Fungal Conservation
Issues and Solutions
Threats to fungi and fungal diversity throughout the world have
prompted debates about whether and how fungi can be conserved. Should it be the site, or the habitat, or the host that is
conserved? All of these issues are addressed in this volume, but
coverage goes beyond mere debate with constructive guidance for
management of nature in ways beneficial to fungi. Different parts
of the world experience different problems and a range of
examples are presented: from Finland in the North to Kenya in
the South, and from Washington State, USA, in the West to
Fujian Province, China, in the East. Equally wide-ranging solutions are put forward, from voluntary agreements, through land
management techniques, to primary legislation. Taken together,
these provide useful suggestions about how fungi can be included
in conservation projects in a range of circumstances.
is Reader in Genetics in the School of Biological
Sciences at the University of Manchester.
is a mycologist at the National Herbarium of
The Netherlands.
is a freelance consultant mycologist in the
spheres of education, survey and conservation.
is Consultant Mycologist to the National
Botanic Garden of Wales and Director of the Cambrian Institute
of Mycology.
MMMM
Fungal Conservation
Issues and Solutions
A SPECIAL VOLUME OF
THE BRITISH MYCOLOGICAL SOCIETY
EDITED BY
DAVID MOORE, MARIJKE M. NAUTA,
SHELLEY E. EVANS AND MAURICE ROTHEROE
Published for the British Mycological Society
CAMBRIDGE UNIVERSITY PRESS
Cambridge, New York, Melbourne, Madrid, Cape Town, Singapore, São Paulo
Cambridge University Press
The Edinburgh Building, Cambridge CB2 8RU, UK
Published in the United States of America by Cambridge University Press, New York
www.cambridge.org
Information on this title: www.cambridge.org/9780521803632
© British Mycological Society 2001
This publication is in copyright. Subject to statutory exception
and to the provisions of relevant collective licensing agreements,
no reproduction of any part may take place without the written
permission of Cambridge University Press.
First published 2001
This digitally printed version 2008
A catalogue record for this publication is available from the British Library
Library of Congress Cataloguing in Publication data
British Mycological Society. Symposium (1999: Royal Botanic Gardens, Kew)
Fungal conservation: issues and solutions: special volume of the British
Mycological Society/edited by David Moore . . . [et al.].
p.
cm.
Includes bibliographical references (p. ).
ISBN 0 521 80363 2 (hardback)
1. Fungi conservation – Congresses. I. Moore, D. (David), 1942– II. Title.
QK604.2.C66 B75 1999
333.95´2–dc21 00-065169
ISBN 978-0-521-80363-2 hardback
ISBN 978-0-521-04818-7 paperback
Contents
page
List of contributors
vii
Preface
ix
1 Fungal conservation issues: recognising the problem,
finding solutions
1
David Moore, Marijke M. Nauta, Shelley E. Evans
& Maurice Rotheroe
2 Current trends and perspectives for the global
conservation of fungi
7
Régis Courtecuisse
3 Conservation and management of forest fungi in the
Pacific Northwestern United States: an integrated
ecosystem approach
19
Randy Molina, David Pilz, Jane Smith, Susie Dunham,
Tina Dreisbach, Thomas O’Dell & Michael Castellano
4 The future of fungi in Europe: threats, conservation
and management
64
Eef Arnolds
5 Fungi as indicators of primeval and old-growth forests
deserving protection
81
Erast Parmasto
6 Recognising and managing mycologically valuable
sites in The Netherlands
89
Leo M. Jalink & Marijke M. Nauta
7 Threats to hypogeous fungi
95
Maria Ławrynowicz
8 Wild mushrooms and rural economies
105
David Arora
v
vi
Contents
9 Threats to biodiversity caused by traditional mushroom
cultivation technology in China
Siu Wai Chiu & David Moore
10 A preliminary survey of waxcap grassland indicator
species in South Wales
Maurice Rotheroe
11 Grasslands in the coastal dunes: the effect of nature
management on the mycota
Marijke M. Nauta & Leo M. Jalink
12 The conservation of fungi on reserves managed by the
Royal Society for the Protection of Birds (RSPB)
Martin Allison
13 Strategies for conservation of fungi in the Madonie
Park, North Sicily
Giuseppe Venturella & Salvatore La Rocca
14 Fungal conservation in Ukraine
D. W. Minter
15 The threatened and near-threatened Aphyllophorales
of Finland
H. Kotiranta
16 Fungal conservation in Cuba
D. W. Minter
17 Microfungus diversity and the conservation agenda in
Kenya
P. F. Cannon, R. K. Mibey & G. M. Siboe
18 Fungi and the UK Biodiversity Action Plan: the
process explained
L. V. Fleming
19 The Scottish Wild Mushroom Forum
Alison Dyke
20 The contribution of national mycological societies:
establishing a British Mycological Society policy
David Moore
21 The contribution of national mycological societies: the
Dutch Mycological Society and its Committee for
Fungi and Nature Conservation
Marijke M. Nauta & Leo M. Jalink
22 Fungal conservation in the 21st century: optimism and
pessimism for the future
David Moore, Marijke M. Nauta, Shelley E. Evans &
Maurice Rotheroe
Index
111
120
136
144
156
162
177
182
197
209
219
223
242
247
256
Contributors
Martin Allison
The Royal Society For The Protection Of
Birds, The Lodge, Sandy, Bedfordshire
SG19 2DL, UK
Eef Arnolds
Holthe 21, 9411 TN Beilen, The Netherlands
:eefarnolds@hetnet.nl9
David Arora
343 Pachecho Avenue, Santa Cruz, Santa
Cruz, California 95062, USA
:arora–david@yahoo.com9
P. F. Cannon
CABI Bioscience, Bakeham Lane, Egham,
Surrey TW20 9TY, UK
:p.cannon@cabi.org9
Michael Castellano
USDA Forest Service, Pacific Northwest
Research Station, Forestry Sciences
Laboratory, 3200 SW Jefferson Way,
Corvallis, OR 97331, USA
Siu Wai Chiu
Department of Biology, Chinese University
of Hong Kong, Hong Kong S.A.R., China
:siuwaichiu@cuhk.edu.hk9
Régis Courtecuisse
Département de Botanique, Faculté des
Sciences Pharmaceutiques
et Biologiques, B.P. 83, F-59006 Lille
Cedex, France
:rcourtec@phare.univ-lille2.fr9
Tina Dreisbach
USDA Forest Service, Pacific Northwest
Research Station, Forestry Sciences
Laboratory, 3200 SW Jefferson Way,
Corvallis, OR 97331, USA
Susie Dunham
Department of Forest Science, Oregon State
University, Corvallis, OR 97331, USA
Alison Dyke
Woodland Ecology and Non Timber Forest
Products, 3F1 46 Broughton Road,
Edinburgh EH7 4EE
:alison.dyke@talk21.com9
Shelley E. Evans
Icknield House, 8 Saxonhurst, Downton,
Salisbury, Wilts SP5 3JN
:Shelley-Evans@myco-services.freeserve.co.uk9
L. V. Fleming
Joint Nature Conservation Committee,
Monkstone House, City Road, Peterborough
PE1 1JY, UK
:Flemin–v@jncc.gov.uk9
Leo M. Jalink
National Herbarium of The Netherlands,
University Leiden Branch, P.O. Box 9514,
2300 RA Leiden, The Netherlands
H. Kotiranta
Finnish Environment Institute, P.O.Box 140,
FIN-00251 Finland
:heikki.kotiranta@vyh.fi9
vii
viii
Contributors
Salvatore La Rocca
Dipartimento di Scienze Botaniche,
Università di Palermo, Via Archirafi 38,
I-90123 Palermo, Italy
Maria Ławrynowicz
Department of Algology and Mycology,
University of Łódź, Banacha 12/16,
PL-90–237 Łódź, Poland
:miklaw@biol.uni.lodz.pl9
R. K. Mibey
Department of Botany, University of
Nairobi, P.O. Box 30197, Nairobi, Kenya
D. W. Minter
CABI Bioscience, Bakeham Lane, Egham,
Surrey, TW20 9TY, UK
:d.minter@cabi.org9
Randy Molina
USDA Forest Service, Pacific Northwest
Research Station, Forestry Sciences
Laboratory, 3200 SW Jefferson Way,
Corvallis, OR 97331, USA
:rmolina@fs.fed.us9
Thomas O’Dell
USDA Forest Service, Pacific Northwest
Research Station, Forestry Sciences
Laboratory, 3200 SW Jefferson Way,
Corvallis, OR 97331, USA
Erast Parmasto
Institute of Zoology and Botany, Estonian
Agricultural University, 181 Riia Street,
51014 Tartu, Estonia
:e.parmasto@zbi.ee9
David Pilz
USDA Forest Service, Pacific Northwest
Research Station, Forestry Sciences
Laboratory, 3200 SW Jefferson Way,
Corvallis, OR 97331, USA
Maurice Rotheroe
Fern Cottage, Falcondale, Lampeter, Dyfed
SA48 7RX, UK
:m.rotheroe@btclick.com9
G. M. Siboe
Department of Botany, University of
Nairobi, P.O. Box 30197, Nairobi, Kenya
David Moore
School of Biological Sciences, 1.800
Stopford Building, The University of
Manchester, UK
:david.moore@man.ac.uk9
Jane Smith
USDA Forest Service, Pacific Northwest
Research Station, Forestry Sciences
Laboratory, 3200 SW Jefferson Way,
Corvallis, OR 97331, USA
Marijke M. Nauta
National Herbarium of The Netherlands,
University Leiden Branch, P.O. Box 9514,
2300 RA Leiden, The Netherlands
:Nauta@nhn.leidenuniv.nl9
Giuseppe Venturella
Dipartimento di Scienze Botaniche,
Università di Palermo, Via Archirafi 38,
I-90123 Palermo, Italy
:gvent@unipa.it9
Preface
Conservation is a major concern at the moment. In the second half of the
twentieth century, naturalists (including mycologists) became aware of a
general decline in natural habitats. Significant decline in the number of
species, and in the occurrence of species of fungi, was detected in the 1960s
in several countries in Europe and was correlated with changes in land
usage and with three consistent themes of environmental pollution:
eutrophication (contamination of water bodies with nutrients draining off
agricultural land), acid rain (mostly downwind of industrial smoke
plumes) and desertification (caused by shifts in precipitation patterns
resulting from climatic change). Since that time the general attitude has
changed to one in which managers try to improve biodiversity.
This book examines the various problems associated with fungal conservation. In different parts of the world there are several threats to fungi and
fungal diversity that prompt thoughts of conservation. However, it is not
self-evident whether and how fungi themselves can be conserved. Perhaps
the emphasis should be placed on conservation of the site, or the habitat,
or the host? All of these issues are addressed and debated here, but this
book goes beyond mere debate by filling the need for practical guidance for
management of nature in ways beneficial to fungi. Different parts of the
world experience different problems, so there are different solutions too.
Here we present a range of different case studies and describe the range of
regulatory and control procedures that are beginning to emerge. Taken
together, these give tips on how to include fungi in conservation projects in
a range of circumstances.
We believe that the combination of reviews of the research literature
with discussion of how to solve problems in the real world makes this book
virtually unique, at least in English. We hope, therefore, that it will be
found to be a succinct and practical guide about the subject – one that will
ix
x
Preface
be used just as much by nature managers, administrators and politicians
who have to make policy to deal with conservation issues, as by amateur
and professional mycologists who want to know more about the status of
the fungi.
The book has its origin in a British Mycological Society Symposium
which was organised by Bruce Ing, Maurice Rotheroe and Shelley Evans
at the Royal Botanic Gardens in Kew on 13 November 1999, entitled
Fungal Conservation in the 21st Century. These contributions have been
integrated with a selection of the most appropriate papers delivered at the
XIIIth Congress of European Mycologists, held in Alcala de Henares,
Spain, 21–25 September 1999. Importantly, the conference contributions
have been complemented by six specially commissioned chapters. Among
these are chapters that enhance the geographic representation of the
book’s contents so that it is truly global in its coverage. We have also
added two chapters that relate how national mycological societies can
contribute to fungal conservation and two editorial overviews. Chapter 1
attempts to provide some signposts for readers to help them get the best
out of the rest of the book. The final chapter is an attempt to convey in
writing some feeling of the discussions that occurred during the meetings.
We asked contributors to respond to the question ‘Are you optimistic or
pessimistic about fungal conservation in the 21st century?’ They reply from
different standpoints, but, thankfully, they all count themselves as optimists. So, read on and find out why!
David Moore
Marijke Nauta
Shelley Evans
Maurice Rotheroe
October 2000
1
Fungal conservation issues: recognising
the problem, finding solutions
DA V I D MO O R E , M A R I JK E M. N A U T A , S H E LL E Y
E. E V A N S & M A U R I C E R O TH ER O E
Nonbiologists may be excused for questioning whether microbial diversity
is really under threat. At a superficial level, micro-organisms seem to be
tolerant of almost any set of conditions thrown at them. Also, they appear
to have reproductive capacities able to generate populations of truly
astronomic numbers in very little time. However, that is a superficial
understanding and any belief that microbial species are not threatened is
simply wrong.
James T. Staley of the University of Washington gave his answer to the
challenge ‘Microbiologists are not concerned with endangered species, are
we?’ in a commentary published in 1997 (Staley, 1997). His simple answer
to this question is ‘Yes, some microbial species are threatened’, but the
argument Staley develops is interesting and has some valuable points for
mycologists to ponder. Even though the commentary was written largely
from the bacteriologist’s point of view, Staley mentions lichens and fungi
so it is clear that he does include even the mushrooms and toadstools
within his definition of micro-organisms. This is useful for us as we attempt
in this brief introductory chapter to highlight and provide cross-references
to the wide variety of aspects of fungal conservation that are included in
this book. We are not alone in the belief that such topics are important!
Indeed, Staley puts the level of importance very high. Micro-organisms
produced the original biosphere of Earth. ‘Not only have they made
conditions suitable for the evolution and existence of macroscopic life
forms, but they also continue to drive and profoundly influence many of
the essential biogeochemical cycles’ (Staley, 1997). Furthermore, most of
the present-day biodiversity among the eukaryotes is microbial, being
generated by the protists, algae and fungi. Bacteria, of various sorts,
provide the biodiversity within the prokaryotes, of course. So the consequence is that ‘the tree of life is largely a tree of microorganisms . . . much
1
2
D. Moore, M. M. Nauta, S. E. Evans & M. Rotheroe
of the diversity on Earth is microbial with the plants and animals appearing as small, terminal branches’ (Staley, 1997).
This fundamental importance of micro-organisms can hardly be doubted, so why is there so little general interest in conservation of microbes?
Staley (1997) puts this down to something he calls ‘kinship’, claiming that
humans share strong kinship with many animals and plants, a kinship
which can blossom into fondness for closely related and ‘warm and fuzzy’
animals. Microbes, though, are generally too small to be noticed much by
humans, even though human lives are daily more closely intertwined with
microbes than with any other organisms. Further, microbes evoke negative feelings because they are associated with disease and spoilage. Finally,
there is a general ignorance about the degree to which our daily lives
depend on the beneficial activities of many microbes – from sewage sludge
through to agriculture, and the making of bread, and antibiotics and other
life-saving drugs. ‘Because microorganisms rank so low on the kinship
scale, the demise of a microbial species is not an emotional issue for
humans’ (Staley, 1997). We do not expect many to rally to a cry to ‘Save
the whale’s intestinal microbes’!
Staley (1997) suggests that the general phenomenon is that a microorganism is threatened when its ecological niche is threatened. Consequently, ‘the most satisfactory manner in which to preserve the organisms
is through protection of the environment and thereby the natural community itself ’ (Staley, 1997). However, Staley acknowledges that ‘we have
described so few species; many species may be threatened whose existence
are still unknown.’ And his final conclusion is that ‘Our knowledge of
microbial diversity . . . is so meagre that we do not yet know if and when
most species are threatened. . . . Our very inability to answer the question
of threatened microbial species cries loudly for the need for microbial
systematists and ecologists to begin to address the exciting challenges
regarding our knowledge of the extent of microbial diversity on Earth’
(Staley, 1997).
That brings us to the first point we wish to highlight from this book. It is
most succinctly stated by David Minter in Chapter 16 (p. 193): ‘In many
parts of the world mycologists are an endangered species. It follows that
fungal conservation can only occur if mycologists are conserved.’ But
other authors express similar opinions. Régis Courtecuisse (p. 10) puts it
this way: ‘Incidental problems and questions around inventories which
have to be considered are (a) promoting the conservation of taxonomists
themselves . . . ’ and Eef Arnolds (p. 77) like this: ‘It is obvious that
conservation of fungi depends on the input of mycologists. But at present it
Fungal conservation, the problem and solutions
3
seems to be also the other way around: the future of mycologists depends
on their input in conservation.’ The root problem is that our level of
ignorance is so great that we do not have the numbers of experts needed to
make serious contributions to knowledge of species sufficiently quickly to
conserve those species. ‘Taxonomists are scarce because of a shift in
academic programmes toward molecular systematics and ecology’ (Randy
Molina et al., p. 39). One might also add that for several years now funding
agencies around the world have been operating a similarly skewed funding
policy. Mycological research is rarely funded, anyway, because a lower
value judgement is placed upon it than is applied to similar research on
lower animals or lower plants. Another aspect, perhaps, of the lack of
kinship to which reference is made above. In the long term these attitudes
must change and the importance of the kingdom of fungi recognised
sufficiently to assure equitable funding for its study. To a very large extent
this is a matter of public education and several of our authors mention this.
Régis Courtecuisse mentions the need for public education (Chapter 2,
p. 14), and David Moore and Siu Wai Chiu claim that ‘Education is the
key’ in China (Chapter 9, p. 118).
It will take a long time for an education policy to result in significantly
more experts with attitudes changed sufficiently for the value of fungal
biology to be fully appreciated. In the meantime we have the real world to
deal with – a real world in which those mycologists who do exist may be
prevented from making a full contribution by poor infrastructure or
political and economic isolation. David Minter, in Chapter 14 (p. 164),
illustrates how effective voluntary help (in this case through provision of
second-hand computers) together with intergovernmental assistance
(through the UK Government’s Darwin Initiative Programme) is enabling
Ukrainian mycologists to complete the databases and surveys that are
essential to effective national conservation policies. He tells a similar story
in Chapter 16 (p. 192), although in this case Cuban mycologists are
suffering the inevitable shortages and isolation resulting from a unilateral
economic blockage imposed by the USA. Again, provision of resources
(another Darwin Initiative Project) enables local mycologists to progress
towards a national fungal conservation strategy.
For more immediate input, particularly to projects under way now, ‘The
depleted ranks of classical fungal taxonomists can be augmented, however,
by a cadre of experienced parataxonomists, people with less formal schooling in mycology, who are trained and gain significant experience in fungal
identification’ (Molina et al., Chapter 3, p. 39). Similar ideas, perhaps,
emerge from the Dutch experience in raising interest which Leo Jalink and
4
D. Moore, M. M. Nauta, S. E. Evans & M. Rotheroe
Marijke Nauta suggest makes it evident that ‘managers need clear instructions’ about mycologically valuable sites (Chapter 6, p. 90).
If the information is provided, landowners, managers and administrators have considerable sympathy for including fungi in their conservation
management. Indeed, it seems a sensible strategy for mycologists to be
proactive in establishing collaborations with those involved in land management and, especially, with groups concerned with conservation of other
organisms (see Martin Allison, Chapter 12, p. 144). There is certainly no
excuse for mycologists being short of cogent arguments for inclusion of
fungi in conservation schemes. Randy Molina et al. (Chapter 3, p. 23)
detail the four themes that need to be emphasised when ‘educating land
managers . . . is vital’. Additional ready-to-use material can be found in
Chapter 17, in which Cannon et al. discuss, largely from the point of view
of population ecology, ‘Why are fungi difficult to conserve’ (p. 198) and
‘Why are fungi important’ (p. 199).
We know very little about fungal population biology; in fact, even less
about fungal population genetics. Randy Molina et al. (Chapter 3, p. 25)
discuss the role of fungi in communities and describe projects aimed at
determining the population genetics of representative species (p. 33). Related to this is the detailed analysis of the population biology of Lentinula
edodes that shows how the traditional cultivation method in China (especially outdoor cultivation accompanied by harvesting at maturity) is
likely to endanger both the cultivars and the wider gene pool of the wild
mushroom (David Moore and Siu Wai Chiu, Chapter 9, p. 113).
The main tools available to the fungal conservationist are outlined first
by Régis Courtecuisse (Chapter 2, p. 10) to be inventories (checklists),
mapping programmes, and Red Data lists. These being the crucial aspects
of fungal conservation, they appear in some guise in all chapters. Particularly helpful discussions can be found in Chapters 3 (p. 35), 4 (p. 70), 5
(p. 83), 6 (p. 90), and 17 (p. 202). Eef Arnolds (Chapter 4, p. 66) also
discusses the species concept – an important issue for any survey, whilst
Molina et al. (Chapter 3, p. 43) describe ‘habitat modelling’ as a tool in
conserving fungal resources. Examples of survey work are given in Chapters 3 (p. 19), 5 (p. 81), 6 (p. 89), 7 (p. 95), 9 (p. 111), 10 (p. 120), 11 (p. 136),
13 (p. 156), 15 (p. 177), 16 (p. 182), and 17 (p. 197).
Surveys and mapping programmes culminate in the production of Red
Data lists. Although ‘Red Data’ in this phrase usually carries with it the
danger connotation commonly linked with the colour red, it’s important to
remember that in this case the word is an acronym, the full phrase being
Rarity, Endangerment and Distribution Data lists. This is important
Fungal conservation, the problem and solutions
5
because the full phrase shows explicitly the amount of information which is
required to make the judgement about whether or not to include a species
in a Red Data list. Red Data lists are discussed to some extent in most
chapters, especially those already highlighted as dealing with surveys.
However, Maria Ławrynowicz (Chapter 7, p. 96) shows how different
national Red Data lists can be integrated to reach wider conclusions, while
Giuseppe Venturella and Salvatore La Rocca (Chapter 13, p. 156), and
Heikki Kotiranta (Chapter 15, p. 177) illustrate how local surveys can be
compared, on the one hand with an international Red Data listing, and on
the other hand with international Red list categories.
Conservation strategies emerge at a variety of levels and provide
examples which might be applicable elsewhere. Molina et al. (Chapter 3,
p. 20) outline the US Federal laws regulating forest management, mentioning the different goals of the different agencies involved. A different set of
conflicts (and their resolution) discussed by Martin Allison (Chapter 12,
p. 153) is that which can arise ‘within conservation management when one
group of animals or plants is favoured above another.’ Vincent Fleming
(Chapter 18, p. 209) details the UK response to the Convention on Biological Diversity – essentially the administrative mechanics of conservation in
the UK. Below the governmental level, David Moore (Chapter 20, p. 223),
and Marijke Nauta and Leo Jalink (Chapter 21, p. 242) show how two
national mycological societies (the British and Dutch mycological societies
respectively) have reacted and developed programmes aimed at conserving
fungi. In Chapter 19 (p. 219) Alison Dyke reports how a purely voluntary
code of practice has been established directly by the groups involved in
wild mushroom harvesting in Scotland. A range of wild harvested fungal
fruit bodies command prices that make them worth shipping over intercontinental distances, as discussed by David Arora (Chapter 8, p. 105), so
this code of practice may be applicable elsewhere. In contrast, the commercial harvest of edible forest mushrooms is controlled by Federal laws in the
United States (Molina et al., p. 46, and see Eef Arnolds, p. 76).
The Scottish Mushroom Forum’s code of practice (Table 19.3, p. 221) is
one of several examples of specific advice and instruction included in this
book. Others are a ‘set of summary statements’ for use when ‘planning and
conducting conservation efforts for fungi’ (Molina et al., Chapter 3,
p. 54); some management guidelines from Leo Jalink and Marijke Nauta
(Table 6.2 and Table 6.3, p. 93); and British Mycological Society codes of
practice (Chapter 20, p. 235).
With these, and other, explicit pieces of advice based upon practical
experience, we hope that this book will make a constructive contribution to
6
D. Moore, M. M. Nauta, S. E. Evans & M. Rotheroe
fungal conservation. It is a global problem and we include examples from
Finland in the North to Kenya in the South, and from Washington State,
USA, in the West to Fujian Province, China, in the East. Our authors
identify threats faced by fungi of all types. Inevitably, even though ‘It is
probably true to say that the majority of fungi would be describable as
‘‘microfungi’’ ’ (Paul Cannon et al., Chapter 17, p. 197) descriptions of
work with larger fungi – truffles and mushrooms – tend to predominate.
The balance, of course, is governed by the research which is being done and
the research interests of those doing it.
Our authors also suggest solutions ranging from voluntary agreements,
through ‘fungus-favourable’ land management practices, and on to primary legislation. We have to stress that this book cannot give ready-made
solutions to all the problems that might arise concerning conservation of
fungi. What we have assembled is a set of descriptions of how far we have
got with conservation of fungi, with some focus on the bottlenecks that
remain, and with a range of guidelines that may help in improving conservation of fungi in the future. The bottom line, though, is quite clearly that
‘Conservation of fungi is, like conservation of other organisms, in the very
first place conservation of their habitats combined with adequate management’ (Eef Arnolds, Chapter 4, p. 72). Save the world and we’ll save the
fungi with it. Conserve the fungi and your one and only planetary home
will be equally safe.
Reference
Staley, J. T. (1997). Biodiversity: are microbial species threatened? Current
Opinion in Biotechnology 8, 340–345.
2
Current trends and perspectives for the
global conservation of fungi
R ÉGI S C OURTECUI SSE
Introduction
For rather more than a year, the specialist group for fungi within the
Species Survival Commission (SSC) of the International Union for the
Conservation of Nature (IUCN) has been revived after interruption in its
activities between 1995 and 1998. As a member of the European Council
for the Conservation of Fungi (ECCF) standing committee, I shared with
my ECCF colleagues the regret that fungi were no longer taken into
account within the IUCN. So we decided to take advantage of the June
1998 Planta Europa meeting, in Uppsala, to establish new contacts with the
IUCN and to revive this specialist group.
Within the new specialist group for fungi, the ECCF serves as the main
framework since it has accumulated much data and experience on the topic
since 1984. But I am also trying to federate further mycologists involved in
conservation outside Europe so that the group will consist of a genuinely
enlarged and international network.
In this chapter I will give a brief description of the present global state of
knowledge concerning fungal conservation and indicate the main priorities
we should consider for the future. Of course, this owes a great deal to the
ECCF heritage, especially through the decisive contributions from some of
its members, such as Eef Arnolds and others. I also received feedback from
some members of the new group but I do not claim that the literature
survey is in any way exhaustive and I do realise that some points may not
be represented here. The specialist group of which I serve as chairman is
currently still ‘under construction’ and we attach great importance to
communication between all people who are interested or involved in
fungal conservation and depend on them for further information.
7
8
Régis Courtecuisse
Current status and problems
Conservation status of fungi
Fungi are very seldom legally protected. Such a situation does exist in
Slovakia, where 52 species have a ‘special legal status’, enabling managers
to prevent damage to their habitats (Lizon, 1999). Where no legal status is
available, some voluntary efforts have been made to produce codes of
practice or have advisory documents published, stressing the importance
of fungal conservation and summarising the recommendations to achieve
that. This has been done, for example, in Switzerland (Egli et al., 1995),
and in the United Kingdom (English Nature, 1998; and see Chapters 18 to
21).
Nevertheless, even without a legal conservation status, Red Data lists
have been published, or are currently in preparation, in many countries,
especially in Europe. This demonstrates the increasing concern of mycologists for this topic. At the same time, more and more nature managers pay
increasingly greater attention to fungi.
So, the idea that fungi may be threatened and deserve special attention
to their conservation is now well established, at least in Europe, where
books and symposia have been devoted to the topic in relation to environmental problems (Frankland, Magan & Gadd, 1996; Rotheroe, 1996a).
But this is not yet true everywhere.
Current problems
The main threats and causes of decline in fungi world-wide can be
categorised as resulting either from global, or specific or local problems.
Air pollution, considered globally, may influence fungi through the greenhouse effect producing a slow climate change, as well as exerting an
indirect effect through modification of vegetation. Such changes might
threaten climate-sensitive species and/or favour the development of more
thermophilic taxa which could in turn act as alien competitors against
native species. The balance between parasites and their hosts might also be
changed (Lonsdale & Gibbs, 1996; Pettitt & Parry, 1996). Some northward
migrations have been demonstrated in recent years, especially in the western part of Europe. Elevation of the sea level could also become a problem
for species inhabiting coastal ecosystems (Rotheroe, 1996b). Another global problem widely recognised to be of paramount importance in the
decline of fungi world-wide is the destruction of habitats and the dramatic
felling of forests, particularly in the tropics, but also representing a poten-
Global conservation of fungi
9
tial problem in more temperate or even sub-boreal to boreal forests.
Some problems with a more local or specific effect on fungi include: air
pollution, which, whatever the scale, causes well-documented species decline; deposition of various pollutants, leading to soil modifications (the
mechanism of which is not clearly understood); accumulation of metals
and other pollutants interfering with fungi, either macrofungi (Ing, 1996)
or microfungi (Boddy et al., 1996; Magan, Smith & Kirkwood, 1996).
Fragmentation of habitats is also a major problem. It has been shown (e.g.
Chaumeton, 1994) that fragmentation of habitats makes it difficult for
some species to maintain normal population. Such a situation often results
from forest cutting, urban extension, evolution of land uses and change of
agricultural practices. In particular, modern agriculture increasingly uses
chemical treatments that can give rise to various kinds of environmental
modifications. This problem is especially well documented in Europe for
grassland fungi, both macromycetes such as Hygrocybe spp. (Rotheroe et
al., 1996) but also soil fungi (Bardgett, 1996). A more specific problem
concerns edible species. When harvesting is done by individuals or families, it seldom leads to concern for survival of the species (Egli, Ayer &
Chatelain, 1990). On the other hand, the commercial harvesting of the
same species causes great concern, as is evident from the numerous publications on the subject, especially in North and Central America (Pilz &
Molina, 1996, 1997; Palm & Chapela, 1997; Rowe, 1997). This is a controversial topic. Some people claim that heavy harvesting is harmless (Arora,
1999a, 1999b) and others claim the opposite (Rotheroe, 1998), depending
mainly on the harvesting tools, but also taking into account the longer
term, for which we have insufficient background knowledge to know what
will really happen in the future. Furthermore, there is currently little
scientific evidence about the impact of harvesting on the mycelium itself
and more studies are needed on that topic. Suggestions for voluntary and
regulatory control have been published also in Europe, such as Switzerland (Keller, 1991), the United Kingdom (Leonard, 1997a,b) or some parts
of France. ECCF members from eastern Europe complain about the
ecological damage caused to their forests by commercial harvesting of
edible fungi that are exported to western Europe (see, for example, Ivancevic, 1998; Pop, 1998). Further problems arise from the harvesting of
edible chanterelles (in part undescribed species) from Africa, tonnes of
which are imported each year into France, for example. A similar problem
may be faced by the decorative tropical polypores that are used in floral
compositions. We need to know more about the actual places and conditions of harvesting of these species.
10
Régis Courtecuisse
What can we do?
Looking at these causes of fungal decline, we might feel discouraged
because, as fungal conservationists, we have no opportunity to manage
global warming or global forest felling. Nor are we likely to have much
influence on the politics that affect agricultural practices or industrial
pollution. It is not even clear how we should handle the harvesting of
edible fungi because important economic interests are involved – commercial interests, of course, but sometimes also at the individual family level,
thus involving a social dimension in the problem. What we can actually do,
at our ‘mycological level’, concerns fungi in the field, and is related to our
knowledge about the ecological and heritage value of fungi. The ‘heritage’
dimension of fungi in this context refers to their potential for indicating the
holistic natural value of a given site or habitat in a way that integrates
many inputs including historical, nature management, biodiversity, and
conservation aspects. It is our task to popularise the value of this heritage
quality of fungi and to use it as a force in nature conservation. For that, we
have some useful tools at hand.
Our tools
First are inventories. In most of the countries of the world, what are
basically lacking are checklists of fungi. This is not only true of tropical or
developing countries, but also of the first world’s great nations. The
situation is even worse at the continental level. A project to initiate a
European checklist has been urged by the IUCN and is currently being
considered by the ECCF for funding in the near future. The first priority in
our tasks as fungal conservationists is to promote fungal inventories.
Incidental problems and questions around inventories which have to be
considered are (a) promoting the conservation of taxonomists themselves,
(b) developing arguments in favour of such inventories, (c) using a reasonable consensus taxonomy which is discerning enough to maintain the
ecological bioindicative value of the taxa recognised.
Second are mapping programmes. Inventories provide an idea of the
fungal diversity: mapping programmes should yield useful information
about the rarity of species, their eventual decline and finally their heritage
value. Mapping is the second important task we must promote as fungal
conservationists. Inventories and mapping programmes are intimately
connected, of course. Mapping programmes are conducted in many countries where inventory data are sufficient, as well as some in which inventory
Global conservation of fungi
11
data are not sufficient, for example in Australia where an interesting and
ambitious project is already running (:http://calcite.apana.org.au/
fungimap/9). In Europe the ECCF is currently organising a mapping
programme of endangered species (Otto & Ohenoja, 1998).
