostoyae). 

Armillaria tabescens, which also grows cespitosely on 

hardwoods, is very similar but can be recognised by its 

lack of a membraneous annulus; A. ostoyae (= A. obdark) 

is darker, with persistent erect scales, annulus bordered 

with brown and prefers to grow on conifers; A. cepistipes 

is more frail and pallid and pallida, fairly hygrophanous; 

A. gallica (= A. bulbous, A. lutea) is less cespitose, 

usually grows as fairly-isolated specimens, happily on the 

ground (on underground woody deposits) and has 

residues of veil, as does the border of its annulus, which 

are largely yellow. 

145

CAP 30-80 (110) mm, hemispheric, then convex or 

convex and mostly umbonate, maintaining a bent edge at 

the extremes, finally distended with fine peripheral area, 

striated in maturity, initially fimbriated due to floccose 

remnants of veil, which is brown reddish, brown-tawny, 

sometimes paler, becoming brown-grey, paler at edge, up 

to cream whitish; scattered with erect scales, denser on 

circumference and dark pink in colour. 

GILLS adnate, then fairly decurrent, bent, not particularly 

close, attenuated at edge and join; whitish, pale yellow, 

often with brownish spots on surface, which eventually 

turns brownish. 

STIPE 60-140 × 10-22 mm, slim, cylindrical or a little 

tapered at base, up to sub bulbous, clustered-cespitose 

growth (generally few connate specimens), striated at tip, 

fibrillated-floccose under the annulus creating a white 

fluff, subcoloured near cap or darker at base, tendency to 

take on brown-olive-blackish tones. 

146 

Armillaria ostoyae (Romagnesi) Herink 

[= A. obdark (Schaeffer) Herink] 

ANNULUS membranous-cottony, fairly persistent, 

striated, white, often brown at extreme edges. 

FLESH fairly consistent on circumference, very fine 

elsewhere, fibrous-leathery in stipe, firm when young, 

then soft, whitish then suffused with flesh colour. 

MICROSCOPY: ellipsoidal spores, sometimes slightly 

compressed, smooth, to thick side, 8-10 × 5-6.6 µm. 

basidia with joints at hinge. 

HABITAT: prefers coniferous woods, but a hardwood 

form seems to exist; autumn, abundant and widespread. 


NOTE - For the differences from similar species, see the 

notes to A. mellea and A. tabescens. Edible, but as with 

the other Armillaria with annuli, a long cooking time and 

eliminating the stipe are necessary to avoid fairly 

unpleasant gastro-intestinal poisoning.

CAP 30-70 mm, initially campanulate-convex, then 

distended, finally flat with broad obtuse umbo, sometimes 

rather depressed in prediscal area, superimposed with 

erect fairly persistent scales; edges curving at the 

extremes, fine, distended in maturity and a little striated; 

brown reddish, brown-beige, paling to cream ochre-ish. 

GILLS bent and decurrent, thinned at the front, not very 

crowded, close, fairly thick, whitish, then pale-flesh 

coloured, browning on surface in maturity. 

STIPE 35-70 × 8-15 mm, slim, cylindrical or subfusiform 

fairly bent, densely connate with other examples, white at 

tip, whitish flesh with splashes of colour, then decidedly 

and extensively browning in a smooth continuum away 

from darker fibrils; full-medullar, finally fistular. 

ANNULUS nearly absent or reduced to transient residues, 

coniform, only visible in very young examples. 

FLESH compact in cap, more fibrous in stipe, white, 

whitish-beige pale; fungal odour, sweet flavour with 

slightly bitter aftertaste. 

Armillaria tabescens (Scop.) Emeland 

MICROSCOPY: ellipsoidal spores with thick wall, 9-10.2 

× 5.4-6.6 µm. basidia without joints at hinge. 

HABITAT: cespitose, in large groups connate, above all 

on logs or at the base of oak trees, then spreading to the 

ground. Not very widespread but abundant in growth 

areas. 


NOTE - the tender consistency of the flesh, even the stipe, 

combined with a “more traditionally fungal” flavour make 

this species an excellent and sought after edible 

mushroom, doubtlessly superior to the annuli-bearing 

Armillaria. It can sometimes seem like a twin of A. mellea 

s.l., but the absence of a membraneous annulus makes for 

a fairly easy distinction. A. ectypa has no annulus either, 

but grows in beds of sphagnum moss or in mountain 

bogs; its size brings to mind the Laccaria. 

147

CAP 80-160 (250) mm, fleshy, from hemispheric to 

convex-pulvinate; edge curving at extremes, then 

regularly distended with white evanescent bloom; matt 

cuticle, finely tomentose, then hairless, not viscous, even 

in humid weather; fairly dark, generally brown-blackish, 

typically discoloured to brown-ochre yellowish. 

TUBES length up to 25 mm, non-marginated-adnate, 

whitish and persistent until maturation is incipient; 

greenish-yellow late on, finally olive, uniform to gill 

edge, very small pores, concolour with tubes. 

STIPE 60-130 × 40-85 (110) mm, often rounded in young 

examples, then slimmer, ventricular, or even cylindrical, 

sometimes curving; ochre-ish, brown-ochre-ish, honey 

coloured, light hazelnut, towards the upper part (from 1/3 

to 2/3) covered with a fine lattice, generally the same 

colour as the skin of the stipe. 

FLESH firm and compact when young and this remains 

in adults, later soft; pure white, uniform, uncoloured 

under cuticle of cap. Weak but very pleasant odour, 

sweet, hazelnut flavour. 

148 

Boletus aereus Bull. : Fr. 

MICROSCOPY: spore fusiform dimensions 13.5-16 × 

4.0-5.0 µm, pale yellow under microscope; olive-brown 

in mass. 

HABITAT: the most thermophilic of porcini mushrooms, 

it prefers sparse oak or chestnut woods, in which it can be 

found from summer’s start till the end of autumn, solitary 

or gregarious, not very common in the north, the species 

is fairly widespread in southern areas . 


NOTE - Commonly known as the “Porcino nero”, after 

the colouration of its cap. Its particular biological needs 

mean that it is widespread particularly in southern areas, 

but can also be found in northern Italy (though not higher 

than 800 m above sea level) in hotter periods of the year. 

Sometimes the distinction from darker forms of B. 

aestivalis can prove challenging. In that case the colour of 

the flesh, pure white only in B. aereus, and the 

decolouration of the pileic cuticle are determining factors.

CAP 100-250 mm, fleshy, from hemispheric to convex, 

finally pulvinate-flat; cuticle finely velvety, never 

viscous, often finely cracked especially at edges or, in 

very dry weather, evenly tessellated in large areolas 

which are barely visible beneath the flesh; uniformly pale 

brown, coffee, hazelnut, reddish brown, often also dark 

brown. 

TUBES up to 30 mm, depressed-adnate, from white milk 

to straw yellow, then yellow-greenish, finally olivish, 

uniform to gill edge; has very small pores, the same 

colour as tubes, unchanging to the touch. 

STIPE up to 150 × 80 mm, initially obese then slimmer, 

cylindrical, often curving, rarely ventricular, rounded at 

base and sometimes rooting a little; bark coloured when 

young, with tones which become intensified and match 

those of the cap, or a little paler. Fine mesh lattice the 

same colour as background, which covers the surface of 

the stipe right to the base. 

Boletus aestivalis (Paulet) Fries 

[= Boletus reticulatus auct. non Schaeff.] 

FLESH firm and compact in young, soon becoming soft 

in cap and a little stringy in stipe, often eroded by small 

larvae; milk white, just brownish under cuticle; peculiar, 

intense and sweet odour; sweet, pleasant hazelnut flavour. 

MICROSCOPY: ellipsoidal fusiform spores with weak 

elevated depression, 12.8-15.1 × 3.8-4.4 µm, pale yellow 

under microscope. Olive brown in mass. 

HABITAT: warm open woody grassland areas, often 

associated with Quercus, Fagus, Castanea, but also to 

conifers (Picea) from late spring to early autumn 


NOTE - This is the “summer porcine” which appears in 

late spring in the clearings and glades of woodlands. The 

characteristics which can help to distinguish it from other 

porcine mushrooms are: a dry, often cracked cuticle; a 

stipe which is coloured even in young specimens, the 

flesh of the cap, which is usually pliable or saggy to the 

touch in mature examples. 

149

CAP 100-250 mm, fleshy, hemispheric then convexpulvinate, 

finally flat; undulating edge in mature 

examples, covered with a whitish bloom when young 

which tends to disappear. Cuticle initially finely velvety, 

soon becoming viscous, wrinkled surface especially in 

marginal zone; chestnut brown, hazelnut, brown dark, 

except at the extreme edges which have a permanently 

white surface, discoloured to pale ochre towards the edge. 

TUBES up to 30 mm, depressed-adnate, milk white then 

yellowish, finally olivish, uniform to gill edge; pores very 

small, concolour with the tubes, unchanging to the touch. 

STIPE up to 150 × 100 mm, obese, then ventricular or 

cylindrical, with rounded base; milk white in young, later 

hazelnut or pale brown; fine mesh lattice, concolour with 

background, spread over most of the surface. 

FLESH firm and compact when young, then a little soft 

in maturation; white, slightly brown-reddish for a few 

millimetres under the cuticle; with typical pleasant odour, 

very pleasant flavour, sweet, like hazelnut. 

MICROSCOPY: ellipsoidal fusiform-spores with weak 


150 

Boletus edulis Bull. : Fr. 

under microscope; olive brown in mass; basidia mostly 

bi- and trisporic. 

HABITAT: ubiquitous in the woods, often associated 

with Fagus, Picea, Abies, but also with many other 

species (chestnut, pine, birch, hazel, etc.) Found in 

temperate or cool periods, from late summer to late 

autumn, Widespread and very common. 


NOTE - This is the “autumnal porcino”, common and 

recognised in almost all the continents, where it manifests 

a substantial stability in its characteristics and it associates 

with a great number of beings. It is recognisable by its 

viscous cap, which tends to lose its colour, especially at 

the edges. B. aereus and B. aestivalis are similar, but 

these have dry caps and more-richly coloured stipes from 

the start; B. pinophilus, which also has a viscous cuticle, 

but shows winey-brown tones on its cap and an obclavate 

stipe.

CAP 80-220 (300) mm, very fleshy, from hemispheric to 

pulvinate-convex, finally flat; edge curving at extreme 

edges, finally distended, scattered with whitish bloom 

which tends to dissolve at maturation; cuticle initially 

finely velvety, soon becoming viscous, rugulous; vinous 

brown, garnet, reddish-brown, copper red, with sporadic 

discoloured areas 

TUBES width up to 30 mm, depressed-adnate, milk white 

to straw yellow, then yellow-green, finally uniform 

olivish to gill edge; pores are very small, concolour with 

the tubes, often slightly rusty in maturation, unchanging 

to the touch. 

STIPE up to 100-200 × 50-120 (150) mm, typically 

obese, then a little longer, but relatively short and stocky 

and almost always dilated-rounded at base; whitish, then 

soon showing hints of reddish brown. Displays a fine 

lattice, concolour with background, which covers most of 

the stipe. 

FLESH firm and compact, then a little soft and watery; 

white, purplish for a few millimetres under the cuticle; 

Boletus pinophilus Pilát & Dermek 

[= B. pinicola Vittadini, non Swartz] 

typically pleasant, but weak odour, very pleasant sweet 

taste. 

MICROSCOPY: ellipsoidal fusiform spores with weak 


under microscope; olivish brown in mass. 

HABITAT mainly associated with Picea, Pinus, Fagus, 

Castanea. Fruits typically two times a year: in late spring, 

at lower altitudes, and from the late summer to late 

autumn in the mountains and at high altitudes. Recurrent. 


NOTE - Usually better known under the name B. pinicola 

Vitt. (an invalid name as it has already been used to 

designate a lignicole fungus, today known as Fomitopsis 

pinicola), it is popularly called the “red porcine”. Its name 

would seem to suggest it is found only in affinity with 

pine trees, however B. pinophilus is, much like edulis, a 

widely-occurring species and is more usually found under 

beech or fir trees. 

151

CAP up to 150 (200) mm, from hemispheric to convex, 

then flat-pulvinate; regular edge, margins slightly larger. 

Plical, finely felted surface, initially coral pink-red, to 

raspberry pink, discoloured with age, gaining yellowish 

tones, often also with olive tones; cracks at areole in dry 

weather. 

TUBES similar thickness to flesh of the cap, when 

carpophore matures, stipe is rounded; from gold yellow to 

olive green; quickly turning blue if cut lengthwise; small 

pores, round then a little angular; from gold yellow to 

olive green, uniform or turning blue if bruised 

STIPE usually narrower in diameter compared to cap on 

maturation of carpophore, up to 40 (50) mm in diameter; 

stocky, with thick base, more often than not cylindrical; 

pale chrome yellow, often with innate raspberry red spots 

towards base; surface unchanging to the touch. Lattice 

usually limited to upper half, often slightly raised, 

concolour with background. 

FLESH initially very firm in cap and also in adult 

carpophore, while stipe tends to become fibrous; pale 

152 

Boletus regius Krombholz 

yellow, more intense yellow above tubes and under the 

bark of stipe, often reddish only at extreme base of stipe, 

rarely or never turning blue at gill edge in zone above 

tubes. Weak, slightly fruity odour; sweet flavour. 

MICROSCOPY: spore pale yellow under microscope, 

fusiform dimensions 11.3-14.5 × 3.5-4.5 µm. Spores olive 

brown. 

HABITAT: isolated or in small groups under hardwood 

trees, mainly Fagus sylvatica and Castanea sativa in 

acidic ground; from early summer to autumn 


NOTE - A beautiful species which is only confusable 

with the Boletus pseudoregius, which however has more 

slanting proportions and flesh which manifests fairly 

intense tones. This species has been implicated in several 

cases of poisoning after undercooked carpophores were 

consumed.

CAP 120-300 mm, very fleshy, from hemispheric to 

convex; cuticle finely velvety when young, soon 

becoming hairless, dry, milk white, pale grey, cream, 

often with olivish hues, finally brownish olive, often just 

pinkish at edges in examples that have grown in humid 

weather. 

TUBES up to 25 mm, rounded near stipe, from yellow, to 

yellow-greenish, finally olivish, blue at gill edge; pores 

very small, initially yellow but soon red, carmine red or 

red-orange; rarely remaining yellow or weakly orange 

even at maturity; blue when touched. 

STIPE 60-150 × 50-100 mm, narrower in diameter than 

cap, very stocky, obese, pear shaped, more often than not 

cylindrical; usually yellow at insertion to cap, red or pinkpurple, 

fuchsia at lower part. Lattice generally concolour 

with background, of fine mesh, isodiametric, limited to 

upper half of stipe; the surface turns blue to the touch. 

FLESH very firm and compact when young, soft when 

old, pale yellow, turning pale blue at gill edge; weak 

particular odour when young, cadaverous or of decaying 

substances when mature, sweet flavour. MICROSCOPY: 

Boletus satanas Lenz 

ellipsoidal spores, with weak elevated depression, 11.4- 

13.4 × 5.1-5.9 µm, yellow under microscope; olive brown 


HABITAT: warm and chalky woods, associated with 

Quercus, commonly with Castanea or Fagus, not at high 

altitudes. Summer-autumn, not very common. 


NOTE - This is one of the largest boletes, characteristic 

for its short and paunchy stipe and for the cadaverous 

odour that it emits when ripe. Its light cap, red pores and 

the latex in its stipe help to distinguish it from other 

species of Luridi that might seem similar. In many cases, 

it can be found with completely yellow pores and stipe, 

and as such it can even resemble certain species of 

Appendiculati (Boletus fechtneri). Notwithstanding, its 

distinctive and unmistakeable odour helps in its 

identification. Toxic when raw, suspect (and certainly 

poorly tolerated by many people) even when cooked 

thoroughly. 

153

CAP 40-80 mm, from convex to flat, convoluted edges, 

often undulated, smooth, silky, matt, white, dirty white or 

dirty cream, often stained ochre or cracked. 

GILLS crowded, close, adnate or non marginated, from 

whitish to pale cream, with the surface undulated or 

crenulated, coloured. 

STIPE 40-70 × 10-20 mm, cylindrical, almost clavate or 

attenuated towards the base, blooming or fully fibrillated, 

then filled, whitish or dirty cream. 

FLESH thick, hard or a little spongy, white, with strong 

smell of flour and sweet, floury flavour. 

MICROSCOPY: spore 5-6 × 2.5-3.5 µm, ellipsoidal, 

smooth, hyaline; basidia 20-25 × 3-5 µm, tetrasporic, 

tightly clavate, filled with siderophile granules; epicyte 

formed from interwoven hyphae, width of 3-5 µm, 

hyphae of subgelled stratum surface, with membranous 

pigmentation; usually with hinges. 

154 

Calocybe gambosa (Fr. : Fr.) Donk 

[= Lyophyllum georgii (L. : Fr.) Kühner & Romagnesi; 

Tricholoma georgii (L. : Fr.) Quélet] 

HABITAT: grows isolated or in groups, more often in 

circles or semicircles (witches’ circles) in grassy areas in 

meadows, or in clearings in coniferous or hardwood areas, 

particularly close to Rosaceae. 


NOTE - This is a much sought after mushroom with many 

pseudonyms, the most famous of which is probably St. 

George’s mushroom (so named because the period in 

which it grows in near that saint’s day). In literature 

several varieties have been identified based on the diverse 

colours that the cap can assume and even sometimes the 

different odours it gives off. For example, C. graveolens 

(= Tricholoma georgii fo. flavida) has a darker cap and a 

rather unpleasant odour.

BASIDIOCARP up to 100 (150) mm, composed of a 

fertile glebe protected by two cortical layers (endo - and 

exoperidia) supported by spongy sterile base; subglobular 

subellipsoidal or with short fairly differentiated 

pseudostipe; initially mostly convex on upper part then 

truncated-flat, external surface of exoperidium smooth, 

ruffled-rugulous and then separated into aureola; from 

milk white to pale grey ochre-ish. Across cracks produced 

by one sees that the endoperidium aureola in maturity is 

grey-brown and finally dark brown, in adults the skin 

tears so extensively and irregularly as to allow the 

dispersal of spores which remain for a long time on the 

ground; very spongy base, slightly marcescible, uniformly 

brown. 

GLEBE white, then yellowish, greenish, olive, finally 

brown-olivish or chocolate brown; fairly soft consistency, 

then dusty. 

Calvatia utriformis (Bull. : Pers.) Jaap 

[= Lycoperdon caelatum Bull. : Pers.] 

MICROSCOPY: spore globular, with warty surface, of 

diameter 4-5 µm, brownish to microscope. Spores olive 

brown in mass. 

HABITAT: isolated or in groups of several individuals, 

often with many examples joined at base, in meadows and 

in mountain pasture, from summer to autumn; recurrent 


NOTE - From summer’s end, walking through mountain 

meadows, one can often come across the seemingly 

indestructible brownish remnants of this large “puffball”. 

These residues can in fact persist on the ground for over a 

year. It may be confused with some Lycoperdaceae, 

which are also edible, such as Langermannia gigantea, 

the largest, with its near-smooth exoperidium, free of 

pseudostipe. C. utriformis is also morphologically very 

similar to Vascellum pratense of which, at first glance, it 

seems to be a giant form. 

155

156 

Camarophyllus pratensis (Pers. : Fr.) Kummer 

[= Hygrocybe pratensis var. pratensis (Fr.) Murril; = Cuphophyllus pratensis (Pers. : Fr.) Bon] 

CAP 25-80 mm, hemispheric then convex, often with 

mostly obtuse umbo, fine margin; hairless, radially dry 

away from fairly fine fibrils; pale apricot orange or fleshy 

ochre, sometimes covered with white bloom; cuticle 

detachable for 3/4 of radius. 