Third are Red Data lists. On the basis of inventory and mapping
programmes, mycologists are able to prepare Red Data lists for various
geographical or political units. This is the third fundamental task for
fungal conservationists. Red Data lists have appeared mainly in Europe, at
the continental as well as national level (Ing, 1993a,b) but also recently in
the USA, for example, with a preliminary list for Oregon (Castellano,
1997). Red Data listing raises many problems, especially the crucial choice
of criteria for putting species on the list or not. Several models are in
current use but there is a strong desire to use a homogeneous system of
criteria, at least at the national level where comparability with neighbouring countries is a priority. We prefer to use the IUCN criteria although the
last edition of these (1994) was clearly made for big African mammals such
as elephants or rhinoceros. Criticism of these criteria has been quite strong
among mycologists and some other naturalists, and attempts are currently
being made to revise and adapt them to fungal characteristics (Arnolds,
1998; Kotiranta, 1998, 1999). Some interesting ideas promoting the use of
unusual but useful criteria are also in current development (Courtecuisse,
1997; and see Chapter 18). A special commission has been established
within the ECCF for that purpose, the work of which is currently ongoing.
Conservation strategies
The results of such programmes, even if still quite fragmentary, enable us
to consider fungal conservation and to include fungi in nature conservation. An important point is that, as widely admitted, fungi cannot be
protected individually, as can sometimes be done with plants or animals,
by publishing lists of legally protected species. The biology of fungi, the
frequent difficulty that even trained people have in identifying them, and
other peculiarities, do not fit with this option.
In situ conservation by conserving natural habitats
The most important tool for fungal conservation is the conservation of
their habitats. This was cited as the first priority by all the contacts I
consulted while preparing this chapter, from Europe and from overseas
and tropical countries as well.
12
Régis Courtecuisse
So the first strategy we must develop is to promote the conservation of
natural habitats, especially in view of the current tendency to replace
native forests by alien trees that are more valuable timber producers, a
problem also in some tropical countries. More locally, some important
places may be in need of protection, partly on a mycological basis (see for
example Rotheroe, 1995).
In proposing a hierarchy of habitats for the purpose of fungal conservation, we must consider whether or not we have enough mycological data. If
such data are available, interesting methods currently being developed can
be used to ‘weight’ species according to their heritage value, inferred from
their status in Red Data lists for example. Such an approach has been
developed in The Netherlands (see Chapter 7) and also in France (Courtecuisse & Ansart-Chopin, 1997; Courtecuisse & Blot-Quénu, 1998).
Where sufficient data are unavailable, a provisional position is to concentrate on habitats that are interesting and rich for other taxonomic groups,
plants for example, with which fungi are frequently associated. If the
congruence of vegetation communities with communities of different types
of fungi is high enough, conserving the plant community will ‘take along’
all the associated fungi, even including the undescribed species. Other
methods for using incomplete inventories are being developed, especially
in France.
In situ conservation on mycological reserves
The concept of mycological reserves is also interesting since it promotes
the conservation of habitats which are especially valuable for fungi and
which are not already considered for conservation otherwise. Such reserves already exist in some European countries and much work is being
done at the moment in France on this concept. Much remains to be done as
to the desirable size of such mycological reserves so that they prove to be
efficient for fungal populations (see, for example, Senn-Irlet, 1998).
In situ conservation with ecological corridors
In relation to problems of habitat fragmentation, the concept of biological
corridors should also perhaps be considered for fungi. As far as I know,
very few studies have been dedicated to this kind of research in mycology.
Only Alexander & Watling (1987) considered the role of Betula within
Sitka spruce forests. Biological corridors have strongly positive effects on
the survival and even on the restoration of populations of various plants
Global conservation of fungi
13
Table 2.1. Current membership of the SSC/IUCN Specialist Group for
Fungi
Chair
Secretary
Ordinary members
Régis Courtecuisse (France)
Claudia Perini (Italy)
Anders Bohlin (Sweden)
Peter Buchanan (New Zealand)
Julietta Carranza (Costa Rica)
Ignacio Chapela (Mexico)
Shelley Evans (UK)
Ana Franco (Columbia)
Bruce Ing (UK)
Teresita Iturriaga (Venezuela)
Heikki Kotiranta (Finland)
Pavel Lizon (Slovakia)
Jean Lodge (USA – Porto Rico)
Tom May (Australia)
Greg Mueller (USA)
Marijke Nauta (The Netherlands)
Lorelei Norvell (USA)
Scott Redhead (Canada)
and animals. It is quite likely that they may also have a positive effect on
fungal populations.
Ex situ conservation
In some cases, especially for saprotrophic species growing in culture, ex
situ conservation programmes could be developed. A good example of this
has been shown for Pleurotus nebrodensis in Sicily (Venturella, 1999) and
more generally by Homolka et al. (1999). This possibility might be of great
importance for species on the verge of extinction or those strictly confined
to very limited or threatened habitats. This concept is rather similar to the
one used for plants, through seed banks.
The SSC/IUCN specialist group for fungi
After reviewing the problems and some possibilities for conserving fungi, I
would like to add some words on the SSC/IUCN specialist group for fungi,
its composition and its role, at least as I understand this.
The composition of this group at the time of writing is shown in Table
2.1. As can be seen, the group lacks members from Africa and Asia. I have
14
Régis Courtecuisse
had contacts with several colleagues from these continents but, at the
moment, no name is readily available. We hope, of course, that the group
will be enlarged in the future.
Future tasks
The main roles of this specialist group are to promote ideas about fungal
conservation and its various implications and also to promote research in
the different possibilities related to the goals and tools I have summarised
here. As chairman of this group, I will encourage research in the crucial
areas of inventories, mapping programmes and Red Data listing programmes. These topics represent the basic research we have to develop in
all countries.
We will also have to develop specific research about the objective
definition of interesting and priority habitats for fungi. Further research is
also necessary and urgent to develop and promote the use of fungal
bioindicators of environmental quality as a tool to be used by nature
managers. The research concerning indicator species of old and primeval
forests (see Parmasto & Parmasto, 1997; Tortic, 1998; O’Dell, Kiester &
Molina, 1999, among other examples) must be continued in this respect.
We should also strengthen the promotion of the fungal role, especially
mycorrhizal species, in rehabilitation of polluted (e.g. Allen, Espejel &
Sigüenza, 1997) or weakened (Janos, 1996) areas. Further fundamental
study is needed on special topics like the conservation of parasitic fungi
which are also important for the holistic functioning of ecosystems, or the
role of natural history collections in fungal and nature conservation (e.g.
May, 1998).
It is necessary, too, to establish, strengthen and regularise contacts with
other naturalists involved in conservation, in order to exchange data and
create synergistic arguments for conservation of globally interesting habitats or places which can be made to administrations and decision holders
at any level, local, regional, national and supra-national. For example, at
the European level, contacts with the Bern convention structure are currently in progress and will hopefully soon lead to the long-awaited integration of fungi into the Bern Convention appendix. At a different level, we
should also reflect on social problems, like the consequences of putting
controls on harvesting of edible fungi, for example.
Finally, one of our tasks will be to build an efficient network for
developing public education. Communication between the members of our
group must be made easier – of course through electronic tools but also
Global conservation of fungi
15
through the journal Fungi and Conservation Newsletter, three numbers of
which were published in the past by the IUCN and which I intend to try to
revive.
Conclusion
As a conclusion, I would like to thank and congratulate the friends and
colleagues who have conducted or chaired the ECCF so far. The work of
these people has provided a strong framework for fungal conservation that
remains to be developed fully in Europe, where the idea was born, and to
be used as a model elsewhere, with inevitable local adaptations. That is
what I will try to do in the course of my position as chair of the ECCF at
the European scale on the one hand and of the SSC/IUCN Specialist
Group for Fungi on a wider basis on the other. I trust that continued
friendly collaboration will yield further positive developments and progress in fungal conservation for the new century.
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3
Conservation and management of forest
fungi in the Pacific Northwestern
United States: an integrated ecosystem
approach
RANDY MOLINA, D AVI D PI LZ, JANE SMIT H,
SUS IE D U N H A M , TI NA D R E I S B A C H , T HO MA S
O’DELL & MICHAE L C ASTELL ANO
Introduction
The vast forests of the Pacific Northwest region of the United States, an
area outlined by the states of Oregon, Washington, and Idaho, are well
known for their rich diversity of macrofungi. The forests are dominated by
trees in the Pinaceae with about 20 species in the genera Abies, Larix,
Picea, Pinus, Pseudotsuga, and Tsuga. All form ectomycorrhizas with
fungi in the Basidiomycota, Ascomycota, and a few Zygomycota. Other
ectomycorrhizal genera include Alnus, Arbutus, Arctostaphylos, Castinopsis, Corylus, Lithocarpus, Populus, Quercus, and Salix, often occurring as
understorey or early-successional trees. Ectomycorrhizal fungi number in
the thousands; as many as 2000 species associate with widespread dominant trees such as Douglas-fir (Pseudotsuga menziesii) (Trappe, 1977). The
Pacific Northwest region also contains various ecozones on diverse soil
types that range from extremely wet coastal forests to xeric interior forests,
found at elevations from sea level to timber line at 2000 to 3000 metres.
The combination of diverse ectomycorrhizal host trees inhabiting steep
environmental and physical gradients has yielded perhaps the richest
forest mycota of any temperate forest zone. When the large number of
ectomycorrhizal species is added to the diverse array of saprotrophic and
pathogenic fungi, the overall diversity of macrofungi becomes truly staggering.
Issues relating to conservation and management of forest fungi in the
19
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R. Molina et al.
Pacific Northwest must be placed in the context of public land resources
and Federal laws regulating forest management. The Pacific Northwest
has the greatest amount of Federal forest in the United States, about 9.8
million hectares in Oregon and Washington alone. The lands are managed
by the US Department of Agriculture (USDA), Forest Service, and the US
Department of the Interior (USDI), Bureau of Land Management and
National Parks Service. Each agency has different overall management
goals. For example, National Parks do not allow commercial extraction of
any forest products. National Park lands are primarily for recreational
purposes and also serve as primary biological reserves. Other Federal
forest lands vary between roadless wilderness areas with virtually no
resource extraction to areas where forests are managed to meet multipleresource objectives, including timber harvest, recreational access, and
conservation of rare species. Several Federal laws and statutes, including
the Endangered Species Act, the National Environmental Protection Act,
and the National Forest Management Act, provide strict rules governing
the planning and implementation of forest management and monitoring
activities. These laws also provide for public input in planning and management. Over the last four decades, legal challenges to the development
and implementation of forest plans have greatly shaped the current directions for managing public forests. These directions and decisions have
direct relevance to the conservation and management of forest fungi.
During the late 1980s and early 1990s a series of lawsuits sharply
curtailed the harvest of old-growth forests in the Pacific Northwest. At the
centre of the controversy was the old-growth dependent northern spotted
owl (Strix occidentalis), a rare species protected by the Endangered Species
Act. Because the economy of many Pacific Northwest communities relied
strongly on timber harvest from Federal forests, this action created political controversy. In 1992, President Clinton held a regional forest conference as a first step toward balancing the economic needs of the region with
the protection of endangered species and old-growth forest habitat on
Federal land. This eventually led to a scientific assessment of the problem
(FEMAT, 1993) and development of the Northwest Forest Plan (USDA &
USDI, 1994a). The plan strives to provide for a sustainable level of timber
harvest and persistence of old-growth forest dependent species based on a
region-wide allocation of land uses, including reserve and nonreserve
areas. As part of the overall scientific assessment process, many rare
species requiring old-growth forest habitat were analysed for protection
under the land reserves and forest management guidelines. In the final
analysis, 234 old-growth dependent, rare, fungal species were listed for
Conservation in the Pacific Northwestern United States
21
additional protection under the Survey and Manage guidelines of the
Record of Decision (USDA & USDI, 1994b). The Record of Decision is
the final legally binding document that defines how the Northwest Forest
Plan is to be implemented. The Survey and Manage guidelines detail the
course of mitigations required by management to protect the species.
Although not a part of the listing process for the Endangered Species Act,
this action still represents the first Federal listing of fungal species for
protection in the United States and highlights the need for a research
programme addressing poorly understood conservation issues for forest
fungi.
The Northwest Forest Plan initiated a significant paradigm shift in
forest management from an emphasis on timber extraction to ecosystem
management. Ecosystem management is a holistic approach to decision
making that integrates biological, ecological, geophysical, silvicultural,
and socio-economic information into land management decisions to preserve biological diversity and maintain ecosystem functioning while meeting human needs for the sustainable production of forest products and
amenities (Bormann et al., 1994; Jensen & Everett, 1994). Planning is
typically conducted at broad scales such as watersheds or regional landscapes. Fungi play significant ecosystem roles in nutrient cycling, food
webs, forest diseases, and mutualisms, and therefore significantly influence
seedling survival, tree growth, and overall forest health. Ecosystem approaches to forest management provide an avenue for integrating our
understanding of the biological and functional diversity of forest fungi
into recent forest management objectives. It is our contention, and indeed
the theme of this chapter, that an ecosystem-based approach provides the
most effective means for conserving this vast array of essentially cryptic
species.
As the numbers and diversity of fruit bodies of forest fungi were declining in Europe (Arnolds, 1991) and Japan (Hosford et al., 1997), the last
two decades in the Pacific Northwest witnessed a dramatic increase in the
harvest of wild, edible mushrooms with export to Europe, Japan, and
elsewhere. In 1992, nearly 2 million kg of edible mushrooms was harvested
in the Pacific Northwest, bringing over US$40 million to the economy
(Schlosser & Blatner, 1995). Forest managers and the public questioned
whether such harvest levels were sustainable or could negatively impact
persistence of harvested species (Molina et al., 1993). The reproductive
biology of these species, their habitat requirements, and effects of mushroom harvest and forest management practices on their continued productivity are poorly understood. Although commercial species are not rare,
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the conservation of them also lends itself to ecosystem management approaches and has prompted new lines of research.
The overall mission of the Forest Mycology Team of the USDA Forest
Service, Pacific Northwest (PNW) Research Station, Corvallis, Oregon, is
to provide basic information on the biological and functional diversity of
forest fungi and to develop and apply this information in an ecosystem
context to both conserve the fungal resource and sustain the health and
productivity of forest ecosystems. The research is divided into three problem areas:
1. Ecology, biology, and functional diversity of forest fungi This
area considers issues related to fungal taxonomy, systematics
and phylogeny, understanding and documenting fungal
communities in space and time, examining effects of natural and
anthropogenic disturbances on the ecology of forest fungi, and
exploring important ecosystem functions of forest fungi,
particularly their roles in ecosystem resiliency and health.
2. Conservation biology of forest fungi This area develops
fundamental approaches to determining population dynamics,
develops habitat-based models to predict species presence and
future potential habitat for rare fungi, develops methods to
survey for rare species, and explores management options to
conserve old-growth associated fungal species under the
guidelines of the Northwest Forest Plan.
3. Sustainable productivity of commercially harvested edible forest
fungi This area addresses challenges associated with measuring
productivity of ephemeral fruit bodies, determines the effects of
mushroom harvest and forest management practices on future
production, and develops protocols to monitor sustainable
productivity of the resource.
This approach addresses fungal resource issues at different scales (species, populations, communities, and landscapes) so that the results are
relevant to ecosystem planning. The beneficial mutualistic mycorrhizal
fungi are given primary attention because of their importance to forest
trees. The remainder of this chapter outlines many issues in these areas,
our research approaches and directions, and application of this research to
the conservation and management of forest fungi. We begin by discussing
basic research components, including ecosystem function, community
ecology, and population biology. We conclude by focusing on the development of tools and concepts to apply this knowledge, including applications
Conservation in the Pacific Northwestern United States
23
in inventory and monitoring. Although this research programme is unique
to the Pacific Northwest region and the Federal policies on public lands,
we believe it provides a relevant framework to approach fungal conservation issues at various scales and for different objectives in other parts of the
world.
Ecosystem functions, fungal communities and populations
Ecosystem functions
Conserving and managing forest fungi requires an understanding of how
fungi function in maintaining healthy ecosystems. Educating land managers about how management practices affect the reproduction, growth,
and overall function of fungi through workshops and literature is vital to
the success of integrating fungal concerns into forest management plans.
In our discourse with managers, we emphasise and discuss four critical
ecosystem functions as follows.
1. Nutrient cycling, retention, and soil structure Almost all fungi
participate in nutrient cycling. This is best known for
saprotrophs (decomposers), particularly those that degrade
wood (cellulose and lignin). Two other roles are equally
important, however, in this ecosystem function. First, fungal
mycelia are large sinks for organic carbon and nutrients in the
soil. In ectomycorrhizal-dominated forests, a significant portion
of the carbon fixed by host trees can go to the mycorrhizal
structures and fungal mycelium to support this symbiotic
association (Smith & Read, 1997). Some fungal mycelia and
ectomycorrhizal root tips turn over rapidly to return nutrients to
the soil, whereas other fungal structures such as rhizomorphs
retain nutrients in relatively long-lived structures or shunt it to
other parts of the mycelium as some sections die. What is
important is that fungi are major sinks for maintaining nutrients
on site, thereby preventing nutrient leaching and loss from soils.
Secondly, fungi exude polysaccharides similar to root exudates,
and these ‘organic glues’ (Perry et al., 1989) are important in
creating and stabilising soil microaggregates and soil
micropores. These soil structures contribute to soil aeration and
water movement. Because most terrestrial organisms are
strongly aerobic, this fungal contribution to soil
microaggregation is critical for overall maintenance of soil
health and productivity.
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2. Food webs Although it is common knowledge that animals
from molluscs to mammals (including humans) eat mushrooms,
the magnitude of this food source and its ecological importance
is underappreciated. The greatest biomass of fungi resides in
hidden mycelia that explore soil and organic substrates, and it is
here that the importance of fungi in food webs and soil fertility is
revealed. Literally tens of thousands of microarthropod species
live in forest soils, and about 80% of those are fungivores
(Moldenke, 1999). Microarthropods provide the biological
machinery to shred the vast quantity of organic matter in forest
soils and prepare it for the final mineralisation processes carried
out by microbes. Given the large amount of fungal hyphae in
forest soils noted above, the magnitude of this soil food web
process becomes readily apparent.
Fruit bodies of fungi (mushrooms and truffles) are also vital
food sources for many wildlife, especially forest mammals. Most
small mammals depend on hypogeous fruit bodies (truffles) for a
significant part of their diets (Maser, Trappe & Nussbaum,
1978). The Pacific Northwest is biologically rich with hypogeous
fungi; several species produce fruit bodies year-round (Colgan et
al., 1999). This knowledge of food web connections directly
relates to the preservation of birds of prey, such as the
endangered northern spotted owl, that eat small mammals which
themselves subsist almost entirely on hypogeous fruit bodies.
Thus, from an ecosystem standpoint, in order for managers to
protect these megafauna, they must understand the direct
connection to a basic food web component, the fungi.
3. Fungal plant pathogens Typically we perceive disease-causing
fungi as agents of destruction and gauge their impact by the
volume of timber destroyed. Although this is an important
consideration, ecosystem management goes beyond timber
management by recognising diseases as important processes that
create forest structure vital to other species and processes. In
addition to providing organic matter for nutrient cycling, for
example, standing dead snags or heart rot in live trees create
needed habitat for primary cavity nesting birds and secondary
cavity users. Forest gaps created in root rot pockets allow for
development of understorey plants, contributing to both species
richness and increasing browse for large mammals such as deer
and elk. Schowalter et al. (1997) provide a framework for
Conservation in the Pacific Northwestern United States
25
integrating the ecological roles of pathogens in managed forests.
4. Mycorrhizal mutualisms The benefits of mycorrhizas to plants
are well documented and include efficient nutrient uptake,
especially phosphorus; enhanced resistance to drought stress;
and direct or indirect protection against some pathogens. From
an ecosystem standpoint, mycorrhizal benefits extend much
further in space and time. In addition to participation in nutrient
cycling and food web processes, mycorrhizal fungi provide
linkages among plants that affect plant community development
and resiliency. Our research has emphasised an understanding of
the mycorrhizal specificity between potential plant hosts and
fungal association, with direct relevance to the ability of plants
to be connected by commonly shared mycorrhizal fungi
(Molina, Massicotte & Trappe, 1992). Indeed, many fungi can
form mycorrhizas with diverse plant species, including across
plant families. Brownlee et al. (1983) and Finlay & Read (1986)
showed that carbon (photosynthates) can move from a donor to
a recipient plant when connected by a common fungus. Simard
et al. (1997) later showed under field conditions that a net flow
of carbon can move from a donor plant to a shaded recipient
plant via mycorrhizal fungus linkages. Thus, understorey plants
may receive partial nourishment from overstorey plants via a
shared mycorrhizal fungus network.
Interplant linkages are equally important for other ecological
reasons. During early plant succession, pioneering plants
maintain or build the mycorrhizal fungal community by
supporting it with photosynthates. Late-seral plants can develop
mycorrhizas with many of the same fungi supported by
early-seral plants and thus benefit from the ongoing active
mycorrhizal processes. We believe that the positive feedback
interactions of these plant–fungal associations in space and time
contribute to ecosystem resiliency after disturbance (Perry et al.,
1989; Molina et al., 1992).
Fungal communities
Little is known about the impacts of forest management activities on
fungal communities in the Pacific Northwest. To manage effectively for the
diversity of forest fungi, it is necessary to identify the fungal species within
forests, describe their community structure in space and time, and under-
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stand the impacts of both human-induced and natural disturbances on
successional dynamics of forest fungi. This approach places fungi into the
same context as that of understanding the dynamics of plant communities,
a context relevant to forest managers. Data from fungal community
studies provide the basis for addressing critical research topics including
(1) conservation biology of rare fungal species; (2) fungi as important
indicators of forest community dynamics; (3) sustainability of edible,
commercially harvested fungi; and (4) development of habitat models for
fungi. In this section, we describe some of the primary considerations in
setting objectives for the study of fungal communities and then describe
our current approaches to determining fungal communities in forests of
the Pacific Northwest. Examples of a few studies are given.
Description of fungal communities
The first step in setting study objectives is defining what the phrase fungal
community means. In its simplest definition, a community is a group of
organisms sharing a particular place. Community structure is characterised by the numbers of species present, their relative abundance, and the
interactions among species within the community. Characteristics of communities emerge from historical and regional processes. Thus, it is important to define clearly the time and space boundaries of the study. For
example, the problem of determining changes in fungal species on a
recently dead tree or log differs from problems associated with examining
the succession of forest floor fungi over several hundred years of forest
development.
If specific functional groups or processes are selected for study, then a
guild concept can help to focus the community definition. A guild is a
group of species that uses various resources or shares a common function,
such as fungi that form ectomycorrhizas or that decompose wood. Studies
of fungi, plant host, and environmental relations often target a particular
guild. Because ectomycorrhizal fungi have been the focus of most fungal
community studies in the Pacific Northwest, they receive primary attention
in this section.
Several factors influence mycorrhizal community structure, including
plant species and habitat conditions, such as tree composition, tree age,
disturbance, and soil nutrients (Molina et al., 1992; Vogt et al., 1992;
Gehring et al., 1998). The objectives for a community study might examine
one causative factor or a combination of factors for effects on community
development. The selection of factors strongly influences the study design
and sampling intensity. For example, if one is interested in the effects of
Conservation in the Pacific Northwestern United States
27
coarse woody debris (CWD) on ectomycorrhizal community structure,
then a simple design of sampling fungi at different distances from large
wood in similar forest stands might suffice. However, if one is also interested in the importance of CWD at different stages of forest development, or
in forests of different tree densities, then a complex design with different
forest treatments is needed, which increases sampling units, time, and
expense.
As for most biotic communities, disturbance phenomena often trigger
the successional dynamics of fungal communities. For example, changes in
plant species composition from forest succession or large-scale disturbances significantly affect ectomycorrhizal species composition and total fruit
body production (Dighton & Mason, 1985; Termorshuizen, 1991; Vogt et
al., 1992; Amaranthus et al., 1994; North, Trappe & Franklin, 1997;
Waters et al., 1997; Baar, Horton & Kretzer, 1999; Colgan et al., 1999).
Fire and timber harvest are the two main disturbance events in Pacific
Northwest forests. Several regional studies have shown a reduction in fruit
body production or levels of ectomycorrhiza formation after forest thinning or fire (Harvey, Larsen & Jurgensen, 1980; Parke, Linderman &
Trappe, 1984; Pilz & Perry, 1984; Waters, McKelvey & Zabel, 1994;
Colgan et al., 1999). Severity and length of time since disturbance, however, are key factors in evaluating the effects of fire or tree death on the
ectomycorrhizal community (Visser, 1995; Miller, McClean & Stanton,
1998; Horton, Cazares & Bruns, 1998; Jonsson, Dahlberg & Nilsson, 1999;
Stendall, Horton & Bruns, 1999; Baar et al., 1999) and should be considered in examining fire disturbance effects on fungal community dynamics.
Ectomycorrhizal community analysis
Once the scope and objectives of the study are clearly defined, the appropriate options for sampling the fungal community are considered. Approaches to ectomycorrhizal community analysis include collection of
fruit bodies, morphological typing of mycorrhizas, and molecular identification of fungal symbionts. Each approach differentially estimates species
dominance. For example, fruit body studies address food resource questions about mammal mycophagy based on fruit body biomass. Identification of mycorrhizal symbionts on root tips best addresses the fungal
community transporting nutrients to the host or fungal symbionts available for microarthropod mycophagy. Selection of an approach depends on
the objectives of the study. Each approach has strengths and weaknesses in
regard to sampling intensity, skills needed, and accuracy in assessing
species occurrence, especially at broad spatial scales. These approaches are
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not mutually exclusive, and most researchers are using a combined approach.
Fruit body surveys poorly reflect the composition of subterranean ectomycorrhizal communities, thereby making interpretations difficult at the
community level (Danielson, 1984; Gardes & Bruns, 1996; Dahlberg,
Jonsson & Nylund, 1997; Kårén et al., 1997; Gehring et al., 1998; Jonsson
et al., 1999). Inherent problems with fruit body surveys include the sporadic production of fruit bodies, infrequent surveys, and the omission of
easy-to-overlook fruit bodies such as resupinate forms found on the underside of woody debris.
Identification of ectomycorrhizal root tips to broad morphological
types (Zak, 1973) became a popular alternative to fruit body collection for
defining ectomycorrhizal communities. This method, refined by Agerer
(1987–1998), Ingleby, Mason & Fleming (1990), and Goodman et al.
(1996), has been applied to glasshouse soil bioassays, as well as field
studies. Morphological typing of mycorrhizas is time consuming and
taxonomic resolution is low.
More recently, the application of molecular techniques, in particular
PCR–RFLP (polymerase chain reaction–restriction fragment length polymorphism) analysis and DNA sequencing, to mycorrhizal community
research has improved our ability to identify morphological types to
family, genus, or species level (Gardes et al., 1991; White et al., 1990;
Gardes & Bruns, 1993; Bruns et al., 1998). Although not a panacea,
PCR-based methods significantly improved the study of ectomycorrhizal
communities and have enhanced our ability to investigate fundamental
aspects of ectomycorrhizal communities in several forest types (Gardes &
Bruns, 1996; Dahlberg et al., 1997; Gehring et al., 1998; Horton & Bruns,
1998; Taylor & Bruns, 1999). They allow comparison of the observed
diversity of fruit bodies with that of root tips and allow us to study known
species that do not grow in culture. Additionally, molecular techniques are
useful for taxonomic placement of fungal symbionts to the family level in
forests where species fruit infrequently or not at all. Similar to challenges
with morphotyping, studying the mycorrhizal community with molecular
techniques is time consuming and may require restricting sample size.
Restricted sampling might inadequately reflect the wide variances typically
seen in ectomycorrhizal root tip community studies.
Current fungal community studies in the Pacific Northwest
Given the size of the region, the diversity of forest types, and the longterm nature of fungal community studies, we have focused on key forest
Conservation in the Pacific Northwestern United States
29
habitats or forest management issues that affect not only fungi but the
overall ecosystem management as well. This approach provides the best
opportunity to interpret mycological findings and integrate results into
forest management plans. It is also cost effective because large-scale forest
manipulations are often prohibitively expensive for stand-alone mycological studies.
Because conservation of old-growth forests is a major concern in the
Pacific Northwest (only about 10% of the original old-growth forest
remains), we have compared the ectomycorrhizal fruit body diversity
(both mushroom and truffle) in old-growth (9400 years old) to younger
stands (30–60 years old) of Douglas-fir. We have found a similar number
of species in old-growth compared to younger stands. However, the species
composition differs, and the number of unique species is greater in oldgrowth versus younger stands (J. Smith and others, unpublished data).
These results provide evidence that some fungal species require old-growth
forest habitat and that protecting the remaining old-growth forests is
important for maintaining fungal species diversity.
Many forest management experiments in the Pacific Northwest focus on
accelerating development of old-growth structure by using various silvicultural approaches such as thinning, green-tree retention, and prescribed
burning to return forest tree composition to that found before European
settlement and wide-scale fire suppression. We currently have several
integrated studies under way in large-scale forest management experiments
that examine the effects of these management treatments on fungal diversity and function. One, done in conjunction with the Ecosystem Study in
the state of Washington (Carey, Thysell & Brodie, 1999), is designed to
manipulate forest stands 60 to 80 years old to hasten their return to
old-growth structure and provide habitat for the northern spotted owl
(Strix occidentalis); the owl is currently absent from these experimental
study sites. The experiment examines silvicultural effects on the diversity of
important food web functions. The focus of this study is on the truffle
community because this is the primary food base for small mammals,
which in turn are the main prey of the northern spotted owl. Our results
(Colgan, 1997; Colgan et al., 1999) describe the community structure of
dominant truffle species in the treated stands and in small-mammal diets.
A key finding was that truffle species dominance shifts under different
forest thinning regimes. Within these managed stands, several truffle species fruit year-round, thereby providing an essential food base during
winter months. These species can thus be considered keystone species
in the ecosystem food web. This type of fungal diversity study allows
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managers to understand the role of maintaining fungal communities and
how specific silvicultural treatments affect their diversity, abundance, and
function.
Many forests in the Pacific Northwest and throughout the Western
United States are experiencing decline, primarily because of fire suppression and silvicultural management. Decades of fire suppression and
selective tree harvest have yielded dense forests of shade-tolerant tree
species prone to water stress and catastrophic insect disease outbreaks
(Agee, 1993). Forest managers are experimenting with prescribed fire and
thinning to improve overall forest health. These experiments also lend
themselves to investigating effects on the fungal community. J. Smith and
others (unpublished data) are examining the effects of burning and thinning on the fungal community in ponderosa pine (Pinus ponderosa) forests. By using primarily PCR–RFLP techniques to document changes in
ecto-mycorrhizal fungal community structure they found that fire communities are dominated by few ectomycorrhizal species. Such species may be
important to ecosystem resiliency after fire disturbance.
In addition to the studies mentioned above, Pilz & Molina (1996)
describe many of the current efforts in the Pacific Northwest aimed at
documenting the biological diversity, function, and community dynamics
of forest fungi. Additional studies are also under way to examine management treatments such as variable thinning, presence of large woody debris,
and soil compaction on fungal diversity and shifts in species dominance.
By understanding the effects of various management practices on fungal
communities, forest managers can integrate fungal diversity and function
into forest management planning.
Population biology
An effort to conserve species, especially rare species, must include some
understanding of the population structure and dynamics of the species in
question. Our understanding of the population biology of fungi, however,
lags far behind that of other taxa for many reasons described below. To
deal with the issue of rare fungal species within the Northwest Forest Plan,
we thought it important to begin a research programme that explores ways
to examine the population structures representative of forest fungal species. Such exploratory research requires large sample sizes not easily
obtained from populations of rare taxa so the initial research projects
described below are focused on species that are common but represent
different life history strategies found in ectomycorrhizal fungi. As tech-
Conservation in the Pacific Northwestern United States
31
niques become refined, we will address similar questions with rare species.
Two long-term goals of our population genetic research on ectomycorrhizal fungi are to gain some understanding of how past and current land
management practices in the Pacific Northwest have altered the dynamics
of fungal populations and to use this information to predict how future
land management scenarios may affect persistence of fungal populations.
Achieving these goals requires an understanding of the spatial and temporal scales at which evolutionarily significant processes occur within and
among fungal populations, which processes are most important at spatial
scales relevant to management, and how spatial heterogeneity in landscape
structure affects these processes (Wiens, 1989, 1997).