GILLS curved, deeply decurrent, concolour with cap or 

paler, highly spaced, noticeably thick, often spoked, 

intercalated with numerous lamellule. 

STIPE 40-100 × 5-12 mm, normally cylindrical or 

attenuated at base, more rarely obclavate, from whitish to 

pale orange, visibly fibrous, fully dry, often invaded by 

larvae or small insects, soon becoming filled. 

FLESH thick and firm fibrous in stipe; whitish or faded 

yellow in cap, white elsewhere, odourless, pleasant 

flavour. 

MICROSCOPY: spore mostly ellipsoidal, 5-6.7 × 4.2-5 

µm; tetrasporophyte basidia, sometimes bisporic, and so 

produce larger spores, up to 8.6 µm. 

HABITAT: grassy areas, meadows, clearings in 

hardwood areas; gregarious; widespread and common 

from summer to late autumn. 


NOTE - This is a good species of edible mushroom, 

which has seen fair commercial success, despite its 

fibrous flesh. Due to its orange colour, it can be confused 

with Hygrophorus nemoreus, but the latter has an even 

more fibrous cap, flesh which both smells and tastes of 

flour, pale gills, which range from creamy-whitish to flesh 

coloured. It also has a smooth sub-bilateral lamellar trama 

and grows in the woods (or, at least in the vicinity of 

trees). Several varieties are reported as different in 

literature due to their differing size and colour (var. 

robustus, of larger size, var. Donadini with denticulate 

edges, var. Vitulinus, more fragile and pale).

CAP 30-70 (120) mm, initially highly differentiated from 

stipe, then convex, then distended with edge convoluted at 

the extremes, eventually irregularly flat-gibbous up to 

fairly deeply depressed-infundibulform, marginal zone 

very thin, with progressively rough, sinuous-lobed, 

wrinkled in parts, shiny appearance; with humidity 

minutely velvety, smooth; when dry, orange, apricot, 

paling yolk-yellow. 

HYMENOPHORE pseudolamellar, very forked from 

compound-branched folds, abundantly spored, 

anastomosed, highly decurrent in adults, close, thick, with 

fairly dull surface, coloured or subconcolour with cap, 

Sometimes with pink shades. 

STIPE 30-60 (90) × 12-25 mm, cylindrical, normally 

progressively flared towards gill insertion, often even a 

little dilated at base, full, firm in maturity softer-rubbery, 

hairless, subcoloured. 

FLESH white, with yellow-pinkish stains on outer zone, 

abundant on circumference, firm and compact in cap, 

fibrous and almost leathery in stipe; pleasant odour, like 

Cantharellus cibarius (Fr. : Fr.) Fries 

apricot or white peach skin or sweet fruit. Initially mild 

flavoured, then sour-astringent-spicy. 

MICROSCOPY: spore from ovoid to ellipsoidal, 

sometimes sublarmiform, smooth or fairly evidently 

grainy, 7.5-9.6 × 4.6-5.6 µm. 

HABITAT: very common and widespread all over, from 

hilly hardwood areas to coniferous mountains; from start 

of summer to late autumn. 


NOTE - A much sought and valued edible mushroom, this 

is one of the few which are also eaten in both southern 

and northern Europe. Literature is full of variants of this 

fungus which are sometimes considered separate species, 

sometimes mere variants. The var. bicolor is more 

precocious and displays a very pale cap and stipe which 

are whitish, in stark contrast to its yolk-yellow gills; the 

var. ferruginascens has tones of olive green and a strong 

tendency to assume a rusty colour when handled; the var. 

amethysteus has violet adnate scales on its cap, and can 

typically be found under beech trees. 

157

CAP 15-45 (65) mm, convex-umbillicate, then flat with 

fairly wide central depression, infundibulform, edge bent 

and fine at the extremes, progressively multi-lobed, fuzzy 

in parts, fissile, revolute in old examples, sinuous; 

fibrillous-scaly orange brown or reddish brown. 

HYMENOPHORE reduced to low venucole with 

numerous branches and anastomoses, mostly rounded at 

margin, rugulous, not infrequently subsmooth especially 

in stipe where it is, somewhat blurred, decurrent 

lengthwise, yellow-orange, yellow or greying with a hint 

of salmon pink in paler forms. 

STIPE 35-70 × 8-12 mm, normally highly irregular, 

dilated at tip and progressively attenuated towards base, 

often bent, canalicular-compressed, gibbous and wrinkled, 

hairless, hollow-tubular, highly characteristic bright 

orange colour or hint of pink salmon,. 

FLESH exiguous all over, of fibrous and fairly tenaciouselastic 

consistency, cream or pale yellowish; fruity odour, 

like of plums or of flowers of the Muscari family, sweet 

flavour. 

158 

Cantharellus lutescens (Pers. : Fr.) Fries 

[= C. aurora (Batsch) Kuyper] 

MICROSCOPY: spore from ellipsoidal to ovoid, smooth, 

guttulate, 9.6-11.4 × 6.5-7.8 µm. 

HABITAT: summer-autumn, in hardwood or coniferous 

woods, often in grassy or mossy areas, more rarely on 

bare ground, widespread all over and abundant in growing 

area. 


NOTE - Had the name C. aurora not been synonymised, 

this would have the nomenclatorial priority, however, 

since C. lutescens is widely used, we adhere to this 

binomial and hope it lasts. Good to eat, Especially after 

drying and then steeping in water and milk. Its fine and 

slightly fibrous-elastic flesh make it appear like another 

species of similar colour: C. tubaeformis, but this latter 

has a hymenophore with fairly distinct pseudogills; less 

vivid colours, ranging from shades of gray to grey-brown, 

and is less odorous.

BASIDIOCARP appearance in young specimens of a 

rounded oval, internally gelatinous, while external surface 

is fragile and waxy at branches. When cut, the parts ready 

for growth are clearly visible. 

EXOPERIDUM is similar to a railing round the elongated 

polygonal meshes, vivid colour, grows up to 70 mm in 

diameter at full development. On the lower part, 

connected to the remnants of the primordial oval, a volva 

is formed, the thin elongated hyphal rhizomorphs are 

clearly visible and whitish. 

GLEBE formed from small granules of green-blackish 

mucilage containing spores, gives off a strong unpleasant 

odour similar to that of Phallus impudicus. 

Clathrus ruber Mich. ex Pers. : Pers. 

MICROSCOPY: ellipsoidal spores, 5 ¥ 6 µm. Spores of 

white-brownish colour. 

HABITAT: grows fairly isolated in the humid parts of 

woods, from summer to autumn. 

EDIBILITY: non edible 

NOTE - This is a rare species, but which grows 

abundantly in its chosen areas. It has the same cadaverous 

odour of Phallus impudicus; an odour which appears even 

before the mushroom does. The same repellent odour is 

often responsible for this species being invaded by flies. 

159

160 

Clitocybe cerussata (Fr. : Fr.) Kummer 

[= C. phyllophila (Pers. Fr.) Kummer; = C. pytiophila (Fries) Gillet] 

CAP 40-90 mm, convex then flat, typically umbonate. 

pure white cuticle, smooth, covered with sericeous fibrils, 

blooming, matt; at most slightly cream colour under 

fibrils. 

GILLS from adnate to subdecurrent, very crowded, pure 

white, sometimes a little cream, concolour with the whole 

surface,. 

STIPE 40-70 × 10-15 mm, from cylindrical to subclavate, 

full, white, slightly fibrillated with base covered with 

white flakes. 

FLESH whitish, firm with no significant odour or flavour. 

SPORES white. 

MICROSCOPY: ellipsoidal spores, 5.0-6.0 × 3.0-4.0 µm, 

smooth, cyanophilia, prevalent in tetrads in dry 

conditions, obtuse base. basidia 16-25 × 4-6 µm, 

tetrasporic. hymenophore almost regular. very loose 

Pileipellis interwoven with structure of hyphal cutis. 

HABITAT: gregarious, in coniferous woods; fairly 

common and widespread, from end of summer. 


NOTE - C. phyllophila belongs to a group of organisms 

which are very similar to each other, and which maybe 

represent different aspects of one, single species. When 

trying to distinguish between C. cerussata (Fr. : Fr.) 

Kummer and C. phyllophila (Pers.: Fr.) Kummer, one 

would mention the habitat of conifers for the former and 

of hardwood trees for the latter, and also their different 

spore colours (pink-cream in C. phyllophila). It is not so 

simple to distinguish C. phyllophila from C. candicans 

(Pers.: Fr.) Kummer or C. rivulosa (Pers.: Fr.) Kummer. 

Normally C. phyllophila is slightly larger than these two 

latter species and has characteristic adnate, slightly 

decurrent gills, and spores arranged in tetrads in dried 

specimens. C. candicans, with its slanting gait, has more 

decurrent gills and single spores. C. rivulosa (= C. 

dealbata sensu auct. plur.) normally grows in meadows.

CAP 60-200 mm, quite deeply infundibulform, with 

central umbo emerging from cavity, not hygrophanous, 

not striated for transparency, pale ochre-ish, alutaceous, 

smooth, matt, silky, Finely felted. 

GILLS fairly deeply decurrent, fairly crowded, whitish, 

stained pinkish-brown, concolour with the whole surface. 

STIPE 60-150 × 13-20 mm, from cylindrical to lightly 

clavate, filled, subcoloured to cap, bearing white fibrils 

lengthwise, tomentum white at base. 

FLESH white, with particular aromatic odour, and 

insignificant flavour. 

SPORES white. 


smooth, single in dry conditions, sublacrimoid base. 

basidia 40-45 × 7-9 µm, tetrasporic. Fairly regular 

hymenophore texture, colourless hyphae, width 3-8 µm. 

Pileipellis interwoven with structure of hyphae cutis, 

Clitocybe geotropa (Bull.) Quélet 

width 3-7 µm, with membranous and minutely encrusting. 

pigment 

HABITAT: terrestrial, gregarious, in hardwood and 

coniferous woods or in meadows and pastures, in the 

shape of "witches’ circles", not widespread but with 

plenty of areas of growth. Autumn and late autumn. 


NOTE - Unmistakeable thanks to its typical shape, 

characterised by a very long stipe compared to cap 

diameter and to its fairly large size, this late-appearing 

species is a much sought edible. 

With its basidia, which are fairly long for a clitocybe, C. 

geotropa is grouped in the subgenus Hygroclitocybe Bon, 

inside which it is unique for its subregular lamellar 

trama, its sublacrimoid spores and its mixed 

pigmentation. 

161

CAP 30-80 mm, fairly deeply infundibulform, with or 

without umbo, with the edges convoluted, then straight, 

often ribbed, not hygrophanous, not striated by 

transparency, ochre-ish pale or alutaceous light reddish, 

finely felted, matt. 

GILLS decurrent, fairly crowded, bent, light yellowish 

brown, often with pink stains, with the whole surface, 

coloured. 

STIPE 20-50 × 5-8 mm, from cylindrical to lightly 

clavate, filled, then fistular, from whitish to light yellow, 

generally lighter than cap, smooth or with white fibrils 

lengthwise, with tomentum white at base. 

FLESH whitish, with pleasant “cyanide” odour and sweet 

flavour. 

SPORES white. 

MICROSCOPY: ellipsoidal spores, 5.2-6.6 × 4-4.6 µm, 

smooth, not cyanophilous, single in dry conditions, with 

confluent base, lacrimoid. Basidia 22-30 × 5-7 µm, 

162 

Clitocybe gibba (Pers. : Fr.) Kummer 

[= Clitocybe infundibuliformis (Schaeff.) Quélet] 

tetrasporic. Hymenophore texture regular with colourless 

hyphae. Pileipellis interwoven horizontally with structure 

of hyphae cutis, with finely encrusted pigment. 

HABITAT: in groups, also numerous, in hardwood and 

coniferous woods 


NOTE - It is known and sought after in Italy as the 

“imbutino” (funnel mushroom), and a much-appreciated 

edible, despite its fairly tough flesh. C. gibba belongs to 

the subgenus Clitocybe whose species are characterised 

by their typically funnel-like caps which are often opaque 

and almost velvety, by their decurrent gills, the 

(sub)regular hymenophoral trama and by the colouring in 

their stipe walls. It is not always easy to distinguish it 

from C. costata, though it can normally be identified by 

its stipe being lighter than its cap and its cap covering 

having a negative reaction to treatment with KOH.

CAP 80-150 mm, convex, with convoluted edges, not 

hygrophanous, from ash grey to grey-brown, smooth, 

finely felted, fibrillated from adnate to lightly decurrent, 

fairly crowded, pale cream, with the whole surface 

coloured. 

STIPE 60-90 × 15-30 mm, clavate, filled, subcoloured to 

cap, striated lengthwise with fine fibrils, base covered 

with white mycelial felt. 

FLESH white, with strong aromatic, slightly unpleasant 

smell and unpleasant flavour. 

SPORES yellowish cream. 

MICROSCOPY: ellipsoidal spores, 6.0-7.5 × 3.-4.5 µm, 

smooth, cyanophilous, prevalent in tetrads in dry 

conditions, obtuse base. basidia 20-25 × 5-7 µm, 

tetrasporic. Hymenophore texture regular with colourless 

hyphae. Pileipellis with structure of hyphal cutis more or 

less parallel, with intracellular pigmentation. 

Clitocybe nebularis (Batsch : Fr.) Kummer 

HABITAT: ubiquitous, in large groups; very common in 

autumn and late autumn. 


NOTE - This is a well known species, which in many 

areas is gathered and eaten with impunity. In any case, 

recent studies (based on nutritional casuistics) seem to 

demonstrate the toxicity of the species, or at least, the illtolerance 

of it by several individuals. Based on the spore 

colour, the cyanophylla of the spore wall (which is 

smooth to optical microscopes, but warty to electronic 

ones), C. nebularis was considered by several authors, 

such as Moser and Bon, as belonging to the genus 

Lepista. We prefer, in accordance with Kuyper, to keep it 

in the genus Clitocybe, along with numerous other similar 

species which display cyanophylla and coloured spores 

which are smooth under an optical microscope. 

163

CAP 60-150 (220) × 30-70 mm, initially glandiform to 

more or less cylindrical then, in maturity, from 

campanulate to conical, more expanded with age, finally 

deliquescent starting at the edges; surface initially silky 

and white, soon becoming covered in overlapping scales 

from whitish to light brown on white background; cap 

often joined and ochre-ish. 

GILLS very crowded, unequal, very wide, free to stipe, 

initially white, then, pinkish at edges, eventually black, 

deliquescent. 

STIPE 100-200-(300) × 10-25-(35) mm, separable from 

cap, slim in maturity, attenuated at tip and fairly bulbous 

at base, white, bearing fine white fibrils, empty with age; 

medial or basal annulus, which is membranous, minute, 

white, sometimes black due to spores. 

FLESH slightly thick, tender in cap and soon becoming 

fibrous in stipe, white; weak and pleasant odour and 

flavour. 

MICROSCOPY: spore from ellipsoidal to ovoid, smooth, 

with centrally germinating pores, brown-black under 

164 

Coprinus comatus (Müll. : Fr.) S.F. Gray 

microscope, 11-14.5 × 6.5-8 mm; tetrasporophyte basidia. 

Black spores. 

HABITAT: from the spring to late autumn in grassy and 

fertile areas in gardens, at the edge of wheel tracks, in 

flood plains, in large groups; frequent. 


NOTE - This is, most probably, the only Coprinus which 

is considered edible and, as such, it deserves a certain 

level of attention; several authors, in fact, consider it 

excellent or, even, «the best edible species»; the exiguity 

of its flesh, however, means that long cooking times 

should be avoided and therefore would best be eaten fried 

or even raw. In any case one should only eat young 

specimens, which still have perfectly white gills. This 

species is unlikely to be confused with similar species: 

Coprinus sterquilinus grows in dung, is solitary or found 

in small groupings, it is also more slender and has larger 

pores; Coprinus vosoustii, is far rarer, has a nondeliquescent 

star shape covering on its cap and much 

larger spores.

CAP 40-80 mm, initially fleshy, campanulate-convex, 

then flat-convex, eventually distended gibbous or with 

wide central umbo, edges convoluted then straight, 

slightly lobed, dry cuticle, non-hygrophanous, matt, silky, 

densely fibrillated or scaled, reddish brown orange, brown 

auburn. 

GILLS adnate-non-marginated, fairly spaced, wide, 

bulging, brown ochre-ish, brown reddish orange then 

rusty reddish due to spores, with eroded surface, 

yellowish on face. 

STIPE 40-90 × 10-15 mm, fairly slim, cylindrical but 

often attenuated at base, more or less supple, full, firm 

citrina yellow, reddish orange in the centre, decorated 

lengthwise with coloured fibrils. 

FLESH firm non hygrophanous, ochre-ish, reddish hint, 

with radish-like odour and acidic flavour. 

MICROSCOPY: ellipsoidal amygdalform spores, 10-12.5 

× 5.5-6.5 µm, densely covered with fine warts. 

Filamentous epicyte, with encrusted pigment. 

Cortinarius orellanus Fries 

HABITAT: isolated or gregarious, mostly under oak 

trees, but also under beech and hazelnut; not widespread, 

but constant in growing areas; from end of summer to all 

of autumn. 

EDIBILITY: deadly 

NOTE - Characterised by its fairly robust shape, its scaly 

orange-brown cap, subdistant coloured gills, yellowish 

stipe and root-like odour, C. orellanus is responsible for 

some of the worst cases cytotoxic poisoning, which have 

a particularly grave effect on the renal system. 

Considering its long latency period, the so called 

Orellanus syndrome is extremely dangerous. Of equal 

toxicity is the C. orellanoides (= C. speciosissimus), 

which has similar colouring, but which is found in conifer 

woods. This latter typically has a conical-umbonate cap 

and a stipe which is decorated with changeable, yellowish 

zig-zag stripes. 

165

CAP 50-200 mm, initially subhemispheric, then convex, 

eventually flat-convex, with fine edges, convoluted at the 

extremes, then straight, very thick and tenacious cuticle, 

viscid in humid weather, shiny, chocolate brown, 

becoming a little reddish, brown purple, more or less 

blended with grey-violet or lilac tones, scattered with 

large silky silver purple strips, residues of veil, marked at 

the edge with visible wrinkles or radial grooves. 

GILLS adnate-non-marginated, fairly crowded, bulging, 

greying or dirty whitish, hint of violet, then clay brown, 

finally rusty due to spores, with jagged surface, whitish. 

STIPE 70-150 (200) × 20-40 mm, very robust, 

progressively dilated to a large non-marginated bulb, a 

little rooting; full, firm and almost bare at tip, hairless, 

whitish then cream, marked lengthwise with silky fibrils, 

the lower two thirds visibly decorated with thick residues 

from veil, violet-blue silver in colour 

FLESH thick, very firm white or cream, hints of violet at 

top part of stipe, with weak, slightly fruity and pleasant 

odour and sweet flavour. 

166 

Cortinarius praestans (Cordier) Gillet 

MICROSCOPY: spore from amygdalform to 

subcitriform, 14.0-17.0 × 8.0-10.0 µm, covered with 

coarse evident warts. Filamentous gelled epicyte, made 

from frail hyphae. Membranous pigment. 

HABITAT: infrequent but abundant in growing area, 

found in hardwood areas. 


NOTE - By far and away the largest of the Cortinari, a 

real giant, plus it has organoleptic properties which make 

it a choice edible in great demand by enthusiasts. Beyond 

its size and masterly shape, the morphochromatic details 

which assist in its identification are the absence of a 

marginated bulb, the whitish colour of the gills which 

contrast with the reddish-brown of the cap, an abundant 

universal veil, at least in young specimens, which leaves 

clear traces on the cap but above all on the stipe, and its 

large spores. C. cumatilis is fairly similar, but is far 

smaller and grows under conifers.

CAP 35-75 (90) mm conical-campanulate then distended 

with broad and slightly accentuated obtuse umbo, fairly 

fleshy for the genus Entoloma; steel blue with hints of 

grey, tendency to take on a less obvious grey-brownviolet 

colour with age, often, with whitish bursts, dry, 

hairless and joint at opening, soon becoming dissociated 

from gathered fibrils, eventually fairly split; often 

rugulous-wrinkled in maturity. Fine Cuticle, separable, 

and only with small stripes. 

GILLS non-marginated-adnate, ventricular, fairly wide, 

with surface irregularly undulated; white ivory, 

sometimes (when seen flat) almost pale cream, soon 

becoming pinkish due to maturing of spores, eventually 

dirty pink. 

STIPE 30-70 × 12-25 mm, normally widening in the 

middle and fairly tapered at base, not infrequently 

slightly canalicular, firm, full, fibrillated lengthwise, 

concolour with cap, often with violet tinges especially at 

Entoloma bloxamii (Berk. & Br.) Saccardo 

[= E. madidum (Fries) Gillet] 

tip, always lighter or white at base; loses a lot of tone 

intensity with age, becoming slightly grey-blueish. 

FLESH white, firm slightly fibrous in stipe flavourful, 

floury odour. 

MICROSCOPY: polygonal, subisometric spores, with 

evident apex 7.5-8.7 (9.5) × 6.2-8.2 (9.5) µm; joint at 

hinge. 

HABITAT: in meadows and in grassy and open areas of 

woods; autumnal species, present from plains to 

mountains, not common, gregarious. 


NOTE - This is one of the tricholomatoid Entoloma with 

the most beautiful colour and shape; similar to E. 

bloxamii for its shape and habitat is E. porphyrophaeum, 

which, however, does not have the blueish colouration but 

is grey-violet or grey-purple-ish and has longer spores of 

8.6-11.6 × 6-7.8 µm. 

167

CAP 50-200 mm, initially campanulate, then convex, 

eventually flat, with or without a low umbo, edges 

undulated, convoluted, lobed, the cuticle more or less 

light grey with silky tinges, metallic or even lead grey, 

grey brownish, blooming, lead or silverish from fine 

fibrils. 

GILLS deeply non-marginated, almost free, fairly spaced, 

bent, then more or less ventricular, typically yellow, soon 

becoming yellowish salmon pink or pink ochre-ish, with 

the surface serrated, coloured. 

STIPE 50-140 × 15-35 mm, slim, cylindrical or supple, 

often widened but often also attenuated at base, firm full 

then filled, white, pruinose at tip, marked lengthwise with 

silky fibrils. 

FLESH white, firm non hygrophanous, with strong, 

unpleasant floury smell, and unpleasant flavour, bordering 

on disgusting. 

168 

Entoloma sinuatum (Bull. : Fr.) Kummer 

[= Entoloma lividum (Bull. ) Quélet] 

MICROSCOPY: spore subisodiametric, with 6 sides, 8.5- 

11.0 × 7.5-8.5 µm. Tetrasporophyte basidia, with joints to 

hinge. Cheilocystidia absent. epicyte framed from 

(ixo)cutis. Intracellular pigment. Joints at hinge. 

HABITAT: in hardwood areas, preferably Quercus and 

Fagus; from end of summer to late autumn; fairly 

common. 


NOTE - Despite its fleshy, inviting appearance and its 

often-pleasant, floury odour, this species is a dangerous 

one, the cause of potentially severe gastrointestinal 

illness. An added risk comes from its similarity with 

Clitocybe nebularis, a mushroom commonly consumed in 

certain parts of Italy, and with which E. sinuatum often 

shares its habitat and growing period. The latter has a 

non-fibrous cap, fairly decurrent, separable gills, a whitish 

cream colour and a very particular, strong odour.

ASCOCARP stipitate and pileate. 

MITRA up to 35 mm, irregular shape vaguely resembling 

a horse saddle, with two-three lobes stretching towards 

the outside, many take on a curly appearance, whitish, 

ivory, up to fairly dark cream. Hymenophore covered 

with visible part of mitra, smooth, undulated. Lower 

surface slightly lighter, tomentose-floccose. Sinuouslobed 

edge, jagged. 

STIPE height up to 120 mm, cylindrical-clavate, widened 

towards at base, deeply ploughed lengthwise, incomplete 

both internally and externally, white or whitish. 

FLESH leathery, but fairly elastic, sub-brittle; whitish, no 

distinctive odour or flavour. 

MICROSCOPY: ellipsoidal or mostly ellipsoidal spores, 

smooth, 18-19 × 10-12 µm, hyaline under microscope, 

monoguttulate, uniseriat in asco; cylindrical asci, non 

amyloid, octasporic; cylindrical thin paraphyses, with 

clavate apex. 

Helvella crispa (Scop. : Fr.) Fries 

HABITAT: on the ground in hardwood or mixed woods 

or, also in grass, single or in small groupings of several 

specimens. Common in summer-late autumn. Sometimes 

even in late spring. 


NOTE - Helvella lactea has a similar shape, but is 

smaller (just 30-40 mm tall), and is completely ice white, 

has a smooth lower surface and fruits on the ground or on 

decaying plant matter (Fraxinus) in autumn. H. lacunosa 

has a much darker, blackish mitra a brownish-grey stipe, 

and is ubiquitous. H. sulcata is considered an extreme 

form of H. lacunose by some authors; it is rarer, bigger, 

and has a regular “saddle-shaped” it is almost completely 

grey in colour. Furthermore, it has a yellowish-ochre-ish 

or yellowish grey mitra, dimensions rather smaller size, 

and prefers southern, sandy ground near H. pityophila. 

169

ASCOMA pileate and stipitate, formed from apothecia 

(cap) and from stipe; total height up to 100 mm. 

APOTHECIA irregularly lobed-curled, rarely 

subselliform, composed of two, three or more variously 

joined lobes, height up to 15-20 mm and of around 30 mm 

in diameter. External surface (hymenial) irregularly 

rippled-undulated, yellowish; lower surface (sterile) 

smooth or lightly rough, whitish. Edge irregularly 

undulated or sinuous. 

STIPE up to 80 × 10-20 mm, subcylindrical or 

significantly widened at base, pleated-ribbed, lacunous 

internally, concolour with cap towards top, but with lilac 

or grey-lilac stains, more marked at base. 

FLESH elastic and fairly tenacious, but fragile, brittle, 

whitish-yellowing; insignificant odour and flavour. 

MICROSCOPY: spores mostly ellipsoidal, 18-20 × 11- 

12.5 µm, guttulate, with smooth wall, hyaline, uniseriat in 

asco; whitish in mass. Cylindrical octasporic, not amyloid 

170 

Helvella pityophila Boud. 

asci. Cylindrical paraphyses, dilatate at tip, sometimes 

forked, with several septums. 

HABITAT: species not widespread all over, fruits alone 

or in small groups generally on sandy ground in humid 

hardwood coniferous woods (mainly Pinus sp.) In both 

alpine and subalpine mediterranean areas; summerautumn. 


NOTE - Very similar to H. crispa (Scop. : Fr.) Fr., it can 

be distinguished only by its colouring, which tends to be 

grey with cream reflections at the apothecia and have an 

absence of grey-lilac tones on its stipe. H. crispa can also 

reach larger dimensions (up to 120 mm in height) and is 

generally found only in hardwood forests. H. lactea Boud. 

is smaller (growing to a height of 40 mm), is completely 

white, becoming ochre-ish-brown in dry conditions; it has 

smaller spores (16-18 × 11-12 µm), and fruits in autumn 

in woods rich in decaying and decomposing plant matter.

CAP 15-50 mm, initially mostly campanulate or 

subhemispheric, eventually convex or convex-flat, 

hairless, shiny-lubricated appearance in humid conditions, 

shiny-dry in dry conditions; in these latter conditions 

shows very adpressed radial fibrils (slowly!). Margin 

fairly regular, initially inflected, fine, tendency to crack 

radially; intense red-cherry or red-carmine colour, paling 

until white to from disc but always with a characteristic 

red annulus in the peripheral area. Fine cuticle, separable 

until circumference. 

GILLS adnate or briefly decurrent, sinuous-ventricular, 

rarely bent, wide, thick, unequal, fairly sparse, 

intercalated by numerous lamellule of different lengths; 

yellow initially, soon becoming red-orange or cherry red, 

the surface remains yellow. 

STIPE 30-80 × 5-8 mm, irregularly cylindrical, 

sometimes compressed-canalicular, often bent, fragile, 

dry, coloured cap at tip, more shades elsewhere, white 

base; hollow. 

Hygrocybe coccinea (Schaeff. : Fr.) Kummer 

FLESH fine, from yellow orange to red orange, also red 

cherry on periphery, watery, odourless, mild insignificant 

flavour. 

MICROSCOPY: elliptical spores at prosurface or 

amygdalform, 8-10 × 4.2-5.2 µm; tetrasporophyte basidia. 

HABITAT: in humid and mossy grassy areas, pastures, 

clearings, gregarious and abundant in growing area, 

frequent; end of summer-autumn even late autumn. 


NOTE - Among other small, red Hygrocybes this one’s 

cap, which is absolutely hairless and never scalytomentose 

sets it apart from H. miniata. Less simple 

however is its distinction from other hairless redcap 

species. H. reae is smaller, has a striated margin, bitter 

flesh and is abundantly viscous-glutinous; H. insipida is a 

near replica of the previously-described mushroom, but 

has sweet flesh and a size closer to H. coccinea; H. 

punicea is noticeably larger, has a campanulate cap, 

yellow stipe and white flesh with clear red fibrils. 

171

CAP 10-30 mm, campanulate or hemispheric-umbonate, 

then distended to a fairly prominent and acute papilla or 

distended and mostly umbonate, abundantly viscousglutinous, 

fragile, with the margin fine and striated with 

transparency; highly polychrome, yellow ochre-ish 

olivish, grass green with sulphur yellow tones, yellow 

orange with green stains, sometimes the yellow pigment 

is absent thus the green tones evolve towards blue-blueish 

ones. Fine cuticle, yet separable at circumference. 

GILLS non-marginated-adnate, spaced, ventricular, with 

regular and joint surface; golden yellow, saffron yellow, 

sometimes with greenish or tawny stains (nearly white in 

absence of yellow pigment); intercalated by numerous 

lamellule of different lengths. 

STIPE 40-80 × 3-6 mm, cylindrical or progressively 

widened at base, often supple-curving, highly viscous due 

to a persistent and thick layer of hyaline gluten; yellow 

greenish, green or green blueish at tip, sometimes with 

tawny or brick coloured veins, soon becoming hollow. 

172 

Hygrocybe psittacina (Schaeff. : Fr.) Kummer 

FLESH fine, white, yet coloured in cap, also fairly 

deeply, on periphery, very watery; no odour and slightly 

earthy flavour, like moss. 

MICROSCOPY: ovoidal spores, smooth, more rarely 

ellipsoidal, (6.5) 7.4-9 × 4.5-5.6 µm. 

HABITAT: in meadows, among grass and moss, highly 

camouflaged, in autumn; gregarious, fairly frequent. 


NOTE - This species is made unmistakeable by its 

colouring and its marked glutinous properties; in any 

case, there are often anomalies in its pigment-distribution 

(see description), which can make it appear with 

unexpected colours. H. perplexa (= H. sciophana) is a 

lookalike, though with wider gills, is a brick-red colour 

with slight green tinge and is fairly rare. Even H. laeta 

can seem similar due to its many colours and viscosity, 

while H. psittacina, on the other hand, has a convex cap, 

and more importantly, very decurrent gills, which are not 

non-marginated or adnate.

CAP 30-60 mm, hemispheric, soon becoming convex, 

viscous, initially white, with tempo takes on fairly 

extensive yellowish; edge with fairly regular flow, finely 

hairy, adorned with floccose-cottony granules soon 

becoming yellow; fine separable cuticle,. 

GILLS decurrent, fairly wide of and fairly spaced, thick, 

intercalated from numerous lamellule of different lengths; 

white, in maturity with vague flesh coloured tinges, then 

tendency to turn yellow like the rest of the carpophore to 

the edge. 

STIPE 30-70 × 8-14 mm, slim, cylindrical, normally 

attenuated at base; cortiniform residues visible at tip 

which, like the edge of cap, bears the same grainy 

flocculence, characteristic, tendency to turn yellow, 

slightly viscous due to humidity, soon becoming dry 

initially full fibrils, soon becoming medullar. 

FLESH very fine towards the edges, compact on 

circumference, white, due to imbibition takes on a citrina 

Hygrophorus chrysodon (Batsch : Fr.) Fries 

tinge; indistinctive flavour, odour not especially 

pronounced, but clearly of the “cossus” type. 

MICROSCOPY: spores lengthwise ellipsoidal, smooth, 8- 

9.2 × 4.2-5 µm. 

HABITAT: in the mountainous hardwood areas, with 

preference for beech but also in the coniferous and mixed 

woods; gregarious, from end of summer to all of autumn, 

fairly common and widespread. 

NOTE - This beautiful hygrophore reaches the peak of its 

loveliness when the granulation which adorns its cap 

margin and the tip of its stipe assume their intense yellow 

colouration. It is liable to be confused with species of 

eburneus-cossus, which however, is highly viscous, non 

flocculent and does not visibly yellow. An exception to 

this last rule however is H. discoxanthus (= H. 

chrysaspis), which, when it dehydrates, becomes a 

distinct rusty fawn colour (the entire carpophore is rusty 

brown when dried). 

173

CAP 15-45 mm, convex, then flat with broad obtuse 

umbo, finally also depressed in prediscal area, with the 

edge bent at the extremes, excess; pure white, only in 

aged specimens vague pale ochre-ish or flesh colour 

stains are visible on circumference, highly viscousglutinous, 

thick, elastic and separable cuticle. 

GILLS adnate or slightly decurrent, sinuous, slightly 

ventricular, then bent, highly thick, not very spaced, 

white, finally lightly flesh coloured; intercalated from 

lamellule of different lengths. 

STIPE 40-100 × 8-15 mm, slim, cylindrical, sometimes 

supple, or a little dilated, attenuated at base, dry and 

floccose at tip, glutinous in the lower 3/4; white, with age 

often with ochre-ish shades or pinkish at base. 

FLESH white, thicker on circumference, not unpleasant 

flavour, while on the contrary odour is pronounced and 

nauseating, with fruity, sweet and sour hints of the 

“cossus” family, similar to shellfish cooking, and defined 

in literature as similar to the odour of woodworm. 

174 

Hygrophorus eburneus (Bull. : Fr.) Fries 

CHEMICAL REACTION: potassium hydrate (KOH) hot 

yellow, yellow orange, base of stipe. 

MICROSCOPY: spores lengthwise ellipsoidal, sometimes 

subcylindrical, smooth, 7.4-8.5 × 4.5-5.6 µm. 

HABITAT: in hardwood areas with preference for beech, 

but not exclusively; from end of summer to all of autumn, 

non very common. 


NOTE – H. eburneus is likely to be a collective species in 

which other entities may be grouped which are currently 

considered as independent species by some authors. Thus 

H. quercetorum is a variant which is just slightly more 

robust; we consider H. cossus to be a simple variety with 

a yellow pale reaction to potassium. H. discoxanthus (= 

H. chrysapsis), despite being similar morphologically, we 

believe merits the dignity of its own species thanks to its 

all-over colouration of rusty-fawn to brown reddish, in 

dry conditions, which is even more evident in dried 

samples.

CAP 50-120 (160) mm, convex with broad and obtuse 

central umbo, then distended and also depressed around 

umbo, very fleshy, firm glutinous, grey-sooty or greygreenish, 

often with vaguely lilac stains (similar to 

Gomphidius glutinosus); generally the disc peripheral 

zone are darker, wide areas decorated in ivory white 

present elsewhere. Thick, stocky, very adherent cuticle. 

GILLS bent-decurrent, whitish, then stained ochre- pale 

flesh colour; thick, wide, venous on background, not very 

spaced, intercalated from numerous lamellule, generally 

small and short in length. 

STIPE 60-150 x 20-50 mm, full, bulky, fusoid, obese in 

middle then subrooting, white, covered by hyaline glutine 

(only in maturity or drying more brown-olivish coloured 

in parts), floccose decorations at tip. 

FLESH thick, compact, firm white; odourless, no 

particular flavour. 

Hygrophorus latitabundus Britzelmayr 

(= Hygrophorus limacinus Scop. ex Fr. ss. Auct.) 

HABITAT: exclusively in Pinus woods, in mountainous 

and coastal areas fairly late; rare, but abundant in growing 

area. 

MICROSCOPY: spores fairly lengthwise ellipsoidal 8.5- 

11.5 x 4.7-7 µm. Tetrasporophyte basidia. 


NOTE - This is a very heavily-set Hygrophorus. It 

belongs to the Olivaceoumbrini with Hygrophorus 

olivaceoalbus, which is decidedly frailer, with a slender 

stipe and associated with spruce and with H. persoonii (= 

H. dichrous), a mushroom which can reach a considerable 

size, but which fruits associated with oak and has a 

particular green reaction to ammonia fumes. Other species 

of the genus are easily recognisable by their different 

morphological features. 

175

CAP 35-100 (150) mm, convex-hemispheric, soon 

becoming distended with broad obtuse umbo, finally also 

depressed in prediscal area, with edge bent at the 

extremes, thinned; initially white or whitish ivory, shades 

of cream or yellow-pinkish on circumference in adults 

some excoriation is also present; slightly viscous due to 

humidity, soon becoming dry generally maintaining a 

fairly shiny look. 

GILLS adnate, fairly decurrent in adults, bent, thick and 

spaced, concolour with cap or fairly uniformly pale 

cream; intercalated from lamellule. 

STIPE 30-70 (90) × 12-25 (30) mm, typically tapered at 

base, firm and fleshy, barely reaches in length the same 

dimensions as the diameter of cap; dry or slightly viscous 

with humidity, white, eventually ochre-ish at base in 

adults; full. 

FLESH abundant on circumference, firm and compact, 

more fibrous in stipe, white; flavourful, sometimes with 

slightly bitter aftertaste, odour is not very clear but 

characteristic and unmistakable, like boiled milk. 

176 

Hygrophorus penarius Fries 

CHEMICAL REACTION: flesh of stipe reacts yellow 

with KOH. 

MICROSCOPY: ellipsoidal spores, sometimes briefly 

ellipsoidal, smooth, 6-7.4 × 4.5-5 µm. 

HABITAT: in autumn, in hardwood areas, especially in 

the oak; abundant in growing area. 


NOTE - Much sought after due to its good organoleptic 

characteristics, and inexpensive thanks to its fleshiness, 

H. penarius has earned a reputation as the best edible 

hygrophore. Other similar white and dry hygrophores are: 

H. Karstenii, found on mountain spruce, with apricot 

gills, H. poetarum, found in beech forests, which has a 

matt cap, pink tinge and a balsamic odour; H. phages, 

grows on beech, is slightly shorter, with a pinkish 

reflection on its gills; finally H. barbatulus, a rare species 

with a pale cream-ochre-ish cap, is fairly hairy on its 

margin and has ochre-cream-ish gills.

CAP: 35-100 (135) mm, convex with edge convoluted 

and excessive at the extreme edges, then from pulvinate to 

fairly distended sometimes lobed, very fleshy; initially 

whitish, soon becoming fairly sporadically spotted with 

wine red patches or frills, more crowded on 

circumference, then with zone fairly wide of delicate 

raspberry pink, sometimes also completely coloured with 

the same tone, sometimes yellowing with age; lightly 

viscous to humid weather, soon becoming dry very fine 

cuticle, separable for 1/2-2/3 of radius. 