Problems inherent to the study of fungal populations at forest
landscape scales
Meaningful analysis and interpretation of genetic spatial patterns requires
that the extent of any selected study area be large enough to encompass
multiple units (discrete populations) affected by gene flow and that individual samples (grain) be dispersed such that variance of process effects within
each unit can be quantified (Wiens, 1989). Several ecological characteristics of fungi impede our ability to select scales appropriate for quantifying
the relative importance of genetic drift, natural selection, and gene flow
within and among populations. Selecting an appropriate extent and grain
for population genetic studies requires an understanding of how individuals are distributed across the landscape and how much area discrete
populations can cover. This presents an obstacle with fungi because the
mass of thread-like mycelium (thallus) that gives rise to fruit bodies is
frequently hidden from view. Attempts to characterise the size and distribution of discrete thalli have identified individuals encompassing hectares (Smith, Bruhn & Anderson, 1992), square metres (Baar, Ozinga &
Kuyper, 1994), and square centimetres (Gryta et al., 1997). Dahlberg &
Stenlid (1990) have also shown that ecological factors such as forest age
can influence the size of individuals and alter the mechanism (sexual or
vegetative) by which new thalli are established. The range and variance of
individual size and distribution unique to each fungal species will dictate
the scale at which population boundaries should be encountered, and set
the minimum size limit for study area extent.
When sampling fungi in the field, it is impossible to discern what
proportion of the fruit bodies collected represent genetically identical
individuals, and genetically distinct mycelium masses may produce many
fruit bodies simultaneously. Discrete thalli can be established either by
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sexual reproduction (genets), selfing, or clonal vegetative propagation
(ramets) (Todd & Rayner, 1980); evolutionary interpretations hinge on
correct estimates of the contribution of each reproductive type to the
genetic diversity observed within populations (Avise, 1994). Estimating
proportional contributions of individual genets and ramets to the total
genetic diversity in a population may not be intuitive. Depending on the
mating system of the species, a maximum of 25% or 50% of the monokaryotic spores produced by an individual fruit body can potentially grow
and fuse to form genetically novel dikaryotic genets so closely related to
the parent mycelium that the two may fuse (Smith et al., 1992). This type of
fine-scale genetic structuring coupled with variance in genet size can bias
allele frequency and gene flow estimates such that potential dispersal
distances are underestimated. Results that are misleading owing to lack of
knowledge of fungal reproductive behaviour can lead to poor development
of management options.
The conceptual foundation of our research approach
Determining the appropriate scale at which to ask management-oriented
questions about fungal population dynamics requires the design of a
nonarbitrary, objective method to determine the scales at which important
evolutionary processes produce particular genetic spatial patterns. Observations from a mismatch of spatial scales (scale of measurements differs
from scale at which the process of interest is acting) can result in erroneous
identification of pattern–processes relations because the types or relative
influences of processes may change as new spatial domains are entered
(Wiens, 1989). For example, at fine scales, biotic factors such as inter- and
intraspecific competition, forest stand age, species composition of understorey shrubs, and type and severity of disturbance history may be strongly
correlated with patterns of genetic variability. At broad geographic scales,
abiotic factors such as climate, rainfall patterns, exposure, and elevation
may strongly influence patterns of genetic variability. Fine-scale processes
may exert control over the total amount of genetic variability on which
large-scale processes can act, and thus may be more relevant to management issues.
Delimitation of fungal population boundaries and identification of
dispersal barriers require the development of sampling techniques that
facilitate accurate gene flow estimates at forest stand, watershed, and
regional spatial scales. These sampling techniques must also account for
heterogeneity in landscape cover, disturbance type, and disturbance intensity at each scale. The ultimate goal of such a multiscale sampling regime is
Conservation in the Pacific Northwestern United States
33
to ensure that the scale of measurements and the response of fungal
populations to some variable of interest (for example disturbance) fall
within the same spatial domain. Variability across fungal species in individual size and distribution, mating system, and dispersal capability makes
it impossible to apply a single sampling protocol to all species. Long-term
advancement in understanding evolutionary processes critical to maintaining genetic variability in fungal populations requires basic research on how
genetic variability in species with different life histories is partitioned at
different spatial scales. This fundamental research will lay a general foundation for predicting how land management strategies may affect rare
species depending on their life histories. Multiscale genetic studies will also
help define scales potentially appropriate for future research on demographics or habitat requirements.
Current research examples
Building a strong knowledge base requires that basic genetic research be
done on model species that will yield large sample sizes and exhibit reliable
fruit body production from year to year. One example is the distribution of
genetic variation across different spatial scales in three model genera:
Cantharellus, Rhizopogon, and Suillus. Research on such common species
will provide initial studies with the statistical power sufficient to correctly
identify evolutionarily important pattern–process relations (Dizon, Taylor
& O’Corry-Crowe, 1995). Golden chanterelles (Cantharellus formosus) are
an economically important, epigeous fungus that forms mycorrhizal associations with various host species over a broad geographic range (Molina
et al., 1993). Rhizopogon vinicolor is an ectomycorrhizal species in the
Boletales that forms hypogeous, truffle-like fruit bodies but is closely
related to epigeous boletes in the genus Suillus, including S. lakei (Bruns et
al., 1989). Both R. vinicolor and S. lakei are common and distributed
throughout the range of Douglas-fir. All three fungal species fruit predictably and abundantly and provide ideal model species with which to initiate
base-line genetic studies because they represent diverse life histories in
respect to spore dispersal. The epigeous fruit bodies of Cantharellus formosus can remain on the landscape continuously producing basidiospores
for over a month (Largent & Sime, 1995), whereas Suillus lakei mushrooms
are more ephemeral. Assuming that wind is the primary mechanism of
spore dispersal for these two species, the potential for long-distance wind
dispersal over time is most likely lower in S. lakei than in C. formosus. In
contrast, R. vinicolor is primarily dispersed by small-mammal mycophagy
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(Maser, Maser & Trappe, 1985) and might be more spore dispersal limited
than the two wind-dispersed species. Research on these three species is
being undertaken currently by independent scientists in study areas that
overlap so that results will be comparable across the life-history types.
Characterisation of genetic patterns at different spatial scales requires
multiple, independent, hypervariable markers that measure variation
within the nuclear genome of the organism of interest (Bossart & Prowell,
1998). Comparative studies of population-level markers have shown that
the use of short, tandemly repeated DNA sequences (microsatellite repeats) is an effective method for detecting variability where other genetic
markers fail (Hughes & Queller, 1993). These valuable molecular markers
do not exist for any ectomycorrhizal basidiomycete fungus but are under
development for these three focal species.
At landscape scales, patterns of genetic isolation and indirect estimates
of gene flow among populations are calculated by using estimates of
variance in allele frequencies within and among sampling units (Wright,
1951; Slatkin & Barton, 1989). Precision of these statistics requires that
sampling units do not encompass multiple random breeding units (genetic
neighbourhoods) (Magnussen, 1993; Ruckelshaus, 1998). Wright (1969)
showed analytically that as more neighbourhoods are included within
sampling units, the genetic variance among sampling units relative to the
total variance declines, and the power to detect spatial genetic structuring
is lost. Progressing to landscape-scale studies requires that we understand
the scale at which random mating occurs in these species. Spatial autocorrelation analysis of genotype locations within forest stands allows estimation of the diameter of areas where random mating occurs (that is, genetic
neighbourhoods) via description of isolation by distance patterns over the
space that a species occupies (Sokal & Oden, 1991; Epperson, 1993). The
information necessary for spatial autocorrelation studies includes mapped
locations of fruit bodies collected over a broad range of distances, the
distance between every pair of sample points, and the genotypes of each
fruit body at multiple, independent loci. Test statistics are calculated for
the null hypothesis that the genotypes are randomly located across the
study area. Spatial autocorrelation studies are currently under way for
C. formosus, S. lakei, and R. vinicolor in overlapping study areas.
Results of this research will aid in the design of a sampling method that
will capture informative variability in genetic patterns and processes at
spatial scales relevant to management. Future research efforts will apply
the sampling techniques, currently in development, to the study of how
heterogeneity in landscape features (for example forest stand age, patterns
Conservation in the Pacific Northwestern United States
35
created by different disturbance regimes) affects rates of gene flow between
populations of fungi. Using this line of research to develop a better
understanding of how fungi disperse across landscapes will facilitate
identification of geographic areas important to the conservation of genetic
diversity.
Applications in inventory and monitoring
Identifying and prioritising rare species
Protecting rare species is often a prominent component in conservation
programmes, regardless of the scale of the conservation effort. Resources
for study of species biology are scarce, and society places a high value on
maintaining rare species. Understanding the causes and consequences of
species rarity is important because it is a basis for setting conservation
priorities. Thus, it is important that mycologists identify fungal species
that are most rare or at risk as one basis for allocating resources to
studying particular species.
As noted in our introduction, the Survey and Manage guidelines of the
Northwest Forest Plan provide protection of rare, old-growth forest dependent fungi. The species list was compiled by three expert regional
mycologists, Joseph Ammirati, William Denison, and James Trappe, and
was based on their extensive experience and personal records from over 30
years of collecting macrofungi throughout the region. The list includes
gilled, cup, polypore, and truffle fungi (Table 3.1). Within the Survey and
Manage programme, each listed species represents a set of hypotheses.
Each species is thought to be (1) rare, (2) associated with old-growth
forests or legacy components of old-growth forests such as coarse woody
debris and (3) not adequately protected by the reserve land allocations and
management actions to protect needed habitat. The Survey and Manage
programme of work addresses these three hypotheses so that appropriate
protection measures can be taken to assure species persistence.
The Survey and Manage conservation programme provides protection
only for rare, old-growth forest associated species; we will address some of
the unique considerations in conducting extensive surveys for those species
in the next section. Two other conservation approaches also have been
useful for identifying and prioritising rare fungal species in the Pacific
Northwest, Natural Heritage programmes and Rarity, Endangerment,
and Distribution lists (RED lists – hence, Red Data lists). These programmes are operated at state levels and cover all species, geographic
36
R. Molina et al.
Table 3.1. Fungal genera and number of species listed under the Survey and
Manage guidelines of the Northwest Forest Plan
Genus
Number of species
Genus
Number of species
Acanthophysium
Albatrellus
Alpova
Arcangeliella
Asterophora
Baeospora
Balsamia
Boletus
Bondarzewia
Bridgeoporus
Bryoglossum
Cantharellus
Catathelasma
Chalciporus
Chamonixia
Choiromyces
Chromosera
Chroogomphus
Chrysomphalina
Clavariadelphus
Clavicorona
Clavulina
Clitocybe
Collybia
Cordyceps
Cortinarius
Craterellus
Cudonia
Cyphellostereum
Dermocybe
Destuntzia
Dichostereum
Elaphomyces
Endogone
Entoloma
Fayodia
Fevansia
Galerina
Gastroboletus
Gastrosuillus
Gautieria
Gelatinodiscus
Glomus
Gomphus
Gymnomyces
Gymnopilus
Gyromitra
1
4
2
3
2
1
1
2
1
1
1
3
1
1
1
2
1
1
1
6
1
2
2
2
2
13
1
1
1
1
2
1
2
2
1
1
1
5
5
2
2
1
1
4
2
1
5
Hebeloma
Helvella
Hydnotrya
Hydnum
Hydropus
Hygrophorus
Hypomyces
Leucogaster
Macowanites
Marasmius
Martellia
Mycena
Mythicomyces
Neolentinus
Neournula
Nivatogastrium
Octavianina
Omphalina
Otidea
Phaeocollybia
Phellodon
Pholiota
Pithya
Plectania
Podostroma
Polyozellus
Pseudaleuria
Ramaria
Rhizopogon
Rhodocybe
Rickenella
Russula
Sarcodon
Sarcosoma
Sarcosphaera
Sedecula
Sowerbyella
Sparassis
Spathularia
Stagnicola
Thaxterogaster
Tremiscus
Tricholoma
Tricholomopsis
Tuber
Tylopilus
1
4
2
2
1
3
1
2
3
1
4
5
1
2
1
1
3
1
3
13
1
1
1
2
1
1
1
28
12
1
1
1
2
2
1
1
1
1
1
1
2
1
1
1
2
1
Conservation in the Pacific Northwestern United States
37
locations, and habitat types within the state (see, for example, Washington
Natural Heritage Program, 1997). Both approaches use similar criteria to
evaluate species and ranking systems that assign risk status. Species evaluation criteria include pattern of occurrence in space and time, vulnerability
of habitat, direct and indirect threats to species decline, degree of protection under current management guidelines, and taxonomic uniqueness.
Rankings are based on number of occurrences (rarity), number of individuals, population and habitat trends, and type and degree of threats. Status
categories under the natural heritage programme include: endangered,
threatened, sensitive, and possibly extinct or extirpated in the state. Two
other categories, review and watch, are assigned to species of uncertain
status for which additional information is needed before it is assigned to
one of the four previous categories.
Red Data lists serve several purposes for fungi (see Arnolds, 1989). They
alert mycologists and conservation biologists to issues surrounding rare
fungal species and provide information to managers and policy makers
responsible for the management and protection of the species. Some of the
listed species are useful as candidates for long-term monitoring of status
and trends of fungal populations. Scientists can also compare lists from
different geographical areas to estimate national and international status
of individual species.
Nearly 25 years ago the first reports concerning a decrease in fungal
species diversity and the abundance of fungal fruit bodies across a wide
geographic area originated from Europe (Schlumpf, 1976; Bas, 1978). This
awareness led to creation of Red Data lists in 11 different European
countries in the next 15 years (Arnolds & De Vries, 1993). Attention to this
phenomenon outside northern Europe was lacking until recently.
In conjunction with the species information generated by the Northwest
Forest Plan process, another regional assessment on the ecosystems of
eastern Oregon and Washington, and the western parts of Idaho (called
the interior Columbia River basin assessment; see USDA & USDI, 1996)
led to the inception of the first Red Data lists of macrofungi for any large
geographic area outside of northern Europe (Castellano 1997a, 1997b).
The Red Data list for Oregon contains 118 species of macrofungi. It is
smaller than the Survey and Manage list for Northwest Forest Plan lands
because it evaluates species across the entire state and across all habitat,
not just late-successional forest habitat. Some species that are rare on
protected Federal land are sometimes more common in younger stands or
on other land ownership. The interior Columbia River basin assessment
yielded a Red Data list of macrofungi for Idaho (Castellano, 1998)
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R. Molina et al.
containing nearly 200 fungal species of special concern. Simultaneously, J.
Ammirati developed a working list of 57 rare macrofungi for Washington
(Washington Natural Heritage Program, 1997). These three listing efforts
are considered preliminary. Additional systematic and strategic sampling
of habitats is critical to uncover previously undiscovered occupied sites for
many of these species. Significant efforts are under way by the USDA
Forest Service and USDI Bureau of Land Management to collect these
fungi in these regions.
As more fungal surveys occur, these lists will likely change. It is important that mycologists update Red Data lists and other lists, such as those for
Natural Heritage programmes, and disseminate the information so that
the mycological community and the public can stay abreast of ongoing
efforts in cataloguing rare species, prioritising them into risk categories,
and monitoring our success in providing for their conservation. As lists are
developed for different parts of the world, we can better understand
distribution patterns of rarity and key habitats needed for their protection.
We also can begin to understand some of the natural and anthropogenic
factors that lead to fungal species rarity.
Methodological considerations in fungal inventories
The Survey and Manage guidelines of the Northwest Forest Plan centre
around the adaptive management principle of first providing for species at
known sites by appropriate management of habitat (manage) and simultaneously searching for new locations (survey) to learn more about the
rarity, range, distribution, and habitat requirements. Gathering range and
habitat information for over 200 rare fungal species in a region the size of
the Northwest Forest Plan is unprecedented in scope for any fungal
inventory programme. Originally, the surveys were to be conducted for at
least 10 years to collect enough new information to develop science-based
management plans for maintaining species persistence. Given the above
survey objectives, several considerations needed to be addressed for effectively implementing regional-scale surveys. This section highlights several
fundamental areas that we considered in conducting broad-scale fungal
surveys and inventories.
Taxonomic expertise
Taxonomic knowledge is the cornerstone of biology and is essential in
efforts to conserve biodiversity. Fungal taxonomy is problematic because
our understanding of species is limited; Hawksworth (1991) estimated that
Conservation in the Pacific Northwestern United States
39
less than 5% of the mycota has been described in the scientific literature.
Even in well-known and economically important fungal groups, new
species are continually described and some species concepts are in flux (e.g.
Redhead, Norvell & Danell, 1997; Hughes et al., 1999; Sime & Petersen,
1999).
Taxonomists are scarce because of a shift in academic programmes
toward molecular systematics and ecology. The depleted ranks of classical
fungal taxonomists can be augmented, however, by a cadre of experienced
parataxonomists, people with less formal schooling in mycology, who are
trained and gain significant experience in fungal identification. Over the
past three years, we have trained many Federal agency biologists and
botanists in the proper identification and handling of fungal specimens.
Such training is essential to conduct the regional surveys. Training also
increases the awareness of field staff and managers about the natural
history of fungi.
Even when accurate identifications are performed, observations of species occurrences are anecdotes unless a voucher specimen is deposited in a
publicly accessible herbarium (Ammirati, 1979). Properly documented
voucher collections allow experts to confirm the identification of potentially rare species. We have entered into formal partnerships with regional
university herbaria to access the many fungal specimens collected from
regional surveys.
Survey approaches
Biological surveys need clearly defined goals. This is also the case when
surveying for both rare and potentially rare species. Clearly stated goals
and prioritisation of those goals allow one to define which data to collect,
where and when to survey, and which species to collect. One can prioritise
by species, ranking from most to least important, or by geography,
ranking different habitats or regions, for example, by where the most
species are expected to occur, or by where there are the least historic data.
In practice, with planning and organisation, multiple goals may be pursued simultaneously. For example, one can visit sites of the rarest species,
and while in the vicinity, survey high-priority habitats.
Prioritising species for survey Ranking species according to rarity is probably the most frequent approach to conservation prioritisation.
We use most of the ranking criteria developed by the natural heritage
programmes mentioned previously. These criteria evaluate the rarity and
distribution of species, with the highest priority assigned to potentially
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R. Molina et al.
extinct species, followed by critically imperilled species (typically known
from five or fewer locations). The criteria are applied at both the global
and local (state) scale, so a species can be extinct in the state, but globally
secure. The rank also is evaluated as to the certainty with which it can be
applied, and the certainty of taxonomic status.
Once species are prioritised, the task of locating begins. Either historic
sites or potential habitat can be targeted; both approaches can be challenging. The rarest species are likely to be known only from historical records
that may lack the precision necessary to find the site. Similarly, habitat for
rare species is usually poorly defined. Two important goals for visiting
historic sites include documenting whether the species still occurs there,
and gathering habitat data. An additional bonus comes, not uncommonly,
when other rare species are found at such historic sites.
Prioritising regions for survey One approach to regional prioritisation is determining appropriate habitat. In many cases, however, species habitat preferences are not known. Herbarium labels are notoriously
vague about habitat, with such descriptors as ‘in woods’, ‘under pine’, or
‘in conifer forests’. Nevertheless, for some species there are relatively
narrow host or habitat requirements that can guide one to areas of interest.
For example, Gelatinodiscus flavidus is a rare ascomycete that fruits only
on the foliage of Chamaecyparis nootkatensis, a conifer with a fairly
restricted distribution. Another example is the number of species on the
Survey and Manage list that fruit in montane forests, usually including
Abies, near melting snowbanks. Where specific habitats can be identified,
prioritising surveys in those habitats can often extend the documented
range of a species.
A second approach that we have used with good success is visiting
data-poor areas. The existing regional database of known site locations is
analysed to identify a data-poor area, that is an area with few known sites
of targeted rare species. After gathering herbarium data for about 135
putatively rare species of fungi, it became apparent that much of the data
resulted from a few mycologists visiting a restricted set of collecting areas
(Castellano & O’Dell, 1997; Castellano et al., 1999). The data were obviously biased, raising the question of how to use it to best advantage. We
have used GAP analysis following the procedures of Kiester et al. (1996) to
analyse regional distribution of rare fungi site locations and select priority
areas for survey. This approach has the advantage that when observations
of a species of interest are made in data-poor areas, they are likely to
extend the known range or habitat amplitude for the species. Many new
Conservation in the Pacific Northwestern United States
41
observations of Survey and Manage fungi have resulted from applying this
strategy (Castellano et al., 1999).
Survey techniques
After selecting where and for which species to survey, one must design an
effective survey method. Again, survey methods must have clear objectives. For example, finding occurrences of rare species requires a different
approach than testing a hypothesis about their habitat preferences. We
have used two primary methods, controlled intuitive and systematic or
random sampling methods.
Controlled intuitive survey Controlled intuitive survey is the
simpler of the two methods. Once the site has been determined, surveyors
travel to the site and survey the area, focusing on habitats or microsites
where they think the target species is likely to occur. This approach relies
on knowledge, experience, observation, and intuition, and may be the
most reliable method for locating rare species.
Systematic or random sampling methods If information on relative abundance of species, or quantitative comparisons of different sites or
habitats are needed, then methods that employ systematic random site and
plot selection must be used that standardise survey intensity and meet
statistical sampling assumptions. Plots that define a survey area are one
way of standardising survey effort. Because fungi tend to occur in clustered
patterns on the landscape, long narrow plots are often used; they are more
efficient at sampling such distributions than circular plots (T. O’Dell,
unpublished data). For truly rare species, however, random and systematic
approaches may prove less useful than controlled intuitive surveys because
of the low probability of encountering the species of interest. Adaptive
sampling methods have been recommended for organisms that are patchily distributed and rare, as is the case for some fungi (Thompson, 1992). In
this approach, sampling intensity is increased when the target species is
encountered. For example, if a rare fungus is found, more plots or time
could be added to the sample in the immediate or nearby vicinity. In this
way, more useful information on its population size and habitat association can be gathered.
Limitations and inferences
Uncertainty surrounds the field sampling of fungi. No matter how much
sampling is done, species will be undetected owing to seasonal and annual
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R. Molina et al.
variation in fruiting, limitations to the amount of area scrutinised, and
inconspicuousness of fruit bodies. Most macrofungi produce fleshy,
ephemeral fruit bodies. Unless sampling occurs during the exact time it is
fruiting, a species might go undetected. In the Pacific Northwest, fleshy
fungi can be collected every month of the year in one habitat or another,
but we tend to sample during limited times to make the most of available
survey resources. Some species, therefore, are overlooked simply because
they fruit at times when most other species do not, and when we are
unlikely to be sampling. During peak fruiting seasons, sampling every two
weeks may fail to detect a significant fraction of species compared with
weekly sampling (Richardson, 1970). Unless an intensive study is being
conducted, however, repeated visits to a site may be impractical. Annual
variation in species occurrence can be tremendous, with as little as 5% of
species observed at sites in one year recurring the next year (O’Dell,
Ammirati & Schreiner, 1999). Many repeated visits to a site are required to
determine the presence or absence of a species at that site.
Although we have noted several difficulties in conducting broad-scale,
systematic surveys for rare fungi, it is important that mycologists are not
discouraged from pursuing the need for fungal inventories. We stress again
the importance of defining clear objectives for the surveys, designing the
appropriate methods to collect the needed information, and scheduling
enough time to conduct the surveys to satisfy the original objectives.
Modelling habitat as a tool in conserving fungal resources
Unlike conservation of rare plants and animals, which may entail reintroducing species into an area of suitable habitat as part of species recovery
and conservation, the primary means of conserving rare fungal species
requires protecting habitat and providing future habitat through land
management planning. Currently, we understand poorly the specific habitat needs of most fungal species so it is difficult to develop land management options that provide for persistence of habitat and therefore the
species. Even as we begin to collect habitat data from our regional surveys,
it remains difficult to discern the key environmental and biotic habitat
variables to which the fungi in question are responding. Because of this
lack of knowledge, we have begun a fungal habitat modelling research
project that will allow us to build suitable habitat indices, construct
hypotheses around key variables, and then test the model for accuracy in
predicting suitable fungal habitat. Because habitat modelling has rarely
been attempted for fungi, in this section we first define habitat modelling
Conservation in the Pacific Northwestern United States
43
and its uses, and then describe unique considerations for modelling fungal
habitat and our research approaches.
Habitat modelling: definition and uses
A habitat model is an abstract representation of the physical and temporal
space in which an organism lives, including the way it interacts with the
surrounding biotic and abiotic environment (Patton, 1997). Habitat
modelling can potentially be a powerful tool in the study of fungi and in
fungal conservation biology for several reasons. Habitat modelling provides insight into the behaviour of a system and an understanding of the
complex interaction between an organism and the environment. As a
method for integrating data from diverse research areas, modelling also
provides a formal organising framework for generating hypotheses, conducting data analysis, and assisting in determining research directions by
helping to define problems and refine questions. Habitat modelling in
particular can be used to develop ideas about the distribution and occurrence of forest fungi, thus contributing to the development of conservation
strategies.
Land managers need to make decisions, regardless of whether they have
sufficient data and understanding of ecosystems. By understanding habitat, we will be able to map geographic areas where particular fungal species
are likely to occur. Once occurrence and species habitat relations have
been established, simulation of habitat availability under different management scenarios and under various disturbances will allow projections
of potentially occupied habitat over time. Although the Northwest Forest
Plan mandates surveys for more than 200 species of fungi, limited resources are available for conducting those surveys. Models will help prioritise data-collection efforts by predicting areas where species exist or where
more intensive sampling may be desirable (Kiester et al., 1996). The GAP
analysis mentioned as a rare species search prioritisation method illustrates a type of modelling that is helping to design regional survey strategies for forest fungi. Using models as ecosystem management tools helps
managers to calculate risks of decisions and actions, and provides a
method of accountability and support to land management decisions. This
will be particularly helpful in developing sustainable harvest strategies for
commercially valuable fungi and in predicting the impact of human intervention on fungal survival and productivity.
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R. Molina et al.
Habitat modelling: challenges and concerns
What are the habitat needs of a given fungal species? We assume that
fungal species are tightly linked biologically to their habitats; that these
habitats can be detected, measured, and quantified; and that we are able to
detect the fungus itself. Some are habitat specialists, restricted to one or a
few host species or a unique vegetation type; others are habitat generalists,
associated with various host plants or vegetation types. Our work investigates habitat requirements and modelling for fungal species that exhibit
contrasting adaptation strategies: for example, aerial spore dispersal for
epigeous species versus small-mammal dispersal for hypogeous species.
Guidelines for habitat model development (US Fish & Wildlife Service,
1981) have been used to generate models for mammals, fish, and birds.
Habitat evaluation procedures provide quantification of habitat based on
suitability and total area of available habitat. Variables are selected according to three criteria: (1) the variable is important to survival and
reproduction of the species, (2) there exists a basic understanding of the
habitat–species relation for the variable, and (3) quantifiable data are
available and the variable is practical to measure. Few studies of fungi,
however, have directly assessed any of these criteria.
We know little about fungal ecology in general, and until recently, we
did not have the tools that allowed us to define the individual, let alone
populations (see section on ‘Population biology’). Many fungal species
disperse passively by wind; however, we do not know the probabilities
associated with spores landing in suitable habitat and developing into a
fungal individual. For many fungal species, nutritional mode is uncertain,
thereby making it difficult to use ‘food source’ as a habitat variable.
Taxonomic problems and database biases encumber the study of mycology, as mentioned in the previous section on survey difficulties. The issue
of scale also arises when examining mycological data and attempting to
define habitat variables. Mycologists often focus on the minutiae and are
unaware of larger scale phenomenon that may affect fungal distribution.
Current modelling approaches
We are compiling information and formulating a concept of habitat for
several species. For forest fungi, we define habitat as all factors (biotic and
abiotic) affecting the probability that a fungus can inhabit, survive, and
reproduce in a given place and at a given time. A review of the published
literature reveals potential habitat factors that fall into four broad categories: vegetation, climate, topography, and soils.
Conservation in the Pacific Northwestern United States
45
Vegetation may be the primary factor contributing to ectomycorrhizal
fungal habitat, because of host specificity. Molina & Trappe (1982) recognise three groups of ectomycorrhizal fungi based on the relations to host
species, ranging from highly host specific to host generalists. Decomposer
fungi also show various levels of host-substrate specificity (Swift, 1982).
We can use, therefore, the presence or absence of particular host species as
an initial indicator for potential habitat. In addition to host specificity,
forest stand age and structure, and disturbances such as timber harvest and
fire contribute significantly to fungal habitat (see section on ‘Fungal
communities’).
Soil organic matter, including humus and coarse woody debris (CWD),
is a substrate for ectomycorrhizal fungi (Harvey, Larsen & Jurgensen,
1976) and decomposer fungi. The CWD may be particularly important as
a moisture-retaining substrate, thereby allowing root tips to support active
ectomycorrhizas in times of seasonal dryness, on dry sites, or after fire
(Harvey, Jurgensen & Larsen, 1978; Amaranthus, Parrish & Perry, 1989,
Harmon & Sexton, 1995). Although no data exist for determining quantities of CWD necessary to support viability of forest fungi in the Pacific
Northwest, availability of CWD seems to be a factor in establishing some
fungi as well as seedlings (Kropp, 1982; Luoma, Eberhart & Amaranthus,
1996). The presence of CWD, therefore, may be associated with habitat for
certain fungal species.
Topographic and soil factors important to fungal habitat include elevation, slope, aspect, soil properties, and local microtopography. To date,
no studies have presented significant correlations with any of these factors,
although mycologists often use such factors intuitively to find forest fungi.
Climate is a complex factor in fungal distribution and therefore habitat.
Seasonal and ecological distribution of fungi as well as fungal productivity
are partly determined by temperature and moisture (Norvell, 1995; O’Dell,
Ammirati & Schreiner, 1999). Wilkins & Harris (1946) contend that moisture may be the most important single environmental factor controlling
fungal reproduction. We know little, however, about the effects of climate
on long-term survival of fungi or on timing of mushroom formation.
Many scientific researchers, commercial mushroom harvesters and recreational forest users have much experiential knowledge as to when and
where particular mushroom species are found. Knowledge- and rulesbased modelling methods and expert systems capture such information;
both quantitative and qualitative information can be synthesised (Starfield
& Bleloch, 1991). In addition, the availability of digital maps and geographic information systems provide opportunities for linking our
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R. Molina et al.
modelling rules with spatial databases and simulation models for other
biological systems. Dreisbach, Smith & Molina (in press) provide more
details on our current modelling approaches.
Sustaining the commercial harvest of edible forest mushrooms
As noted in the introduction, conservation of fungi entails more than
protecting rare species. Conservation includes sustaining populations of
all fungal species, both rare and common. As described by Arnolds (1991),
common fungi can become rare because of habitat loss or large-scale
anthropogenic disturbances such as air pollution. The commercial harvest
of common, edible mushroom species illustrates a human disturbance that
might diminish fungal resources. This section describes our research programme aimed at understanding this resource so that we can continue to
enjoy economic gain and culinary delights while sustaining the survival
and reproduction of these valuable fungi. We outline edible mushroom
harvesting issues and concerns, discuss results from various recent research
projects, describe forest management implications of the findings, and
consider future directions in edible mushroom research and monitoring.
Scope of issues
Edible mushrooms have been widely collected from the forests of the
Pacific Northwest since the 1860s when European settlers began hunting
for mushrooms similar to species they had collected in their homelands.
Some Native American tribes harvested mushrooms for various uses, but
we lack evidence of widespread consumption (Kuhnlein & Turner, 1991).
Regionally, amateur mushroom clubs and organised mushroom events
date to the 1950s (Brown et al., 1985), and now there are about 34
mycological societies based in the region (Wood, 2000). Academic mycological activities date back to the 1920s for the region. During the 1980s
and early 1990s, commercial mushroom harvesting expanded dramatically. The increased public demand for wild edible mushroom-harvesting
opportunities has affected all forest landowners in the Pacific Northwest,
especially Federal forests.
Federal lands encompass a large portion of suitable mushroom habitat
in the Pacific Northwest, and are managed for multiple use, including
commercial forest products. Since the 1950s timber has been the major
forest product, but environmental concerns and adoption of the ecosystem
management philosophy defined in the introduction have led to dramatic
Conservation in the Pacific Northwestern United States
47
Table 3.2. Important nationally and internationally marketed wild
mushrooms harvested from the forests of the Pacific Northwest
Local common names
Latin name
Use
American or white matsutake,
pine or tanoak mushroom
Morels
Pacific golden chanterelle
Tricholoma magnivelare (Peck)
Redhead
Various Morchella species
Cantharellus formosus Corner
(previously called C. cibarius,
Redhead et al. 1997)
Cantharellus subalbidus Smith
& Morse
Hydnum repandum L. ex Fr.
Hydnum umbilicatum Peck
Boletus edulis Bull.:Fr.
Tuber gibbosum Harkn.
Leucangium carthusianum (Tul.
& C. Tul.)
Food
White chanterelle
Hedgehogs
King bolete
Oregon white truffle
Oregon black truffle
Food
Food
Food
Food
Food
Flavouring
Flavouring
reductions in timber harvesting from Federal lands during the 1990s. In
response to reduced Federal timber harvests, many forest workers
broadened the range of commercial products that they harvested from
forests. Other social and ethnic groups joined them for various reasons
(Arora, 1999; Love, Jones & Leigel, 1998; Sullivan, 1998). The mushroom
industry has benefited from and been encouraged by the development of
international markets and increased use of wild mushrooms by gourmet
chefs (Schneider, 1999). The harvest of nontimber forest products in
general has become a widely recognised industry in the Pacific Northwest
and globally (Ciesla, 1998; Lund, Pajari & Korhonen, 1998). In 1992,
nearly 11 000 people, employed full- or part-time, contributed over
US$41.1 million to the economies of Oregon, Washington, and Idaho by
harvesting and selling nearly 1.82 million kg (4 million pounds) of wild
edible mushrooms (Schlosser & Blatner, 1995). Likely, the harvest has
grown since then. Schlosser & Blatner (1995) state that over 25 species of
mushrooms and truffles are commercially harvested in the Pacific Northwest. Table 3.2 lists the most economically important species. With the
exception of some morels, all of the fungi in Table 3.2 are ectomycorrhizal.