GILLS: from horizontal-adnate to lightly bent-adnate, 

sometimes non-marginated; very crowded for the 

Hygrophorus genus, thick, of irregular width but always 

fairly scarce; white or whitish-flesh coloured, in maturity 

fairly densely spotted with winy red colour, intercalated 

from lamellule in report of ca. 1:1. 

STIPE: 35-70 × 10-25 mm, irregularly cylindrical, often 

slightly attenuated at base, floccose at tip, white then 

Hygrophorus russula (Schaeff. : Fr.) Quélet 

[= Tricholoma russula (Schaeff. : Fr.) Gillet] 

fairly extensively marked with red-purpling spots; dry, 

firm, full. 

FLESH: white, very firm can take on light pinkish stains 

especially towards the foot of stipe; highly variable 

flavour also contained in the same report, from mild (ca. 

50%) to bitter up to clearly bitter, no significant odour. 

MICROSCOPY: ovoid or ellipsoidal spores (6) 7-9.3 × 

4.5-5.2 µm. 

HABITAT: in hardwood areas with preference for oak, in 

autumn and also late autumn, gregarious in large groups, 

rarely solitary. 


NOTE - In some zones it is highly prized and is stored in 

oil. The most trustworthy characteristic in distinguishing 

it from the similar species, H. erubescens, is the particular 

crowdedness of its gills; other valid distinguishing 

features are its habitat, its indisposition to yellowing, and 

its slightly-smaller pores. 

177

CAP 20-50 mm, initially convex, then from flat to 

depressed in centre, with the edges convoluted at the 

extremes, then distended, a little undulated, scalloped and 

furrowed; completely vivid purple-amethyst fairly dark, 

paling, with dry conditions, towards grey-blueish, ochrecream 

or dirty white with lilac-ish tinges; cap covering 

matt, smooth or felted, slightly scaled in the centre; 

hygrophanous. 

GILLS mostly adnate to stipe or a little decurrent, 

fairly spaced, width of and thick, anastomosate at base; 

vivid violet, pruinose; with irregular surface. 

STIPE 40-60 (100) × 4-8 mm, cylindrical, often 

undulated, often wider at tip, striated lengthwise with 

white fibrils on dark violet-amethyst background; whitish 

towards base, strongly paling in dry conditions and when 

cut lengthwise reveals whitish flesh. 

178 

Laccaria amethystina (Huds.) Cooke 

FLESH fine in cap, pale violet; flesh of stipe is fibrous, 

elastic, white and firm with sweet odour and flavour. 

MICROSCOPY: spores 9.0-10.0 × 8.5-10.0 µm, globular, 

with tall spines up to 2 mm; tetrasporophyte basidia. 

Cheilocystidia abundant, 30-80 × 6.0-10.0 µm, wiry, 

irregular, sometimes branched, hyaline. Pileipellis 

hyphae cutis cylindrical arranged radially. 

HABITAT: grows humus rich soil, from hills to 

mountains, gregarious or in groups, in hardwood and 

coniferous woods. 


NOTE - This species cannot be confused with other 

Laccaria. Only very old and particularly dry specimens, 

which have lost their characteristic violet colour, run any 

risk of confusing one as to their identity.

CAP 10-35 mm, initially convex, then flat and depressed, 

striated for transparency almost up to centre; edges from 

round to crenulated. hygrophanous cuticle, ochre-beige, 

finely scaly with dry weather with almost marginal area, 

brown-tawny, finely warty-grainy in humid weather. 

GILLS mostly adnate or lightly decurrent, fairly spaced, 

wide, salmon pink or red-brownish, with the surface 

whole coloured. 

STIPE 30-60 × 1-3.5 mm, cylindrical, widening at base, 

from full to hollow, covered with fine white fibrils 

lengthwise on brown-red background, pruinose to apex, 

with mycelium white at base. 

FLESH fine, red-brownish, watery. Pleasant odour, sweet 

fungus flavour. 

Laccaria fraterna (Sacc.) Pegler 

MICROSCOPY: spores 8.5-10.5 × 7.0-9.5 µm, from 

subglobose to mostly ellipsoidal, echinulate, hyaline, with 

spines reaching 1 µm in length; bisporic and monosporic 

basidia. 

HABITAT: basidiomata, gregarious or cespitose, near 

Eucalyptus, Pinus, Cupressus sp. pl., in Mediterranean 

environment. Summer-autumn. 


NOTE - It is almost impossible to determine a Laccaria 

with only the morphological data such as that evidenced 

by L. fraterna, to go on without the aid of a microscope. 

This species can be recognized by its mostly bisporic 

basidia, its spore size, the length of its spines and its 

typical Mediterranean habitat. 

179

CAP 10-35 (45) mm, initially hemispheric, then convex 

ed finally flat and depressed in the centre; edges striated 

for transparency, undulated and denticulate; from brownorange, 

brown reddish, to pinkish red in humid weather, 

paler to beige or faded ochre-ish with the dry; matt 

surface, smooth, very fine radial fibrils, a little dandruff in 

centre, hygrophanous. 

GILLS mostly adnate or lightly decurrent, wide, spaced; 

light flesh colour light at first, then brown-pink, on the 

whole surface. 

STIPE 35-100 × 3-5 mm, cylindrical, often slightly 

widened at base, full then hollow, elastic; surface from 

brown-red to brown dirty, smooth or just striated, covered 

longitudinal with whitish fibrils. 

FLESH watery, from brown-grey to whitish, fine; weak 

herbal odour; sweet fungus flavour. 

180 

Laccaria laccata (Scop. : Fr.) Berkeley & Broome 

MICROSCOPY: spores from subglobose to ellipsoidal, 

hyaline, 8.0-10.0 × 6.5-8.0 µm; spines 1-2 µm in length. 

basidia clavate, 28-45 × 8-15 µm, tetrasporic, hinged; 

length of sterigmata up to 10 µm. regular lamellar web. 

Cheilocystidia cylindrical, fairly supple, 25-60 × 3-7 µm. 

Cuticle formed from parallel hyphae, interwoven, width 

of 7-15 µm, with some hyphae scattered perpendicularly. 

Hinges present. 

HABITAT: isolated or gregarious, in hardwood or 

coniferous woods or on their outskirts, in uncovered 

areas, on carpets of needles, among moss 


NOTE - L. laccata var. laccata with its small and delicate 

carpophores this can be distinguished from other varieties 

above all due to its ellipsoidal spores, which have a 

length/width ratio of over 1-2.

CAP 50-150 mm at start, convex, then flattened, finally 

depressed-infundibulform, yellow-orange, orange-ochreish, 

with more saturated concentric zone, edges 

convoluted, then more or less undulated-lobed, hairless 

surface, a little viscous in humid weather, then dry, 

pruinose, more clear at edges. 

GILLS lightly decurrent, crowded, rigid, forked, pale 

orange then carrot red in fractures, finally, very slowly, 

dirty greenish. 

STIPE 30-60 × 15-25 mm, filled then hollow, pruinosefelted, 

pale orange, more or less bearing red-orange 

dimples. 

FLESH pale, carrot red in fractures to the latex, then very 

slowly becoming greenish (in 24 ore), with fruity odour 

and mild flavour. 

LATEX: carrot red, uniform or paling, with mild flavour. 

Lactarius deliciosus (L. : Fr.) S.F. Gray 

MICROSCOPY: spores from mostly ellipsoidal to 

ellipsoidal, 8.1-10.3 × 6.5-8.0 µm, with warts joint with 

fairly thick ridges, forming an almost complete lattice; 

tetrasporophyte basidia, subclavate; Pileipellis, an 

ixocutis is present 

HABITAT: exclusively associated with Pinus; frequent in 

pine. From summer to the first signs of winter. 


NOTE - Of those species which associate with conifers 

and have orange or red latex, L. deliciosus is easily 

recognisable by its cap with orange-ochre-ish patches, its 

stipe covered with red-orange dimples, its unchanging 

latex and its habitat of pine trees. Of the large, edible 

milk-caps this one is fairly well known, and, probably, the 

one best suited to the dinner table. 

181

CAP 30-100 mm, initially convex then flat, finally 

depressed in the centre, vivid orange, then rusty orange, 

greening all over with age, not zoned or with crowded 

zoning at edges, slightly evident, rounded edges then 

open, the surface slightly viscous in humid weather, then 

dry, pruinose-rugulous. 

GILLS adnate-decurrent, very crowded, forked, ochreorange-ish, 

stained green-brownish in fractures. 

STIPE 30-60 × 10-25 mm, filled, soon becoming hollow, 

concolour with cap, generally not scrobitulate, pruinose, 

bearing a white rim at tip, eventually greening all over. 

FLESH cream-orange, slowly turning to winey red, then 

dark green, with fruity or carrot like odour and mild or 

lightly acrid and bitter flavour. Guaiac, winy grey. 

LATEX: slightly abundant, orange, turning red-winey in 

15 minutes, with mild flavour, then a little acrid and 

bitter. 

182 

Lactarius deterrimus Gröger 

MICROSCOPY: spores from subglobose to mostly 

ellipsoidal, 8.0-11.4 × 6.6-8.7 µm, with warts joined with 

thin ridges, forming a very incomplete lattice; 

tetrasporophyte basidia, subclavate; Pileipellis, it has an 

ixocutis. 

HABITAT: symbiont of Picea abies; highly widespread 

and common in fir abetaie and red fir; from summer to 

late autumn. 


NOTE - The peculiar characteristics of this species are the 

non patchy cap and stipe, though which soon become 

stained green. The stipe is not dimpled, though has a 

typical white circle at its tip; the latex moves slowly from 

orange to winey red and its habitat is near Picea abies. In 

spite of its name, which means “the worst”... it is 

considered a good edible, even if less well-esteemed than 

L. deliciosus.

CAP 50-150 mm, initially convex, then flat-depressed, 

finally infundibulform, white-cream, soon becoming 

dotted with brown-ochre ones, eventually with some rusty 

shades, the edges are fine and convoluted when young, 

then lobed-undulated, the cuticle is adnate, dry, hairless, a 

little rugulous, with the tendency to crack. 

GILLS from adnate-decurrent to slightly decurrent, very 

crowded and close, thin, with numerous lamellule, white 

with pale cream and flesh colour tinges, brownish in 

fractures. 

STIPE 60-10 × 15-25 mm, short or slim, cylindrical or 

attenuated at base, often eccentric, full, firm smooth, 

white, with dirty cream or brown-ochre-ish stains from 

the base. 

FLESH thick, firm, white to gill edge, but soon turning 

cream, with a very acrid flavour and no significant odour. 

LATEX: uniformly white if isolated, but lightly yellow 

olivish, drying on the gills, with very peppery flavour. 

Lactarius piperatus (Scop. : Fr.) S.F. Gray 

MICROSCOPY: spores from subglobose to oblong, 7.2- 

10.4 × 5.2-7.5 µm, with warts joined in ridges and 

forming un incomplete lattice; bisporic or tetrasporic 

basidia, subclavate ; Pileipellis, epithelium. 

HABITAT: in hardwood and coniferous woods, very 

common, grows in groups, often early; summerautumn. 


NOTE - L. piperatus can be told apart from L. 

pergamenus by the negative reaction of its latex with 

potassium hydroxide and by its gills and flesh which do 

not assume grey-greenish stains. L. vellereus and similar 

species are usually more robust and stubby, and have 

spaced out gills; L. controversus, a symbiont of poplar 

trees, has decidedly pinkish gills. 

183

CAP 40-80 mm, from convex to flat-depressed, firm 

fleshy, with the edges convoluted at the extremes, then 

rounded, undulated-lobed, finely felted. Cuticle lightly 

viscous in humid weather, then dry, pruinose, slightly 

zoned at most, light orange, orange-ochre-ish, orangegreying, 

finally with non uniform greenish stains. 

GILLS crowded, thin, from adnate to lightly decurrent, 

forked, lilac-pinkish, grey-lilac-orange, grey- vinous red, 

red-brownish in fractures then stained dark green. 

STIPE 20-40 × 10-20 mm, firm filled, soon becoming 

hollow, cylindrical or a little attenuated at the base, 

pruinose, white-greying at the summit, pink-violet, pinkgreyish, 

grey-violet in the lower part, smooth or with 

small darker dimples 

FLESH firm whitish, then brick pink, red-brick. Fruity 

odour, mild then lightly bitter flavour. 

184 

Lactarius sanguifluus (Paulet) Fr. 

LATEX slightly abundant, red-vinous, uniform, mild then 

bitter flavour. 

MICROSCOPY: spores 7-9 × 6-7 µm, mostly ellipsoidal, 

crested-reticular, with little mesh completely closed. 

HABITAT: thermophilic species, grows exclusively 

under Pinus. Summer-autumn. 


NOTE - A thermophilic species which is fairly common 

to maritime pines. It can be recognised by its orangetoned 

cap and its lilac-pink gills. It is confusable with 

Lactarius vinosus Quél., which, however, has a cap with 

patches of reddish-violet tones, and gills which are 

initially winey-red to violet, often with almost total 

greening.

CAP 40-110, initially convex then flattened, finally 

depressed, more or less umbillicate, viscous if humid, dry 

and rough in dry weather due to fine, short innate fibrils, 

from flesh-pink to pink-orange-reddish, with dark pinkreddish 

concentric zoning, the edges are convoluted, 

bearing tangled woolly hairs. 

GILLS adnate-decurrent, crowded, close, forked, cream 

pinkish. 

STIPE 25-50 (80) × 10-20 mm, full, then hollow, whitishcream, 

often with a pink labrum at tip, often with some 

pinkish dimples. 

FLESH thick, hard whitish, sometimes with flesh colour 

stains, with fruity or pelargonium odour and very acrid 

flavour. 

LATEX white, uniform, but slowly yellowing on a tissue 

or sheet of white paper. 

MICROSCOPY: spores from mostly ellipsoidal to 

ellipsoidal, 8.3-9.8 × 6.2-7.5 µm, with warts connecting to 

form several closed meshes; tetrasporophyte basidia, 

subclavate; pileipellis, an ixocutis is present. 

Lactarius torminosus (Schaeff. : Fr) S.F. Gray 

HABITAT: in hardwood areas near birch; fruits from the 

end of summer to the end of autumn; fairly common. 

NOTE - This can be distinguished from similar species by 

its white, uniform latex, its fairly red-pinkish and hairyfelt 

cap, and by its symbiosis with Betula. L. Pubescens is 

a smaller and nearly white or white-pinkish replica. L. 

tesquorum and L. mairei, have a more yellowish colour, 

and tend to grow in more southern climes and are not 

linked to birch trees. Even if some guidebooks, a little 

recklessly perhaps, nominate this as an edible species, 

which it may be after a long cooking time, it is certainly 

poisonous, and causes gastrointestinal distress. This 

characteristic, which is rapidly discovered by those 

unfortunate enough to confuse it with the saffron milkcaps 

(edible milk-caps with a red or carrot-orange latex), 

is the origin of its well-deserved Latin name, meaning 

“causing colic”. 

185

CAP 50-120 mm, initially convex, then flat, finally 

depressed, from yellow-reddish to tawny-orange, with the 

centre darker. Adnate, dry, matt, pruinose surface 

initially, from smooth to lightly rugulous, velvety 

convoluted edges, then distended often cracked radially 

when dry. 

GILLS from adnate to a little decurrent, crowded, forked 

to stipe, cream, pale yellow-ochre colour, stained dark 

brown in fractures. 

STIPE 30-80 (100) × 15-30 mm, firm, full then filled, 

pruinose, cylindrical or a little attenuated at base, 

concolour with cap, but lighter at tip, browning in the 

manipulated points. 

FLESH thick, firm compact, whitish-cream at gill edge, 

browning in cap, with an odour characteristic of fish 

(herring). 

LATEX: very abundant, dense, white, brownish drying 

when exposed to air, with a sweet flavour. 

186 

Lactarius volemus (Fr. : Fr.) Fries 

MICROSCOPY: spores from globose to subglobose, 9.2- 

11.2 × 8.4-10.6 µm, with clearly joint warts in ridges 

forming an almost complete lattice; bisporic or tetrasporic 

basidia, subclavate ; pileipellis, epithelium. 

HABITAT: mostly hardwood areas, more rarely in 

coniferous areas; not widespread, faithful to its growing 

areas; not very common. 


NOTE - Apart from its dry cap, L. volemus can be 

identified by its flesh, which has a greenish reaction to 

ferrous sulphate solution, its voluminous latex which 

seeps from any cracks in its skin, and its characteristic 

odour of herrings (very similar to that evinced by the 

Russula in the amoena group). It is edible, though 

appreciated by all, and is best cooked on the grill so as to 

preserve a vaguely smoky aftertaste.

Langermannia gigantea (Batsch : Pers.) Rostkovius 

BASIDIOCARP prosurface is globular, to irregularly 

roundish, with a circumference ranging 50 to 600 mm 

and, in exceptional cases, can reach even more 

remarkable dimensions. Sessile, with sterile radiciform 

base. 

EXOPERIDIUM formed from a single layer to a white 

and velvety, smooth then yellowish-cream bark; at 

maturity tears into irregular strips, fairly coarse, leaving 

the fine endoperidium free, whitish, then greying-sooty or 

grey-brown-ochre-ish, of papyrus consistency, friable, 

gradually dehiscent due to erosion, starting from the 

summit. 

GLEBE white, firm and compact when young, with 

weak, fungal odour and pleasant flavour, then soggy, 

from yellow-ochre to brown-olivish, powdery in maturity. 

MICROSCOPY: spores spherical or mostly ellipsoidal, 

3.6-5.6 µm, with short, finely warty peduncle. Brownish 

spores. 

HABITAT: in grassy areas, in pastures, grown in parks 

and gardens, from the end of summer throughout autumn. 

Solitary, gregarious. Infrequent. 


NOTE - This mushroom is spectacular for the sizes it can 

attain and there are reports of specimens over a metre in 

diameter and which have achieved the considerable 

weight of 20-25 kg. Species in the genus Calvatia, which 

have an exoperidium in two layers, can also achieve 

considerable size, among them, C. lilacina, which is 

coloured pale brownish-purple, prefers cultivated soil or 

footpaths, while C. utriformis, a white species whose 

exoperidium is ornamented with pyramidal warts, tends to 

grow in alpine pastures. 

187

188 

Leccinum aurantiacum (Bulliard) S.F. Gray ss. Pilát 

CAP 40-120 (150) mm, subglobular, then tightly 

parabolic, margin finally distended regularly, visibly 

hanging from excess of cuticle; finely velvety-felted, 

sometimes with adnate scales, just greasy in humid 

weather; uniformly red, red-orange, red-tawny, 

decolouring to brick or to yellow orange at maturity, dry. 

TUBES rounded-depressed or almost free to stipe, tall, 

even beyond 30 mm; grey-whitish, then grey, grey 

greenish dirty due to the maturation of spores, dark grey 

at gill edge, then through a temporary violet colour. Very 

small, round pores,, concolour with tubes; staining grey 

brownish to the touch. 

STIPE 60-130 (150) × 15-30 (45) mm, progressively 

attenuated towards tip; whitish, thickly covered with 

scales becoming more crowded and coarse towards the 

base, initially white, then becoming darker brownreddish,, 

almost black in old age or to the touch; basal 

area is often stained with a green-blue colour. Quickly 

becoming more woody-fibrous in consistency, full. 

FLESH fairly firm in cap, very fibrous in stipe; whitish, 

slowly turning grey-lilac, then dirty violet, finally orange 

subtle, insignificant odour and flavourful. 

MICROSCOPY: fusiform spores 13.5-16 × 3.8-5 µm, 

light brown under microscope; grey brownish olive in 

mass. 

HABITAT: in humid woods, associated with Populus 

tremula, in summer-autumn, recurrent. 


NOTE - This is the most common of the ”porcinelli 

rossi”, (“little red porcines”). It is very easy, however, to 

confuse it with the other red Leccinums: L. quercinum, 

which is bulkier, has more precociously coloured reddish 

scales, and associates with other hardwoods than Populus; 

L. vulpinum and L. piceinum are symbionts of conifers. 

L. versipelle is orange and is a symbiont of Betula. In our 

own gathering fieldwork we often had the impression that 

the correlations between carpophore characteristics as 

described in literature and symbiosis with host plants does 

not always follow precise rules, but rather preferential 

tendencies.

CAP 50-125 mm, initially convex, then obtusely conical 

and finally flat, sometimes with large umbo; convoluted 

margin, sometimes a little affected, curving towards the 

base at the extremes, distended sinuated-undulated late 

on; smooth, matt surface, slightly greasy in humid 

weather; violet, lilac, blueish colour, tendency to become 

brown tawny mostly towards the centre of the cap, 

decolouring to ochre-ish-violet or ochre-ish-pinkish with 

age. 

GILLS adnate-uncinate or just decurrent, crowded and 

with numerous wide lamellule; grey-lilac, lilac blueish 

colour, then with brownish tones. 

STIPE 50 –90 × 15-30 mm, cylindrical or widening 

towards the base, clavate, sometimes bulbous; base is rich 

with mycelial residues which typically encompass the 

substrate when the carpophore is picked; fibrous and 

elastic; lilac, lilac-grey-violet, violet colour, covered with 

a white fluff especially at the tip. 

FLESH firm, soon becoming soft and a little watery; light 

grey colour, with violet tones; odour is characteristically 

strong, aromatic, hot and indefinable. 

Lepista nuda (Bull. : Fr.) Cooke 

MICROSCOPY: ellipsoidal, lightly warty spores, 

dimensions 7-8.5 × 3.5-5 µm; salmon pink in mass. 

HABITAT: ubiquitous, on the substrate humus of plants, 

in large groups, often in lines or circles; from autumn to 

winter, often even in spring. 


NOTE - This is a sought-after edible with a delicate 

aroma. It can be confused with L. sordida, which is 

slightly thinner and usually has a more intense and “dirty” 

colour, bordering on violet-dark grey. A L. nuda var. 

lilacea has been described, bearing intense violet 

colouring all over, with a slightly smaller body. L. 

glaucocana has a much paler cap, tending toward lilacgrey, 

and has a weaker and less-pleasant odour. L. 

personata is very similar to L. nuda, but combines a 

greying, white-coffee coloured cap with a lovely violet 

stipe. Several other mushrooms can display a general 

blue-violet colouration, (e.g.: Cortinarius violaceus and 

Entoloma bloxamii) but also have varying generic 

characteristics; (see their respective genus descriptions). 

189

BASIDIOCARP high 30-80 mm, 20-60 mm in diameter, 

subglobular, obvoid or piriform. Exoperidium formed 

from dense converging spines and composites, width of 3- 

6 mm, initially white, then dark brown, which can be 

removed fairly easily leaving a reticulated endoperidium. 

Endoperidium is, light brown papyrus. 

GLEBE from grey-brown violet to chocolate brown, with 

an indistinct pseudocolumella. Areolated subglebe, with 

various brown and lilac stains. 

MICROSCOPY: globose spores, 4.0-5.0 µm, warty. Light 

brown, elastic scalp, with small rounded pores, fairly 

numerous. Para-capitulum is absent. Exoperidium has 

large irregularly shaped thick walled spherocytes, dark 

190 

Lycoperdon echinatum Pers. : Pers. 

brown. Spore powder is chocolate brown or with lilac 

tones. 

HABITAT: solitary or in small groups, in calcerous soil, 

mainly in beech woods. 


NOTE - This is a very-easily recognisable species thanks 

to the entirety of its morphological features. When found 

in forests of chestnut trees it is even likely to be taken for 

a chestnut shell! Lycoperdon foetidum also possesses a 

brownish exoperidium and a reticulated endoperidium 

after its spines have dropped, but these are significantly 

shorter; L. umbrinum and L. molle too display a brownish 

exoperidium but their endoperidia are smooth.

BASIDIOCARP height from 30-90 mm, 20-40 mm in 

diameter, subglobose, piriform, subcylindrical or almost 

pestle shaped, white, from cream to light brown. 

Exoperidium is formed from conical spines, length of 1-2 

mm, fragile, whitish or cream, then light brown, 

surrounded with a circular row of more persistent spines. 

When the spines fall, they move apart to form a 

characteristic polygonal grid pattern on the endoperidium. 

Endoperidium is, grey-brown papyrus. 

GLEBE white, then brown or olivish brown, with a well 

developed pseudocolumella. Subglebe is strongly 

developed, cellular, from olivish brown to grey-brown. 

MICROSCOPY: globose spores, 3.5-4.5 µm, warty. Scalp 

is yellow-brown, formed from hyphae sized 3-8 µm, with 

relatively thin walls; fairly numerous pores. 

Paracapitulum, usually abundant. Exoperidium has fine 

Lycoperdon perlatum Pers. : Pers. 

[= L. gemmatum Batsch] 

walled spherocytes of 20-30 µm,. Spore powder is 

yellow-brown, olivish brown or grey-brown. 

HABITAT: ubiquitous, in hardwood and coniferous areas. 


NOTE - On a macroscopic level, L. perlatum can be 

identified by the shape of its basidiocarp and by the 

characteristics of its exoperidium, and microscopically, by 

its small spores. Occasionally it can be found on dead 

wood but confusion with Lycoperdon piriforme is highly 

unlikely. This latter displays, in fact, white glebae even at 

maturity and after shedding its exoperidium, its 

endoperidium has a smooth appearance. L. nigrescens has 

darker and more-persistent spines plus larger and lessornamented 

spores. 

191

BASIDIOCARP height 15-60 mm, piriform, clavate, 

rarely subglobose, connected by thick white rhizoids. 

Dehiscence comes through a fairly wide, open operculum. 

Exoperidium is white, then fairly dark brown, wartygranular, 

with adpressed scales, soon becoming hairless at 

tip. Endoperidium is papyrus colour and matt. 

GLEBE with distinctive pseudocolumella, white then 

olivish, finally grey-brown, with a strong unpleasant 

odour and sweet flavour. Subglebe is firm and areolate 

which remains white. 

MICROSCOPY: globose spores, 3.5-4.0 µm, almost 

smooth. Capitulum is brown, elastic, not pored, largo 3- 

192 

Lycoperdon pyriforme Schaeff. : Pers. 

7.5 µm, with thick walls 0.7-1.0 µm. Paracapitulum is 

abundant. Exoperidium has large spinyspherocytes, walls 

thick, irregular shape. Spore powder is olive brown. 

HABITAT: in large groups on decaying wood, often on 

burnt wood, in woods, parks and gardens. 


NOTE – Very easy to recognise, by its lignicole habits, 

floury, inconsistent exoperidium, its cespitose growth, 

and its uniform, white subgleba.

CAP 50-70 mm, hemispheric, convex, then flat, often 

with large obtuse umbo, the margin is often lobed and 

undulated, fairly convoluted, then distended. Cuticle is 

smooth or fibrillated radially, shiny, lardaceous to the 

touch in humid weather, brown-grey, dark brown-ochre, 

with the edge lighter, often almost whitish. 

GILLS adnate or uncinate, fairly crowded, relatively 

close, from whitish to cream, often stained pink. 

STIPE 30-120 × 8-20 mm, from cylindrical to clavate, 

full, elastic-fibrous, whitish, fibrillated with the tip 

covered with a white bloom. 

FLESH elastic, whitish, with fungal odour and sweet 

flavour. 

MICROSCOPY: spores 5.5-7 × 5.0-6.5 µm, subglobose, 

smooth, hyaline; tetrasporophyte basidia, clavate, with 

siderophile granulation; epicyte formed from fairly 

Lyophyllum decastes (Fr. : Fr.) Sing. 

[= Lyophyllum aggregatum (Schaeffer) Kühner] 

parallel, braided hyphae, with pigment partially brownish, 

hinged. 

HABITAT: grows generally collated and in groups or in 

circles in hardwood and conifer woods, especially in open 

areas, at the edges of paths, in parks or in gardens among 

grass. 


NOTE - The species in the section Difformia with 

tricholomatoid silhouette, brownish-grey colours and 

globose spores, constitute a fairly homogenous group of 

species, sometimes difficult to categorise. When the 

carpophores are connate at the base, but not branching, 

and their growth is not cespitose then we are looking at 

either L. decastes or L. loricatum. The former has a finely 

fibrillated, relatively thin cuticle, the while the latter has 

a thick, tenacious, glabrous cuticle. 

193

CAP 100-250 mm, initially spherical, ovoid, then 

hemispheric-campanulate, finally flat, with large obtuse 

umbo, the cuticle is decorated with concentric plical 

scales, hazelnut ochre-ish in colour, often fairly brownish 

or reddish, on a light background, the margin is large and 

fringed. 

GILLS free, crowded, white then ochre. 

STIPE up to 200-350 (500) × 10-20 mm, slim, 

cylindrical, with base dilated to form a clear bulb, filled, 

then hollow, bearing mottled brownish bands which leave 

hints of the cream flesh below, subsmooth above the 

annulus. Double annulus, mobile, whitish externally, with 

the lower leaf brownish. 

FLESH white, uniform, with pleasant fungal odour,, and 

hazelnut flavour. 

194 

Macrolepiota procera (Scop. : Fr.) Singer 

MICROSCOPY: ellipsoidal smooth, hyaline spores, with 

germinating pores, 12.5-17.8 × 8.5-11 µm. clavate, 

tetrasporic basidia. Polycystidea is absent. Cheilocystidia 

clavate. Epicyte formed from trichoderma. Partially 

dominant pigment. Rarely joint at hinges. 

HABITAT: isolated or gregarious in hardwood and 

conifer woods or in meadows; very widespread and 

common, from summer throughout autumn. 


NOTE - By its antonomasia, this is the “drumstick”, a 

high-quality, much-sought edible. It is easy to identify, 

just look for its large size and outline which explain its 

German name, meaning “parasol”, its stripy stipe and its 

characteristic, movable double annulus.

CAP 80-150 mm, initially campanulate, then convex, 

finally flat, the cuticle is excoriated up to the 

circumference with wide, coarse, crowded, overlapping 

scales, fairly light brownish in colour, with a fairly tight 

central cap, of colour brown reddish. 

GILLS free, whitish, cream, then dirty pink, reddening 

when rubbed, with a floccose surface. 

STIPE 100-160 × 10-15 mm, stocky, cylindrical, 

progressively dilated to form a bulbous submarginated 

bulb, hollow, smooth, white, gradually brown-reddish 

with time or when rubbed. Annulus is membranous, 

robust, whitish, mobile. 

FLESH white, turning orange vivid at gill edge, then to 

red-vinous, with an odour of raw potato and sweet 

hazelnut flavour. 

MICROSCOPY: ellipsoidal or ovoid, smooth, hyaline, 

with germinating pores, 8.8-11.2 × 6.8-8.0 µm. clavate, 

tetrasporic basidia. Polycystidea is absent. Cheilocystidia 

clavate or piriform. Epicyte formed from a trichoderma of 

Macrolepiota rhacodes (Vittadini) Singer 

fairly erect hyphae. Membranous brown pigment. Joint at 

hinges. 

HABITAT: isolated or gregarious, in parks, gardens, and 

also in the woods, mostly of mixed conifers. from the end 

of summer to autumn, recurrent. 


NOTE - M. rhacodes is characterised by its peculiar pileic 

scaling, which is formed of tortoiseshell scales, 

concolour with the cap background, and above all for its 

reddening flesh. The variety, bohemica differs by having 

scales of a different colour to those of its background and 

for its habitat on the periphery of woodlands. M. 

venenata, heavily toxic, can be distinguished by the radial 

arrangement of its pileic scales and by the total absence 

of joint hinges, a characteristic which is incredibly 

difficult to verify. Therefore, in case of doubt, one would 

be best advised to avoid consuming any Macrolepiota 

with reddening flesh. 

195

CAP 20-50 mm in diameter, initially hemispheric, 

campanulate, then flat, umbonate in the centre, the margin 

is acute, smooth, often lightly crenulated, the surface is 

smooth, hygrophanous, from orange-ochre to brownish 

with humid weather, but light cream-hazelnut when dry, 

spaced, interspersed from lamellule, sinuous, wide, from 

whitish to cream. 

STIPE 30-70 (100) × 3-5 mm, full, cylindrical, often a 

little widened at the two extremities, slim, tenaciouselastic, 

whitish dirty-cream at tip, brownish on the lower 

part, from finely pruinose to velvety for the whole length, 

with brownish mycelium. 

FLESH whitish, elastic, fine, hygrophanous, with a 

characteristic odour, like almonds, and sweet hazelnut 

flavour. 

196 

Marasmius oreades (Bolt. : Fr.) Fries 

MICROSCOPY: spores from ellipsoidal, fairly elongated, 

to amygdalform, (7.0) 8.0-10.5 (11.5) µm; 

tetrasporophyte basidia, tightly clavate; pileipellis 

hymeniform. Hinges present. 

HABITAT: in meadows, in large groups found in lines or 

circles; from the spring throughout autumn. 


NOTE - This, fairly common mushroom, grows 

abundantly, from the spring to autumn, in meadows, 

forming “witches’ circles”. It is a choice edible, much 

sought after by collectors and can be stored and eaten dry. 

M. collinus (regarding whose edibility there are some 

doubts), is very similar, though can be told apart by its 

slender, smooth stipe and crowded gills.

BASIDIOCARP pileate, devoid of or with rudimentary 

stipe, reduced to a point of attachment on the cap. 

CAP of width up to 300 mm, depth of 10-20 mm and 

projection of around 100-150 mm; irregularly circular 

shape to gill edge, with several individuals around a 

common base. Surface plicea is undulated, brown or 

brown-reddish, markedly zoned, rough due to the 

presence of adpressed flakes. undulated margin, complete. 

HYMENOPHORE tubes and pores, follow the trend of 

the undulated surface plicea. Tubes about 15 in thickness 

mm, monolayer, from whitish to dark ochre-ish, 

blackening if handled. pores 0.2-0.3 mm in diameter, 

round. 

FLESH fibrous and tenacious, but not hard (in the context 

of a monomorphic constitution), cream white, with fungal 

odour and sweetish flavour. 

MICROSCOPY: spores subglobose, in the shape of 

chestnuts, monoguttulate, not amyloid, smooth and 

hyaline, dimensions 5.5-6.5 × 4.5-5.5 µm; basidia 20-25 

(45) × 5-10 µm, cylindrical-claviform, tetrasporic, 

without joints at basal hinge; cystidia absent. 

Meripilus giganteus (Pers. : Fr.) P. Karsten 

[= Polyporus giganteus Pers. : Fr.] 

HABITAT: saprophyte or parasitic to conifer or Fagus 

trees, in thick groups with specimens overlapping, from 

summer to autumn; not widespread. 

NOTE - At first sight this is liable to be confused with 

Polyporus squamosus, as it grows exclusively on 

hardwood trunks, the latter, however, has a very scaly cap 

surface and larger, angular pores; it is also marked out by 

its strong odour of cucumber, especially in the younger 

specimens. 

The patchy appearance of the cap surface Meripilus 

giganteus is common also to several species of Trametes, 

as is their leathery consistency due to its trimeric hyphal 

structure; of these the ones which spring to mind are, in 

particular, T. zonatella and T. versicolor, the former 

having a creamy ochre-ish-brown colour, the latter being 

instead grey-brown, or with decidedly blueish tones. Both 

produce cylindrical-allantois spores and prefer to live on 

broken hardwood substrates. 

197

ASCOCARP pileate and stipitate, of 120-130 mm in 

height and 40-45 mm in diameter. 

MITRA elongated-conical, with slightly pointed tip, with 

alveoli clearly separated from intersection of lengthwise 

and transversal ribs arranged parallel to them. 

HYMENOPHORE found on all exposed parts of the 

mitra, smooth, brick-brown colour, which blackens on the 

ribs with age. Edge is regular, separated from the stipe by 

a vallecula. 

STIPE subcylindrical, sometimes swollen at base, rarely 

furrowed, rough, white suffused with pinkish tones, 

hollow or filled internally. 

FLESH leathery-elastic, whitish, with light spermatic 

odour and sweetish flavour. 

198 

Morchella conica Persoon var. costata Ventenat 

MICROSCOPY: ellipsoidal spores, 19-25 × 12-13 µm, 

smooth, with some small guttules arranged on the external 

surface of the polar zone, uniseriate in asco; asci are 

cylindrical, not amyloid, octasporic; slightly cylindrical 

paraphyses extended at tip, septate and sometimes forked. 

HABITAT: single or in small groups in conifer woods, in 

spring; not common. 


NOTE - M. conica var. costata can be distinguished from 

similar species by its characteristic darkened parallel ribs, 

and also the pinkish colouration of its stipe. Its habitat and 

microscopic features are not particularly useful in its 

identification.

Morchella esculenta (Linnaeus) Persoon var. esculenta 

ASCOCARP pileate and stipitate, up to 100 to 250 mm in 

height and 50-80 mm in diameter. 

MITRA irregularly ovoid, subspherical, globular, with 

longitudinal and transversal ribs in relief, variably carved 

between them forming irregularly polygonal alveoli. 

Hymenophore is smooth, from yellow cream to light 

ochre; edges of the ribs are coloured. insertion to stipe is 

without vallecula. 

STIPE widened at base, sometimes semibulbous, often 

partially ruffled, irregularly cylindrical, cream ochre-ish, 

granular, internally cavernous. 

FLESH elastic, ochre, with light spermatic odour. 

MICROSCOPY: smooth ellipsoidal spores, 18-25 × 14- 

16 µm, hyaline under microscope, thick walls, with some 

small guttules on the extreme surfaces, monoseriat in 

asco; asci are cylindrical, not amyloid, octasporic; 

cylindrical clavate paraphyses at tip, septate, simple or 

branched. 

HABITAT: single or in small groups on mouldy, sandy 

ground to sandy or sandy-clay ground in fresh hardwood 

areas, under Fraxinus, Ulmus, Alnus, in humid places. 

Fairly common, in spring. 


NOTE - There are some other varieties of M. esculenta, 

which can be distinguished by their morphological 

features; var. rigid which has a visibly more globular 

mitra and is larger (up to 300 mm in height); var. vulgaris 

which has a conic-elongated mitra, (sometimes) smaller 

dimensions (from a minimum of 50 mm in height); var. 

rotunda which has an oval, subglobular mitra, which, in 

size, can match var. rigid (which is difficult to distinguish 

from the latter). Morchella crassipes is very similar to M. 

esculenta (of which it may be just a giant variety), has a 

conical-rounded ochre-greyish mitra, clavate stipe which 

can be as large as 70 mm in diameter; Morchella 

esculenta var. umbrina, instead, diverges from the usual 

appearance by the brown colour of its mitra, which, 

however has rib edges which are light ochre-ish. Grows 

close to hardwoods such as Fagus or Fraxinus. 

199

200 

Morchella esculenta (L.) Persoon var. vulgaris Persoon 

ASCOCARP pileate and stipitate, from 80 to 150 mm of 

height and 50-60 mm of diameter. 

MITRA ovoid-conica, with rounded or subpointed apex, 

with irregular, polygonal or roundish alveoli, deep. 

HYMENOPHORE found on the entire visible part of the 

mitra, smooth, light brown, whitish on the ribs. Edge is 

regular, separated from the stipe by a vallecula. 