Biological and management concerns
The magnitude of the commercial mushroom harvest in the Pacific Northwest has raised controversy about conservation of the mushroom resource,
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particularly regarding harvest effects on species viability and ecosystem
function. As part of a comprehensive research programme for managing
commercial mushroom harvests, we have conducted several public workshops to gather information on the primary concerns of resource managers, the mushroom industry, and the general public. Those concerns fall
into four categories; we present them as a series of questions so readers can
envision the scale and complexity of required mycological research.
Production and distribution
How many fruit bodies are being produced? How are they
distributed across the landscape or within a given area? How
does production differ during a season and from year to year?
What is the actual or potential commercial productivity of a
given area? What proportion of forest habitat is available and
accessible for harvesting? What factors determine
productivity and how might they be managed?
Mushroom harvesting
How can the sustainability of mushroom harvesting be
assured? What proportion of the crop can be harvested
without unacceptable impacts on the fungus itself or other
resources? What techniques will mitigate those impacts; does
mushroom harvesting increase or decrease subsequent
production? Is spore dispersal reduced by removal of
immature mushrooms, and, if so, does it impair reproductive
success and degrade genetic variability? Is fungal mycelium
and subsequent mushroom production affected by search and
harvest techniques such as raking, moving woody debris, or
digging? Do numerous harvesters trampling the forest floor
harm mushroom production? How important are
commercially valuable species as food for wildlife, and is
human competition for the resource significant?
Forest land management
How do various timber harvesting methods (clear cutting,
thinning, host species selection) affect subsequent mushroom
production? What is the impact of soil compaction or
disturbance from logging activities? What are the relations
between fire and subsequent mushroom production, especially
for morels? How does the intensity and timing of fire influence
edible mushroom production? What impacts do grazing,
fertilisation, or pesticide application have on mushroom
Conservation in the Pacific Northwestern United States
49
production? Can mushroom production be improved through
habitat manipulation (for example: planting tree seedlings
inoculated with specific fungi, thinning understorey brush for
sunlight and rainfall penetration, prescribed burning, or
irrigation)? Can production be increased across the landscape
by managing forests to attain tree age class, structure, and
composition optimal for mushroom production?
Biology and ecology
What are the important reproductive events in the life cycle of
a particular fungus species? How are new colonies or
populations established and maintained? What causes them to
diminish or perish? How important is spore dispersal to
reproductive success, population maintenance, genetic
diversity, or adaptability to unique microhabitats? How much
genetic diversity exists within and among populations? Are
there endemic, narrowly adapted, or unusual populations of
otherwise common species? What are the growth rates of
fungal colonies in soil and degree of mycorrhizal development
by specific fungi on root systems? To what degree do other
mycorrhizal or saprotrophic fungi compete with marketable
fungi for colonisation sites on host roots or space in the forest
soil?
Various measures have been adopted on Federal forest lands to ameliorate impacts until more is known about the long-term consequences of
intensive commercial mushroom harvesting. Maintaining appropriate forest habitat is the most important and practical means of sustaining healthy
populations of fungi. Essential habitat characters include appropriate host
tree species for ectomycorrhizal fungi, coarse woody debris for wood
inhabiting fungi and noncompacted soils for both saprotrophic and ectomycorrhizal fungi that fruit on the forest floor. Interim conservation
measures that can be used until needed research and monitoring is conducted include no-harvest areas, fallow harvest rotations, limited harvest
seasons or times, limited harvest permit numbers, limited harvest quantities, and incentives for leaving nonvaluable older specimens to disperse
their spores. The approaches used in any given situation depend on land
tenure, land management goals, mushroom values, availability of law
enforcement, and especially, support for regulations among harvesters
themselves.
Answering the questions above requires scientific investigations at vari-
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ous ecological scales (soil microniches to regional landscapes) and by using
various investigative methods. Some of our approaches are noted below.
Research findings and management implications
In the Pacific Northwest, edible mushroom research is currently concentrated on the three most valuable and widely collected edible forest mushrooms: the American matsutake, morels, and chanterelles. The other
species in Table 3.2 also deserve attention. The Forest Service and Bureau
of Land Management sponsor many research projects in conjunction with
the Pacific Northwest Research Station, but significant research is also
being conducted by state universities and mycological societies. Collaboration among public land management agencies, private and corporate
forest owners, research institutions, nongovernment organisations, researchers, managers, harvesters, and other stakeholders has greatly
broadened the scope and enhanced the quality of past and present research
projects. A summary of these research projects can be found in Pilz,
Molina & Amaranthus (in press).
Many of the research projects being conducted in the Pacific Northwest
are still active or not yet published. Our discussion, therefore, provides a
cursory overview of emerging information as it applies to forest management and, where possible, citations for further information.
One of the first and most frequently asked questions about harvest
sustainability was whether the greatly increased levels of harvesting would
reduce subsequent levels of fruiting. Two approaches have been used to
address this issue: experiments to examine harvesting impacts and
monitoring projects to detect trends in productivity. Research conducted
by the Oregon Mycological Society (Norvell, 1995) concurs with that
found by Egli, Ayer & Chatelain (1990) in Switzerland, that picking
mushrooms per se does not diminish subsequent fruiting over the period of
a decade or more. Search methods, however, can have some impact.
Recent studies (D. Pilz and others, unpublished data) of raking forest litter
to find young matsutake suggest that raking into the mycelial layer can
interfere with fruiting for several years, but that recovery is hastened by
replacing the duff. These studies, however, do not address the long-term
and broad-scale impacts of intensive commercial harvesting, forest management activities, or regional threats to forest health. Assessing these
factors will require long-term monitoring and silvicultural research.
Several research projects in the Pacific Northwest have obtained baseline data on edible mushroom productivity: specifically, the seasonal (sum-
Conservation in the Pacific Northwestern United States
51
med within a fruiting season) productivity per unit area (by using systematic sampling designs) of given forest stands or habitat types. These
productivity values, derived from biologically diverse mixed-conifer
stands, tend to be lower than edible mushroom productivity estimates
reported elsewhere in the literature (Ohenoja & Koistinen, 1984; Ohenoja,
1988; Slee, 1991; Kalamees & Silver, 1993). Values range from less than
1 kg to as high as 20 kg per hectare per year for chanterelles and American
matsutake (Pilz, Molina & Liegel, 1998; Pilz et al., 1999). Typical values
for commercially harvested stands range from 3 kg to 10 kg per hectare per
year (Pilz & Molina, in press). A common goal of productivity studies
conducted by the PNW Research Station is the development of statistically
valid, practical and cost-effective monitoring protocols for edible mushrooms (Arnolds, 1992; Pilz & Molina, in press; Vogt et al., 1992).
Sustaining harvests of edible mushrooms that are ectomycorrhizal depends on understanding the relations between the silvicultural characteristics of a stand and mushroom productivity. Tree species, age, growth rates,
and density can all potentially influence the quantity of carbohydrates
available to ectomycorrhizal fungi for growth and fruiting. Understanding
these relations could lead to predictive models that would allow silviculturalists to ascertain the consequences of their stand prescriptions for
mushroom productivity. The influence of thinning young stands on edible
mushroom productivity is of particular interest in the Pacific Northwest
because many forest tree plantations will be commercially thinned in the
coming decades, especially on Federal lands where mushroom harvesting
is allowed. Preliminary evidence indicates that heavy thinning (many trees
removed) can substantially reduce mushroom productivity immediately
after the harvest, but that lighter thinning (fewer trees removed) has a
lesser impact on productivity (D. Pilz & R. Molina, unpublished data).
The rate at which mushroom productivity rebounds as the remaining trees
reoccupy a site has yet to be determined. Mushroom productivity might
rebound to higher levels as the remaining trees grow more vigorously, but
the complicating influence of soil compaction from ground-based thinning
equipment will need to be factored into the interpretation of results.
Several genetic studies of edible mushrooms are showing greater species
diversity than previously documented by alpha taxonomy. For instance,
the west coast Pacific golden chanterelle, previously considered the same as
the European Cantharellus cibarius Fr. has been shown actually to consist
of three or more species, none exactly the same as C. cibarius (Redhead et
al., 1997; Dunham et al., 1998). Similarly, morel studies in eastern Oregon
have revealed five species of morels fruiting on a burned site, some not yet
52
R. Molina et al.
described (D. Pilz and others, unpublished data). Several mushroom species are harvested as ‘king boletes’ in the Pacific Northwest (Boletus aereus
Fr., B. barrowsii Smith, and B. edulis), and Boletus edulis itself might
actually comprise a species complex here. This level of species diversity is
not surprising given the diversity of climates, habitats, soils, and ectomycorrhizal host tree species found in the region. The diversity does,
however, present additional challenges to forest managers because unique
ecological adaptations likely exist for each species, and they could respond
differently to forest management choices. These are just a few examples of
the management implications of recent edible mushroom research. Additional research and long-term monitoring are needed, however, to ensure
the sustainability of widespread, intensive commercial mushroom harvesting and healthy populations of edible forest fungi.
Research and monitoring for sustainability
Most of the abundant and commercially harvested forest mushrooms,
with the exception of some morels, are ectomycorrhizal species. The plentiful carbohydrate nutrition they derive from forest trees probably contributes to their abundant fruit body production. Their mycelial colonies are
long-lived, and harvesting does not seem to impair fruiting in the short
term. Indeed, humans world-wide have harvested edible forest mushrooms
since ancient times (Arora, 1999; and see Chapter 8). So, a perfectly
reasonable question is, ‘Why bother monitoring their productivity in the
first place?’
Managers of Federal lands in the United States are required to monitor
the commercial harvest of natural resources to determine sustainable
harvest levels and avoid degrading the environment. In addition to this
legal mandate, some of the factors that have contributed to declines of
edible mushrooms in Europe (Arnolds, 1991) and Japan (Hosford et al.,
1997) are also beginning to impact forests in the Pacific Northwest. These
factors include pollution from growing urban areas, thinning ozone, climate change, introduced pathogens, and intensive timber management.
Lastly, in any system that has functional thresholds, changes in quantity
can translate into changes in kind when the thresholds are exceeded.
Commercially harvesting edible forest mushrooms for international markets arguably entails greater and more widespread harvesting pressure
than existed historically, and changes such as reduced (or enhanced) spore
dispersal or trampling could have unforeseen long-term consequences.
The effort and resources applied to monitoring should be commensurate
Conservation in the Pacific Northwestern United States
53
with resource value, socially acceptable levels of risk, current understanding of how resilient a renewable resource is to harvesting, and perceived
threats to the resource. Pilz & Molina (1998, 2000) have proposed a
regional programme that combines short-term silvicultural research with
long-term but modest monitoring activities. The proposed programme has
three components. The research component is intended to develop predictive models relating silvicultural activities to mushroom productivity.
Ideally, a widely applicable model can be developed from data on several
mushroom species and a range of sites and forest types.
The two monitoring components entail commercially harvested sites
and sites where neither mushroom nor timber harvesting are allowed. On
the commercially harvested sites, landowners would give co-operating
harvesters exclusive access to discrete areas in exchange for information on
what is harvested each year. On the natural sites, landowners could cooperate with mycological societies to estimate productivity without harvesting the sampled mushrooms. With various landowners, sites, and
co-operators scattered around the region, long-term monitoring of potential trends in fruiting could be accomplished with a minimum of investment by any one participating agency or organisation. The use of standardised sampling methods, prearranged analyses, and a web site for posting
annual results would provide all participants with timely results and
regular encouragement to remain involved. The flexible co-operative organisation of the monitoring programme would allow co-operators to join
or leave the programme as their interest and budgets dictate, although the
best information will come from sites that are monitored for many years.
Regardless of further developments in research and monitoring in the
Pacific Northwest, commercial harvesting of edible forest mushrooms
likely will continue. The resource promises to be renewable and sustainable
as long as we maintain healthy forests, understand the impacts of our
actions, and remain alert to possible trends.
Concluding remarks
As noted throughout this chapter, our knowledge is sparse regarding the
ecology and natural history of forest macrofungi. There are especially
large gaps in our understanding of habitat requirements needed by species
for maintaining healthy populations within changing landscapes. Thus,
the challenge of conserving this biologically rich array of species remains
considerable. Fungal conservation issues in the Pacific Northwest as outlined are unprecedented in scale and scope, and so our research pro-
54
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gramme includes a broad suite of ecological topics that provide a broad
knowledge base to apply mycological principles to forest management
plans. We use an ecosystem approach to plan and conduct our research at
different scales because it provides opportunities to work closely with
other forestry and ecological disciplines so that results can be woven into
integrated solutions for broad-scale forest management. Fungi are critical
ecosystem components, and their immense biological and functional diversity perform essential processes that lead to ecosystem resiliency and
health. Our ability to demonstrate and communicate this primary fungal
characteristic to other biological scientists, land managers, and the public
is key to promoting fungal conservation issues.
We offer the following set of summary statements for mycologists to
consider when planning and conducting conservation efforts for fungi.
∑ Set clear objectives for the study or inventory project. Knowing
how the information will be used, and by whom, will guide the
assemblage and proper communication of the findings.
∑ Select the appropriate methodology to meet the objectives. Data
should be collected at the same scale at which the organism or
process of interest is operating.
∑ Address mycological concerns within the context of local and
regional land and natural resource management issues. This
allows for fungi to be considered as part of the land management
planning process.
∑ Explore opportunities to work with research and management
colleagues across science disciplines. When possible, conduct
mycological investigations within integrated studies that include
other taxa and shared objectives such as overall maintenance of
biodiversity.
∑ Emphasise the characterisation of habitat for mycological
studies. Protecting fungal habitat at various scales of space and
time will provide the most likely solution to the long-term
conservation of fungi.
∑ Develop monitoring projects to inventory fungi periodically at
long-term study sites to detect trends in populations. Enlist the
help of local mycological societies in data collection.
∑ Develop quantitative models to help managers make decisions.
∑ Publish literature and conduct workshops to educate land
managers and the public continuously about the importance of
Conservation in the Pacific Northwestern United States
55
fungi in maintaining healthy ecosystems. They are critical allies
and partners in helping to set the conservation agenda for fungi.
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4
The future of fungi in Europe: threats,
conservation and management
EEF ARNOLD S
Introduction
During the 9th Congress of European Mycologists in Oslo, 1985, quantitative data were presented for the first time on changes in the mycota, in
particular in The Netherlands (Arnolds, 1988). These data supported the
fears of many mycologists and naturalists, namely that macrofungi were
decreasing, at least some species in some areas. This information was the
motive for the establishment of the European Committee for Protection of
Fungi, which later changed its name into the European Council for Conservation of Fungi (ECCF). In this chapter I will review the developments
in fungus conservation in Europe during the past 15 years and look
forward towards challenges in the future. I restrict myself to macrofungi
since microfungi are rarely considered in relation to conservation (but see
Chapter 17).
Effective conservation of fungi depends on the compilation and integration of data from different disciplines, here summarised in the scheme
shown in Fig. 4.1. Central in this scheme are Red Data lists because they
reflect our knowledge on the status of fungal species and are meant as a
basis for measurements and to present our conclusions to a wider audience. Obviously, progress in conservation depends on progress in other
mycological disciplines, such as taxonomy, distribution and ecology.
Therefore I shall pay some attention to these disciplines as well.
European Council for Conservation of Fungi
The ECCF has played a central role in further development of methods to
study changes in the mycota, in stimulating the compilation of Red Data
lists and in spreading the message that fungi are in need of conservation
64
Fungi in Europe
65
Fig. 4.1. Schematic representation of relations between conservation of
fungi and other disciplines of mycology.
measures, like many other organisms. It is the international informal
forum for exchange of ideas, and particularly valuable ECCF meetings
were those in Lódz (1988, proceedings edited by Jansen & Ławrynowicz,
1991), Vilm, Germany (1991, proceedings edited by Arnolds & Kreisel,
1993), Le Louvrin, Switzerland (1993) and Vipeteno, Italy (1997, proceedings edited by Perini, 1998). Official representatives attended these meetings as well as other people involved in conservation from many European
countries. In between these meetings, business meetings of the ECCF were
held at intermittent Congresses of European Mycologists. Since the Oslo
congress, conservation has become a constant and substantial topic at
European congresses. For instance, one third of the papers in the proceedings of the Kew congress were devoted to conservation or related subjects
(Pegler et al., 1993).
The consideration of conservation aspects was stimulated by several
factors, among which were:
66
Eef Arnolds
∑ Mycologists and ecologists were concerned about the possible
decline of fungi and the likely consequences for the functioning
of ecosystems.
∑ Knowledge of ecology and geographical distribution of fungi has
strongly increased in some countries, which enabled an analysis
of trends of various species.
∑ Public and political interest in nature conservation has strongly
increased, culminating in international treaties on biodiversity
and protection of endangered organisms (e.g. Bern convention).
Taxonomy
Conservation depends on good taxonomic knowledge since species are the
main units to be dealt with. Much progress has been made in recent years
in taxonomy of European fungi, both by thorough classical revisions
based on morphological characters and by the application of new
methods, such as molecular techniques and compatibility tests. Despite all
the recent effort, the work is not yet finished. Numerous new species are
described each year and many critical groups are awaiting modern revisions.
A practical problem in conservation is the existence of different morphological species concepts in different parts of the world. In some regions
relatively broad concepts are prevailing, for example in Scandinavia and
Germany, in others relatively narrow concepts are favoured, for example
in France by taxonomists like Bon, Henry and others. I mention only one
out of many examples: the blackening taxa of Hygrocybe are described by
Boertmann (1995) as one species with two varieties and by Courtecuisse &
Duhem (1994) as seven distinct species.
In some cases the application of modern techniques can support a
particular point of view, but in other cases the problems are becoming
rather more complicated. For instance, Aanen & Kuyper (1999) found in
the group of Hebeloma crustuliniforme twenty compatibility groups, representing biological species, but they were able to recognise at most four
species on the basis of morphological characters. Biological species are
impractical units in field mycology when it is impossible to identify them
by morphological characters.
Divergence of species concepts is a major problem when comparing
species diversity, checklists and Red Data lists of different countries. It
means that mycologists do not speak a common language and this may
produce irritation in the nonmycological world. This problem is increased
Fungi in Europe
67
further by substantial differences in nomenclature. In my opinion it is a
challenge to European taxonomists to compile a common European
checklist with recommended names. Such a survey should be compiled on
the basis of national checklists, which are also quite useful for the compilation of national Red Data lists. Dennis et al. (1960) in Great Britain
published the first modern checklist. It is a taxonomic list with references
to synonyms, plates and descriptions. A variant with more relevant information for conservationists is the ecological checklist of Sweden, providing concise information on habitat and distribution patterns (Hallingbäck,
1994). An extended combination of a taxonomic and ecological checklist
appeared in The Netherlands (Arnolds et al., 1995). It is the basic document for mapping, the national Red Data list and conservation in that
country. The list of The Netherlands contains almost 3500 species, including 2700 basidiomycetes. A recent checklist of basidiomycetes in Greece
enumerates only 811 species (Zervakis et al., 1998). Apparently the mycota
of Greece is not yet fully explored, as the authors also point out. Nevertheless, such a list is a major achievement as a basis for further exploration of
the regional mycota and as a contribution to knowledge at the European
level. Any checklist is better than no checklist.
Distribution and mapping
Reliable data on present and former distributions, as well as frequency of
fungi, are indispensable for compilation of Red Data lists. The actual
distribution is a clue to the rarity of species and their degree of endemism.
Data from the past are needed to establish possible increase or decrease.
Before 1970 no systematic recording of fungi took place in Europe. This is
a serious problem in drawing conclusions on declining species and therefore in assigning Red List categories to fungi on an objective basis.
In the 1960s a European mapping programme was initiated for 100
selected species. The results were interesting but not really representative
because of methodological problems and uneven recording in various
parts of Europe. In many cases the maps reflected the distribution of
mycologists rather than of fungi (Lange, 1974). Nevertheless this attempt
had a stimulatory effect. Mapping programmes have been initiated since
1970 in several European countries. The methodology differs considerably
from one country to another, for instance concerning grid units, collected
data and format of the database. Up to now, complete distribution atlases
have been published for Germany (Krieglsteiner, 1991, 1993) and The
Netherlands (Anon., 2000), without comments on the patterns of the
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Eef Arnolds
species. In addition, in The Netherlands an atlas was published treating
370 species extensively, with data on changes in frequency, habitat and
periodicity (Nauta & Vellinga, 1995). The latter type of atlas is, of course, a
much more valuable source of information in relation to conservation. In
many other countries maps have been published of selections of species,
for instance of threatened polypores in Finland (Kotiranta & Niemelä,
1996). Mapping was carried out until recently mainly in northwestern
Europe, but at present projects have also been launched in Greece and
Ukraine, for instance.
National mapping programmes are the essential bricks for construction
of any European survey. It is sometimes felt to be a great problem that the
databases are not directly compatible because of, for instance, differences
in scale of map grids. In my opinion this is a technical problem that can be
solved by modern database management. It is inevitable that different
methods are developed, depending on national goals and possibilities. The
participation of well-trained amateur field mycologists in mapping programmes is extremely useful, as the results in The Netherlands and Germany demonstrate.
In 1999 the ECCF launched a new pilot project for producing European
maps of 20 selected, threatened species, based on a compilation of existing
data. Such projects are essential to bring conservation of fungi to the
attention of European authorities and may, moreover, stimulate interest in
national mapping. Ideally, all distribution data should be included in a
European database, but such an initiative depends on availability of
researchers and funding.
Ecology
Ecological knowledge of fungi is necessary for identification of valuable
habitats, conservation of these habitats and the realisation of proper
management. Data on autecology of species mainly come from compilation of field surveys and are summarised in some checklists (Arnolds et al.,
1995; Hallingbäck, 1994). Mycocoenological studies have revealed many
data on the composition of fungus communities, periodicity and constancy
of fruit bodies. Some forest communities prove to be very rich in species,
for instance up to 800–1200 species in some deciduous virgin forest types in
Bialowiecza, Poland (Falinski & Mulenko, 1995). Experimental work has
recently shed some light on the population dynamics of fungi, including
the distribution, size and age of individual mycelia (e.g. Dahlberg &
Stenlid, 1990). Field experiments have shown that an impoverished ec-
Fungi in Europe
69
tomycorrhizal biota, caused by air pollution, can be partly restored by
removal of polluted litter layers (Baar & Kuyper, 1998).
Nevertheless, many fundamental questions remain to be solved. One of
the intriguing questions is whether macrofungi, known as rare species, are
really rare or only organisms that rarely produce fruit bodies. In conservation practice we accept the former hypothesis, but nobody is sure about it.
Theoretically this question can be answered by analysis of mycelium
populations with molecular techniques. It has already been demonstrated
that the underground population of ectomycorrhizas on roots is not
necessarily correlated with the frequency of fruit bodies of various species
in the same plot. This may also be true for saprotrophs.
Another question is to what extent the enormous species diversity of
fungi also represents functional diversity. For field mycologists many
species seem to occur in the same habitat on the same substrate, using the
same type of habitat exploitation. For instance: the numerous ectomycorrhizal species in a homogeneous forest plantation, the diversity of Aphyllophorales on branches and twigs, the surprisingly high number of fungi
on a homogeneous substrate like dung. And why are some dunginhabiting fungi exceedingly rare everywhere, whereas others are widespread and common?
A third question concerns the reproduction strategies of fungi. Are the
billions of spores really necessary to maintain populations in the long run
or is there a great surplus? How effective is fungus colonisation by spores,
for instance after long-distance transport by air currents? Unsolved problems like these are relevant for development of a strategy on mushroom
harvests and for judging the importance of the ecological infrastructure of
a landscape for fungi. The answers to these questions are, in fact, crucial
for conservation policies and research in these fields seems to be most
rewarding.
The changing mycota
In various publications changes in the mycota are described on a local or
national scale, including both evidence of increasing and decreasing species (e.g. Arnolds, 1988; Senn-Irlet, 1997). For conservation purposes data
on decrease are the most relevant. Important causes of decline of macrofungi are:
∑ Habitat destruction, e.g. clear-cut of old-growth forests, digging
of peat in peat bogs, building recreational facilities on sand
70
Eef Arnolds
dunes and in alpine areas.
∑ Changes in land use, often due to EU policy, e.g. replacement of
native forest by plantations of exotic trees; improvement,
abandonment or afforestation of old, semi-natural meadows.
∑ Intensification of agriculture, e.g. higher input of fertilisers, deep
drainage, both measures with negative effects on adjacent nature
areas as well.
∑ Air pollution, in particular causing acidification and nitrogen
enrichment (eutrophication). The decline of ectomycorrhizal
fungi in densely populated areas of Europe is mainly caused by
these factors (Termorshuizen & Schaffers, 1991), but
acidification and eutrophication are without doubt also harmful
to some saprotrophs in and outside forests.
Harvesting of edible mushrooms is a much discussed subject in connection with decline (e.g. Leonard & Evans, 1997). In experimental plots in
Switzerland (Egli et al., 1990) and the Pacific USA (Pilz & Molina, 1996;
and see Chapter 3) no negative effect of intensive picking of fruit bodies
has been observed. Nevertheless, representatives of eastern Europe and
Portugal expressed great concern on the situation in their countries during
ECCF meetings. It is quite possible that destructive harvesting and side
effects of collecting, such as trampling, have a negative impact in some
regions. Also accurate data on quantities harvested and exported are
lacking. It would be a good possibility for European co-operation to
initiate a project on the ecological, economic and social aspects of mushroom harvests.
Red Data lists
Red Data lists enumerate species that are considered as threatened or
susceptible in a certain area. They are necessarily based on formal or
informal data on distribution, ecology and trend of species, so in fact on a
compilation of mycological knowledge. In agreement with guidelines of
the International Union for Conservation of Nature (IUCN) (Anon.,
1995a), usually five categories of threatened species are distinguished:
extinct, critically endangered, endangered, vulnerable and susceptible or
rare.
Since the first Red Data list of fungi appeared in 1982, many European
countries have published formal or informal Red Data lists, occasionally
also revised editions. Many Red Data lists are difficult to trace since they
71
Fungi in Europe
Table 4.1. National Red Data lists of macrofungi in Europe
Country
Authors
Year
Number of
species
Austria
Austria (2nd edn)
Czech Republic
Denmark
Estonia
Finland
Finland (2nd edn)
Germany East
Germany West
Germany (2nd edn)
Great Britain
Greece
Hungary
Latvia
Lithuania
Macedonia
Netherlands
Netherlands (2nd edn)
Norway
Norway (2nd edn)
Poland
Poland (2nd edn)
Spain and Portugal
Sweden
Sweden (2nd edn)
Switzerland
Ukraine
USSR (former)
Yugoslavia
Krisai
Krisai
Kotlaba et al.
Vesterholt & Knudsen
Anon.
Rassi & Väisänen
Rassi et al.
Benkert
Winterhoff
Benkert et al.
Ing
Diamandis
Rimóczi
Vimba & Peterans
Kutorga et al.
Karadelev
Arnolds
Arnolds & van Ommering
Bendiksen & Hoiland
Bendiksen et al.
Wojewoda & Ławrynowicz
Wojewoda & Ławrynowicz
Calonge
Anon.
Aronsson et al.
Senn-Irlet et al.
Shelyak-Sosonka
Borodin et al.
Ivancevic
1986
1999
1995
1990
1995b
1987
1992
1982
1984
1992
1992
2000
1998
1996
1999
2000
1989
1996
1992
1997
1986
1992
1993
1991
1995
1997
1996
1984
1998
211
542
120
898
76
161
325
309
1032
1402
453
150
535
38
740
67
944
1655
649
831
800
1013
153
514
528
232
56
17
97
were published in rather obscure journals or pamphlets. A survey of
national Red Data lists, hopefully up to date, is presented in Table 4.1. A
total of approximately 3000 species are included in one or more lists.
Regional lists have been omitted, such as the well-documented lists for
most German states.
The status of Red Data lists varies from official documents, approved by
the government to informal, provisional papers, compiled by a group of
mycologists or even a single individual. Also, the applied methods and
criteria vary considerably. Most Red Data lists are necessarily based on
expert judgement because of lack of reliable data on distribution of species
in the past. Only in The Netherlands has an attempt been made to use
72
Eef Arnolds
quantitative criteria (Arnolds & van Ommering, 1996), as prescribed by
IUCN (Anon., 1995a). In that country a large set of 700 000 computerised
data was available. Nevertheless, it was only possible to use the IUCN
criteria after adaptation to the specific characteristics of the data set. One
third of the species could not be considered because of lack of data. It was
also concluded that a Red List based on expert judgement may produce
comparable results, in some cases even more realistic results (Arnolds,
1997). I consider the international trend for scientific perfection in Red
Data lists as a risk to progress in compilation of Red Data lists of fungi and
therefore for conservation of the mycota. It cannot be denied that available databases in almost all European countries are too incomplete to meet
the quantitative criteria of IUCN.
The number of species on national Red Data lists varies between 17 in
former USSR and 1655 in The Netherlands (Table 4.1). It is evident that
these figures are by no means representative of the real threats to the
mycota in various countries. Indicative is the difference in the number of
listed species in similar countries such as Latvia (38 species) and Lithuania
(740 species). The compilation of the lists depends, among other things, on
the available data, the applied criteria and the aims of Red Data lists. For
instance, in some countries all species on the list are automatically protected by law, but in other countries lists do not have such consequences.
In some countries two editions of a Red Data list have already been
published. The second edition invariably contains more species than the
first. In general, the increase in species is mainly caused by increased
knowledge.
In the framework of necessary European co-operation, standardisation
of the procedures is badly needed. International criteria should be developed by the ECCF that bridge the gap between rigid IUCN criteria (Anon.,
1995a), specific mycological needs and practical possibilities. A preliminary Red Data list for Europe, based on available national lists, was
published by Ing (1993). It contains 278 species.
Threatened habitats of fungi
Conservation of fungi is, like conservation of other organisms, in the first
place conservation of their habitats combined with adequate management.
Conservation of appropriate habitats is also essential for the protection of
phanerogams, birds and other organisms that are more popular and better
known than fungi. In my opinion we may leave the initiative for the
conservation of most habitat types to these specialist groups and national
Fungi in Europe
73
and international organisations with a general conservation goal, such as
the IUCN and WWF. If a tropical rainforest is protected for the rhinoceros or tiger, fungi will also automatically profit from the maintenance of
the ecosystem. In Europe many efforts are already under way for conservation of habitats such as peat bogs, sea dunes and alpine meadows. The
mycological community can support these attempts by delivering additional data on the biodiversity of fungi in those areas and the occurrence of
endangered species.
However, mycologists should pay special attention to those habitats
that are rich in (threatened) fungal species and that are not adequately
characterised by other organisms, in particular plants. In this connection I
mention primeval forests, forests on very poor soils and old grasslands.
These examples are certainly incomplete, also because of lack of relevant
information from large parts of eastern and southern Europe.
Primeval, virgin and old-growth forests share an undisturbed development of vegetation and soil extending over several centuries. In Europe
they are found mainly in remote areas in the north and east, elsewhere in a
few strict reserves. Characteristic features are abundance of large dead logs
in all stages of decay, uneven age of trees, the occurrence of gaps and
uneven soil surface caused by holes left by uprooted trees. Characteristic
phanerogams are few, but primeval forests are well characterised by a
number of lichens, bryophytes and fungi, along with some groups of
animals. In Scandinavia and the Baltic states extensive programmes are
carried out to locate and preserve valuable old forest relics. Polypores and
other lignicolous Aphyllophorales are used as indicator organisms (e.g.
Kotiranta & Niemelä, 1993; and see Chapters 5 and 15). Old-growth
forests are even rarer and more endangered in the belt of deciduous forests
and in the Mediterranean area. Also in these cases characteristic fungi may
be used as indicator species, e.g. Hericium coralloides for undisturbed
beech forests and Podoscypha multizonata for very old oak trees.
Both coniferous and frondose forests on very oligotrophic soils, in
particular poor in nitrogen, are extremely rich in ectomycorrhizal fungi.
Many species are restricted to these forest communities, such as the
Cladonio-Pinetum (e.g. Wöldecke & Wöldecke, 1990), Dicrano-Quercetum
(Jansen, 1984), Luzulo-Fagetum leucebryetosum and Carici-Fagetum (Jahn
et al., 1967), the latter type growing on calcareous slopes. Apparently, the
slowly growing trees on these sites can only survive with the aid of their
numerous mycorrhizal partners. These forest types are very sensitive to
eutrophication, caused either by gradual accumulation of nitrogen in
forest soils due to air pollution, or by deliberate application of fertilisers in
74
Eef Arnolds
order to increase timber production. Special attention must be given to
conservation of these habitats in central and western Europe since nitrogen
deposition exceeds critical values in many regions. In strongly polluted
areas, such as The Netherlands and northern Germany, oligotrophic
forests have become extinct or greatly impoverished and so is their mycota
(Arnolds, 1991; Wöldecke, 1998).