STIPE subcylindrical, but swollen at base where it is 

often deeply ribbed or furrowed, rough, white-ochre-ish, 

filled internally. 

FLESH elastic, whitish, with light spermatic odour and 

sweet flavour. 

MICROSCOPY: ellipsoidal spores, 19-25 × 12-13 µm, 

smooth, with some small guttules arranged on the external 

surface of the polar zone, uniseriat in asco; asci are 

cylindrical, not amyloid, octasporic; paraphyses lightly 

cylindrical elongated in height. 

HABITAT: single or in small groups near hardwood 

trees, especially Ulmus, in spring; not common. 


NOTE - This is confusable with other species of the same 

genus, both for its colour and appearance, which are not 

infrequently influenced by its growth conditions.. One of 

its peculiar characteristics is the colour of its 

hymenophore which is light brown in contrast to its white 

ribs.

ASCOCARP stipitate and pileate, up to 200 mm in height 

and around 30 in diameter. 

MITRA conical-pointed, often rounded, sub-cerebriform, 

with longitudinal and transversal ribs slightly in relief 

forming the irregular alveoli. Hymenophore is smooth, of 

brown-brick colour, darker on the edges of the ribs. 

Lower surface is smooth or minutely rough, white cream. 

STIPE slim, subcylindrical, often ribbed lengthwise, 

rough, white cream. 

FLESH tenacious, elastic, but to con fragile, sweet, with 

spermatic odour; white cream colour. 

HABITAT: in humid places, on sandy or clay-sandy 

ground, under trees such as Alnus, Crataegus, Fraxinus 

and Populus, but also in vineyards, more rarely under 

other hardwood; in spring, in groups of several 

specimens, recurrent. 

MICROSCOPY: ellipsoidal spores, smooth, 20-25 × 14- 

17.5 µm, hyaline under microscope, often with small 

Morchella semilibera De Cand. : Fr. 

guttules on the external surface at the ends, uniseriat in 

asco; asci are cylindrical, not amyloid, octasporic; 

cylindrical paraphyses with widened and often guttural 

apex, septate. 


NOTE - This has in the past been considered as part of 

another genus, the genus Mitrophora, distinct from the 

Morchella by the way the stipe was inserted into the 

mitra. In the Mitrophora the vallecula is lengthened, and 

joins onto the lower surface at around 1/3-2/3 of the 

length of the mitra itself. Today this difference is not held 

to be sufficient to warrant maintaining two diverse 

taxonomic genera. 

M. fusca, which is smaller (up to 80 mm in height) (and 

has spores just 8-9 µm wide) while M. gigas can reach 

heights of 200 mm, with a clavate stipe which measures 

up to 50 mm at its wide base. 

201

CAP 20-50 mm, from fairly campanulate to flat-convex, 

with or without an obtuse umbo, hygrophanous, striated to 

transparency, hairless; pale pink with light lilac stains. 

GILLS 25-30, from ascending to fairly horizontal, from 

adnate to non-marginated, slight crowded, bulging, from 

smooth to venous, whitish, suffused with very light violet, 

concolour with the surface. 

STIPE 40-70 × 3-8 mm, cylindrical, a little widened at 

base, smooth, hollow, tenacious, pruinose at tip, dirty 

whitish, suffused with violet, lighter at tip, with base 

fairly densely covered with whitish fibrils. 

FLESH fine, whitish, with raphanoid odour and flavour. 

MICROSCOPY: ellipsoidal spores, amyloid, 7.0-9.5 × 

3.5-4.5 µm. Tetrasporophyte basidia, clavate, with hinges. 

Cheilocystidia is fusiform clavate, fairly elongated, 

generally smooth but covered with sparse diverticula at 

the edges of the cap. Pleurocystidia similar. Epicyte 

formed from filamentous smooth hyphae, very thin. 

202 

Mycena pura (Pers. : Fr.) Kummer 

Hyphae of stipe is smooth, with fusiform or ellipsoidal 

caulocystides, smooth. 

HABITAT: in hardwood and conifer woods, between 

fallen needles and leaves; very widespread and frequent at 

every latitude. From the end of spring. 


NOTE - Of all the species belonging to the genus 

Mycena, M. pura, with its different varieties and forms, is 

probably the one which has the widest range of cap 

colouring. Following the guidelines proposed by Maas 

Geesteranus, any taxon’s colour variations should be 

considered as mere forms rather than varieties. Among 

these we mention, for example, the completely white fo. 

alba, the blue-capped fo. ianthina, which has a hint of 

violet or grey, and the fo. lutea which can be identified by 

the yellow colour of its cap but also its pale stipe bearing 

just a hint of violet.

CAP 30-50 mm, from conical to campanulate, then 

convex or flat-convex, mamelon, pale pink, pink-lilac, 

sometimes vinous hints, with the edge lighter, 

hygrophanous, smooth, hairless, a little unctuous, striated 

to transparency. 

GILLS 32-36, wide, ascending, bulging, adnate or nonmarginated, 

fairly light pink, whitish towards the surface. 

STIPE 70-110 × 5-10 mm, cylindrical, hollow, smooth or 

covered with very fine fibrils, white, hint of pink. 

FLESH white pinkish, hygrophanous, with fairly clear 

raphanoid odour. 

MICROSCOPY: spores mostly ellipsoidal, smooth, 

hyaline, amyloid, 7.5-10.2 × 3.8-4.5 µm. basidia clavate 

tetrasporic, hinged. Cheilocystidia and pleurocystidia 

fusiform, clavate or spherical-pendunculated. Epicyte 

Mycena rosea (Bulliard) Gramberg 

formed from parallel interwoven hyphae, the surface layer 

is geleld. Hinges present. 

HABITAT: isolated or gregarious, on decomposing 

residues in hardwood and of conifer woods; non very 

common, fruits preferably in autumn. 


NOTE - M. rosea belongs to the subsection Purae (Konr. 

& Maubl.) Maas G. of section Calodontes (Fr. ex Berk.) 

Quélet and differs from M. pura, of which it is often 

considered a variety, not only by the colour of its cap, 

which is a particular shade of pink, but also by its shape, 

which is conical-campanulate at the extremes or 

parabolic, and by its more-delicate stipe. 

203

BASIDIOCARP initially content in a whitish peridium, 

globular, often and filled with a gelatinous substance, 

external surface is membranous, initially smooth, soon 

becoming percolated by elevated veins; the peridium has 

clear mycelial rootlets and the tip tears at maturity to 

allow the exit of a phallic receptacle, made from the stipe 

and cap. 

STIPE 120-210 × 20-35 mm, cylindrical, progressively 

tapered at tip, white, fragile and spongy, often curving, to 

surface thickly dimpled. 

CAP vaguely resembling a mitra, wider in diameter than 

stipe, honeycombed, shows a round opening at tip and 

margin; whitish colour but at the covered with a rotten 

green or olive green dark mucilage paste at extremes, 

which constitutes the glebe and contains its spores. It is 

not unusual for portions of gelatinous peridium to adorn 

the summit of the cap. After the dissolution of the 

greenish-whitish glebe, the cap eventually takes its 

cellular-ribbed form. 

FLESH fragile in cap, fragile and spongy in stipe. Odour 

at first, until the carpophore is closed within the peridium 

204 

Phallus impudicus L. : Pers. 

is not unpleasant, raphanoid, with a tendency to smell of 

flesh or faeces or with hints of gorgonzola cheese; very 

unpleasant. the odour is noticeable several feet away and 

clearly indicates the presence of the mushroom even 

before it is seen 

MICROSCOPY: ellipsoidal spores, smooth, brownish 

under microscope, dimensions: 4-5 × 1-2 µm. 

HABITAT: solitary or in groups in humid recesses dei 

woods, often hidden among the undergrowth and close to 

decomposing vegetable residues; not rare. 


NOTE - An unmistakeable mushroom with its odour 

which recalls fecal or strongly-rotten organic matter. An 

odour which attracts the flies which are necessary to the 

diffusion of its spores. P. hadrianii, which is rarer, is very 

similar but has a pink peridium, is slightly smaller and 

grows in sandy, arid earth. Mutinus caninus has a moreslanting 

stipe which is coloured orange-pink where it 

intersects with the cap, which is the same diameter as the 

stipe, and is pointed.

CAP: 50-100 (150) mm, convex, then more distended 

with convoluted margin at the extremes, finally flat, very 

eccentric, in the shape of a fan or a shell, thinned in the 

marginal zone which is sometimes a little striated; 

hairless, greasy-shiny, grey, grey steel blueish, greybrown, 

sometimes with fairly violet stains, cuticle is 

separable. 

GILLS: very decurrent, not very crowded, with forks 

multiple and to various levels, thin, unequal, white or 

white-silver on whole surface, sometimes pale whitecream. 

STIPE: 10-35 × 10-20 mm, sometimes nearly absent, very 

accentric or lateral, irregularly cylindrical, of pruinose 

appearance, white, or slight hint of grey; dry, firm, full. 

FLESH: white, abundant in correspondence with stipe 

insertion, soon becoming thinned, fairly elastic-tenacious, 

sub-leathery in adults, especially around the stipe; odour 

vaguely of mould in older specimens, sweet flavour. 

Pleurotus ostreatus (Jacq. : Fr.) Kummer 

MICROSCOPY: spores tightly ellipsoidal, subcylindrical, 

smooth, 8-11.6 × 3.2-4.2 µm. 

HABITAT: on living or dead wood of various hardwood 

trees, in woods or in parks, numerous examples are found 

with overlapping caps; late autumn or winter, common in 

growing areas, but not widespread. Easily cultivated 


NOTE - There is a variant of this which grows on conifer 

trees, the var. columbinus, which has a light blue tinge all 

over. This species is very well known for the ease with 

which they can be cultivated. Very commonly sold, it can 

be found on sale in all European and most international 

markets. It also used to be very readily found in the wild 

too, from meadowlands to the mountains, but it is 

becoming less and less easy to find them in their natural 

state. They are a choice edible and almost impossible to 

confuse with poisonous species. 

205

BASIDIOCARP coralloid, of 80-100 mm in width and 

50-120 mm of height, verticillated, made from a base up 

50 mm wide from which many small branches reach out, 

in various numbers. The principal branches create a 

pointed tip pink vinous in colour, which contrast with the 

rest of the branches which are white or whitish. 

HYMENOPHORE indistinct, found on the smooth 

surface of branches and roughly half their height. 

FLESH fairly compact but fragile (monomorphic hyphal 

structure), white, with pleasant odour and flavour. 

MICROSCOPY: spores cylindrical-ellipsoidal, 14-17 × 

4.5-8 µm, with fairly pronounced apiculture, irregularly 

furrowed lengthwise, not amyloid, hyaline under 

microscope; basidia 45-60 × 9-11 µm, cylindricalclaviform, 

tetrasporic, joint to hinges at base. Cystidia 

absent. 

HABITAT: on the ground in hardwood and of conifer 

woods, fruits in groups which sometimes cover a large 

area, in summer-autumn; not widespread. 


206 

Ramaria botrytis (Pers. : Fr.) Ricken 

NOTE - Ramaria botrytis has a winey pink colour which 

is commonly found in other species such as R. 

subbotrytis, in which, however one can see hues verging 

on corral pink; this latter has similar-looking carpophores, 

fruits in hardwood and conifer forests and has smaller, 

rough spores, from 8-11 × 3-4 µm. R. holorubella seems 

to be a variant found in conifer woods, and has a fairlywell 

rooted basal trunk which is firmly embedded in the 

ground. More common is R. rufescens, always whitereddish, 

with a tendency to yellow at the lower part of its 

branches. It has distinct basal trunk dimensions and can 

be found from summer to autumn in woodlands. It has 

amygdaliform spores, 8-10 × 3,5-4 µm. 

R. formosa has basidiocarps with an attenuated base with 

decreasing branches at the top; these tend to be of three 

colours: yellow on the uppermost extremities, salmon 

pink on the branches and white at the base. It has are 9-13 

× 5-6 µm, warty-crested spores, and lives on the ground 

in hardwood forests.

BASIDIOCARP initially in the shape of a cauliflower, 

then coralloid, width of 100-200 mm and height 100-150 

mm, fairly verticillated, formed from an irregularly 

cylindrical, solid and fairly developed basal trunk from 

which numerous branches grow, eventually fairly long, 

and divide further, until ending in two short spikes at the 

upper end. The branches are yellow or apricot colour 

while the trunk is whitish 

HYMENOPHORE not distinct, found on the surface on 

the upper half of the branches. 

FLESH compact (monomorphic hyphal structure), white, 

sometimes marbled, no particular odour and weak, 

slightly sour flavour. 

MICROSCOPY: spores cylindrical-ellipsoidal, 9-13 × 4- 

5.5 µm, with clear apiculture, with partially joined warts, 

not amyloid, hyaline under microscope; cylindricalclaviform, 

tetrasporic basidia, with basal joints at hinges. 

Cystidia absent. 

HABITAT: on the ground in hardwood or 

mixed woods, in summer-autumn; occasional. 

Ramaria flavescens (Schaeff.) Petersen 


NOTE - There are numerous yellow species in the genus 

Ramaria; some of these are fairly poisonous; among 

which R. formosa typically stands out due to three 

colours: white on its trunk, pink on its branches and 

yellow at the branch tips. Other species should be eaten 

with caution as they can have a laxative effect. R. flava, 

has least developed basal trunk and can also be identified 

by its large spores (10-15 × 4-6 µm) and the unpleasant 

odour of its flesh. R. aurea possesses a less-developed 

trunk, from which arise several columns, which in turn 

branch out into fairly-long, golden yellow appendages; it 

lives on the ground in woods of Fagus and has warty 

spores of the same shape, but smaller (9-11 × 3,5-5 µm). 

R. largentii too has several analagous features to R. 

flavescens but is a very vivid yellow-orange colour and 

possesses an unpleasant odour, similar to car tyres, and as 

such is usually considered inedible. 

207

BASIDIOCARP coralloid, about 150 mm in width and 

200 mm in height, usually appears robust, very branched, 

with wide base, usually wider than it is tall, very distinct, 

white or whitish, from which fairly thick, erect, 

cylindrical branches, vivid pink in colour when young, 

then, slowly becomes, pink-ochre-ish, salmon, ending 

with a lemon yellow pointed tip, finally concolour with 

branches. The bifurcation of the branches (saddles) very 

close to the structure of a U. 

HYMENOPHORE not distinct, found on the surface of 

the branches. 

FLESH fairly thick, white, soft, marbled with humidity, 

brittle when dry, often a little red-brownish at gill edge 

and when handled; odour is light or insignificant, slightly 

bitter, sour flavour. 

MICROSCOPY: spores cylindrical-ellipsoidal, 9.0-13.0 x 

5.0-6.0 μm, warty, hyaline under microscope; tightly 

clavate, tetrasporic basidia, with joints at hinges; 

monomorphic structure, made from septate hyphae, with 

joints at hinges. Spores are yellow. 

208 

Ramaria formosa (Pers. : Fr.) Quélet 

HABITAT: grows on the ground in hardwood areas, 

especially oak; summer-autumn, fairly common a little 

widespread. 


NOTE – This is one of the largest Ramaria (carpophores 

over 300 mm in height have been found!) and can be 

easily recognised by its three colours and its parallelshaped 

branches. It is toxic and provokes gastrointestinal 

disturbances including strong and continual diarrhoea. R. 

neoformosa has divergent branches, with prevalent 

bifurcations similar to Var; R. flavescens, with which it 

often shares the same habitat. Its colour makes it the 

closest species but it has divaricating, scattered branches, 

mixed, U- and V-shaped saddles and its branch tips are a 

corn-yellow colour; R. fagetorum has a longer, tighter 

base and prevalently V-shaped angles; R. cettoi can be 

told apart by the dark reddish colour of its branches and 

its sweet, pleasant odour and flavour.

BASIDIOCARP initially subglobular and with 

resembling a cauliflower due to the presence of branches, 

then coralloid, about 150-180 mm in width and in height, 

made from a solid, but not very developed basal trunk, 

white or whitish-yellow in colour, from which various 

columns grow which are then divided ending with one or 

two short points, yellow-orange, fire orange, sometimes 

scarlet red in colour. 

HYMENOPHORE not individualised, found on 

the upper half of the branches. 

FLESH compact (having a monomorphic hyphal 

structure), white, with a fairly pronounced odour of tyres 

or “dental surgery” and sweet-sour flavour 

MICROSCOPY: spores ellipsoidal or cylindricalellipsoidal, 

12-14.5 × 3.5-5 µm, with clear apiculture, 

warty, not amyloid, hyaline-yellow under microscope; 

claviform, tetrasporic basidia, with basal joint on hinges. 

Cystidia absent. 

Ramaria largentii Marr & Stuntz 

HABITAT: on the ground in conifer woods, in summerautumn; 

fairly common. 

EDIBILITY: not edible 

NOTE - Ramaria aurea is similar but forms smaller 

carpophores with a golden-yellow colour and has 

branches which spread out from a rather undeveloped 

base. It fruits on the ground in Fagus woods during the 

summer-autumn period and produces small spores 9-11 × 

3,5-5 µm, which are covered with partially-united warts. 

R. flava too has yellowish colouration and has not-very 

differentiated branches which develop from its basal 

trunk; it grows in conifer woods and has 10-15 × 4-6 µm 

spores which are mostly warty. Other species in this 

genus have similar colours; their morphological 

identification can mostly be based on the colour of their 

branches, but to be sure we must recourse to microscopic 

analysis, particularly of spore characteristics. 

209

BASIDIOCARP initially like a cauliflower, soon 

coralloid, up to 200 mm in width, height can reach more 

than 150 (200) mm, made from a basal trunk which is at 

most 40 mm in from which various branches grow and 

are subdivided again, ending in two short points. The 

trunk is whitish ivory towards the base, with ochre-ish 

shades elsewhere; branches are yellow or weakly 

yellowish, sometimes with flesh coloured stains, darker 

due to the maturation of the spores, fairly often bearing a 

sinuous pattern. 

HYMENOPHORE not distinct, distributed over the upper 

half of the branches. 

FLESH compact, then soft (in a monomorphic hyphal 

structure), white, with weak or insignificant odour of dry 

grass, and no distinct flavour. 

MICROSCOPY: spores irregularly ellipsoidal, sometimes 

flattened on one side, 9-12 × 4.5-5.5 µm, with partially 

joined warts, not amyloid, hyaline-yellow under 

microscope; claviform, tetrasporic basidia, without basal 

joints at hinges. Cystidia absent. 

210 

Ramaria pallida (Schaeff.) Ricken 

HABITAT: on the ground in conifer and mixed woods, in 

summer to autumn; common. 


NOTE - The light colour of R. pallida is similar to several 

Clavulina, such as C. rugosa, which has far-less 

developed carpophores, with no defined basal trunk and 

featuring thick, sometimes rough and flattened branches. 

Furthermore, it has very diverse globose, round and 

smooth spores 9-13.5 × 7.5-10 µm, and typically fruits on 

woodland floors. R. gracilis grows in conifer woods and 

has coralloid, white-ochre carpophores no larger than 60 

mm, which have a pinkish colour to them. Its spores, 

which are ellipsoid and finely verrucose, are noticeably 

smaller and 5-7 × 3-4 µm in size. With colours tending to 

grey-violet is R. fumigata, which produces arborescentcoralloid 

carpophores; fruits in hardwoods forests in 

summer and autumn and produces 9.5-11.5 × 4-5 µm 

spores covered with partially-united warts.

CAP 50-80 (100) mm, hemispheric or truncatedhemispheric, 

then convex, finally flat or flat depressed, 

with obtuse margin, slightly grooved in old age; initially 

of firm, hard, becoming more fragile during maturity of 

the carpophore. Surface appears greasy-shiny, viscous 

with humidity, fairly corrugated, from cinnabar red to fire 

red, vivid orange, often with extended sulphur yellow 

zone, sometimes completely yellow; fine cuticle, 

separable for 1/3 of the radius. 