A third, strongly threatened habitat type of immense mycological significance is that of old, unfertilised, semi-natural and natural grasslands.
These communities are rich in specialised saprotrophs, such as many
species of the genera Hygrocybe, Entoloma, Clavaria and Geoglossum.
They are often indicated as ‘Hygrocybe grasslands’. Old-growth grasslands, parallel to old-growth forests, could be an alternative term. The
most characteristic species depend on continuous grassland use over decades or centuries, undisturbed soil and management by grazing or cutting.
Some areas are rich in rare phanerogams as well, but many old-growth
grasslands do not possess a spectacular flora of green plants. These communities were widespread in traditional cultural landscapes, but are
threatened nowadays all over Europe by two opposite trends: intensification of agriculture and abandonment or afforestation. Special attention is
paid to the conservation of these grasslands in Scandinavia (e.g. Boertmann, 1995) and Great Britain (see Chapter 10).
All three mycologically important habitat types are very suitable for
European co-operation concerning inventories, monitoring and attempts
to improve conservation and management. It should be possible to obtain
European funding for research on these subjects. But who is taking the
initiative?
Fungi in conservation practice in Europe and North America
Important progress has been made in gathering knowledge on the needs of
fungal conservation, thanks to only a small number of professional and
amateur mycologists who are devoted to the subject. Most people involved
are all-round mycologists, usually also involved in research on taxonomy
and/or ecology and teaching. Specialists in fungal conservation do not yet
exist as they do in other disciplines of biology. Nevertheless, numerous
data on the subject are available for anyone interested in publications in
scientific and popular journals, proceedings of meetings and Red Data
lists. However, the practical impact of this information has been limited up
to now. The occurrence of threatened fungi is still rarely used as an
important criterion for selection of nature reserves or the application of a
Fungi in Europe
75
certain management. There is little progress in legal protection of fungi,
either in national or international legislation. Also, funding of research in
favour of fungal conservation lags far behind funding of projects on birds,
mammals, phanerogams, butterflies, dragonflies and so on.
The impact of fungi in international nature conservation is still minimal. For instance, the influential International Union for Conservation
of Nature (IUCN) has numerous expert groups, covering all living organisms. There are even different expert groups for Indian and African
elephants. The entire Kingdom Fungi is represented by only one expert
group that has published a few newsletters, but during the last five years
it has undertaken no action at all. The organisation Planta Europa was
established in the early nineties to co-ordinate and stimulate conservation
of plants throughout Europe. Mycologists were not involved and were
even unaware of this initiative until some representatives of the ECCF
were invited to attend the second congress in Uppsala and to present
information on fungus conservation there (Arnolds, 1999). During that
congress we learnt much about European legislation on nature conservation, for instance about the importance of the Bern Convention. This
convention from 1979, signed by 28 European countries, regulates protection of threatened animals and plants and their habitats. Some mosses
and lichens are included, but fungi are completely lacking. At present
attempts are being made to have some species protected under these
regulations.
Another recent international development is the European Community
Directive on the Conservation of Habitats, Fauna and Flora: in short,
Habitats Directive. Member states can be obliged to protect areas with
great significance for biodiversity. Again, fungi are not considered and
mycologists are not involved in this initiative.
In North America mycological knowledge about taxonomy, distribution as well as ecology, is less complete than it is in Europe. Conservation
was not on the agenda of mycologists for a long time until harvesting of
wild mushrooms became a booming business in the late eighties. In particular in the Pacific Northwest, many studies are currently investigating
the influence of harvesting and forest management on the mycota. An
anthology of these studies was recently published by Pilz & Molina (1996;
and see Chapter 3). The future of old-growth forests is an important
political question in this region, which even directly concerned President
Clinton. Timely warnings by mycologists have emphasised the crucial
importance of fungi in these ecosystems and their suitability as bioindicators. They have successfully influenced management practices and research
76
Eef Arnolds
efforts, as exemplified by this quotation from the report by Pilz & Molina
(1996):
The final record of decision is the document that specifies how the Final
Supplemental Environmental Impact Statement (an official governmental document) will be implemented. Table C-3 in that document lists 234
fungus species deemed of ‘special concern’. These species were analysed
and classified into four survey categories. Survey strategy 1 requires
management of known sites for all species shown in Table C-3. Sites with
rare and endemic fungus species will have 64.8 hectares (160 acres)
temporarily withdrawn from management activities until those sites can
be thoroughly surveyed and site-specific management measures prescribed. One particular fungus, the ‘noble polypore’ (Oxyporus nobilissimus), will have 240 hectares (600 acres) of habitat preserved at each
known fruiting location. Survey strategy 2 requires surveys before the
occurrence of any ground-disturbing activities. Only two fungus species
require this strategy: Oxyporus nobilissimus and Bondarzewia montana.
Survey strategy 3 requires conducting extensive surveys to find highpriority sites for species management. All 234 fungus species are included
in survey strategy 3, and protocols are currently under development. The
general regional surveys of strategy 4 are designed to provide further
information about little known species not yet designated as rare and
endemic. These regional surveys must begin before 1997 and are expected
to take 10 years to complete.
In Europe, mycologists can only dream about efforts on this scale for
fungus conservation and research on threatened species!
Conclusions
In the last two decades conservation of fungi has become one of the
important duties and fields of research for European mycologists, in
particular field mycologists working on macrofungi. At the same time the
number of professionals involved in these activities has decreased in most
countries. This results in growing responsibilities for fewer people. The
practical results of efforts related to fungus conservation are rather limited
so far. Financial support for research and public information is also poor.
The countries of Fennoscandia are positive exceptions in both respects. I
think that the general situation is caused by (1) failure to spread the
message sufficiently outside circles of mycologists, (2) inadequate attention
to international developments, (3) lack of acquaintance with and unclear
status of organisations, such as ECCF, (4) lack of international co-operation in mycological affairs.
Fungi in Europe
77
I am convinced that fungi deserve, and can gain, more attention in
nature conservation, environmental policy and management of nature
reserves. Three properties are in favour of this group of organisms: (1)
their enormous species diversity and functional importance as decomposers and symbionts, (2) the economic and recreational significance of
picking edible mushrooms, and (3) most important of all, the magic of
mushrooms: their beauty and variety in shape and colour, their sudden
appearance, their role in imagination. Mycologists should make use of the
popularity of fungi among people. The present public interest in biodiversity and conservation can and must be used to obtain funding for research
and international co-operation, related to conservation of fungi.
The ECCF is a very useful, but informal, expert group without any
official status and without funding. It would be better for it to be linked to
an authoritative organisation, such as IUCN. Then it would be automatically regarded as an authoritative body and a partner for international
treaties. In this connection one might also think of the establishment of a
European Mycological Society. Such a society should be the organiser of
future Congresses of European Mycologists, another institution without a
sound basis in the international scientific arena. It could also play an
important role in co-operation and integration of mycological research,
e.g. concerning a common checklist, mapping and monitoring. The publication of a European Journal for Mycology might be useful in this respect.
It is obvious that conservation of fungi depends on the input of mycologists. But at present it seems to be also the other way around: the future of
mycologists depends on their input in conservation.
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5
Fungi as indicators of primeval and
old-growth forests deserving protection
ER A S T P A R M A S T O
Introduction
Conservation of fungi, particularly populations of rare and endangered
fungal species, is only possible when their habitats are protected. Consequently, efforts in fungal conservation will be successful only when cooperation with other environmentalists interested in nature conservation is
established. Joint, well-motivated proposals to create new protected areas
or to regulate a conservation regime in existing reserves will then accepted
more easily than otherwise. The role of mycologists is not only to study the
mycota of nature reserves, but also to take the initiative in creating new
reserves.
Protection of primeval and old-growth forests
Primeval forests are very rare in Europe. Some kind of forest management
has been practised almost everywhere except in those forest stands that are
low in productivity and poor in species composition growing on oligotrophic Sphagnum bogs in northern areas. Seminatural old-growth forests
with minimal human impact have survived mainly in nature reserves.
In Estonia, of the 138 hemerophobic vascular plant species found, 90 are
growing only or mainly in forests (Kukk, 1999; Trass, Vellak & Ingerpuu,
1999); there are 79 hemerophobic bryophytes and 88 lichens in the Estonian forests. In the boreal zone, old unmanaged forest is the main
refugium for rare and endangered species, including fungi. Protection of
different forest site types is effective for conservation of species as well as of
communities.
In the European Union, the main target of nature protection has been
different groups of higher organisms (primarily birds) and a few selected
habitat types (wetlands, Mediterranean communities). Primeval or
81
82
Erast Parmasto
old-growth forests have been included in nature reserves in earlier times;
most of the European forests belong to nongovernmental owners and are
intensively managed. In several countries of eastern and Central Europe,
ownership has changed during the last decade. The forests are being
privatised. Due to the economic difficulties in these countries, the scale of
forest cutting has increased considerably. At the same time, due to the
influence of European ideology, nature protection has received much more
attention than earlier. Among other states, in Estonia the NATURA 2000
plan for the creation of a network of protected sites has obtained priority.
Following the political decision of the Estonian Parliament, the intention
is to increase the area of strictly protected forests to at least 4% (equivalent
to 80 000–90 000 ha).
Inventory of forests in Estonia
At the beginning of 1900s about 14% of the area of Estonia was covered
with forest stands and this increased to 50.1% (2 187 700 ha) during the
century (Ratas & Raukas, 1997; Kohava, 2000). The cause of the increase
is a combination of afforestation of land unsuitable for agriculture in the
last fifty years and the drastic decline of agriculture that has occurred in the
last decade. Half of the forest area (about 1 000 000 ha) belongs to the
State; 2.7% of the total area of Estonia is dedicated to four national parks
and nature reserves make up about 1.4%. In total, about 11% of Estonia
has some kind of nature protection (Randla, 1996). About 3% of forests is
in nature reserves, but this area also includes fens, peat bogs and young
forests.
A further half of Estonian forests has been or will be returned to private
owners. Due to the economic difficulties in the country and lack of interest
in sustainable forestry by most of the owners, coupled with the liberal
government policy, this second half is subjected to unlimited cuttings even
now.
Twenty-two different types of site and 71 types of forest have been
distinguished in Estonia including boreal and nemoral forests. Because of
intensive forest management, old-growth natural forests can rarely be seen
in this country; intact primeval forests cover only 0.2% of the protected
forest area (Randla, 1996). In Tartumaa County, 60% of the old-growth
forests found in 1993–1994 were clear-cut before 2000. Virgin or primeval
forests exist only in a few strictly protected nature reserves.
In the last two to three years, two new co-ordinated projects have been
initiated. The intention of an Estonian–Danish project is to create a Forest
Fungi as indicators of old-growth forests
83
Conservation Area Network in state forests, while the other is an Estonian–Swedish project, Inventory of Woodland Key Biotopes (Habitats).
One of the aims is to increase the strictly protected forests from 3% to 4%
of the total forest area including all the main but also the rare forest types.
Inventory of forests deserving protection
General approach
A preliminary list of what are presumed to be valuable forest stands in
state forests has been compiled for the Network Project using the Forestry
database at the Estonian Forest Survey Centre, digitised forest maps and
aerial photographs. Rare forest types with an age of at least 70 years are
growing on 20 455 ha; all state forests pre-selected for further study together cover an area of about 267 300 ha (K. Viilma & J. Öövel, unpublished). These stands were studied in nature by foresters and sites selected
by them were then studied by a group of biologists including botanists (as
well as bryologists and lichenologists), a mycologist, an entomologist
and (when additional data were needed) ornithologists. In 1998–2000,
29176 ha of forests were considered to be deserving of protection. The
forest authorities exclude any management in the selected stands until a
final decision on their fate has been made.
In nongovernmental forests, about 170 000 ha has been pre-selected to
find key biotopes; during the first year of the pilot field inventory, about
670 habitats (covering 1247 ha) deserving protection were found (Andersson, Ek & Martverk, 1999). Trained foresters carried out the inventory.
Several field courses have been organised to train these personnel, especially including identification of Estonian Red Data list species and indicator
species. This inventory of key habitats will be continued in all forests
regardless of ownership over the next few years.
Mycological studies
For inventory of forests deserving protection, and of key biotopes, fungi
may be used as good indicators of the species richness of the biota. In
contrast to vascular plants, the species richness of fungi, many microorganisms and insects depend heavily on the forest management type, age
of forests and amount of coarse woody debris. In boreal forests, the
number of wood-rotting species of fungi is up to four times higher on old
decomposed fallen trunks compared with freshly felled trunks (Niemelä,
84
Erast Parmasto
Renvall & Penttilä, 1995). The number of rare fungal species has been
demonstrated to be four times less in managed forests than in key biotopes
in Norway (Stokland, Larsson & Kauserud, 1997). My own data indicate
that in Estonia there are about five times fewer rare fungi in managed
forests. At late stages of wood decay, the trunks maintain ‘exceptionally
diverse species combinations’ of saprotrophic fungi (Renvall, 1995). The
highest numbers of wood-inhabiting fungal species, including Red Data
list species, were found in old forests with numerous dead logs in Central
Norway (Høiland & Bendiksen, 1997). More than a hundred rare species
of Polyporaceae and Corticiaceae sensu lato have been found only in
unmanaged old forests in Estonia (E. Parmasto, unpublished).
Diversity of species composition of fungi is a sign of general biodiversity
in a site. In many cases, stands in Estonia with a rich fungal biota were
found to have several rare species of vascular plants, insects and birds.
To characterise the level of biodiversity of a site, the presence of Red
Data list species has been used for the Woodland Key Biotopes (Habitats)
Project in Latvia, Sweden and Estonia. In The Netherlands, the mycological value of sites has been evaluated using a weighted sum of Red Data list
species occurring in a site (Jalink & Nauta, 1999). However, there is a big
difference between the Red Data lists in Estonia and in most other countries. In Estonia, 91 species of fungi have been included in the Red Data
list, in Finland 325, Sweden 528, Norway 758, Denmark about 900, Poland
1013, The Netherlands 1655, and former German Federal Republic 1729.
In several countries, the aim of including species in a Red Data list has
been the continuation of a ‘floristic’ inventory of species under the modern
slogan of ‘Biodiversity Studies’.
A limited number of rare or very rare species recognisable by nonspecialists has been included in the Red Data list in Estonia. The intention
is to give these species some practical protection, which is not mere formal
protection status ‘on paper’, and to inform people that fungi are worthy of
protection equally with other species of living beings. When the Estonian
Red Data list of fungi is used for evaluation of old-growth forests or key
habitats, the number of forest stands worthy of protection will be much
smaller than in countries where ten times more species have been included
on their equivalent list.
Use of indicator species
Karström in Sweden, and Kotiranta and Niemelä in Finland (references
below) initiated the use of fungi as indicators of old-growth natural forests
Fungi as indicators of old-growth forests
85
in 1992–1993. This has been mainly based on qualitative observations not
supported by quantitative studies. In Sweden, only five indicator species
associated with old-growth forests were used by Karström (1992a,b).
Kotiranta & Niemelä (1993, 1996) list 34 species characteristic for old
forests and 23 for virgin forests. In Latvia, ten indicator species were listed
by Meiere (1996); in former Yugoslavia, 36 indicator species have been
selected by Tortic (1998). A list of 43 indicator species for Estonia was
published by Erast and Ilmi Parmasto in 1997; the number of suitable
species has increased to 50–55 in 1999.
Selection of indicator species is based on different species distribution
influenced by different ecological conditions (Bondartseva, 1999), and
correlation between presence of indicator species and general high species
diversity. Høiland & Bendiksen (1997) have demonstrated that the highest
species diversity of fungi is found in forests with access to logs with high
levels of decay and large dimensions lacking in managed forests. The
old-growth forest indicator fungi are more or less easily recognisable
macrofungi associated mainly or only with old-growth natural (primeval)
forests minimally affected by forest management and other kinds of human impact. Indicator species indicate species richness in such areas; that
is, they are indicators of natural biodiversity (Parmasto & Parmasto,
1997). The list of indicator species is specific for any region, and may be
specific for different types of forest site.
In a list of indicator species, only a few, if any, agarics (gilled mushrooms) and boletes have been included. To describe the species composition of agarics and boletes in a forest stand, the place must be visited more
than once a year (cf. Kirby, 1988). Tofts & Orton (1998) published a survey
on a 20-year study of the mycota of a nature conservation area in Britain
(about 3000 ha). During the 20 years, the accumulative curve of species
found is a straight ascending line. This is one reason why polypores and
other wood-rotting fungi must be preferred in a short-time inventory;
species with perennial fruit bodies are the best for this purpose.
The number of indicator species selected for fieldwork ought to be high
enough for there to be a reasonable chance they can be found in natural
forests that are candidates for protection. Five species, as proposed by
Karström (1992a,b), is obviously too low a number. These species are quite
rare and distributed randomly. A greater number may be suitable for
experienced mycologists but may not be practical for foresters without
special training. For the latter, species with brightly coloured or otherwise
morphologically visually striking fruit bodies are preferred. To limit the
number of species, very rare species are excluded.
86
Erast Parmasto
Table 5.1. List of indicator species for Estonia
Aphyllophorales, polypores
Ceriporia purpurea
Ceriporia reticulata
Diplomitoporus flavescens
Fomitopsis rosea
Ganoderma lucidum
*Hapalopilus salmonicolor
Leptoporus mollis
Oligoporus placentus
Perenniporia medulla-panis
Perenniporia subacida
Phaeolus schweinitzii
Phellinus chrysoloma
Phellinus ferrugineofuscus
Phellinus ferruginosus
Phellinus nigrolimitatus
Phellinus populicola
Physisporinus vitreus
Pycnoporellus fulgens
Rigidoporus crocatus
Skeletocutis odora
Skeletocutis stellae
Aphyllophorales, other groups
Aleurodiscus amorphus
Amylocorticium subincarnatum
Asterodon ferruginosus
Clavariadelphus pistillaris
Clavariadelphus truncatus
Dentipellis fragilis
*Gomphus clavatus
Hericium coralloides (syn.: H. clathroides)
*Hydnellum sp. (one species*)
*Lindtneria trachyspora
Multiclavula mucida
Phellodon sp.
Phlebia centrifuga
Phlebia subochracea
Pseudomerulius aureus
Sarcodon sp. (excluding S. imbricatum)
Serpula himantioides
Sistotrema raduloides
*Steccherinum robustior
Tomentella crinalis
Xylobolus frustulatus
Boletales
*Boletus satanas
Gasteromycetes
Geastrum sp. (8 rare, one common species)
Heterobasidiomycetes
*Eocronartium muscicola
*Tremiscus helvelloides
Ascomycetes
*Sarcosoma globosum
*Sowerbyella sp. (3 species)
*Xylaria polymorpha
* Species included in the Estonian Red Data list.
In Estonia, indicator species have been employed since 1995 for evaluation of forests in two nature reserve areas, and for the general inventory
of natural forests since 1998. During the years studied by mycologists, 175
of these were recorded as deserving protection. In most cases, three to five
indicator species were found in one selected forest stand, the record
number being 14.
Now the use of indicator fungal species is included in the list of standard
methods in use by the Forest Conservation Area Network Project. In the
Woodland Key Biotopes (Habitats) Project, indicator species combined
with the Red Data list species are in use.
In the list shown as Table 5.1, 44 species and five genera of fungi are
mentioned. The selection is based on 50-year observations in Estonian
Fungi as indicators of old-growth forests
87
forests. The five genera included are mainly comprised of more or less rare
indicator species that are not easily distinguishable by a nonmycologist.
Acknowledgements
I acknowledge the valuable personal communications of Kaili Viilma, M.
Sc., and Jürgen Öövel (Forest Conservation Area Network Project,
Tartu). This research was supported in part by the Estonian Science
Foundation (grants nos 2145 and 4086).
References
Andersson, L., Ek, T. & Martverk, R. (1999). Inventory of Woodland Key
Habitats. Final report. National Forestry Board, Estonia: Tallinn & County
Forestry Board: Östra Götaland, Sweden.
Bondartseva, M. A. (1999). Wood-destroying basidial macromycetes in
ecosystems of the different state: strategies of propagation and survival. In
Abstracts, XIII Congress of European Mycologists, p. 16, Alcalá de Henares
(Madrid).
Høiland, K. & Bendiksen, E. (1997). Biodiversity of wood-inhabiting fungi in a
boreal coniferous forest in Sør-Trøndelag County, Central Norway. Nordic
Journal of Botany 16, 643–659.
Jalink, L. M. & Nauta, M. M. (1999). Towards a better protection of mycological
valuable sites in The Netherlands: how to detect and manage them. In
Abstracts, XIII Congress of European Mycologists, p. 61, Alcalá de Henares
(Madrid).
Karström, M. (1992a). Steget före-presentation. Svensk Botanisk Tidskrift 86,
103–114.
Karström, M. (1992b). Steget före in det glömda landet. Svensk Botanisk
Tidskrift 86, 115–146.
Kirby, K. J. (1988). The conservation of fungi in Britain. Mycologist 2, 5–7.
Kohava, P. (2000). Estonian Forests 1999. Eesti Metsakorralduskeskus: Tallinn.
[In Estonian.]
Kotiranta, H. & Niemelä, T. (1993). Uhanalaiset käävät Suomessa. Vesi-ja
Ympäristöhallinnon Julkaisuja-Sarja B17, 1–116.
Kotiranta, H. & Niemelä, T. (1996). Uhanalaiset käävät Suomessa. Toinen,
uudistettu painos (with English summary: Threatened polypores in Finland.
Second revised edition). Edita: Helsinki.
Kukk, T. (1999). Vascular Plant Flora of Estonia. Teaduste Akadeemia Kirjastus:
Tartu & Tallinn. [In Estonian, with a summary in English.]
Meiere, D. (1996). Polypores as bioindicators in Mezole. In Fungi and Lichens in
the Baltic Region. Abstracts, 13th International Conference on Mycology
and Lichenology, Riga (ed. E. Vimba), p. 38–39. University of Latvia: Riga.
Niemelä, T., Renvall, P. & Penttilä, R. (1995). Interactions of fungi at late stages
of wood decomposition. Annales Botanici Fennici 32, 141–152.
Parmasto, E. (1999). Fungi as indicators of primeval forests deserving protection.
In Abstracts, XIII Congress of European Mycologists, p. 102, Alcalá de
Henares (Madrid).
88
Erast Parmasto
Parmasto, E. & Parmasto, I. (1997). Lignicolous Aphyllophorales of old and
primeval forests in Estonia. 1. The forests of northern Central Estonia with a
preliminary list of indicator species. Folia Cryptogamica Estonica 31, 38–45.
Randla, T. (1996). Nature conservation. In Estonian Environment: Past, Present
and Future (ed. A. Raukas), pp. 86–88. Ministry of the Environment of
Estonia: Tallinn.
Ratas, R. & Raukas, A. (1997). Main Outlines of Sustainable Development in
Estonia. Ministry of the Environment: Tallinn.
Renvall, P. (1995). Community structure and dynamics of wood-rotting
Basidiomycetes on decomposing conifer trunks in northern Finland.
Karstenia 35, 1–51.
Stokland, J. N., Larsson, K.-H. & Kauserud, H. (1997). The occurrence of rare
and red-listed fungi on decaying wood in selected forest stands in Norway.
Windahlia 22, 85–93.
Tofts, R. J. & Orton, P. D. (1998). The species accumulation curve for agarics
and boleti from a Caledonian pinewood. Mycologist 12, 98–102.
Tortic, M. (1998). An attempt to a list of indicator fungi (Aphyllophorales) for
old forests of beech and fir in former Yugoslavia. Folia Cryptogamica
Estonica 33, 139–146.
Trass, K., Vellak, K. & Ingerpuu, N. (1999). Floristical and ecological properties
for identifying of primeval forests in Estonia. Annales Botanici Fennici 36,
67–80.
6
Recognising and managing
mycologically valuable sites in
The Netherlands
LE O M. J ALI NK & MARI JKE M. NAU TA
Introduction
Until recently, nature conservation in The Netherlands has focused on the
protection of plants and birds; management plans for nature reserves
contain no measures aiming at maintenance or improvement of the mycota. The reasons for this were mainly practical: although a great deal is
known about the distribution, ecology and management of higher plants
and birds, there is hardly any knowledge concerning fungi. However, site
management that is good for birds and higher plants is not necessarily
beneficial for other groups of organisms, such as insects or fungi. Moreover, sites with valuable vegetation are not necessarily rich in fungi, and
sites with an abundance of very rare fungi can occur among vegetation of
little interest.
Since the Dutch Mycological Society started a project for the recording
of macrofungi in 1980, much more information about the distribution and
ecological preferences of fungi has become available. The Dutch Mycological Society’s database now contains more than a million records.
However, the data are not of direct use to managers of nature reserves. The
data must be filtered and interpreted to be of practical value.
In an attempt to raise interest in mycota in conservation management
plans the Dutch Mycological Society organised a meeting entitled ‘Fungi
and Nature Conservation, Implications for Management’ in 1993, to
which nature managers were also invited (Kuyper, 1994). The abstracts of
this symposium have been published in Dutch and are widely distributed
between organisations and individuals involved in nature management.
Apart from expressing concern for fungi the impact of these abstracts has
been rather limited. The abstracts were not translated into practice and
hardly any change in management was observed.
Interviews established that among the reasons for unchanged practices
89
90
Leo M. Jalink & Marijke M. Nauta
were that wardens, rangers and managers of reserves have many obligations and limited time, the mycological value of their own reserves is
little known and no site-specific instructions were given. Evidently,
managers need clear instructions, and mycologically valuable sites should
be indicated on maps as precisely as possible. As an example of a site in
which more was achieved, a map of the nature management plan for the
‘Huys ten Donck’ estate does have mycologically valuable sites indicated
on it.
One of the positive results of the meeting and its abstracts was that some
organisations for nature management requested that the mycologically
valuable sites in their nature reserves be indicated, and asked for advice
concerning management of these sites. In order to comply with these
requests, the Committee for Fungi and Nature Conservation of the Dutch
Mycological Society has initiated the project ‘Mycological Reserves’.
The project ‘Mycological Reserves’
The aim of the project is to improve understanding of, and concern for,
fungi by site managers, namely rangers and wardens of nature reserves,
owners of estates, and municipality and county councils. This is achieved
by indicating mycologically valuable sites, not only within but also outside
nature reserves, and by giving advice on site management in order to have
fungi better protected.
An index has been developed to define the mycological importance of a
site. The mycological value is the weighted sum of Red Data list species
occurring in a site. The weighting factor is related to the threat category on
the Red Data list (Arnolds & Van Ommering, 1996). A species considered
to be seriously threatened is assigned four points, threatened three, vulnerable two and susceptible one point. The total of these species-specific
threat scores is then a measure of the mycological value of the site.
The first part of the project aims at an overview of the richest sites in The
Netherlands. In search of the Dutch ‘mycological crown jewels’, the project makes use of the database of the Dutch Mycological Society, although
this is organised according to map grid-unit, and no site names are included. Another problem is that as new checklists are published (e.g.
Arnolds, Dam & Dam-Elings, 1995), they bring about changes in interpretation of threat categories for taxa in the database and give new meaning to
others, thus creating some confusion in the database. A solution to these
problems is a recently devised digital checklist containing data such as Red
Data list status, and occurrence in The Netherlands, with coded ecological
Mycologically valuable sites in The Netherlands
91
preferences for every species found in The Netherlands (Anon., 1997). By
combining the distribution database with the digital checklist, a new
database was produced with relevant statistics per square kilometre including: number of records, number of species, number of Red Data list
species per category. This automatically calculates the mycological value
as defined above. Further, figures are given for two periods: from the
oldest record in the database (1850) until 1985, and then 1986 to the
present.
Subsequently, a map could be created, indicating the square kilometre
grid units with high mycological values. The association of the name of a
site with data expressed per square kilometre grid unit can only be done
with the help of people in the field. There can be several named sites per
grid unit, as well as single sites occupying more than one square kilometre.
Combining detailed ordnance survey maps with the known ecological
preferences of the observed fungi, it should be possible to relocate named
sites with a valuable mycota. Fortunately, the members of the Dutch
Mycological Society are very co-operative in finding the sites and their
appropriate names.
Results
As a result, several mycologically valuable sites have been rediscovered
which had not been visited at all in the last 20 years, an amazing fact in
such a well-investigated country. In other sites, once known as hot spots
for rare fungi, the mycota has declined, and sometimes the causes for this
could be indicated. A list of the 200 best sites has been published (Jalink,
1999). This list has already proved to be valuable since it has attracted the
interest of the owners and managers of sites included on it and in some
cases this has proved to be an incentive for improved management to the
benefit of fungi of the sites.
All information is now being verified, by encouraging fellow mycologists to visit the forgotten or poorly investigated sites, and report on their
state of management. Analysing the results confirms again that the mycota
in general is declining, something which has also been shown before
(Arnolds & Jansen, 1992; Nauta & Vellinga, 1993, 1995). Of the top 25
sites, many have a lower mycological value after 1985 when compared with
the period before (Table 6.1). The best site in The Netherlands according
to the definition used here is ‘Nijenrode’, an old estate on clay along the
river Vecht, which is fortunate to have a manager who is an enthusiastic
and knowledgeable mycologist. But many other sites are not managed for
92
Leo M. Jalink & Marijke M. Nauta
Table 6.1. The top 25 mycological sites in The Netherlands
Site name
Nijenrode
Neerijnen
Staverden:
Leemputten
Bunderbos
Tiendweg
Gunterstein
Groeneveld
Schaelsberg
Notenlaan
Amelisweerd
Eikenhorst
Twickel
Stiphout
Duin en
Kruidberg
Valkenswaard
Fort Rhijnauwen
Zuiderheide
Amelisweerd
Stokhem
Abbertse bos
Beatrix kanaal
Koningshof
Michielsgestel
Huys ten Donck
Voornes duin
Mycological
value index
1850–1998
Change between
pre-1985 and
post-1985?
623
518
402
Number of Red Data list
species
1850–1985
1986–1998
=
–
129
145
134
240
122
64
373
372
357
344
342
335
306
296
268
263
248
++
–
–
–
=
–
–
–
=
–
47
124
115
120
54
116
115
80
80
12
88
171
21
53
11
131
33
33
49
31
114
38
239
238
232
230
223
217
213
206
202
201
196
–
=
–
–
–
–
–
=
83
52
71
71
62
80
80
65
70
83
52
26
67
58
38
55
41
5
28
0
24
47
?key: –, decline 950%; -, decline 10–50%; =, change −10% to +10%; +, increase
10–100%; ++, increase 9100%.
the benefit of fungi, and sometimes site management is detrimental to
fungi.
Considering the geographical distribution of sites with the highest
number of Red Data list species, it is shown that many of these sites are in
the province of Utrecht and occur along the coast. The probable reason for
this distribution is the presence of calcareous soils combined with old
estates and avenues or relatively old woods, which can be found on clayey
soil in the province of Utrecht, and in the inner dunes along the coast and
in south Limburg.
Mycologically valuable sites in The Netherlands
93
Table 6.2. Guidelines for mycological management: protecting fungi on
valuable road side verges
Do
Don’t
remove leaf litter
mow and remove the hay
use mycorrhizal trees when replanting, such as Quercus, Betula, Tilia
use nutrient-poor loamy or sandy soil if in-filling is necessary
use fertilisers
use lawnmowers or garden shredders
add nutrient-rich (organic) soil
spread organic debris and mud from ditches on the road side verge
remove all trees at once (but instead replant gradually and preferably
with the same kind of trees)
Table 6.3. Guidelines for mycological management: protecting fungi in old
forests in estates and parks
Do
Don’t
remove leaf litter from mossy patches
leave old trees
use mycorrhizal species if additional tree planting is necessary
remove dense undergrowth of species like Rubus, Acer, Prunus
serotina
spread wood chips (from a garden shredder) on the forest floor or on
mossy patches
remove dead trees
plant too many trees that produce slowly decomposing litter
(Aesculus, Quercus, conifers)
run heavy vehicles on the forest floor
Management
Concerning the management of mycologically valuable sites, the Committee for Fungi and Nature Conservation is working on practical guidelines
of mycological management, grouped by habitat. As examples, the guidelines for two habitats are given here (Tables 6.2 and 6.3), with an overview
of management which is thought to be beneficial for fungi (the Dos), and
management which is often carried out with good intentions but thought
to be harmful for fungi (the Don’ts). These overviews will be published in
the form of leaflets and handed out to any person or organisation that is
responsible for a valuable site.
The guidelines are not only meant for nature managers, but also for
owners of estates, and municipality and provincial councils. Indeed, for
everybody who has to deal with management of nature in whatever way.
Although most nature managers have good intentions, bad mistakes are
94
Leo M. Jalink & Marijke M. Nauta
sometimes made. For example, using wood chips on a roadside verge with
rich Hydnaceous mycota, or dead branches on moss-rich vegetation to
prevent people from walking on it. Both measures result in eutrophication,
followed by a rapid increase of grasses and nitrophilous plants and declines of both moss flora and mycota.
Discussion
Of course, this method of estimating the mycological value of sites has its
limitations. Such an exercise can only be carried out if a carefully evaluated
Red Data list already exists, and if there are enough distribution data.
Also, the method largely ignores the biodiversity of organisms smaller
than about 1 mm, but not enough is known at present to allow a careful
evaluation of the mycological value of sites on the basis of their microfungi. Consequently, such consideration is confined to the macrofungi.
But by locating and giving guidelines for better management of mycologically valuable sites it is at least a start towards better protection of them.
References
Anon. (1997). BioBase 1997, Register Biodiversiteit. Centraal Bureau voor de
Statistiek: Voorburg/Heerlen.
Arnolds, E., Dam, N. & Dam-Elings, M. (1995). Standaardlijst van Nederlandse
macrofungi 1995. In Overzicht van de paddestoelen in Nederland (ed. E.
Arnolds, T. W. Kuyper & M. E. Noordeloos), pp. 754–828. Nederlandse
Mycologische Vereniging: Wijster.
Arnolds, E. & Jansen, E. (1992). New evidence for changes in the macromycete
flora of The Netherlands. Nova Hedwigia 55, 325–351.
Arnolds, E. J. M. & van Ommering, G.(1996). Bedreigde en kwetsbare
paddestoelen in Nederland. Toelichting op de Rode lijst. IKC Natuurbeheer:
Wageningen.
Jalink, L. M. (1999). Op zoek naar de mycologische kroonjuwelen van Nederland
1: De 200 meest waardevolle kilometerhokken. Coolia 42, 143–162.
Kuyper, T. (1994). Paddestoelen en natuurbeheer: wat kan de beheerder?
Wetenschappelijke Mededelingen van de Koninklijke Nederlandse
Natuurhistorische Vereniging 212.
Nauta, M. M. & Vellinga, E. C. (1993). Distribution and decline of macrofungi in
The Netherlands. In Fungi in Europe, Investigation, Recording and
Conservation (ed. D. N. Pegler, L. Boddy, B. Ing & P. M. Kirk), pp. 21–46.
Royal Botanic Gardens: Kew.
Nauta, M. M. & Vellinga, E. C. (1995). Atlas van Nederlandse paddestoelen. A. A.
Balkema Uitgevers B. V.: Rotterdam.
7
Threats to hypogeous fungi
MARIA ŁAWRYN OWIC Z
Introduction
The European Council for the Conservation of Fungi, during its fourteen
years of activity, focused attention on threats and conservation problems
of fungi. As a result of meetings, workshops and discussions, several
regional and national Red Data lists as well as a preliminary European
Red Data list (Ing, 1993), have been published.
A decline in the number of species of macrofungi has been reported from
different European countries (Arnolds & Jansen 1991). Hypogeous species
must also be included among the threatened fungi although hypogeous
fungi form a special ecological group of macromycetes because their fruit
bodies originate and remain underground until they break down naturally.
Elaphomyces is the most common genus in the northern part of Europe
and includes several species that are under threat. Tuber is the best-known
genus of hypogeous fungi in the Mediterranean area. Maturity of the fruit
bodies of many species is indicated by a strong pleasant smell. Indeed, the
aroma of truffles has attracted attention since ancient times and there are
some highly valued species of the genus Tuber, such as T. melanosporum, T.
brumale and T. aestivum, which have been exploited for consumption since
prehistoric times. In the history of western civilisation, truffles were a
frequent subject of research by natural philosophers. Because of the high
culinary value assigned to these fungi even early scientists like Theophrastus, Dioscorides and Pliny tried to investigate their nature and biology
(Pegler, Spooner & Young, 1993).
Numerous hypogeous species are involved in ectomycorrhizal associations with trees. They also have particular requirements concerning type
of soil and climatic conditions. They grow in a large variety of habitats
from virgin forests to sites in city centres (Hawker, 1954; Ławrynowicz,
1988). As they occur under the ground, mycologists often fail to record
95
96
Maria Ławrynowicz
them; hypogeous fungi require different methods of searching and special
techniques to enable them to be discovered.
The existing data concerning the occurrence and distribution of hypogeous fungi is still limited to scarce collections in various regions of
Europe (Ławrynowicz, 1989, 1990). As rare species, they are placed in the
regional and national Red Data lists of endangered fungi.
Red Data lists
Threatened species of macromycetes placed in the national Red Data lists
have been summarised by Arnolds & de Vries (1993) and these also include
hypogeous species.
Red Data lists of hypogeous fungi have been compiled on the basis of
data collected from Germany (Benkert et al., 1996), Poland (Wojewoda &
Ławrynowicz, 1992), The Netherlands (Arnolds & van Ommering, 1996),
Denmark (Vesterholt & Knudsen, 1990), Finland (Rassi & Väisänen,
1987), Sweden (Larsson, 1997), Norway (Bendiksen & Høiland, 1992), the
Czech Republic and Slovakia (Kotlaba, 1995) and HELCOM (Helsinki
Commission) – a draft working paper dated 27 November 1995 entitled
Red Lists of Macrofungi in the Baltic and Nordic Regions.
These Red Data lists include hypogeous fungi comprising 14 genera of
ascomycetes and 17 basidiomycete genera (Table 7.1). Altogether, 118
species are recorded, 56 ascomycetes (Table 7.2) and 62 basidiomycetes
(Table 7.3) are on the lists in one or other threat category. This means that
about 50% of all existing hypogeous fungi are threatened in these parts of
Europe.
The number of species in each country varies from 93 in Germany to 6 in
the Czech Republic and Slovakia taken together (Tables 7.2, 7.3 and 7.4).
In fact, the numbers of species included in these lists indicate the quality
and quantity of data available and state of our knowledge, rather than the
true extent of the real threat to these fungi. The example from Poland
illustrates this point: the number of ascomycetes is higher (Table 7.4)
because the taxonomic and ecological revision of this group has already
been made (Ławrynowicz, 1988), but basidiomycetes are still awaiting
study to this depth.
The final columns of Tables 7.2 and 7.3 show the number of countries
that list the threatened species. The majority of species are recorded on the
Red list of only a single country of the six that were examined (24 of the 56
ascomycetes and 31 of the 62 basidiomycetes). There are no species indicated on all national lists, although two ascomycete species and one
97
Threats to hypogeous fungi
Table 7.1. Numbers of threatened species in different genera of hypogeous
fungi
Ascomycetes
Number of
species
Basidiomycetes
Number of
species
Balsamia
Barssia
Choiromyces
Elaphomyces
Fischerula
Genea
Geopora
Hydnobolites
Hydnotrya
Pachyphloeus
Picoa
Sphaerosoma
Stephensia
Tuber
3
1
1
11
1
6
2
2
5
3
1
1
1
18
Alpova
Arcangeliella
Chamonixia
Elasmomyces
Gastrosporium
Gautieria
Hydnangium
Hymenogaster
Hysterangium
Leucogaster
Martellia
Melanogaster
Octavianina
Rhizopogon
Sclerogaster
Stephanospora
Wakefieldia
3
2
1
2
1
3
4
16
8
2
1
6
1
9
1
1
1
Total
56
Total
62
species of basidiomycete are found on six of the lists (Table 7.5). It is worth
noting that these three species: Pachyphloeus citrinus, Hydnobolites cerebriformis and Chamonixia caespitosa all require natural forest conditions.
The number of extinct species is fairly high (25). These species (Table 7.6)
need to be checked and analysed first.
As we can see from this analysis of Red Data lists, the strategy or
inclusion of hypogeous fungi varies between two extremes. The first is that
some authors place almost all known hypogeous fungi appearing in the
country among the threatened fungi on the basis of their rarity. The second
case is that no hypogeous fungi are included on the Red Data list. These
extreme viewpoints emphasise the deficiency of our current knowledge and
lack of data. The following reasons account for this: (1) the real status of
occurrence has not been recognised because only a few, if any, people in
the country specialise in this group of fungi and a random collection
cannot be established as the basis for the evaluation; (2) it is not possible to
verify all the existing data to confirm reliability; (3) taxonomic difficulties
and nomenclatural problems in this group require the support of experts.
98
Maria Ławrynowicz
Table 7.2. Threat categories of hypogeous ascomycetes in national Red
Data lists
Country
Species
D
PL
NL
DK
FIN
S
Hydnobolites cerebriformis
Pachyphloeus citrinus
Balsamia platyspora
Elaphomyces anthracinus
Hydnotrya michaelis
Tuber aestivum
Tuber maculatum
Elaphomyces leveillei
Genea verrucosa
Genea hispidula
Pachyphloeus melanoxanthus
Tuber rapaeodorum
Choiromyces meandriformis
Elaphomyces aculeatus
Genea klotzschii
Pachyphloeus conglomeratus
Stephensia bombycina
Tuber borchii
Tuber foetidum
Tuber puberulum
Tuber rufum
Balsamia vulgaris
Elaphomyces papillatus
Elaphomyces reticulatus
Geopora sumneriana
Hydrobotrya cerebriformis
Hydnotrya tulasnei
Sphaerosoma fuscescens
Tuber dryophilum
Tuber excavatum
Tuber mesentericum
Tuber scruposum
Balsamia fragiformis
Barssia oregonensis
Elaphomyces asperulus
Elaphomyces cyanosporus
Elaphomyces granulatus
Elaphomyces muricatus
Elaphomyces mutabilis
Elaphomyces striatosporus
Fischerula macrospora
Genea fragrans
Genea lespiaultii
Genea sphaerica
Geopora cooperi
Hydnobolites fallax
3
0
1
3
2
1
2
3
1
3
1
3
0
3
2
3
4
2
0
3
4
3
4
3
3
3
3
3
1
0
1
3
2
0
0
0
2
3
1
2
1
0
3
3
2
3
0
1
3
0
1
2
4
2
1
1
3
3
3
3
4
4
2
0
3
4
4
3
1
4
4
4
2
2
4
3
0
0
1
4
3
3
3
3
3
2
3
0
1
3
2
1
1
1
2
1
4
2
0
3
2
4
2
1
4
3
4
4
2
3
3
4
CZ-SK NCs*
4
4
3
1
4
4
1
3
N
6
6
5
5
5
5
5
4
4
4
4
4
3
3
3
3
3
3
3
3
3
2
2
2
2
2
2
2
2
2
2
2
1
1
1
1
1
1
1
1
1
1
1
1
1
1
99
Threats to hypogeous fungi
Table 7.2. (cont.)
Country
Species
D
Hydnotrya ploettneriana
Hydnotrya suevica
Picoa carthusiana
Tuber brumale
Tuber exiguum
Tuber foetidum ss. Hawker
Tuber macrosporum
Tuber michailowskiianum
Tuber murinum
Tuber nitidum
Total 56
PL
NL
DK
FIN
S
N
CZ-SK NCs*
1
1
1
1
1
1
1
1
1
1
1
3
0
1
0
1
0
1
1
4
43
25
21
14
5
12
9
1
* NCs, Number of countries with the species on the national Red Data list.
Ecological remarks
Observations of hypogeous fungi from the Red Data list in Poland lead to
the conclusion that it is not enough to protect the locality of hypogeous
fungi. Rather, it is necessary to provide them with special conditions,
taking into account the requirements of hypogeous fungi, which are often
different from that of epigeous macromycetes. Two particular examples
are worthy of note.
Tuber mesentericum. The most northeasterly locality for this
species in Europe is protected in a nature reserve near
Czȩstochowa (Ławrynowicz, 1988). The locality is situated on
top of a hill in a beech–oak–hornbeam forest on calcareous
soil. Ascocarps of T. mesentericum were collected on a
walking trail used every year by thousands of tourists and
pilgrims passing by. The vegetation on the trail is reduced to
10% of its normal cover, and after rains drainage water can
easily run down the trail taking debris with it and sometimes
uncovering the upper part of fruit bodies. This place has been
monitored twice a year, but the project is kept secret in order
to avoid its destruction.
Barssia oregonensis. The only locality in Europe where this
species is known to occur is protected in the Tatra National
Park (Ławrynowicz & Skirgiełło, 1984). The locality is
situated in a mossy Picea abies forest, cut through by a tourist
100
Maria Ławrynowicz
Table 7.3. Threat categories of hypogeous basidiomycetes in national Red
Data list
Country
Species
D
PL
Chamonixia caespitosa
Melanogaster tuberiformis
Gautieria morchelliformis
Alpova diplophloeus
Gastrosporium simplex
Hydnangium carneum
Hymenogaster vulgaris
Hymenogaster niveus
Hymenogaster tener
Hymenogaster arenarius
Hymenogaster olivaceus
Melanogaster variegatus
Octavianina asterosperma
Rhizopogon vulgaris
Arcangeliella stephensii
Elasmomyces mattirolianus
Elasmomyces krjukowensis
Gautieria otthii
Hymenogaster populetorum
Hymenogaster decorus
Hymenogaster muticus
Hymenogaster luteus
Hymenogaster rehsteineri
Hymenogaster megasporus
Hymenogaster hessei
Hymenogaster verrucosus
Hysterangium stoloniferum
Hysterangium calcareum
Hysterangium separabile
Melanogaster ambiguus
Rhizopogon hawkeri
Alpova klikae
Alpova rubescens
Arcangeliella asterosperma
Gautieria mexicana
Hydnangium pila
Hydnangium neuhoffii
Hydnangium cereum
Hymenogaster bulliardii
Hymenogaster cinereus
Hymenogaster thwaitesii
Hysterangium thwaitesii
Hysterangium clathroides
Hysterangium crassum
Hysterangium nephriticum
Hysterangium rickenii
Leucogaster badius
Leucogaster nudus
3
3
2
3
3
2
0
3
1
1
3
3
3
3
3
3
3
2
NL
DK
3
0
2
3
FIN
S
N
CZ-SK NCs*
2
2
2
2
4
4
2
1
3
0
1
4
2
3
3
4
1
1
3
4
4
1
3
2
0
2
3
4
2
2
2
3
3
2
3
3
3
3
3
3
3
3
3
3
0
2
3
3
4
3
4
4
4
0
4
2
3
4
2
4
4
2
2
2
0
2
3
2
1
2
4
1
3
1
3
3
4
6
5
4
3
3
3
3
3
3
3
3
3
3
3
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
101
Threats to hypogeous fungi
Table 7.3. (cont.)
Country
Species
D
Martellia soehneri
Melanogaster broomeianus
Melanogaster intermedius
Melanogaster macrosporus
Rhizopogon briardii
Rhizopogon luteolus
Rhizopogon luteorubens
Rhizopogon marchii
Rhizopogon parksii
Rhizopogon rubescens
Rhizopogon villossus
Sclerogaster compactus
Stephanospora caroticolor
Wakefieldia macrospora
Total 62
PL
NL
DK
FIN
S
N
CZ-SK NCs*
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
3
3
2
3
2
2
2
3
3
3
2
3
1
50
8
21
7
5
7
10
5
*NCs, Number of countries with the species on the national Red Data list.
Table 7.4. Numbers of hypogeous ascomycetes and basidiomycetes on Red
Data lists of different countries
Ascomycetes
Basidiomycetes
Total D
PL
NL
DK
FIN S
N
CZ-SK
56
62
25
8
21
21
14
7
5
5
9
10
1
5
43
50
12
7
Table 7.5. Distribution of hypogeous fungi between the national Red Data
lists of different countries
Number of lists recording the same species
Ascomycetes
Basidiomycetes
Total 1
2
3
4
5
6
56
62
11
17
9
11
5
1
5
1
2
1
24
31
102
Maria Ławrynowicz
Table 7.6. Extinct species of hypogeous fungi
Species
Country
Ascomycetes
Balsamia platyspora
Elaphomyces aculeatus
Elaphomyces papillatus
Genea hispidula
Genea klotzschii
Genea sphaerica
Pachyphloeus citrinus
Pachyphloeus conglomeratus
Picoa cartusiana
Sphaerosoma fuscescens
Tuber aestivum
T. borchii
T. excavatum
T. exiguum
T. foetidum
T. macrosporum
T. mesentericum
T. rapaeodorum
NL
D
D
D
D
D
D
D
D
D
PL
NL
NL
D
NL
D
D
NL
Species
Basidiomycetes
Alpova rubescens
Chamonixia caespitosa
Gautieria morchelliformis
Hydnangium carneum
Hydnangium neuhoffii
Hymenogaster hessei
Rhizogogon vulgaris
Country
D
PL
NL
NL
PL
NL
NL
route and with numerous walking trails running through the
forest. Ascocarps of Barssia grow on both sides of the trail,
partly in the upper layer of soil among mosses and liverworts
in areas trodden by tourists wandering off the trail to avoid
wet places.
As these examples imply, disturbance may encourage hypogeous fruit
body formation. Some hypogeous fungi are permanent components of
urban ecosystems, inhabiting ruderal places, growing under trees in parks
and streets, close to buildings (Ławrynowicz, 1988). Lilian Hawker (1954)
wrote: ‘Even those forms, such as Tuber puberulum, that grow within the
layer of partially decayed leaves, are not found where the leaves drift to
form a thick layer. Other species, such as Tuber excavatum, are found only
in places such as edges of woods, artificial mounds or banks or near the top
of slopes, where leaves do not accumulate. Slight disturbance of soil may
stimulate some species, presumably through improved aeration. Thus,
large numbers of young fruit bodies of T. puberulum were found on
returning to a patch which when searched, and therefore disturbed, some
months earlier had yielded only few. Tuber aestivum was found in quantity
Threats to hypogeous fungi
103
in the grounds of the University of Bristol one year after the ground had
been dug, after having been undisturbed for many years.’
For more than 300 years efforts have been made to cultivate truffles
extensively in the south of Europe. The most famous truffières are in
France. Here we can see the exact place where truffles produce their
ascocarps since the vegetation becomes extremely reduced. Tuber melanosporum, which yields fruit bodies over many years and sometimes in great
quantity, provides an optimistic example for the future of hypogeous
fungi.
On the basis of analysis of Red Data lists, together with field observations of several threatened hypogeous fungi, we arrived at the following
conclusions about steps necessary for the conservation of hypogeous
fungi.
Conservation of habitat. It is necessary to protect the locality of
a threatened fungus on a large scale, including potential
habitats of threatened species.
Protection of subterranean features of the habitat. The features in
need of attention are no accumulation of litter, scanty
vegetation cover, and exposure to direct sunlight and
occasional running water.
Monitoring of sites. It is necessary to collect the data concerning
frequency and abundance of threatened fungi to know the
trends and scale of their changes. Habitat requirements of
threatened species should be carefully observed also.
Habitat management. By afforestation or by changing the soil
structure it is possible to create new sites for hypogeous fungi.
Silent protection. Information on the occurrence of truffles
should not be distributed among the local inhabitants because
the locality is likely to be too intensively harvested.
References
Arnolds, E. & Jansen A. E. (1991). Conclusions of the First Meeting of the
European Committee on the Protection of Fungi, Łódź, August 10–13, 1988.
In Conservation of Fungi and Other Cryptogams in Europe (ed. A. F. Jansen
& M. Ławrynowicz), Science Tracks 18, pp. 90–103. Łódź Society of Science
and Arts: Łódź, Poland.
Arnolds, E. & de Vries, B. (1993). Conservation of Fungi in Europe. In Fungi of
Europe: Investigation, Recording and Conservation (ed. D. N. Pegler, L.
Boddy, B. Ing & P. M. Kirk), pp. 211–230. Royal Botanic Gardens: Kew.
Arnolds, E. & Van Ommering, G. (1996). Bedreigde en kwetsbare paddestoelen in
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Nederland. IKC Natuurbeheer: Wageningen.
Bendiksen, E. & Høiland, K. (1992). Red list of threatened macrofungi in Norway.
Report for 1992–6, pp. 31–42. Directorate for Nature Management: Oslo.
Benkert, D., Dörfelt, H., Hardtke, H. J., Hirsch, G., Kreisel, H., Krieglsteiner,
G. J., Lüderitz, M., Runge, A., Schmid, H., Schmitt, J. A., Winterhoff, W.,
Wöldecke, K. & Zehfuss, H. D. (unter Mitarbeit von Einhellinger, A.,
Gross, G., Grosse-Brauckmann, H., Nuss, I. & Wölfel, G. K.) (1996). Rote
liste der Grosspilze Deutschlands. Vegetationskunde 28, 377–426. Bundesamt
für Naturschutz: Bonn-Bad Godesberg.
Hawker, L. E. (1954). British hypogeous fungi. Philosophical Transactions of the
Royal Society of London, Series B 237, 429–546.
Ing, B. (1993). Towards a Red List of Endangered European Macrofungi. In
Fungi of Europe: Investigation, Recording and Conservation (ed. D. N. Pegler,
L. Boddy, B. Ing & P. M. Kirk), pp. 231–237. Royal Botanic Gardens: Kew.
Kotlaba, F. (1995). Červena kniha ohrazenych a vzácnych druhov rastlin a
zivoc̆ichov ČR a SR 4. Huby (Makromycéty), pp. 30–119. Priroda:
Bratislava.
Larsson, K. H. (1997). Rödlistade svampar i Sverige – Arfakta [Swedish Red Data
Book of Fungi]. Art. Databanken, Sveriges Lantbruksuniversitet: Uppsala.
Ławrynowicz, M. (1988). Flora Polska. Grzyby (Mycota) 18: Ascomycetes;
Elaphomycetales, Tuberales. Państwowe Wydawnictwo Naukowe:
Warszawa-Kraków.
Ławrynowicz, M. (1989). Chorology of the European hypogeous Ascomycetes, I.
Elaphomycetales. Acta Mycologia 25, 3–41.
Ławrynowicz, M. (1990). Chorology of the European hypogeous Ascomycetes.
II. Tuberales. Acta Mycologia 26, 7–75.
Ławrynowicz, M. & Skirgiełło, A. (1984). Barssia, a new genus in Europe. Acta
Mycologia 20, 277–279.
Pegler, D. N., Spooner, B. M. & Young, T. W. K. (1993). British Truffles. A
Revision Of British Hypogeous Fungi. Royal Botanic Gardens: Kew.
Rassi, P. & Väisänen, R. (1987). Threatened Animals and Plants in Finland.
Ministry of Environment: Helsinki.
Vesterholt, J. & Knudsen, H. (1990). Truede Storsvampe I Denmarken Rødliste.
Foreningen til Svampekundskabens Fremme: København.
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Cracow.
8
Wild mushrooms and rural economies
DA VID ARORA
Introduction
Wild mushrooms have been praised by Roman and Chinese emperors
alike while providing an important everyday food source for rural people
around the world. But in America and other English-speaking countries, a
deep and exaggerated distrust of mushrooms has denied them a cherished
place at the dinner table. As Stephen Jay Gould has pointed out (Gould,
1997), this prejudice is expressed even in our everyday language: urban
crime and taxes are said to ‘mushroom’ while prosperity and the arts
‘flower’!
Until recently very few Americans dared to eat wild mushrooms, and
even fewer picked them. But in the 1980s, rising demand in the wealthier
mushroom-loving countries caused them to look abroad for new sources
of wild mushrooms. At the same time, Americans’ palates grew bolder and
more sophisticated. Entrepreneurs rushed in to fill the rapidly growing
demand for gourmet foods, and out-of-work rural Americans and recent
immigrants (particularly from southeast Asia) saw picking wild mushrooms as a chance to make a decent living in a familiar environment, the
forest, while maintaining a measure of personal dignity and cultural autonomy.
Thousands of people in the Pacific Northwest now pick and sell wild
mushrooms. Most of them pick locally or opportunistically for a little
extra cash or as one of several seasonally based strategies for survival. But
the notoriously fickle nature of mushrooms – they may be overwhelming
abundant one year and then frustratingly scarce the next – has created the
need for a skilled cadre of pickers and buyers (many do both) willing to go
where the rainbows lead them. And the rare combination of abundant
public lands and private vehicles makes it possible for whoever so chooses
to do just that.
105
106
David Arora
The global mushroom trade
Wild mushrooms have long been gathered intensively in many parts of the
world. But with the globalisation of trade, mushrooms are now being
picked in more places than ever before and they are travelling farther and
faster. The global trade in matsutake alone is estimated at US$3 to 5
billion annually; for chanterelles it is about US$1.5 billion.
In many developing countries wild mushrooms have become an important source of income for people in remote forested regions where there are
few other opportunities to make money. Impoverished farmers in Bulgaria, for instance, have bought new tractors with money gained from
selling boletes to Italy, and villagers in Zimbabwe pay school tuition fees
for their children by selling mushrooms from their native miombo woodlands, including chanterelles that they can ship out of season to Europe.
One particularly dramatic success story is in Champa, on the eastern
flank of the Himalayas where the Chinese provinces of Yunnan, Sichuan,
and Tibet come together. Virtually all of the Tibetan villagers in this
rugged forested region spend the summer months picking, buying, and
selling wild mushrooms, or servicing those who do.
Despite a short growing season, the mushrooms provide families with
anywhere from 50% to 100% of their annual income. In two months, some
Tibetan families living in matsutake-rich forests are able to earn more than
ten times the annual average wage of a worker in developed regions of
China like Shanghai. And in contrast to North America, where the pickers
are widely scattered and the material benefits of the mushroom harvest are
difficult to distinguish, the wealth generated by mushrooms in Champa is
dramatically evident to outsiders because virtually all of the local money
comes from the mushroom trade. Villages near matsutake beds are dotted
with new two-storey wooden houses built in the traditional Champan style
but several times larger and more ornate than anything known before.
Small shops and other businesses have begun to blossom as the new
homeowners look for other ways to invest their money.
Even in forests where less valuable mushrooms predominate (e.g.
chanterelles or gypsy mushrooms, a locally sold species) the villagers have
new houses. They are less grandiose than the matsutake ‘mansions’, but
are still a dramatic improvement over the hovels that characterise deforested areas.
Rural boomtowns financed by distant urban elites are certainly nothing
new. But what is extraordinary about this area in Champa is that intact,
healthy forests are perceived as the key to rural development rather than as
an impediment to it. Several villages have developed their own mushroom
Wild mushrooms and rural economies
107
management plans, timber harvest has been scaled back, and the indigenous culture is far more intact than in nearby areas that cater to tourism.
A wild mushroom primer
Japan and western Europe are the major importers of wild mushrooms,
but many food stores in America now carry a small selection of fresh and
dried fungi, and it has become de rigueur for up-market urban restaurants
to feature morels, chanterelles, boletes (porcini), lobster mushrooms,
candy caps, and other kinds, depending on their availability. Here is a
rundown on some of the kinds you may encounter.
Matsutake are the most expensive wild mushrooms after truffles. The
Asian species, Tricholoma matsutake, sells for as much as US$200 a piece
in Tokyo markets. The white North American version, T. magnivelare, has
the same complex, spicy fragrance but is somewhat cheaper. Found across
North America and south as far as Guatemala, it is most abundant in the
coniferous forests of the Pacific Northwest and the tanoak woodlands of
northern California. Nearly all of the crop goes to Japan, but in good years
you can find it fresh in America in Japanese restaurants and markets, and
in a few gourmet food stores. The Japanese pay phenomenal prices for
young specimens, but opened ones are considerably cheaper yet taste just
as good.
Chanterelles (Cantharellus cibarius and close relatives) are the best
known wild mushrooms on the west coast. The lovely, yellow-orange
mushrooms have a fruity fragrance and chewy texture, and can be refrigerated for up to a month after picking. They are found throughout the
world and are nearly always sold fresh. Up-market restaurants in America
serve them practically year-round. The autumn crop from the coniferous
forests of the Pacific Northwest is legendary. A slightly different species is
abundant in the oak woodlands of California during the winter. In the
summer, fresh chanterelles originate from Nova Scotia, Saskatchewan and
eastern Europe.
Morels (Morchella species) are prized in America as well as western
Europe. Their hollow, honeycombed caps are distinctive but blend in
uncannily with their surroundings. Morel hunting is a popular pastime in
the Midwest, but the really big crops occur after forest fires in the West.
The several species that are harvested differ somewhat in flavour and
colour. Fresh morels are most plentiful during the spring and summer, but
the Mexican crop comes in during the autumn and even into December.
Turkey and North America are the major exporters of fresh morels to
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Europe, while the Himalayan countries have been exporting large quantities of dried morels since World War II.
The king bolete (Boletus edulis and close relatives) is the most popular
wild mushroom of Europe. It goes under many names there: porcini, cepe,
steinpilz, borovik, etc. West coast king boletes are among the largest in the
world, attaining weights of 10 pounds or more, but are very perishable;
only limited quantities of young specimens are shipped fresh. Some are
frozen whole but most are sliced and dried. Fresh king boletes are served in
some up-market restaurants; they usually originate from the coastal and
mountain areas of the West, and vary considerably in quality. Dried
boletes are famous for their concentrated flavour and wonderful aroma.
They are available year-round and may come from almost anywhere,
China, Europe, South America, South Africa, etc., but beware of less
aromatic imitations.
Hedgehog mushrooms (Hydnum umbilicatum and H. repandum) are
similar to chanterelles in colour and texture, but with a layer of soft spines
on the underside of the cap instead of ridges. Mainly used by restaurants
during the spring when chanterelles are scarce.
Black trumpets or black chanterelles (Craterellus cornucopioides) tend to
be difficult to see but look like clusters of black petunias erupting from the
ground. The main harvest area is in northern California, but market share
is being lost to North Africa and Eastern Europe, which are much closer to
the main place of consumption, which is France. This flavourful mushroom has yet to catch on in America, perhaps because of its dark colour.
Slow, prolonged cooking is best; excellent fresh or dry.
Lobster mushrooms are becoming increasingly popular in restaurants.
This is actually two fungi in one: a fluorescent red-orange fungal parasite
(Hypomyces lactifluorum) that completely engulfs a brittle white species of
Russula. Available fresh in the summer and autumn from the Pacific
Northwest and Mexico.
Yellow foot (Cantharellus tubaeformis and relatives) is a mild-flavoured
chanterelle favoured by American chefs for its clean, slim, petite appearance. It is picked on the west coast in winter and spring.
Cauliflower mushroom (Sparassis crispa) is a large, fragrant mushroom
looking more like a brain than a cauliflower. It grows at the bases of
conifers and is best in soups and casseroles.
Candy caps (Lactarius fragilis) has become so popular in restaurants
that demand exceeds supply. Chefs are enthralled with its powerful aroma.
A single cooked or dried candy cap will envelop the entire restaurant in an
aroma like maple syrup.
Wild mushrooms and rural economies
109
Hen of the woods (Grifola frondosa) forms large clusters of small,
overlapping caps which look like a ruffled-up hen. Grows at the bases of
oaks and other hardwoods in eastern North America, especially New
England, but a smaller, cultivated form is also available under the Japanese name, maitake. Both are excellent.
Giant puffballs (Calvatia gigantea and relatives) are enormous ball-like
mushrooms of prairies and open ground. Sometimes sold by the slice, but
the flesh must be pure white to be edible.
A fully ripe Oregon white truffle (Tuber gibbosum) is just as aromatic as
its famous European counterpart but is a bargain at less than one-tenth the
price. The problem is that these underground fungi are hunted with rakes
instead of being sniffed out by dogs or pigs as is the traditional method in
Europe. The result is that many are picked before their full aroma and
flavour develops.
Oregon black truffles (Leucangium carthusiana) are a different fungus
altogether from the European black truffle, softer and fruitier with a
shorter shelf life. Both Oregon truffles are associated with young Douglasfir. Most of the harvest takes place on tree farms in the winter and spring.
Cultivated mushrooms such as the shiitake (shiang-gu), portabello,
enoki and oyster mushroom are often advertised in restaurants as ‘wild
mushrooms’, which they are not. A better label for them is ‘exotic’ or
‘gourmet’ mushrooms, catch-all phrases for virtually any mushroom besides the common supermarket variety.
Sustainability of harvests
Some people have wondered whether the mushroom harvest is sustainable
at current levels. While this is a complex subject beyond the scope of this
chapter, studies show that as long as the ground isn’t dug up deeply,
intensive picking has little influence on future crops and may even have a
slight stimulatory effect. This is not terribly surprising because the commercially valuable species tend to be more plentiful in second-growth
forests or those influenced to some degree by human beings (which is
probably why we came to value them in the first place). In Italy, where
competition for porcini has been fierce for hundreds of years, they had a
bumper crop in 1998.
The good news, then, is that wild mushrooms, like pine nuts or huckleberries, can be harvested without significantly damaging our forests, and
that buying them in a gourmet restaurant probably comes at less cost to
biodiversity than wine, beef, or almost any other item on the menu. The
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bad news is that overly restrictive policy and unrealistic permit fees imposed by the US Forest Service and other agencies are turning an already
marginal existence into a well nigh impossible one. They make mushroom
picking profitable only as an occasional opportunistic activity that does
not encourage lifestyle conservation or land stewardship. Such policies
portend a future in which packets of dried mushrooms labelled ‘Buy Wild
Mushrooms – Help Conserve Rain Forests’ refer only to forests that are
not our own.