GILLS rounded, subfree, fairly wide, very thin and 

fragile, fairly crowded; whitish, then pale cream flat 

appearance, finally also extensively yellow, typically with 

a yellow surface (these features are rarely evident and 

often completely absent). 

STIPE 35-80 × 12-25 mm, from cylindrical to 

subfusiform dry corrugated-rugulous, white, not 

infrequently evident and with fairly sulphuric yellow 

shades; full, soon becoming filled-medullar. 

FLESH fairly hard in young specimens, soon becoming 

fragile and brittle, almost friable in adults; white, 

Russula aurea (Persoon) 

sometimes yellow for a fairly extended subcuticular area, 

of flavourful and non distinctive odour. 

CHEMICAL REACTION: Guaiac quickly blue greenish. 

FeSO weak, pink pale ochre-ish. 

4 

MICROSCOPY: spore from mostly ellipsoidal to ovoid, 

7.5-9.0 × 6.2-7.5 µm, warty, partially reticulated. Light 

yellow in mass. 

HABITAT: ubiquitous and mostly widespread; present 

from summer and fruits until late autumn. 


NOTE - This species is easy to recognise when it shows 

its typical characteristics; its colouring, with a yellow 

surface surrounding its gills, and the fragility of adult 

specimens’ flesh; however, it should be noted that this is a 

very “capricious” species, which sometimes appears 

completely yellow, and sometimes does not have the 

yellow marking around the gills. Among edible species it 

is, with good reason, considered one of the best, 

notwithstanding the frail flesh and gills of adult 

specimens. 

211

CAP 50-140 mm, fleshy and compact, subglobular with a 

fairly pointed, gradually expanded summit, finally also 

depressed, margin is bent at the extremes, subacute; 

cuticle separable to half of the radius, lubricated-shiny, 

even greasy in humid weather, from violet lilac to violet 

blueish, fairly variegated of green, grey-green, sometimes 

completely cyclamen pink (fo. lilacea), completely green 

olive or pear green (fo. peltereaui), eventually dimpled 

towards the edge (fo. cutefracta). 

GILLS bent-adnate or a little decurrent, sometimes 

biforked at insertion, close, fairly crowded, lardaceous, 

whitish, sometimes fairly clear cream, eventually also 

stained brown ochre. 

STIPE 30-90 × 15-35 mm, cylindrical, rugulous, 

completely white or suffused with lilac pink, vaguely 

greying due to imbibition, eventually a little stained of 

brown, filled with a compact medulla, then cavernousfilled. 

FLESH highly compact, almost hard normally lilac under 

the cuticle, white elsewhere; mild flavour, slightly 

sensitive odour, in mature specimens, after rubbing, it is 

212 

Russula cyanoxantha (Schaeffer) Fries 

possible to experience an unpleasant metallic note, like 

FeSO . 

4 

CHEMICAL REACTION: Guaiac strong and rapid. 

Aniline late on, orange on the gills. FeSO negative, then 

4 

slowly grey-green. 

MICROSCOPY: spores ellipsoidal-oboval or reniform in 

parts, (6.4) 7.2-9 (9.5) × 5.8-7 µm, warty, hemispheric 

0.4-0.5 µm, isolated or grouped in part from slightly 

amyloid connections. Spores are pure white. 

HABITAT: ubiquitous species, common from the start of 

the season under hardwood and conifer, from the 

mediterranean up to limited tree vegetation. 


NOTE - According to Bon, the types with cream, elastic 

instead of waxy gills, a dark violet cap and stipe usually 

featuring a lilac-pink colouring, belong to an independent 

species named R. langei. Specimens with a subuniform 

violet-lilac cap (fo. lilacea) can seem similar to R. grisea, 

differentiated only by the cream spores, fragile gills, spicy 

taste and red-orange reaction to FeSO 4 .

CAP 60-140 (185) mm, fleshy and firm, hemispheric, 

then convex, with obtuse umbilical dimple in the 

background, slowly expanding, eventually deeply 

infundibulform; margin bent at the extremes, thinned; 

semiadnate dry and matt cuticle, thorny or a little felted, 

covered with residues of soil and leaves difficult to 

separate, whitish at first, soon becoming stained ochre 

brown, then of rusty brown, starting with the most 

exposed parts. 

GILLS a little decurrent on stipe, falciform-arcuate, 

subacute at the front, forked in parts, width of 6-9 mm, 

intercalated from lamellule of diverse lengths, fairly 

spaced, fragile although rigid, cream, with the whole 

surface coloured, soon becoming stained rusty brown on 

the most exposed parts. 

STIPE 25-48 × 15-35 mm, hard, highly stocky and robust, 

flared at the top, or cylindrical subequal, subsmooth or 

finely corrugated surface, pruinose, white, then stained 

rusty brown, rarely with glucose belts under the gills, the 

medulla is compact, only eventually a little springy and 

wormy. 

Russula delica Fries 

FLESH very thick, firm and fragile, white, clearly 

browning on the surface and washed with ochre tones to 

air after the gill edge; strong and unpleasant odour, like 

fish or salty, with fruity hints when young, peppery 

flavour, less on the gills. 

CHEMICAL REACTION: Guaiac strong and rapid. 

FeSO pale pink. 

4 

MICROSCOPY: spores from obvoid to subglobose, 8.5- 

11.2 × 7-9 µm, echinulate, crested-catenulate, partly 

connected, subreticulate, with obtuse spines; creamwhitish 


HABITAT: very common species under hardwood and 

conifer, with preference to dry, calcerous ground. 


NOTE - R. chloroides can be distinguished by its funnelshaped, 

fairly regular cap and its very close and crowded 

gills. R. palespora is a highly characteristic species for its 

refreshing, amarescent flavour which is never harsh. Its 

gills are ochre with an orange lustre at maturity, with full, 

cream spores. 

213

CAP 50-100 (120) mm, fleshy, initially firm in the shape 

of a helmet, then convex, becoming flat, finally also 

lightly depressed; with obtuse and joint margin; cuticle is 

separable up to half of the radius, lubricated and shiny, 

full vivid red, apple red, arterial blood red, sometimes 

with blackish shades in the centre or with a sharply 

demarcated ivory-cream zone. 

GILLS rounded or almost free at insertion, from lightly 

convex to straight, width of 6-9 mm, thin and fragile, 

finally spaced, sparsely forked, interveined, sometimes 

whitish, with cream straw tinges very clear view of gill 

edge; intercalated by sporadic lamellule. 

STIPE 50-80 (100) × 10-20 mm, slim, slightly claviform 

when young, then fairly cylindrical, often lightly thinned 

at tip, visibly corrugated, white, slightly stained yellowbrown 

in certain conditions, finely striated and a little 

satin for the rest; medullar, soon becoming incomplete. 

FLESH fragile, a little watery, white, slightly yellowing 

when handled; peppery flavour in every part, 

imperceptible or lightly fruity odour at gill edge. 

214 

Russula emetica (Schaeff. : Fr.) Persoon 

CHEMICAL REACTION: Guaiac, subnul. 

MICROSCOPY: spores mostly oboval 8.8-10.5 × 7.4-8.8 

µm, echinulate, with large conical-obtuse warts, 

sometimes briefly crested, reticulated-connected. Spores 

are whitish. 

HABITAT: under conifers in mountains, mostly Picea, 

but also under birch; with preference to sphagnum or 

other types of moss. 


NOTE - Among similar species, we can distinguish: R. 

grisescens Sphagnicolous, around half the size of R. 

emetica, flesh greying with humidity, positive reaction to 

Guaiac, spores very much smaller and finely ornate; R. 

nanatypical of Alpine microsilva. R. mairei found in fresh 

hardwood forests, with a predilection for the Beech has 

flesh that turns blue on treatment with Guaiac and gills 

that reveal a glaucous tinge which is inconsistent but 

characteristic. R. silvestris grows in dryer ground on 

mossy carpets at the bottom of oak, chestnut or pine trees.

CAP 55-140 (200) mm, fleshy and rigid subglobular, 

progressively expansive, then flat, grooved-tubercolate 

margin for 20-30 mm, acute; thorny surface, cuticle is 

separable for a third of the radius, viscous, persistently 

shiny when dry, yellow-brown colour, honey like, paler in 

peripheral area, then bearing brown tawny spots, also 

tawny blackish in the contused parts. 

GILLS rounded-free, partly connate at insertion, width of 

8-16 mm, unequal, fairly thick, sparsely forked, not very 

crowded, interveined, fragile, ivory cream, bearing watery 

drops in humid weather, with residual rusty stains on gill 

edge, browning with age. 

STIPE 60-120 × 20-40 mm, cylindrical, fairly flared at 

tip, corrugated, rusty at tip, stained brown greying 

elsewhere, medullar-incomplete for three or four cells 

soon becoming confluent with wide caverns, the skin is 

fragile and thick, papered with brownish lumps and soft 

internally. 

FLESH rigid and fragile, whitish on gill edge, not 

yellowing, very stained soon becoming rusty brown; very 

clear peppery flavour, unpleasant odour, complex and 

Russula foetens Pers. : Fr. 

difficult to define, with fruity hints in the second level and 

only to perceptible traits. 

CHEMICAL REACTION: Guaiac positive. KOH 

negative. 

MICROSCOPY: spore subglobose, 8-9.8 × 7-8.2 µm, 

with flat spines and apex, incompletely amyloid. 

HABITAT: gregarious, widespread and abundant, under 

hardwood and conifers. 


NOTE - R. laurocerasi can be identified by its average or 

small stature, the absence of gluten, a strong smell of 

bitter almonds; its rounded spores, embellished with 

spectacular winged crests. Also R. illota gives off 

effluence of bitter almonds, though to a lesser degree and 

mixed with rather less pleasant hints than R. foetens. The 

stipe and the gill edge of its gills display a characteristic 

blackish-brown series of markings. Its most similar 

lookalike, however is, R. subfoetens, recognizable by its 

slightly smaller stature, the surface of its cap which is 

usually without gluten, its yellowing flesh and its positive 

reaction to KOH. 

215

CAP 60-100 (130) mm, very fleshy and hard, globular or 

with lightly pointed summit, pulvinate, finally distended 

and a little depressed, with fleshy and obtuse rigid 

margin, curving at the extremes; cuticle is separable for a 

third of the radius, thick, tenacious-elastic, appears 

greasy, shiny, rarely matt or even pruinose, brown, brown 

honey or chestnut. 

GILLS attenuated, then rounded, fairly crowded, with 

some forking, subacute at the front, nearly straight, width 

of 5-8 mm, interveined, sublardaceous, straw cream, then 

stained brown on the surface; some lamellule are present. 

STIPE 40-100 × 15-35 mm, bulky, from cylindrical to 

subclavate, clearly corrugated, stained brown and often 

plicate at base, ivory-cream, then with fairly extensive 

rusty stains; full, compact medulla, finally cavernous. 

FLESH of notable thickness and hardness, firm, white, 

shades of yellow under the cuticle, clearly washed with 

ochre brown when exposed to air and with age; sweet 

flavour and no distinctive odour. 

CHEMICAL REACTION: FeSO vivid orange. aniline 

4 

on the gills, slowly yellow. Guaiac fairly rapid and 

intense. 

216 

Russula mustelina Fries 

MICROSCOPY: spores oboval 7-9.7 × 5.8-7.8 µm, 

warty, crispate, weakly connected, subreticular. Pale 

cream ochre-ish in mass. 

HABITAT: highly common alpine forests where it grows 

in abundance and fairly underground; end of summerautumn. 


NOTE - An edible mushroom which is widely 

commercialised thanks to its firm flesh and pleasant 

flavour. It can be identified by its tawny-brown to honeybrown 

colouration, evocative of R. foetens or Boletus 

edulis, its pale cream spores, and a vivid orange reaction 

to FeSO 4. It loves to grow fairly buried in temperate 

periods, from the end of summer to autumn, until the first 

signs of winter. In some years it can be found abundantly 

while in others it can still be found, yet far less 

numerously.

CAP 40-70 (100) mm fleshy and firm then more fragile, 

at first convex-subhemispheric, progressively flat, finally 

mostly depressed, with thinned but obtuse margin, briefly 

grooved in maturity; cuticle separable to almost half of 

the radius, wet and shiny collectively, red purple, violet 

vinous, dark violet, often blackish towards the disc, other 

times brown violet, partly brownish, streaks of greengrey. 

GILLS attenuated or lightly rounded, obtuse at the front, 

interveined, sparsely forked, fairly crowded, width of 4-8 

mm, fragile, whitish, then dirty cream, interspersed with 

infrequent lamellule. 

STIPE 30-90 × 10-20 mm, cylindrical, sometimes a little 

fusiform, rugulous, white at base, with red shades 

elsewhere, carmine under a whitish bloom thickened and 

fugacious to the touch, often completely white, a little 

greying due to imbibition, the medulla is compact, then 

softened and partly incomplete. 

FLESH violet under the cuticle, a little greying due to 

imbibition; intense fruity odour, like mature pears, clearly 

peppery flavour. 

Russula queletii Fries 

CHEMICAL REACTION: Guaiac positive, slow. FeSO 4 

pale pink-orange. 

MICROSCOPY: spores ellipsoidal-oboval 7.3-9 (9.8) × 

6-7.3 (8.2) µm, from echinulate to spinulose, to warty 

conical-obtuse. Light cream in mass. 

HABITAT: highly common in the subalpine area under 

Picea in calcerous non-humid ground, sometimes also 

under white fir and Pinus. 


NOTE - R. cavipes is smaller and grows under Abies alba, 

and more rarely under Picea, it has a humid and bright 

cuticle, more widely-spaced gills which are whitish when 

young, its stipe is never red-violet and its spores are pale 

cream; Furthermore, it has a subnul reaction to Guaiac 

testing and a positive, reddening, reaction to ammonia. It 

has a peppery flavour and an intense, pleasant odour of 

geraniums. R. sardonia, can be identified by its pale, 

sulphuric gills, insignificant odour and its arenaceous pine 

habitat. 

217

CAP 30-100 (120) mm, from convex to flat, finally 

depressed, fine firm margin, curving, regular, smooth, yet 

slightly striated, briefly, in adult specimens; vivid red, 

cherry red, carmine red, without violet tones, paler at 

margin, fading to pale pink, sometimes with white ivory 

patches; surface rough due to fine granules, dry and matt, 

lightly viscous and shiny with humidity. 

GILLS adnate and fairly decurrent, initially bent, then 

horizontal or a little ventricular, width of 3 to 10 mm, 

thick, fairly crowded, sometimes biforked, joined on the 

background by fine veins, irregularly intercalated from 

some lamellule; whitish then cream-ochre-ish, fine edge, 

sometimes coloured red. 

STIPE 30-80 × 10-30 mm, fairly stocky, cylindrical or 

attenuated at base, rigid, full, eventually filled-cavernous; 

normally completely suffused with red or red-pinkish, up 

to almost concolour with cap, on white-ochre-ish 

background, fairly yellowing starting from the base; 

surface is finely rugulous-reticulated. 

218 

Russula sanguinea (Bulliard) Fries 

FLESH firm compact, firm, whitish, carmine red under 

the cuticle, yellowing fairly slowly; weak fruity odour; 

acrid, hot and also bitter flavour. Slow but positive 

reaction to Guaiac. 

MICROSCOPY: spores pale ochre in mass, oboval with 

conical-obtuse warts about 1 µm high, joined in parts with 

infrequent ridges, 7.8-9.4 × 6.5-8.2 µm. 

HABITAT: in conifer woods, principally under pine, 

fairly common, in summer-autumn. 


NOTE - Russula sanguinea, due to its morphochromatic 

features, is similar to: R. persicina which, however, 

favours hardwood forests; R. helodes, typical of high 

mountain bogs and is linked to conifers; R. rhodopus, 

with its lacquered red cap, and habitat of acidic grounds 

in red fir forests. In the wild one also comes across some 

chromatic variants, generally ranked by shape or variety.

CAP 45-100 (140) mm, fleshy and firm, subglobular, then 

irregularly flat, eventually depressed, with thinned yet 

obtuse margin; cuticle is separable for two fifths of the 

radius, often a little retracted towards the margin, soon 

becoming dry and matt, pink lilac, brown vinous, 

sometimes with undefined pale areas, cream flesh colour, 

occasionally stained green-grey, with streaks darker than 

the background. 

GILLS vaguely decurrent and biforked mostly at the 

insertion, subacute at the front, crowded and relatively 

close, delicately interveined, sublardaceous when young, 

whitish, with rusty stains and finally yellow when 

handled. 

STIPE from subcylindrical to fairly progressively 

attenuated towards the base, corrugated, rare hints of pink 

on the side, rusty low down, with some yellow-brown 

stains, full, then a little filled with age. 

FLESH compact, white, clearly yellowing when touched 

and partly stained brown; completely sweet flavour and 

indistinctive odour. 

CHEMICAL REACTION: FeSO red-orange. Guaiac 

4 

positive. aniline, yellow. 

Russula vesca Fries 

MICROSCOPY: spores oboval or a little elongated, 6.4-8 

× 5.3-5.8 µm, with isolated warts; any links between them 

are very thin and sporadic. characteristic needle-like 

forms protrude from rigid and thick cell walls. 

HABITAT: highly common in mildly acidic or neutral 

ground under various types of hardwood and under 

conifers in mountains; from the late spring. 


NOTE - R. vesca can be recognised by its slightly lilac or 

winey-brown flesh, the tendency to turn yellow-brown on 

its deteriorating parts, its sweet flavour, white spores and 

a red-orange reaction to FeSO 4 . One should guard against 

giving excessive importance to characteristics which are 

inconsistent and none too specific: the tendency of the 

cuticle to retract towards the margin (“habillé trop court” 

according to a metaphor given by French authors) for 

example. In the case of greening or partially discoloured 

samples, the completely sweet flavour, white spores and 

an energetic reaction to FeSO 4 should help to distinguish 

this against other, macroscopically similar species of 

Griseinae. 

219

220 

Sarcosphaera crassa (Santi ex Steudel) Pouzar 

[= S. eximia Durieu & Léveillé; S. coronaria (Jacquin) Schroeter] 

ASCOCARP made from a subspheric, sessile apothecium. 

APOTHECIUM initially semi-underground, globular, up 

to 160 mm in diameter, top is open only for a fairly small 

operculum (sometimes in a non apical position), then 

more and more open and protruding from the ground up 

to appear domed and epigean. Hymenophore is smooth, 

lightly undulated, initially violet, then darker, tending to 

turn brown violet. Smooth external surface, white greyish. 

edge soon becoming cracked, lacianated, with erratic 

points due to the lacerations on the carpophore during 

growth. 

FLESH fragile, leathery, whitish, thick. 

MICROSCOPY: regular ellipsoidal spores, with well 

rounded extremes, smooth,18-20 × 7-8 µm, hyaline under 

microscope, biguttulate, uniseriat in asco; asci are 

cylindrical, amyloid, octosporic; cylindrical paraphyses 

with slightly widened tip, septate and forked. 

HABITAT: ubiquitous, on the ground among needles, 

leaves, grass or moss, in humid places; rarely isolated, 

more often found in large groups, from the spring to 

summer, rarely in autumn. Quite common. 


NOTE - S. crassa has amyloid asci such as the species in 

the genus Peziza; the genus Sarcosphaera is, however, 

distinguished from the genus Peziza by its semi-buried 

growing habits. This prerogative does not really seem a 

convincing justification for the creation and maintenance 

of this separation, not least of all because several “true” 

Peziza species, such as P. ammophila and P. 

pseudoammophila, fruit almost completely underground 

in the sand of coastal dunes in autumn.