Acknowledgement
This chapter is excerpted from a longer piece on the wild mushroom
harvest in America published by the author in California Wild in the
autumn of 1999 and I thank Maurice Rotheroe for his help with this
adaptation.
Reference
Gould, S. J. (1997). Dinosaur in a Haystack. Penguin Books: Harmondsworth,
Middlesex.
9
Threats to biodiversity caused by
traditional mushroom cultivation
technology in China
SI U W AI CHIU & D AVI D MOORE
Introduction
China is the world’s major mushroom producing country. Agaricus production is mainly for export but Lentinula edodes (shiitake or shiang-gu) is
the traditional local product, and now the major crop. Lentinula edodes is
indigenous to China. It was first cultivated there more than 800 years ago,
and today, China accounts for about 70% of world production. In 1997,
Chinese production was recorded as 91 500 metric tonnes of the dried crop
(drying produces the characteristic taste of the mushroom), ten times that
in fresh weight. Shiang-gu (the Chinese name) is presently about the
second or third most popular cultivated mushroom in the world, being
consumed throughout China, Taiwan, Japan and Korea, and with increasing world-wide popularity. One-third of the Chinese crop is exported. As
this amounts to the equivalent of about 300 000 tonnes of fresh mushrooms, the industry is an important earner of foreign exchange as well as
making a very significant contribution to the income of peasant-farmers
especially of the mountainous regions in China (Chang & Chiu, 1992). In
these regions the land is poor in fertility and too distant from reliable
transport to make conventional farming of green crops profitable.
Traditional technology
The traditional log-pile cultivation method is still the one that is most
frequently used. For this, locally felled logs (oak, chestnut, hornbeam,
maple and other trees) over 10 cm diameter (probably about 20 to 30 years
old) and 1.5 m to 2 m long are normally cut in spring or autumn of each
year. Felling at this time minimises pre-infestation by wild fungi or insects.
Holes drilled in the logs (or saw- or axe-cuts) are packed with spawn, and
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the spawn-filled holes are then sealed with wax or other sealant to protect
the spawn from the weather. The logs are stacked in laying yards on the
open hillside, or under shelters built of bamboo or other materials, in
arrangements which permit good air circulation and easy drainage and
temperatures between 24 °C and 28 °C. The logs remain here for five to
eight months for the fungus to grow completely through the log before
transfer to the raising yard to promote fruit body formation. This is
usually done in winter to ensure the lower temperature (12–20 °C) and
increased moisture that are required for fruit body initiation. The first
crops of mushrooms appear in the first spring after being moved to the
raising yard. Each log will produce 0.5 kg to 3 kg of mushrooms, each
spring and autumn, for 5 to 7 years. We now have enough information for
a ‘back of an envelope calculation’: if each log produces 5 kg mushrooms
per year; total annual production today is probably in excess of one million
metric tonnes so we need the equivalent of 200 000 2-m logs. This amounts
to between 50 000 and 100 000 trees every year just to maintain current
production levels. Traditional usage of natural wood logs has been pursued to the extent that as availability of mature trees has declined attention
has turned to younger trees and other tree species. This, combined with
other demands for timber and land, has contributed to a loss of 87% of the
native forests in China (Anon., 1997). China now faces the problem that
the rate of deforestation is much greater than the rate of reforestation in
the remaining 13% forest cover (Mackinnon et al., 1996; Loh et al., 1999)!
There are regulations about planting and prohibitions on felling young
trees, but these are difficult to monitor and ‘conservation awareness’ is
especially low among those poor peasants living in the remote mountainous regions.
Conservation issues
The conservation pressures of Lentinula production result from the scale of
the industry and from the biological consequences of traditional practices.
There has been a minimum four-fold increase in production of Lentinula
over the past twenty years or so, and every prospect that demand for the
crop will continue to increase. The conservation pressure the industry
exerts is more likely to increase than to ease. The scale is difficult for
Europeans to imagine. The crop is still grown mostly by open outdoor
cultivation on a very large number of small farms. One well-informed
commentator (Luo, 1998) has claimed that there are 10 million mushroom
farmers in China! These mushroom farms are distributed over the whole of
Threats to biodiversity in China
113
the central highlands in China. This means that the Lentinula growing
region covers an area about equal to the entire land area of the European
Union.
The main conservation issues raised by the traditional cultivation practices are:
∑ the obvious impact on wooded hillsides caused by felling of
mature trees (of which only the largest logs are used);
∑ the fact that outside cultivation is used and the mushroom crop is
harvested after basidiospore release has started with the
consequent (unknown) danger(s) resulting from cross
contamination between cultivated and natural populations of
Lentinula edodes.
Population biology
Over the past several years we have investigated the cultivation physiology
of Lentinula edodes with a view to providing a good scientific basis for
using alternative cultivation substrates (Tan & Moore, 1992, 1995), but we
have put greater efforts into studying the population biology of Lentinula
edodes by examining both the cultivated (Chiu et al., 1996) and natural
populations (Chiu et al., 1998a,b, 1999a,b). The research covers a geographical area which is around 1700 km north to south and 700 km west to
east (Fig. 9.1), but includes detailed surveys down to individual logs, and
deals with phenotypes varying from morphology and palatability to DNA
sequences.
Nineteen strains of Lentinula edodes which are used for spawn production for farms throughout mainland China were characterised with three
arbitrarily-primed polymerase chain reaction (AP-PCR) profiles, seven
random amplified polymorphic DNA markers (RAPD) profiles and five
restriction patterns (restriction fragment length polymorphism patterns of
the PCR-amplified ribosomal DNAs; rDNA-RFLPs). For AP-PCR, 4–14
DNA bands were amplified for a particular strain while 1–9 DNA bands
were amplified using RAPD. Among them only three of the strains tested
showed different amplification profiles with most of the primers used. The
others showed small differences in three or fewer DNA amplification
profiles. All strains showed identical rDNA-RFLPs.
This study indicates that cultivated strains of shiang-gu in China are
genetically very homogeneous. In part this probably results from the
concentrated artificial breeding work that has been done using a limited
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range of imported Japanese cultivars, originating in a series of breeding
experiments with wild isolates collected from China, Taiwan and New
Guinea (Mori, Fukai & Zennyoji, 1974). However, genetic homogeneity is
also evident in cultivated strains of Agaricus bisporus (Loftus, Moore &
Elliott, 1988) and Volvariella volvacea (Chiu, Chen & Chang, 1995). Loftus
et al. (1988) could find no RFLP polymorphisms between three commercial cultivars of Agaricus bisporus. They noted that the growth conditions
and flushing (times of fruiting) patterns for each of the three strains were
also very similar, despite the fact that the three strains were marketed by
different companies as original and independent products. These authors
commented on how remarkable was ‘ . . . the coincidence of their genetic
similarity . . . ’. Since the same remarkable genetic similarity between
allegedly different commercial cultivars has been encountered in Volvariella volvacea (Chiu et al., 1995) and now in Lentinula edodes, it is
entirely feasible that genetic homogeneity in cultivated mushrooms does
not result from any peculiarity of mushroom genetics but most probably
from behaviour patterns of mushroom growers around the world.
Although a very narrow gene pool is used in cultivated strains, our
survey of diversity of rDNA sequences indicates that China harbours the
greatest germplasm resource of the mushroom Lentinula edodes (Chiu et
al., 2000). The internal transcribed spacer (ITS) regions of rDNA in
Lentinula edodes are rather conserved and therefore can be used to trace
lineage relationships. Hibbett et al. (1995) and Hibbett, Hansen & Donoghue (1998) have identified five lineages in Asia–Australasia with two
lineages appearing in China. As the sample size of all previous studies is
rather small, we carried out a large-scale and more detailed screening in
China, focusing attention on three provinces that are not among those in
which traditional cultivation is popular (to avoid the danger of contamination with the ‘commercial gene pool’). These are central Hubei, north
central Shaanxi and the most Southwest Yunnan (Fig. 9.1). Several fruit
bodies from single fallen logs and fruit bodies from different logs were
collected and used for culture isolation. Biomass grown in pure culture was
used for small scale DNA ‘mini-preparations’ (Chiu et al., 1996), followed
Fig. 9.1. (opposite) The experimental area. The three provinces from
which field isolates were collected (Shaanxi, Hubei and Yunnan, the latter
sharing borders with Myanmar, Laos and Vietnam) are shown shaded in
the upper map. In the larger scale map in the lower part of the figure the
location of traditional shiang-gu cultivation provinces, Guangdong,
Fujian and Jiangxi (and provincial capital cities), is indicated. Commercial cultivars were collected from institutes/universities in Beijing, Hubei,
Shanghai, Guangdong, Fujian and Hong Kong.
Threats to biodiversity in China
Modified from Magellan Geographix Map in CorelDRAW 8
115
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Sui Wai Chiu & David Moore
by specific polymerase chain reaction using a fungal-specific primer set
(ITS4 and ITS5). Direct sequencing was performed with the purified
PCR-amplified rDNA fragments.
The results reveal that a dominant group I rDNA lineage appears in
these three provinces. Compared with the rest, the Group I lineage has the
widest geographical distribution, extending from North Korea and Japan
to southern China and Thailand. This could indicate an ancient history for
this lineage but as many Japanese and Chinese cultivars belong to the
Group I lineage it is more probably the effect of breeding work and
subsequent adoption of those cultivars for commercial cultivation in these
regions. Similarly, wild isolates found to show group I rDNA sequences
that were collected in the traditional shiang-gu growing areas of Fujian
and Guangdong provinces may indicate cross contamination of the wild
population with cultivar genes or even escape of the cultivar to the wild.
The collection made by the authors in remote areas such as reserve areas of
national status or mountainous areas far beyond human access could
answer this problem. In the three provinces Hubei, Shaanxi and Yunnan,
group I rDNA lineage sequences were found in the wild isolates. Thus,
group I rDNA lineage occupies the largest territory and China might be
the home for this lineage. Isolates of the group IV rDNA lineage previously reported in Nepal and Hubei were found in Shaanxi province instead.
However, novel rDNA sequences (not belonging to either of the five
lineages previously described) were found in the three provinces. Thus,
several additional new rDNA lineages are present in China. Besides, as
isolates of different rDNA lineages appeared in one single log, it is evident
that genetic recombination is possible among the different lineage populations existing in the remote mountainous areas of China.
To investigate small-scale population structure, a field study was carried
out in a remote broad-leaved Fagus longipetiolata forest in Shaanxi province. Following spatial mapping, 24 fruit bodies were collected for tissue
isolation into axenic culture. Twenty-four genets distributed on fallen tree
trunks within a distance of 120 m were identified and clustered into 7
groups using the unweighted pair–group method algorithm using data
based on colony morphologies, abilities to degrade aromatic poly-R478
dye, somatic incompatibility reaction patterns and DNA fingerprints.
Among the parameters used, the somatic incompatibility reaction, a polygenic phenotype, was the most differentiating, identifying 22 incompatible
classes. Two sets of fruit bodies of different genets were so close together
that they would otherwise have been described as aggregate fruits of
presumed identical origin. Eighteen genets found on the same 5.6 m long
Threats to biodiversity in China
117
tree trunk divided roughly into two clusters, matching their spatial distribution, and a nearby branch bore another distinct cluster. More heterogeneity was encountered between isolates the greater the distance separating them on the original site. Genets on the same tree trunk showed more
compatible somatic reactions among themselves, and their DNA fingerprints showed higher similarity. Nevertheless, considering the totality of
phenotypic characters, each fruit body is a genet in L. edodes.
Such features are concluded to result from a reproductive strategy that
depends on basidiospore dispersal. Within each cluster of isolates from the
collection site genets seemed to have arisen from multiple sib-mating
events. Thus, a cluster may represent a lineage of L. edodes. Individualism
in L. edodes is based on a strong somatic incompatibility system. Strong
competition from contaminating individuals arriving as air-borne
basidiospores could explain decreased and fluctuating crop yields which
are now frequently observed in later flushes from the outdoor wood log
cultivation system. Further, it would also explain why multispore spawn is
not favoured in artificial cultivation of this economically important edible
mushroom. This also applies to Agaricus and Pleurotus mushrooms which,
together with Lentinula form the triumvirate of the most popular cultivated species in the world, with China being the main producer and
exporter.
Improved technology
The traditional approach to shiang-gu production is demanding and exploitative in its use of both land and trees. For these reasons more
industrial approaches are being applied to shiang-gu growing. Hardwood
chips, sawdust and other solid agricultural wastes packed into polythene
bags as ‘artificial logs’ provide a highly productive alternative to the
traditional technique, and the cultivation can be done in houses (which
may only be plastic-covered enclosures) in which climate control allows
year-round production. The industry raises other conservation issues that
our recent analyses of molecular and conventional genetic markers have
addressed. A very limited gene pool is exploited in the cultivated strains in
China, yet there is an enormous biodiversity in the species in the wild.
Analysis of local populations reveals that L. edodes strains show strong
somatic incompatibility reactions and individual territories can be small (a
few hundred mm). The widespread nature of the species and absence of
other means of dispersal indicate that basidiospores are the major, even
only, method of natural distribution. As the crop is harvested after
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initiation of basidiospore release and the crop is mostly grown outdoors,
cross contamination between wild and cultivated strains is inevitable. This
puts the natural gene pool under threat. Protection of the natural environment is still the best strategy for conserving the biodiversity of this important commercial resource. Collection of wild strains for preservation in a
culture collection would directly conserve the wild germplasm. Making
such a gene bank readily accessible to the public and industry would also
generate a commercial resource for exploitation in both cultivation and
breeding programmes. It may also reduce the pressure caused by nonprofessional collection from nature.
Our studies suggest that a move to indoor cultivation, less dependence
on multispore spawns and exploitation of a wider range of natural genotypes would better safeguard both cultivated and natural populations of
the fungus and avoid denuding hillsides of mature trees. The concept of
sustainable management is too novel for the mushroom farmers of a third
world country. But villages in China are accustomed to working cooperatively to establish a shared facility, and the advantages of indoor
cultivation to the farmer (consistency of yield, much shorter production
cycle, use of solid industrial/agricultural wastes in the substrate) can be
readily appreciated. An essential step is to educate the public to introduce
these ideas. Education is the key.
References
Anon. (1997). The Chinese Forests. Forestry Department: Beijing, People’s
Republic of China.
Chang, S. T. & Chiu, S. W. (1992). Mushroom production – an economic
measure in maintenance of food security. In Microbial Technology:
Economical and Social Aspects (ed. E. J. DaSilva, C. Ratledge & A. Sasson),
pp. 110–141. Cambridge University Press: Cambridge, UK.
Chiu, S. W., Chen, M. J. & Chang, S. T. (1995). Differentiating homothallic
Volvariella mushrooms by RFLPs and AP-PCR. Mycological Research 99,
333–336.
Chiu, S. W., Chiu, W. T., Lin, F. C. & Moore, D. (2000). Diversity of rDNA
sequences indicates that China harbours the greatest germplasm resource of
the cultivated mushroom Lentinula edodes. In Science and Cultivation of
Edible Fungi, vol. 1 (ed. L. J. L. D. Van Griensven), pp. 239–243. A. A.
Balkema: Brook Field.
Chiu, S. W., Ma, A. M., Lin, F. C. & Moore, D. (1996). Genetic homogeneity of
cultivated strains of shiitake (Lentinula edodes) used in China as revealed by
the polymerase chain reaction. Mycological Research 100, 1393–1399.
Chiu, S. W., Wang, Z. M., Chiu, W. T., Lin, F. C. & Moore, D. (1999a). An
integrated study of individualism in Lentinula edodes in nature and its
implication for cultivation strategy. Mycological Research 103, 651–660.
Threats to biodiversity in China
119
Chiu, S. W., Yip, M. L., Leung, T. M., Wang, Z. W., Lin, F. C. & Moore, D.
(1998a). A preliminary survey of genetic diversity in a natural population of
shiitake (Lentinula edodes) in China. Proceedings, Fourth Conference on the
Genetics and Cell Biology of Basidiomycetes, Nijmegen, The Netherlands,
27–30 March, 1998, p. 175. The Mushroom Experimental Station: Horst,
The Netherlands.
Chiu, S. W., Yip, M. L., Leung, T. M., Wang, Z. W., Lin, F. C. & Moore, D.
(1998b). Genetic diversity in a natural population of shiitake (Lentinula
edodes) in China. Abstracts of the Sixth International Mycological Congress,
Jerusalem, Israel, 23–28 August 1998, p. 34.
Chiu, S. W., Yip, P. & Moore, D. (1999b). Genetic diversity of Lentinula edodes
collected from Hubei Province, China. Poster in Abstracts, Seventh
International Fungal Biology Conference (ed. J. H. Sietsma), University of
Groningen, The Netherlands, 22–25 August, 1999.
Hibbett, D. S., Fukumasa-Nakai, Y., Tsuneda, A. & Donoghue, M. J. (1995).
Phylogenetic diversity in shiitake inferred from nuclear ribosomal DNA
sequence. Mycologia 87, 618–638.
Hibbett, D. S., Hansen, K. & Donoghue, M. J. (1998). Phylogeny and
biogeography of Lentinula inferred from an expanded rDNA dataset.
Mycological Research 102, 1041–1049.
Loftus, M. G., Moore, D. & Elliott, T. J. (1988). DNA polymorphism in
commercial and wild strains of the cultivated mushroom, Agaricus bisporus.
Theoretical and Applied Genetics 76, 712–718.
Loh, J., Randers, J., MacGillivray, A., Kapos, V., Jenkins, M., Groombridge, B.,
Cox, N. & Warren, B. (1999). The Living Planet. WWF International:
Gland.
Luo, X.-C. (1998). Mushroom Genetic Resource, Evaluation and Utilization in
China. In Proceedings of the ’98 Nanjing International Symposium, Science
and Cultivation of Mushrooms (ed. M. Lu, K. Gao, H.-F. Si & M.-J. Chen),
pp. 142–152. JSTC-ISMS: Nanjing, China.
Mackinnon, J., Sha, M., Cheung, C., Carey, G., Zhu, X. & Melville, D. (1996). A
Biodiversity Review of China. WWF International: Hong Kong.
Mori, K., Fukai, S. & Zennyoji, A. (1974). Hybridization of shiitake (Lentinus
edodes) between cultivated strains of Japan and wild strains grown in
Taiwan and New Guinea. Mushroom Science 9, 391–403.
Tan, Y.-H. & Moore, D. (1992). Convenient and effective methods for in vitro
cultivation of mycelium and fruiting bodies of Lentinus edodes. Mycological
Research 96, 1077–1084.
Tan, Y.-H. & Moore, D. (1995). Glucose catabolic pathways in Lentinula edodes
determined with radiorespirometry and enzymic analysis. Mycological
Research 99, 859–866.
10
A preliminary survey of waxcap
grassland indicator species in
South Wales
MAURICE ROTHEROE
Introduction
Over the past several decades conservationists have become increasingly
concerned about the disappearance of traditionally managed meadows
and pastures. These grasslands and their characteristic communities of
native plants have been lost through treatment with artificial fertilisers and
selective herbicides, through ploughing and reseeding and by encroaching
development. It was against this background in 1987 that the Countryside
Council for Wales (CCW) began a systematic survey of semi-natural
grasslands in Wales (currently nearing completion), using the National
Vegetation Classification (NVC) (Rodwell, 1991, et seq.) to define plant
communities. As well as providing basic information on the distribution
and extent of unimproved grassland, a further aim of this CCW survey was
to identify the best examples of this habitat in Wales and to afford them
official recognition and protection as Sites of Special Scientific Interest
(SSSIs). These are areas of land or water which are designated as being of
nature conservation importance for their biological, geological or physiographic features. Designation affords the site statutory protection and
aims to secure its sympathetic management. Currently, there are more
than 1000 SSSIs in Wales.
In parallel with vascular plants, a series of species of macrofungi is
largely confined to, and therefore indicative of, traditionally managed
grasslands. Such habitats are known to mycologists as Hygrocybe grasslands (Feehan & McHugh, 1992) or waxcap grasslands (Rotheroe et al.,
1996). The genus Hygrocybe (waxcaps) is not the only fungal indicator of
this grassland community. Waxcaps are often accompanied by fairy clubs
(clavarioid fungi); pink-spored species of the genus Entoloma (pink-gills);
and earth-tongues (family Geoglossaceae). Many of the taxa in these
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Waxcap grassland indicator species in South Wales
121
groups feature in the British and European Red Data lists of threatened or
endangered fungi (Ing, 1992, 1993). Several are also listed in many of the
national Red Data lists in Europe. For example, Arnolds & de Vries (1993)
have stated that all European species of Clavaria (fairy clubs) are included
on a Red Data list somewhere in Europe, while 67% of another grassland
fairy club genus, Clavulinopsis, feature on at least one European Red Data
list. Figures for other genera they gave are: Hygrocybe, 89% and Entoloma
sensu lato, 97%.
Criteria for the selection and notification of biological SSSIs largely
ignore the fungal element of the community, principally because of the
absence of data on species distribution and abundance (Hodgetts, 1992).
This has led to concern that mycologically rich sites may be underrepresented within the SSSI series, a situation that may have been exacerbated by the fact that certain waxcap grasslands can be floristically impoverished (Keizer, 1993). Responding to this data deficiency, the British
Mycological Society (BMS) initiated a survey throughout the British Isles
of the fungi characterising unimproved meadow and pasture grassland
(Rotheroe et al., 1996). This began in 1996 and is an ongoing survey.
The main aim of the South Wales project was to record fruiting of
waxcap grassland indicator species at a number of grassland sites in
Carmarthenshire, a county somewhat under-recorded mycologically.
Most of the survey sites were selected on the basis of their rich vascular
plant flora. The additional sites, both inside and outside the county, were
included for their known mycological interest and for comparative purposes. The survey also aimed to examine any relationship between mycological interest and NVC type or management regime. The survey was
commissioned by CCW and this chapter represents a summary of the
report to that agency (Rotheroe, 1999).
Site selection
Ten target sites were identified in conjunction with CCW officers. The
selected locations embraced a number of different NVC types, including
examples of Mesotrophic Grassland (symbolised MG); Mire (M); Calcifugous Grassland (U); and Calcicolous Grassland (CG). These sites are
listed in Table 10.1. Visits were also made to seven additional sites (listed in
Table 10.2), but NVC surveys have not been carried out at these additional
locations.
Past records from sites 1, 9, 11, 13 and 17 enabled cumulative lists to be
compiled. Cumulative data were also obtained from recording visits by
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Maurice Rotheroe
Table 10.1. Details of the ten Carmarthenshire sites selected for survey
Site
number Location
Code
Grid
reference
NVC types
represented (data
supplied by CCW)
1
‘Talley’ SSSI?
TAL
SN6–3–
2
Pwll Edrychiad SSSI
PED
SN584162
3
Caeau Nant Garenig SSSI CNG
SN673124
4
Cae Blaen-dyffryn SSSIB
CBD
SN604445
5
CBB
SN558439
6
Caeau Blaen-bydernyn
SSSI
Carreg Cennen SSSI
CCN
7
8
Whitehill Down SSSI
Rhosydd Castell-du SSSI
WHD
RCD
Waun-las (Middleton)
Caeau Caradog
WLS
CCG
SN670191 CG1e, CG2c,
MG6b
SN290135 MG5a, M24b/cA
SN655116 MG5c, M24b/c,
M25b
SN528178 MG6b
SN694460 MG5a, MG5c,
U4a
9
10
MG5a, MG5c,
M25b
MG5a, MG5c,
M25b@
MG5, M25b,
M24b/cA
MG5aA, MG6b
(in annex)
U4c, M25b
?The owner of this site requested anonymity. @Also includes some secondary
calcareous grassland around rocky outcrops. AAlso includes small stands of
MG5c. BAlso includes Annex to SSSI.
Key to grassland communities and subcommunities (for fuller details see
Rodwell, 1991 et seq.): Calcicolous Grassland, CG1 & CG2 (Festuca ovina–
Carlina vulgaris grassland and Festuca ovina–Avenula pratensis grassland). Both
are listed above as CG. Mires, M23a and M25b (Juncus effusus/acutiflorus–
Galium palustre rush–pasture: Juncus acutiflorus subcommunity and Molinia
caerulea–Potentilla erecta mire: Anthoxanthum odoratum subcommunity).
Mesotrophic Grassland, MG5a and MC5c (Cynosurus cristatus–Centaurea nigra
grassland: Lathyrus pratensis subcommunity and Danthonia decumbens
subcommunity). Mesotrophic Grassland, MG6 (Lolium perenne–Cynosurus
cristatus grassland). Calcifugous Grassland, U4a; U4b & U4c (Festuca ovina–
Agrostis capillaris–Galium saxatile grassland typical subcommunity, Holcus
lanatus–Trifolium repens subcommunity and Lathyrus montanus–Stachys betonica
subcommunity).
other workers to Gilfach Farm, Rhyader, Radnorshire (Ray Woods, pers.
comm.). This site is abbreviated by the code letters GILF.
Management regimes
The management regimes that were in place are listed in Table 10.3 as far
as they are known. This information has been obtained from CCW, the site
owners and from personal observations.
123
Waxcap grassland indicator species in South Wales
Table 10.2. Additional sites surveyed
Site
number Location
11
12
13
14
15
16
17
Llanerchaeron Estate (NT), Cardiganshire
Llanrhystud Chapelyard, Cardiganshire
Hafod Estate, Cardiganshire
Dinefwr Park (NT), Carmarthenshire
Maestir Churchyard, Lampeter,
Cardiganshire
St David’s College lawns, Lampeter,
Cardiganshire
Garn Ddyrys, Tumble, Breconshire
Code
Grid
reference
LLAN
LRST
HAFD
DINF
SN480600
SN547692
SN756731
SN617225
MAES
SN553494
STDC
GARN
SN579482
SO258117
Field procedures
Each of the 17 sites was visited at least twice and some sites were visited
more often. Each recording visit lasted between two and four hours,
depending upon the size of the site. Recording effort was concentrated in
relevant stands of vegetation which were identified from CCW’s Phase II
vegetation maps. Once a homogeneous stand representing one of the
target NVC types was located, a ‘mowing’ transect was carried out, by
walking back and forth across the area as if mowing a lawn. However, in
fields and areas which from the outset could be regarded as lacking any
fungal fruiting (usually the mire sites), a less rigorous system of transects
was used. Relevant NVC type and management regimes, from Phase II
reports and from field observations, were also noted. Fruit bodies of
common or easily identifiable fungi were left in situ but for many identifications it was necessary to collect representative samples for closer macroscopic and microscopic examination in the laboratory. Dried voucher
material was retained for all but the commonest species and this material
was deposited in the herbarium of the National Botanic Garden of Wales
(NBGW) or in Herb K(M) (Royal Botanic Gardens, Kew). The data
obtained from all the fieldwork were entered into the BMS database.
Evaluation of conservation value
Conservation of this unimproved grassland habitat has been the subject of
intensive study in recent years. The results of these studies provided a
well-documented context in which to evaluate the findings of this survey.
The data generated in the South Wales survey were analysed using the
124
Maurice Rotheroe
Table 10.3. Management history at survey sites
Site
number
Location and vegetation
Management history
1
‘Talley’ SSSI (MG5a, MG5c,
M25b)
2
Pwll Edrychiad SSSI (MG5a,
MG5c, M25b)
3
Caeau Nant Garenig SSSI (MG5,
M25b, M24b/c)
4
Cae Blaen-dyffryn SSSI and Annex
(MG5a)
5
Caeau Blaen-bydernyn SSSI (U4c,
M25b)
Carreg Cennen SSSI (CG1e, CG2c,
MG6b)
Pasture plus two hay meadows,
cattle-grazed. Farmed by organic
methods.
Pasture, cattle and sheep-grazed, with
clear evidence of rabbit grazing. No
evidence of fertiliser treatment.
Pasture and some Molinia-dominated
areas cut for hay. Sheep and pony
grazing.
Pasture, sheep-grazed. Unfertilised
(Annex outside SSSI had some recent
treatment).
Pasture, cattle and sheep-grazed.
Unfertilised.
Pasture, cattle and sheep-grazed.
Neutral grassland ‘heavily modified’
according to SSSI notification
documentation.
Hay cutting with aftermath grazing by
cattle. Artificial fertiliser used in certain
parts. Some areas show past
disturbance ‘probably ploughing’.
Pasture, cattle-grazed. No evidence of
fertiliser treatment.
Pasture, cattle and sheep-grazed, but
sheep withdrawn from large areas for
12 months prior to survey. Previous
owners claim no artificial fertiliser or
ploughing in living memory.
Pasture, horse, pony and sheep-grazed,
very heavily in some areas. MG5a has
suffered substantial modification.
Lawn, grazed by rabbits and
occasionally sheep escaping from
adjoining fields. Mown twice a year.
No fertiliser or treatment within living
memory.
Ungrazed, mown twice a year. No
fertiliser.
Pasture, relatively heavy sheep-grazing.
No fertiliser or treatment in living
memory.
Deer park, deer and cattle-grazed.
Owners claim no fertiliser treatment,
but some past treatment is possible.
Ungrazed, cut ‘regularly’. Believed no
fertiliser.
Ungrazed, cut ‘regularly’. Believed no
fertiliser.
Pasture, sheep and rabbit-grazed,
relatively heavily in many areas. No
information on fertiliser treatment, but
this is unlikely to have taken place.
6
7
Whitehill Down SSSI (MG5a,
M24b/c)
8
Rhosydd Castell-du a Plas y Bettws
SSSI (MG5c, M24b/c, M25b)
Waun-las (Middleton) (MG6b)
9
10
Caeau Caradog (MG5a, MG5c,
U4a)
11
Llanerchaeron Estate (NT) (MG5?)
12
Llanrhystud Chapelyard (?)
13
Hafod Estate (?)
14
Dinefwr Park (NT) (?)
15
Maestir Churchyard,
Lampeter, Cardiganshire (?)
St David’s College lawns,
Lampeter (?)
Garn Ddyrys, Tumble (?)
16
17
Waxcap grassland indicator species in South Wales
125
Table 10.4. Separation of conservation values into four classes according to
Rald (1985)
Conservation value
Number of
Hygrocybe species
recorded on a
single visit
Total number of
Hygrocybe species
expected to be
recorded
National importance (I)
Regional importance (II)
Local importance (III)
No importance (IV)
11–20
6–10
3–5
1–2
17–32
9–16
4–8
1–3
methodologies proposed by Rald (1985) and Rotheroe et al. (1996),
Rotheroe (1997, 1999). The Rald formula is based on the cumulative
number of Hygrocybe species recorded at a location, with numbers recorded on a single visit enabling tentative predictions of the full mycota to
be made. Rald separated the conservation values into four classes (Table
10.4).
Rotheroe (1999) proposed that an indicator-species profile of a site
would refine the evaluation of conservation value. This was a modified
version of various weighting systems elaborated by a number of Scandinavian workers (John Bjarne Jordal, pers. comm.). The profile can be expressed in a shorthand using the following code letters: C (clavarioid fungi
– fairy clubs); H (Hygrocybe sensu lato – waxcaps); E (grassland species of
the Entolomataceae – pink-gills); G (Geoglossaceae – earth-tongues).
Thus a site with survey records of eight waxcaps, two fairy clubs, two
earth-tongues and one species of Leptonia, would be described as: C2, H8,
E1, G2. This shorthand system has been used in the present study. In the
interests of simplicity, Dermoloma and Porpoloma are merged with H. This
is legitimate, since they have the same ecological affinities and Porpoloma
was, until recent taxonomic revisions, traditionally included in the genus
Hygrophorus (as was Hygrocybe), while the current classification places
Dermoloma in the tribus Hygrocybeae. The author therefore interpreted
Rald in this fashion and also referred to the ‘CHEG profile’ as a means of
making easy quantitative assessments for comparison of different sites and
to suggest their relative conservation value. In using these numerical
formulae a variety record is given equal weighting to that of a species. Thus
the numbers refer to taxa, rather than to species.
126
Maurice Rotheroe
Table 10.5. Highest number of Hygrocybe (waxcap) taxa recorded on
single visits made in 1998
Conservation value
Indicator species profile?
Site
number
Location C
H
E
G
17
13
6
11
9
1
16
2
15
4
14
5
12
7
10
3
8
GARN
HAFD
CCN
LLAN
WLS
TAL
STDC
PED
MAES
CBD
DINF
CBB
LRST
WHD
CCG
CNG
RCD
17
15
14
13
10
9
9
8
7
6
6
4
4
3
3
0
0
0
0
1
4
3
6
3
3
1
2
0
0
0
0
0
0
0
4
0
0
1
0
0
3
1
2
0
0
0
0
0
0
0
0
1
1
3
4
2
1
2
1
2
0
1
1
0
1
0
0
0
Assigned according to
Rald (1985)@
National importance
National importance
National importance
National importance
Regional importance
Regional importance
Regional importance
Regional importance
Regional importance
Regional importance
Regional importance
Local importance
Local importance
Local importance
Local importance
?Rotheroe (1999) and see text. @See Table 10.4.