CAP 40-120 mm, from hemispheric to convex, then flat; 

margin from convoluted to curving towards the base, then 

distended cuticle overflowing on the hymenophore; 

surface is viscous in humid weather, otherwise slimy, 

totally separable, smooth and shiny in dry weather; from 

brown reddish, to tawny, to brown yellowish. 

TUBES up to 10 mm, from adnate to weakly decurrent; 

yellow, then golden yellow and finally yellow-olive at 

complete sporal maturation; pores are initially small and 

round, secretes opalescent yellow drops, a little angular in 

maturation, concolour with tubes, sometimes browning in 

patches. 

STIPE 40-90 × 10-25 mm, cylindrical, a little widened at 

base, sometimes supple or curving; covered with a very 

minute, pale yellow granulation, sometimes milky, 

coloured on background and only browning later on; skin 

is chrome yellow, pale lemon yellow in colour, often 

brownish patches at base. 

FLESH firm when young, then softer; whitish, yellow 

pale near the tubes and under the skin of the stipe; 

uniform to the area; weakly fenolic odour, sweetish 

flavour. 

Suillus granulatus (L. : Fr.) Roussel 


pale yellow under microscope; cuticle of the cap is 

formed from un gelatinised trichoderma, made from 

cylindrical hyphae which soon transform becoming cutis. 

Spores are brown-ochre. 

HABITAT: largely widespread species, considered to be 

highly associated with Pinus and to two needles. Fruits 

mainly in hills and mountains, from summer to late 

autumn; common. 


NOTE - This is perhaps the most well-known Suillus, 

very common, and grouped together with similar species 

by the common name of “pinarolo” in Italy. It might be 

confused with S. collinitus, due to its identical edible 

properties, but it can be told a part by its colour, which is 

normally more brown-red, the absence of radial fibrils on 

its cap cuticle and its more-minute stipe decorations. The 

spores of S. granulatus are also a fair degree smaller as 

well. 

221

CAP up until 120 (150) mm, initially hemispheric then 

convex, pulvinate, rarely; margin curving at the extremes 

towards the base, regular, acute, a little excessive, often 

decorated with whitish remnants of the partial veil; 

smooth cuticle, very viscous, separable, brown, yellow 

brown, brown violet, chocolate brown, often with darker 

radial fibrils when dry. 

TUBES up to 12 mm, adnate or decurrent, `yellow, then 

chrome yellow, finally yellow brownish; small, round 

pores, only angled in advanced maturation, concolour 

with tubes, uniform when pressed. 

STIPE 40-70 × 12-30 mm, longer than the diameter of the 

cap when young, then the same length as the diameter of 

the cap or shorter, cylindrical, often a little widened 

towards base, full; with a broad membranous annulus, 

whitish in colour, then violet brown, it is possible to find 

adherent volviform residues at the base, from whitish to 

white-grey-violet. A fine yellow lattice can be found 

above the annulus, below the annulus there are yellow 

points which then become concolour with cap. 

222 

Suillus luteus (L. : Fr.) Roussel 

FLESH initially firm soon becoming soft and watery in 

cap, more fibrous in stipe; white, then yellowing; uniform 

to gill edge. Pleasant, fruity odour, sweet flavour. 


pale yellow under microscope. Spores brown olive rusty 

in colour. 

HABITAT: only in groups in Pinus woods, recurrent 

from late summer to late autumn. 


NOTE - This is a very common Suillus, the only one 

which has an annulus adorning its stipe and which grows 

with two-fascicle (P. nigra, P. sylvestris, more rarely 

anche P. pinaster) and three-fascicle (P. radiata) Pinus 

trees. Grows tendentially in hills and mountains and is not 

usually found under coastal pines. As it matures and ages, 

the cuticle has a tendency to become dehydrated and at 

that stage it often assumes an appearance very similar to 

that of S. collinitus. Several forms and varieties have been 

described: fo. albus, completely white, and fo. volvaceus, 

with a short stipe whose annulus has the appearance of a 

volva.

CAP 30-60 mm, a little fleshy, convex campanulate, soon 

becoming flat, with an obtuse umbo, lightly convoluted, 

then distended and thinned margin , sometimes cracked. 

Cuticle is dry, separable, without remnants of veil at the 

edge, decorated with fine radial fibrils which take the 

appearance of small scales, grey-whitish when young, 

then grey. 

GILLS non-marginated and decurrent with teeth, fairly 

crowded, thin and fragile, with numerous lamellule, white 

with greyish hints, not yellowing with age or when 

handled, the surface is undulated-crenulated, often a little 

serrated. 

STIPE 35-70 × 5-20 mm cylindrical, slightly curving, 

lightly widened at base, white, silky, often with remnants 

of veil, more visible in the apical part of young examples. 

FLESH compact then soft in cap, firm in stipe then 

fibrous, white not yellowing visibly after collection, with 

strong odour and flavour of fresh flour. 

Tricholoma argyraceum (Bull. : Fr.) Gillet 

MICROSCOPY: spores 5-6 × 2.5-3.5 µm, ellipsoidal, 

guttulate. clavate, tetrasporic basidia. Epicyte made from 

parallel, collated hyphae, fairly erect. 

HABITAT: rare and late species, grows in small numbers, 

on broken ground normally near hardwood (hornbeam 

and hazel). 


NOTE - This is a species belonging to the Section 

Scalpturatum, and is characterized by its odour and 

flavour, which recall fresh flour, and by the remnants of 

veil on its stipe. It differs from T. scalpturatum (Fr.) Quél. 

in its grey-silver cap and its non-yellowing flesh, and can 

be distinguished from the Terreum mushrooms, in that 

these do not have either a flavour or odour of flour. 

223

CAP 50-100 mm, convex or campanulate, then flat and 

with a large umbo, the cuticle is dry, lightly viscous and 

shiny with humidity, almost smooth or velvety on 

circumference in dry conditions, decorated with 

concentric brass brown scales or often with brown reddish 

fibrils darker in the centre, the margin is convoluted 

lengthwise, then distended, lobed and irregular, golden 

yellow with age. 

GILLS non-marginated or subfree, fairly crowded, intense 

yellow or citrina yellow, tendency to darken with age, the 

whole surface is lightly undulated. 

STIPE 60-90 × 8-15 mm, subcylindrical or with lightly 

clavate base, sometimes short and bulging, stocky, bent, 

concolour with cap, with some sparse brown light reddish 

floccules towards base. 

FLESH ochre-ish-yellowish or brass colour under the 

cuticle of the cap or in stipe, with a pleasant, lightly 

floury or a little aromatic odour and sweet floury flavour, 

bitter if chewed. 

MICROSCOPY: spores mostly ellipsoidal or 

amygdalform, hyaline, 6.0-7.5 × 3.5-4.5 µm. clavate, 

224 

Tricholoma equestre (L. : Fr.) Kummer 

tetrasporic basidia. Epicyte made from fairly erect, 

parallel interwoven hyphae, 

HABITAT: in conifer and hardwood areas where it seems 

to prefer poplars. 


NOTE – This species is fairly variable in its colour. Dry 

and completely yellow examples are confusable, at first 

glance, with T. sulphureum, which, however has spaced 

gills, a less intense yellow and unpleasant odour of coal 

gas. Known and valued as a choice edible up until a few 

years ago, they are today suspected of having been 

involved in several cases of poisoning (and even death) 

after abundant consumption and in undercooked meals; 

the episodes under investigation all occurred in a certain 

area in France and today we are awaiting further 

verification. In any case, as a precautionary measure, and 

in the absence of certainty, the gathering and consumption 

of this species (and of all the entities belonging to its 

immediate group – Ed.) is forbidden (by law) throughout 

the entire Italian and French territories.

CAP 60-150 mm, convex, campanulate, with large obtuse 

umbo, then flat, grey-ochre-ish, dark grey, slate grey, 

sooty blackish, with greenish or violet stains on a yellow 

background just visible towards the centre, but stands out 

at the margin which is often clearly grey- citrina yellow, 

the cuticle is fibrillated radially, a little viscous and shiny 

in humid weather, otherwise silky, the edges are slightly 

supple then lobed, cracked, often revoluted with age. 

GILLS non-marginated, a little crowded, fairly wide, 

slightly thick, sinuate, white, then ash grey with yellowish 

stains, the edge is irregular and sometimes serrated. 

STIPE 50-110 × 8-20 mm, robust, cylindrical or fusiform, 

satin-fibrillated, whitish, always with yellow stains 

especially towards the top, the apex is dandruff white and 

stains yellow-brownish-olive to the touch 

FLESH firm in cap and fibrous in stipe, white, a little 

yellowish in the stipe, fairly greyish below the cuticle of 

the cap with a pleasant floury odour and taste. 

Tricholoma portentosum (Fr. : Fr.) Quélet 

MICROSCOPY: ellipsoidal, hyaline spores, 5.5-7.0 × 4.0- 

5.0 µm. Clavate, tetrasporic basidia. Epicyte made from 

an ixocutis of fairly erect, parallel interwoven hyphae. 

HABITAT: in conifer and hardwood areas, in autumn 

even late on. 


NOTE - When the cap fades, leaving just a glimpse of the 

yellowish-ochraceous colour, it can be confused with T. 

sejunctum, which, however, has a bitter, floury taste. Also 

T. virgatum has a fibrillated-virgated cap but its flavour 

and odour are quite unpleasant. One should pay attention 

to avoid taking it for the toxic T. josserandii, which has a 

dry, velvety cap, and a characteristic insect-like odour. 

225

CAP 30-90 mm, campanulate, conical, flat, often 

irregular, obtusely umbonate, matt cuticle, almost smooth 

or slightly woolly-felted at first, then bearing thick fibrils, 

almost uniform, smoke grey, dark brown or almost black 

in colour, the margin is convoluted or curving for a long 

while, often with an overflowing edge. 

GILLS non-marginated-adnate or hooked, slightly 

crowded, whitish or a little light greying, the surface 

lightly crenulated with age. 

STIPE 30-70 × 8-12 mm, cylindrical, stocky, full and 

fibrous, fragile, hollow or a little fistular with age, 

smooth, silky, completely white or with light greying 

fibrils which darken slightly in young specimens. 

FLESH fibrous, fragile, white, greyish under the cuticle, 

with a light fungal odour, and a herby or light floury 

flavour . 

MICROSCOPY: spores mostly ellipsoidal, subglobulose, 

hyaline, guttulate 6.0-7.5 × 4.5-5.5 µm. clavate, 

226 

Tricholoma terreum (Schaeff. : Fr.) Kummer 

tetrasporic basidia. Epicyte made from fairly erect, 

parallel interwoven, hyphae. 

HABITAT: generally abundant in growing areas, found in 

large groups; in conifer woods (pine or fir), from the end 

of summer until the first frosts. 


NOTE - This Terreum species iso ne of the most soughtafter 

mushrooms for human consumption and it is 

popularly known a “Moretta” (little brunette) in Italy. It is 

often confused with: T. myomyces which is smaller and 

more fragile with remnants of a silvery veil at the tip of 

its stipe and a floury odour; T. triste whose gill surface is 

darker and which has a stipe with brown-blackish shades; 

or else it is mixed up with grey-toned species of the 

scalpturatum and atrosquamosum groups, but which have 

almost no odour, non-wide, white or more or less grey 

gills, and whose stipe is whitish with no veil; in the field 

they can be told apart very easily.

CAP 50-120 mm, hemispheric, convex, then a little flat, 

finally depressed, vivid red-brown, the cuticle is very 

viscous or glutinous, in dry conditions it tends to stain and 

discolour towards the margin with shades of ochreorange, 

the margin is convoluted at the extremes, with 

very evident ribs. 

GILLS hooked, sinuate-adnate, not very crowded, close, 

white, stained reddish in adult specimens, the surface is a 

little sinuous. 

STIPE 50-120 × 10-20 mm, cylindrical or fairly clavate 

or fusiform full then hollow, white dandruff at tip, stained 

brown-reddish to from the annular line which is more 

evident in young specimens, with fibrils coloured or often 

lighter, a little viscous with humid weather. 

FLESH firm white, with strong smell of flour or 

cucumber and floury flavour. 

MICROSCOPY: spores subglobose, suboval hyaline, 

guttulate 5.5-6.5 × 4.5-5.0 µm. clavate, tetrasporic 

Tricholoma ustaloides Romagnesi 

basidia. Epicyte made from an ixotrichoderma of fairly 

erect, interwoven, parallel hyphae. 

HABITAT: under hardwood (oak, chestnut, beech, 

hornbeam), in autumn; fairly common. 


NOTE - This species is characterised by a slimy cuticle 

that, on drying, tends to leave traces of mucus grouped 

around the edges. It is often confused with T. ustale, 

which, however has a smooth margin, a stipe devoid of a 

delimited annulus zone and the flesh of the base of the 

stipe right up to the gill edge is a reddish-brown. It can 

also be confused with T. fracticum and T. striatum with 

smooth caps bearing radial fibrils and which principally 

grow under mountain conifers. T. ustaloides, however, it 

can easily be recognised in the field by its flavour and 

odour. 

227

CAP 30-140 mm, conical, hemispheric, then flat, often 

with obtuse umbo, vinous red, pink-red intense purple on 

a vivid yellow or golden yellow background, the cuticle 

bearing adpressed woolly decorations in the centre and 

small scales which are adpressed gradually and radially 

towards the edges, the margin is convoluted for a long 

time. 

GILLS from non-marginated to mostly adnate, partly 

anastomosed, wide, moderately crowded, intense sulphur 

yellow or golden yellow, the surface is finely floccose or 

fimbriated. 

STIPE 50-120 × 10-25 mm, cylindrical or fusiform often 

sinuous, full, hollow with age, concolour with cap or 

lighter, decorated with scales, more fleeting towards the 

base which eventually becomes yellow, the apex is whiteyellowish. 

FLESH soft, thick in the centre, yellow-cream, with sour 

mould or light odour, like wood, and sweet light flavour, 

like hazelnut or a little bitter. 

228 

Tricholomopsis rutilans (Schaeffer : Fr.) Singer 

MICROSCOPY: spores mostly ellipsoidal, hyaline, 

guttulate 7.0-8.5 × 5.5-6.5 µm. clavate, tetrasporic 

basidia. Epicyte made from fairly erect, parallel, 

interwoven hyphae. 

HABITAT: grows collated or in groups on rotting parts of 

conifers especially fir and pine. 


NOTE - This is an unmistakeable species thanks to its 

lignicole habitat and its yellow gills which contrast with 

the wine colour of its cap. T. decora can be found in the 

same habitat, but has a lighter cap which bears no winey 

tones and is generally more slender. T. flammula is a very 

small species, with a cap diameter that reaches only up 

until 15 mm, purplish-brown fibrillated scales and a 

yellow stipe; T. ornata has a yellow-olivish cap with 

fairly sparse, intense reddish-brown scales, a pale yellow 

stipe with fibrils, and grows on woody debris.

CAP 50-120 (150) mm, fleshy, from hemispheric to 

convex, pulvinate, finally flat; edge very soon becoming 

regularly distended or a little undulated; cuticle is velvety 

when young, dry, sometimes dimpled; with very variable 

colourations, from pure yellow, to yellow with citrina 

tinges, to alutaceous brown with greenish hints, brown 

orange in dry weather, rusty brown, brown reddish, up to 

liver red in humid weather. Scratching the cuticle with the 

finger when it is in its dry state it is possible to observe a 

rusty Brown subcuticular layer. 

TUBES up to 15 mm, adnate and sometimes 

subdecurrent, chrome yellow, then with greenish hints, 

finally olivish, slowly turning blue at gill edge; pores are 

concolour with tubes, round, soon becoming open, then 

large and angular, turning blue to the touch. 

STIPE 50-80 (100) × 10-20 (25) mm, cylindrical, curving 

at base, supple, almost always dilated at tip and attenuated 

at base; very pale yellow, tendency to turn brown slowly 

during maturity; fine points or lengthwise ribs are often 

found which form a sort of lattice. 

FLESH firm and compact, soon becoming soft in cap and 

fibrous in stipe, pale chrome yellow, typically ochre at 

Xerocomus subtomentosus (L. : Fr.) Quélet 

base of stipe, more evident in humid weather; slowly 

turns blue at gill edge; weak slightly acidic odour; sweet 

flavour. 

MICROSCOPY: ellipsoidal fusiform spores with superior 

depression, 10.6-13.2 × 4.3-5.0 µm, pale yellow under 

microscope, olive brown in mass. 

HABITAT: in relatively low numbers, fairly indifferent to 

substrate, isolated or in small groups; recurrently 

associated with oak and chestnut; summer-autumn. 


NOTE - This is one of the most noted and common 

Xerocomus species. The chromatic variability of its cap is 

mainly due to climatic-environmental conditions: 

examples with a reddish-brown cap are common after it 

rains, even in the same places where one usually finds 

carpophores with olive-brown caps. The variants with 

yellow caps should probably be considered as intraspecial 

varieties. X. ferrugineus is very similar, but 

prefers to grow on siliceous ground and, generally, at 

higher altitudes. 

229

230

European Commission 

EUR 24415 EN – Joint Research Centre – Institute for Environment and Sustainability 

Title: Chemical elements in Ascomycetes and Basidiomycetes – The reference mushrooms 

as an instrument for investigating bioindication and biodiversity 

Authors: R. M. Cenci, L. Cocchi, O. Petrini, F. Sena, C. Siniscalco, L. Vescovi 

Luxembourg: Publications Office of the European Union 

2011 – 232 pp. – 18 x 24 cm 

EUR – Scientific and Technical Research series – ISSN 1018-5593 

ISBN 978-92-79-20395-4 

doi:10.2788/22228 

Abstract 

Fungi in the wild are among the principal agents in biogeochemical cycles; those cycles of 

matter and energy which enable ecosystems to work. 

By investigating the biodiversity of Italian fungal species and concentration levels of chemical 

elements in them, it may be possible to employ these fungi as biological indicators for the 

quality of forest, woodland and semi-natural environments. The data archives of EUR 

Reports record the dry-material concentrations, of 35 chemical elements, including heavy 

metals, in over 9000 samples of higher mushrooms. These samples represent around 200 

genera and a thousand species. As the archive has attained statistical stability it has been 

possible to define the concept of a “reference mushroom”. The use of a “reference 

mushroom” may bring benefits – perhaps only as a methodological approach – in various 

fields of mycological and environmental research; from biodiversity and bioindication, 

through taxonomy right up to health and sanitation issues. 

The sheer volume of the collected data may prove to be useful as a comparison for data 

collected in the future; such results would also allow a better and more-exhaustive 

interpretation of the effects of environmental-protection laws which have been enacted over 

the years in order to reduce or remedy current climate-change phenomena and the 

environmental damage caused by human activity. Studies pertaining to the frequency of 

occurrence and the ecology of the various fungal species found on Italian soil have tended to 

link the reference habitats used to European classification guidelines (Natura 2000, CORINE 

Land Cover, CORINE Biotopes and EUNIS). Thereby the foundations have been lain for the 

use of mushrooms as biological indicators for the measurement of soil and ecosystem 

quality. 

How to obtain EU publications 

Our priced publications are available from EU Bookshop (http://bookshop.europa.eu), 

where you can place an order with the sales agent of your choice. 

The Publications Office has to worldwide network of sales agents. You can obtain their 

contact details by sending to fax to (352) 29 29-42758. 

231

The mission of the JRC is to provide customer-driven scientific and technical 

support for the conception, development, implementation and monitoring 

of EU policies. As to service of the European Commission, the JRC functions 

as to reference centre of science and technology for the Union. Close to the 

policy-making process, it serves the common interest of the Member States, 

while being independent of special interests, whether private or national. 

232 

LB-NA-24415-EN-C

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