Nomenclature
In general the classification of the Dictionary of the Fungi (Hawksworth et
al., 1995) has been used in this study. However, there is some disagreement
currently over aspects of nomenclature for certain groups. For the Entolomataceae (pink-gill species), this study uses the nomenclature of Noordeloos (1992), which reduces some genera to subgeneric level, notably
Leptonia and Nolanea, rather than the classification of the Dictionary of
the Fungi, which retains these at generic level. For Hygrocybe, the nomenclature follows Boertmann (1995), with a number of small modifications
proposed by Henrici (1996).
Analysis of results
Table 10.5 shows the total number of waxcap species recorded on a single
visit to survey sites, and Table 10.6 the cumulative totals of indicator
Waxcap grassland indicator species in South Wales
127
Table 10.6. Cumulative totals of indicator species recorded to the end of
December 1998
Conservation value
Indicator species profile?
Site
number
17
13
11
9
6
1
2
16
14
15
5
4
12
7
10
3
8
Location C
H
E
G
GARN
HAFD
LLAN
GILFA
WLS
CCNB
TAL
PEDB
STDCB
DINFB
MAESB
CBBB
CBDB
LRSTB
WHDB
CCGB
CNGB
RCDB
28
26
23
23
22
19
16
11
9
9
8
6
6
4
3
3
0
0
0
7
7
0
7
4
14
3
3
0
1
0
2
0
0
0
0
0
5
0
2
0
0
0
0
1
3
0
2
0
0
0
0
0
0
0
3
4
6
3
4
4
2
3
2
1
2
1
0
0
1
0
0
0
Assigned according
to Rald (1985)@
National importance
National importance
National importance
National importance
National importance
National importance
Regional importance
Regional importance
Regional importance
Regional importance
Local importance
Local importance
Local importance
Local importance
?Rotheroe (1999) and see text. @See Table 10.4. AGilfach Farm, Rhayader,
Radnorshire. BSites for which no previous records existed, so the cumulative
totals shown are for 1998 only.
species recorded up to the end of December, 1998. The conservation values
shown in these Tables were conceived in a European context and are
effectively being extrapolated to Wales. This seems to be justified, since
similar analyses in the UK have been applied during the BMS waxcap
grassland survey and the results do appear to reflect the conservation
importance of the respective sites examined (personal observations).
Correlations with NVC types
Table 10.7 summarises the number of waxcap grassland indicator species
recorded from the various NVC grassland types at the sites surveyed. The
full list of species is shown in Table 10.8. No NVC surveys had been carried
out at the additional sites listed, nor at Waun-las Farm, but it was
suggested that most of the grassland at this latter location is referable to
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Maurice Rotheroe
Table 10.7. Numbers of indicator species recorded at different grassland
types categorised using the National Vegetation Classification (NVC)
NVC grassland type
Number of species recorded
Calcicolous grassland CG
Calcifugous grassland U4a
Calcifugous grassland U4b
Calcifugous grassland U4c
Mesotrophic grassland MG5a
Mesotrophic grassland MG5c
Mesotrophic grassland MG6
Mire M23a
Mire M24
Mire M25b
10
2
4
7
19
18
34
2
0
1
MG6 (Jamie Bevan, pers. comm.). Records from this site are therefore
listed under that heading. NVC types for the other additional sites are
omitted.
The greatest numbers of species were clearly recorded from the neutral
grassland types MG5 and MG6. Fewer species were recorded from calcicolous and calcifugous grassland, although to some extent this may
simply reflect under-representation of these habitats within the survey
sites. The mire types are clearly very impoverished from a mycological
perspective.
Rare, threatened or endangered species
A list of those species recorded at the study sites which feature in the
British (Ing, 1992), European (Ing, 1993), and Welsh (Rotheroe, 1998) Red
Data lists is shown in Table 10.9. Also indicated are the three species
covered by the UK Biodiversity Action Plan (BAP) (Anon., 1999).
Meteorological considerations
The survey area suffered abnormally heavy precipitation in 1998. Total
annual rainfall, as reported from two local meteorological stations, was
28% higher than the norm for the past 30 years (John Wildig & John
Powell, pers. comm.). One of these stations is upland, at an elevation of
301 m; the second, 100 km to the south, is almost at sea level. Most of the
Carmarthenshire sites lie approximately halfway between the two. Further
details are given in Rotheroe (1999).
Waxcap grassland indicator species in South Wales
129
Table 10.8. Species of macrofungi recorded from the various grassland
types
Species
Grassland type (NVC)
Clavaria vermicularis
Clavulinopsis corniculata
Clavulinopsis fusiformis
Clavulinopsis helvola
Clavulinopsis luteoalba
Dermoloma cuneifolium
Entoloma anatinum
Entoloma conferendum
Entoloma corvinum
Entoloma cruentatum
Entoloma hebes
Entoloma infula
Entoloma porphyrophaeum
Entoloma serrulatum
Hygrocybe calyptriformis
Hygrocybe ceracea
Hygrocybe chlorophana
Hygrocybe cinereifolia
Hygrocybe citrinovirens
Hygrocybe coccinea
Hygrocybe colemanniana
Hygrocybe conica
Hygrocybe conica v. chloroides
Hygrocybe glutinipes
Hygrocybe insipida
Hygrocybe irrigata
Hygrocybe laeta
Hygrocybe nitrata
Hygrocybe persistens
Hygrocybe pratensis
Hygrocybe pratensis v. pallida
Hygrocybe psittacina
Hygrocybe psittacina v. perplexa
Hygrocybe punicea
Hygrocybe quieta
Hygrocybe reidii
Hygrocybe russocoriacea
Hygrocybe splendidissima
Hygrocybe virginea
Hygrocybe virginea v. fuscescens
Hygrocybe virginea v. ochraceopallida
Hygrocybe vitellina
Microglossum olivaceum
Ramariopsis kunzei
MG5a
CG, MG5a, MG6
MG6
CG, M25b, MG5c, MG6, U4c
MG5c
MG5a, MG6
MG5c
MG6, U4b
M23a
MG5c
MG5c
MG5c, U4b
MG6
MG6
MG5a, MG5c, MG6
MG5a, MG6, MG6
CG, M23a, MG5a, MG5c, MG6, U4c
MG6
MG5a, MG6
MG5a, MG6, U4c
CG, MG6
MG5a, MG5c, MG6
MG6
MG6
CG, MG5a, MG5c, MG6
MG5a, MG5c, MG6
MG5c, MG6, U4a, U4b, U4c
MG6
MG6
MG5a, MG5c, MG6, U4a, U4b, U4c
MG5a, MG5c
MG5a, MG5c, MG6, U4c
MG6
MG6
CG, MG5a, MG6
CG, MG5a, MG5c, MG6, U4c
CG, MG5a, MG5c, MG6
MG6
CG, MG5a, MG6
CG, MG6
MG5a, MG6
MG5c
CG
MG6
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Maurice Rotheroe
Table 10.9. Rare, threatened or endangered species recorded at grassland
sites in the present study
Species
Threat category?
Location(s) where recorded
Clavaria zollingeri
BRDL, WRDL,
BAP
ERDL
ERDL
BRDL, WRDL
WRDL
BRDL, WRDL,
priority BAP sp.
GILF, HAFD, LLAN,
MAES
LLAN, STDC
STDC
LLAN
TAL
CBD, GARN, GILF,
HAFD, LLAN, LRST,
MAES, TAL
GILF
GARN, HAFD, STDC,
WLS
CCN, GARN, HAFD,
LLAN, PED, TAL, WLS
DINF, GILF, WLS
GILF, HAFD, LLAN,
TAL, WLS
GARN, GILF, WLS
GILF, LLAN
CCN, HAFD
Geoglossum fallax
Geoglossum glutinosum
Entoloma bloxamii
Entoloma cruentatum
Hygrocybe calyptriformis
Hygrocybe cantharellus
Hygrocybe fornicata
ERDL
ERDL
Hygrocybe insipida
ERDL
Hygrocybe intermedia
Hygrocybe irrigata
ERDL
ERDL
Hygrocybe nitrata
Hygrocybe ovina
Hygrocybe psittacina v.
perplexa
Hygrocybe punicea
ERDL
ERDL
ERDL
Hygrocybe quieta
ERDL, WRDL
Microglossum olivaceum
BRDL, ERDL,
WRDL, priority
BAP sp.
BRDL, WRDL
WRDL
ERDL
WRDL
Porpoloma metapodium
Ramariopsis kunzei
Trichoglossum hirsutum
Trichoglossum walteri
ERDL
CCN, GARN, GILF,
HAFD, LLAN, MAES,
WLS
CBD, GARN, HAFD,
PED
GARN, LLAN, MAES,
PED
DINF, HAFD, LLAN
CCN, HAFD
GARN, MAES
STDC
?BRDL, British Red Data list (Ing, 1992); ERDL, European Red Data list (Ing,
1993); WRDL, Welsh Red Data list (Rotheroe, 1998); BAP, UK Biodiversity
Action Plan (Anon., 1999).
All the sites visited were showing the effects of continuous heavy rainfall.
This resulted in sodden ground and anaerobic soil conditions which persisted for long periods during what is regarded as the ‘normal autumn fungus
season’. Because of the waterlogged conditions, sites grazed by cattle or
horses suffered additional damage through ‘poaching’ of the soil, probably
disrupting underground mycelial networks. Previous work has shown that
Waxcap grassland indicator species in South Wales
131
mire communities hold only marginal mycological interest (Rotheroe,
1997) and sites supporting these communities were particularly adversely
affected by the excessive rain.
Comments on some individual sites
Many of the locations surveyed are already nature reserves or SSSIs, but
none has been notified as such because of any mycological consideration.
However, all the following sites should, in the author’s view, be listed as
Important Areas for Fungi. This category refers to a new initiative sponsored by the charity Plantlife to compile an inventory of mycologically
important sites in Britain. It is an ongoing project. Summarised below are
some of the best of the sites covered by the survey.
‘Talley’ SSSI
This site gains a Regional Importance ranking in Tables 10.5 and 10.6,
featuring in sixth and seventh position amongst the top ten sites. However,
there are reasons for regarding it as being of National Importance. It is
remarkable for the comparatively large number of Entoloma species recorded there, the cumulative total being 14, which is double that of any
other location in this survey. Among them is Entoloma cruentatum, which
was only the second British record when it was first collected there in 1996,
although is also now known from two other locations in Britain.
Pwll Edrychiad SSSI
A site that appears to be potentially of National Importance, although the
Rald formula rates it as being only of Regional Importance. It is the only
site in Carmarthenshire where the olivaceous earth-tongue (Microglossum
olivaceum) has been recorded. This rare fungus is known from only six
other localities in Wales. It has a westerly distribution in mainland Britain
and Wales appears to be one of its main population centres. It was fruiting
at Pwll Edrychiad in October 1998 in large numbers (over 200 fruit bodies
in two colonies) on small tracts of open, rocky calcicolous grassland. Two
visits probably revealed only a fraction of this SSSI’s full mycota.
Carreg Cennen SSSI
Carreg Cennen was included in the survey for its stands of calcicolous
grassland (a scarce resource in Carmarthenshire), although these form
132
Maurice Rotheroe
relatively small, narrow tracts. Paradoxically, however, most of the indicator species were recorded on the surrounding, extensive MG6b grassland areas. This was one of the best sites visited and easily qualifies as being
of National Importance, with a cumulative indicator score of C4 H19 E4
G0. It seems likely that this site has a far more extensive and rich mycota
than has been revealed from a single year’s data.
Waun-las
Having been surveyed in the previous two years, Waun-las was already
known to have a rich grassland mycota. On the first visit in October 1996, a
total of 16 different species of Hygrocybe were recorded. It was noted in
1998, however, that grazing had been withdrawn during the preceding 12
months. The grass was therefore longer and ranker than previously. Not
only were the number of waxcap species recorded in 1998 reduced, but the
number of fruit bodies in evidence showed a considerable drop compared
with previous years. Previous grazing by sheep in this area had maintained
a relatively short turf, with good bryophyte cover. It is understood that
shortly after the final recording visit to the site in 1998, sheep grazing was
resumed (Janet Moseley, pers. comm.). The current owners, the National
Botanic Garden of Wales, have already made approaches to CCW, suggesting that the mycological richness of these unimproved grasslands
warrants their being notified as an SSSI. They acknowledge, however, that
the grasslands are unremarkable floristically and appear to be impoverished MG6b Lolium–Cynosurus grassland. This, taken together with the
observations at Carreg Cennin above, indicate that a rich mycota can
persist in partially improved neutral grassland provided that grazing pressures are sufficient to maintain short-cropped swards.
Llanerchaeron Estate
This was an additional site, included because its small unimproved lawn
has previously shown it to be one of the best waxcap sites in Wales. It
deserves SSSI status, but certainly qualifies as an Important Area for Fungi.
Hafod Estate
This estate has a large area of traditionally managed grassland, with rather
intensive grazing by sheep. It is not highly regarded for its floristic qualities. However, the CHEG rating confirms its mycological importance.
Every year new species of the indicator group are recorded there.
Waxcap grassland indicator species in South Wales
133
General conclusions
In South Wales, 1998 was an atypical year so far as the autumn fungus
season was concerned. Fruiting of macrofungi in grasslands was delayed
and rainfall much higher than average resulted in waterlogged conditions
at many of the study sites. Nevertheless, some valuable county data were
obtained and comparisons could be made with sites known to be mycologically important for grassland indicator species. Four Carmarthenshire sites are judged to be of high conservation value. These are: Waun-las
Farm, Middleton; Carreg Cennen SSSI; ‘Talley’ SSSI; and Pwll Edrychiad
SSSI. The first two are regarded as being of putative National Importance
and the other two might also qualify for this ranking. Two of the remaining sites, Caeau Caradog and Whitehill Down SSSI, may well have a much
richer mycota than was observed in this survey.
Grassland types
The richest grasslands were clearly the mesotrophic types. Plentiful data
from MG5 Cynosurus–Centaurea grassland was perhaps expected from
past experience of this community. However, it may be surprising to higher
plant ecologists to discover that some of the most mycologically productive habitats were those of the MG6 Lolium–Cynosurus type.
It is clear from this survey that certain less modified forms of MG6
grassland can be rich in fungi despite appearing impoverished from a
vascular plant perspective. At both Carreg Cennen and Waun-las a clear
distinction can be made from mycological data between MG6 grassland
which has been heavily fertilised or improved in some way and the same
grassland type which has been less modified. This distinction is often a
dramatic one.
This is perhaps a consideration which the users of NVC for conservation
evaluation should address. The conservation importance of some MG6
sites may be considerably underestimated unless they are visited by a
mycologist during the late summer and autumn fruiting season.
Calcicolous grasslands are poorly represented in Carmarthenshire and
the survey results do not reflect the potential of this NVC type. Mires M23,
M24 and M25 were mycologically poor.
Management recommendations
At those sites which have been identified as being mycologically important
the main recommendation is to maintain the status quo, that is to refrain
134
Maurice Rotheroe
from fertiliser application or other chemical treatments and to continue at
existing stocking levels. The mycological interest of some of the remaining
sites might be enhanced if fertiliser treatments were withheld and stocking
levels increased slightly. If a choice is available, then grazing by sheep, even
at quite heavy stocking levels, as at Hafod, appears to be the most
successful regime to encourage fungal fruiting, although cattle grazing at
relatively low intensity (such as occurs at ‘Talley’ SSSI) can clearly maintain favourable conditions too. Personal observations suggest that rabbit
populations should be encouraged. This grazer produces close-cropped
areas, which are extremely productive for certain indicator fungal species.
Future work
Within Carmarthenshire further mycological recording is recommended,
particularly at Carreg Cennen SSSI, Pwll Edrychiad SSSI, Caeau Caradog
and Whitehill Down SSSI. This should ideally take place from spring,
through summer and into the autumn, over a period of several years if the
full mycotas of the sites are to be demonstrated.
A broader programme of survey throughout Wales would also be
advantageous so that a fuller assessment of the distribution of mycologically important grasslands of the Principality might be made. This would
be a valuable contribution to the process of inventory-taking in order to
evaluate fungal biodiversity. It would also make a significant contribution
to various obligations under the UK Biodiversity Action Group.
The methods of evaluation of conservation value used in this survey
have been shown to be workable and informative in this project. A more
wide-ranging analysis of the data from the BMS waxcap grassland survey,
using these and other similar methods, would produce a clearer, quantitative picture of the important grasslands for fungi in the British Isles, and
their distribution. This could be a valuable contribution towards an inventory and comparative study of waxcap grasslands in Europe.
Acknowledgements
This chapter is based on a report published by the Countryside Council for
Wales (Rotheroe, 1999) and the author is grateful for their permission to
publish this revised version and for their financing of the project. He is also
indebted to the National Trust and the Hafod Trust who provided financial support for similar projects. He is grateful for the valuable suggestions
on modifications to the text from Vincent Fleming and for the help, advice
Waxcap grassland indicator species in South Wales
135
and support from the following individuals: Jamie Bevan, David Boertmann, Penny David, Shelley Evans, Gareth Griffith, John Bjarne Jordal,
Janet Moseley, John Powell, John Savidge, Michael Smith, David Stevens,
Nigel Stringer, John Wildig and Ray Woods.
References
Anon. (1999). UK Biodiversity Group Tranche 2 Action Plans: Volume III – Plants
and Fungi. English Nature: Peterborough.
Arnolds, E. & de Vries, B. (1993). Conservation of fungi in Europe. In Fungi of
Europe: Investigation, Recording and Conservation (ed. D. N. Pegler, L.
Boddy, B. Ing & P. M. Kirk), pp. 211–230. Royal Botanic Gardens: Kew.
Boertmann, D. (1995). The Genus Hygrocybe. Fungi of Northern Europe 1.
Svampetryk: Greve, Denmark.
Feehan, J. & McHugh, R. (1992). The Curragh of Kildare as a Hygrocybe
grassland. Irish Naturalists Journal 24(1), 13–17.
Hawksworth, D. L., Kirk, P. M., Sutton, B. C. & Pegler, D. N. (1995). Ainsworth
& Bisby’s Dictionary of the Fungi. Eighth edn, 616 pp. CAB International:
Wallingford, Oxon.
Henrici, A. (1996). Waxcap-grassland Fungi. British Mycological Society:
Stourbridge, UK.
Hodgetts, N. (1992). Guidelines for Selection of Biological SSSIs: Non-vascular
Plants. Joint Nature Conservation Committee: Peterborough.
Ing, B. (1992). A provisional Red Data List of British Fungi. Mycologist 6,
124–128.
Ing, B. (1993). Towards a red list of endangered European macrofungi. In Fungi
in Europe: Investigations, Recording and Conservation (ed. D. N. Pegler, L.
Boddy, B. Ing & P. M. Kirk), pp. 231–237. Royal Botanic Gardens: Kew.
Keizer, P. J. (1993). The influence of nature management on the macromycete
flora. In Fungi of Europe: Investigation, Recording and Conservation (ed. D.
N. Pegler, L. Boddy, B. Ing & P. M. Kirk), pp. 251–269. Royal Botanic
Gardens: Kew.
Noordeloos, M. (1992). Entoloma s.l. Libreria editrice Giovanna Biella: Saronno.
Rald, E. (1985). Vokshatte som indikatorarter for mykologisk vaerdifulde
overdrevslokaliteter. Svampe 11, 1–9.
Rodwell, J. S. (1991 et seq.). British Plant Communities. Cambridge University
Press: Cambridge, UK.
Rotheroe, M. (1997). A Mycological Study of NVC Grassland Communities.
JNCC Contract Report No. F76–01–50. Joint Nature Conservation
Committee: Peterborough.
Rotheroe, M. (1998). The Rare Mushrooms of Wales. Unpublished report:
Countryside Council for Wales.
Rotheroe, M. (1999). Mycological Survey of Selected Semi-Natural Grasslands in
Carmarthenshire. Countryside Council for Wales: Bangor.
Rotheroe, M., Newton, A., Evans, S. & Feehan, J. (1996). Waxcap-grassland
survey. Mycologist 10, 23–25.
11
Grasslands in the coastal dunes: the
effect of nature management on the
mycota
M A R I J K E M . N A U T A & L EO M . J A L I N K
Introduction
Many regions of the dunes along the coast of The Netherlands are now in
use as part of the drinking water supply chain and are therefore well
protected. The Amsterdam Waterworks Dunes, situated south and west of
Haarlem, deliver drinking water to the city of Amsterdam. The site is
owned and managed by the municipality of Amsterdam. In this dune field
of 34 km the mycota of two, locally decalcified, natural grasslands has
been studied since 1986 (Becker & Baeyens, 1992; Nauta & Jalink, 1996).
The management consists of summer grazing by cattle in one site (Eiland
van Rolvers), and yearly mowing and removal of the hay in the other
(Groot Zwarteveld).
History
The area of the Amsterdam Waterworks Dunes consists mainly of young
dunes, formed during several periods of blowing sand from the eleventh
century onwards. The soils differ in calcium content, depending on the
period in which they were formed, and this can often be seen from the
vegetation. For instance, the Hippophae rhamnoides scrub, developed in
the central part of the Amsterdam Waterworks Dunes from around
1500–1600 onwards, has usually developed on more or less calcareous soil.
From early days humans have used the dunes. There is evidence that from
1500 onwards, wet dune slacks have been grazed in summer by cattle and
sheep.
In the early nineteenth century the dune area also became popular for
growing potatoes. Though this was done only for a short period, the fields
can still be recognised at several places in the dunes. These fields were
136
Grasslands in the coastal dunes
137
fertilised with organic debris and ploughed and are now the most decalcified sites.
In the course of centuries, a sweet-water reservoir has been formed
under the dunes, limited in the west by sea water. The area of the Amsterdam Waterworks has served as a water supply for the city of Amsterdam
from 1851. Before that time the dunes were so wet that it was possible to
skate from Zandvoort to The Hague in winter. After this time a gradual
drying has occurred, not only because of the withdrawal of drinking water,
but also because of the reclamation of the Haarlemmermeer polder and the
removal of sand on the eastern side of the dunes. During World War II
many woods were cleared in the dunes, resulting in enhanced drifting of
sand.
Because of the noticeable drying of the dunes, infiltration with river
water started in 1957, not because of any noble nature maintenance
argument, but to secure the water supply for Amsterdam. For this purpose
many canals were made and the unpurified water from the river Rhine was
led into the dunes in the west, to be purified by the dunes and withdrawn in
the east. The unpurified water caused not only a re-establishment of the
groundwater water table, but also an enormous increase of certain grasses
like Calamagrostis epigejos and nutriphilous plants like Cirsium. The
original wet dune slacks and grasslands in the dunes were soon overgrown
with high vegetation of Calamagrostis epigejos and Carex arenaria. Therefore, the Water Works Company took the decision to purify the infiltrated
river water and later began a management programme involving mowing
of several grasslands, in the hope this would restore the original vegetation.
Groot Zwarteveld and Eiland van Rolvers
Groot Zwarteveld (in English ‘large black field’) was in former times a wet,
grass-rich dune slack. It was in use by farmers from 1850 for cattle grazing
and potato growing. The field owes its name to the thin layers of black peat
at the surface, which are no longer visible. In a period with strong winds in
the past the valley was blown out and the sand drifted eastward. The soil is
locally decalcified.
The potato fields were deserted again around 1900, but the cattle grazing
continued until World War II, when the farmhouse was destroyed. Until
1960 the field was described as ‘dry and dusty’. Because of the infiltration
with river water, the groundwater water table reached its original level of
7 m above the usual reference point which is the elevation of Amsterdam
138
Marijke M. Nauta & Leo M. Jalink
(referred to as NAP) again in 1975, involving the previously mentioned
increase of Calamagrostis. The area, amounting to 12 ha, has been mown
two-yearly since 1975, and the wettest area is mown every year. A high
groundwater level also results from a reduced withdrawal of water, and a
rain-water lens (a lens-shaped body of rain water floating on the groundwater) is usually present all year round. Rain water is more acidic than
groundwater. The infiltrated river water also influences the western part.
In the wettest part some Sphagnum vegetation occurs.
Eiland van Rolvers (Rolvers’ Island) as it is called today, was formerly
the eastern part of Groot Zwarteveld. The Rolvers family, who grazed
their cattle in the area, inhabited it. It consisted of a wet dune slack and
drier dunes with Hippophae. From 1975 to 1984 part of the area was mown
every second year, and since 1985 it has been grazed in summer by seven
yearlings. The area covers 36 ha, and also has a rain-water lens.
Research
The vegetation in both areas has been examined since 1975 and this has
shown positive effects of mowing on the composition of the vegetation. As
an experiment, in a part of the area the mowing was replaced by grazing in
1985. Grazing was thought to be more natural and less expensive than
mowing. In the same period grazing was introduced in several other dune
areas as a result of a changing attitude to dune management. At the request
of the Amsterdam Waterworks volunteers started to prepare an inventory
of all sorts of organisms in 1985 in order to evaluate the effect of both
forms of management. For the mycota, the emphasis of the research was
laid on the fungi indicative of nutrient-poor grasslands, like waxcaps
(Hygrocybe), species of Entoloma, clavarioid fungi (Clavaria, Clavulinopsis) and earth-tongues (Geoglossum, Trichoglossum, Microglossum). No
permanent plots were installed but the whole grassland area was investigated several times a year. Unfortunately no data on the mycota are
available from previous times.
Waxcap grasslands
Grassland fungi are the most striking aspects of the mycota in the two
areas. Although the numbers of taxa show large annual fluctuations, there
has been a general increase in the number of taxa of Hygrocybe in both
areas (Fig. 11.1). Within 20 years both areas have developed into good
waxcap grasslands. Waxcap grassland is a term invented by Schweers
Grasslands in the coastal dunes
139
Fig. 11.1. Number of taxa of waxcaps, Hygrocybe, found in Eiland van
Rolvers and Groot Zwarteveld each year.
(1949) who noticed that in nutrient poor grasslands in The Netherlands a
certain mycota was present composed of species of Hygrocybe, certain
Entoloma species and earth-tongues. Grassland fungi belong to one of the
most reduced and threatened groups in The Netherlands (Nauta & Vellinga, 1995).
On Eiland van Rolvers a total of 13 taxa of Hygrocybe were found. The
Groot Zwarteveld, with a total of 17 taxa of Hygrocybe, is one of the best
waxcap grasslands in The Netherlands. Clavarioid fungi such as
Clavulinopsis luteo-alba, C. helveola and C. laeticolor occur as accompanying species. Besides an increase in number of taxa in both sites, a spectacular increase in the number of fruit bodies of Hygrocybe and Clavarioid
fungi was observed. Unfortunately there is no clear pattern in number of
taxa or fruit bodies of grassland Entoloma (Fig. 11.2).
Differences
There are also a number of differences between the two investigated areas.
More taxa of Hygrocybe occur in the mown area (Groot Zwarteveld), and
they are in general slightly more acidophilic (Table 11.1). In the mown
area a sudden increase is shown in 1993 in the species Hygrocybe
acutoconica, H. helobia, H. insipida, H. laeta, H. psittacina. In the grazed
area (Eiland van Rolvers) the increase in number of taxa has taken place
140
Marijke M. Nauta & Leo M. Jalink
Fig. 11.2. Number of taxa of grassland Entoloma found in Eiland van
Rolvers and Groot Zwarteveld each year.
more gradually, but in 1991 H. conica and insipida were suddenly found.
In the grazed area more taxa and especially far more fruit bodies of
earth-tongues were found. The same applies to the clavarioid fungi.
An explanation of the differences can be found in the difference in
landscape, management and vegetation. Eiland van Rolvers shows a more
varied vegetation on more broken ground, also with dune-scrub,
Crataegus and Hippophae. In particular, the small hilly area is drier than
Groot Zwarteveld. The richness in clavarioid fungi is connected with the
drier parts of the terrain; the richness in earth-tongues is caused by the
presence of gradients in the soil below the hills and at the margin of scrub.
Groot Zwarteveld is more decalcified because of the former use as potato
fields, which is manifested for example in the occurrence of Sphagnum in
the small pools, and the area is slightly wetter. This can explain the
difference in species of Hygrocybe.
Cattle do not graze rushes and the moist parts of Eiland van Rolvers are
partly overgrown by rushes. Protected by dense rush vegetation, seedlings
of birches have grown up, resulting in the development of birchwood on
Eiland van Rolvers and an increase of important and threatened
ectomycorrhizal fungi, like Lactarius vietus.
Grazing also has a direct enriching effect on the mycota, namely fungi
on dung. In the grazed area many coprophilous fungi were found which
have been under threat in The Netherlands. For example, Panaeolus
141
Grasslands in the coastal dunes
Table 11.1. Number of years in which taxa of Hygrocybe occurred in
Groot Zwarteveld and Eiland van Rolvers
Taxa of Hygrocybe
acutoconica
cantharellus
ceracea
coccinea
conica var. conica
conica var. conicopalustris
conica var. chloroides
glutinipes
helobia
insipida
laeta
luteolaeta
miniata var. miniata
miniata var. mollis
pratensis var. pallida
psittacina
russocoriaceus
virgineus var. virgineus
virgineus var. fuscescens
Groot
Zwarteveld
5
1
12
2
13
2
2
4
6
6
5
8
13
3
5
1
10
Eiland van
Rolvers
8
3
6
2
6
5
13
1
1
7
1
6
1
fimiputris is a species that seriously declined during the sixties and seventies
(Nauta & Vellinga, 1995), but has recently begun to recover. It is a species
that occurs on fibre-rich cattle dung, a substrate that had become very
scarce in The Netherlands in the past few decades. Thanks to an increase in
cattle grazed on nutrient-poor grasslands in the dunes, and nature reserves
more inland, several formerly threatened species are showing an increase.
Other interesting finds on the dung are Panaeolus speciosus (only recently
discovered in The Netherlands; Van de Bergh & Noordeloos, 1996), and
the rare Peziza bovina.
Mowing or grazing?
From the mycological point of view, both mowing and grazing have
proved to be very valuable management methods. Both result in a rich
mycota, although the species spectrum is different. As a result of our
studies, the management of the Municipal Waterworks of Amsterdam has
been advised to continue both methods. In general, mowing gives a larger
area of grassland and, consequently, more habitat for grassland fungi.
142
Marijke M. Nauta & Leo M. Jalink
Grazing results in a mosaic of vegetation types ranging from grassland,
scrub and small woodlands. This gives opportunities to a wider range of
fungi, including ectomycorrhizal fungi of woods and special groups like
dung fungi. As far as cost is concerned, grazing is generally cheaper than
mowing, especially since the hay is relatively useless in terms of nutrition
for cattle, but grazing does not prevent development of scrub and forest.
However, if grazing is stopped now, the area will probably become rapidly
overgrown again.
On Groot Zwarteveld a stable situation has now developed, with a
short, grassy dune slack vegetation, whilst on Eiland van Rolvers the
vegetation is still under development. Because of the expansion of rushes
and development of the Betula wood, some of the grassland in the western
part is becoming overgrown. In the central and eastern part the vegetation
is far more varied, with nicely developed scrub of Salix repens and Hippophae on the more calcareous parts.
Future
A difficult question in nature management concerns the future. Nowadays
there is a strong tendency to let nature rule itself. In the case of grasslands
in the dunes there have been some recent developments caused by human
intervention which have not been beneficial for biodiversity, for instance.
The large fields of Calamagrostis and Carex arenaria were caused by
humans, and bringing it back to a situation that existed around 1500 does
not seem unnatural. Of course, for many groups of (lower) fungi and other
organisms it is not known what the effect will be. But in general it is
thought that biodiversity will benefit. If natural processes are allowed to
do their job in the dunes the climax vegetation is the formation of dune
scrub and woods. The result of this would probably also be a biota rich in
fungal species.
References
Becker, A. & Baeyens, G. (1992). Wasplaten en andere paddestoelen in een
vochtige vallei van de Amsterdamse Waterleidingduinen. De Levende Natuur
93, 111–117.
Nauta, M. M. & Jalink, L. M. (1996). Ontwikkelingen in de mycoflora onder
invloed van begrazing op het Eiland van Rolvers. Gemeentewaterleidingen
Amsterdam: Amsterdam.
Nauta, M. M. & Vellinga, E. C. (1995). Atlas van Nederlandse paddestoelen. A. A.
Balkema Uitgevers B. V.: Rotterdam.
Grasslands in the coastal dunes
143
Schweers, A. C. S. (1949). De Hygrophorusweide, een associatie. Fungus 19 (2),
17–18.
Van de Bergh, F. & Noordeloos, M. E. (1996). Panaeolus speciosus P. D. Orton:
nieuw voor Nederland. Coolia 39, 74–77.
12
The conservation of fungi on reserves
managed by the Royal Society for the
Protection of Birds (RSPB)
MARTIN ALLI SON
Introduction
The Royal Society for the Protection of Birds (RSPB) is the largest wildlife
conservation charity in Europe with over one million members. It manages
over 150 nature reserves throughout the UK, covering more than 108 000
hectares. Away from the reserves the Society safeguards sites and species
of conservation importance through research, lobbying and education,
whilst outside the UK the RSPB is working with Birdlife International
Partners on major conservation projects in 18 countries in Europe, Africa
and Asia. The RSPB, together with voluntary organisations, has produced
a practical guide to conserving the UK’s biodiversity that has been adopted at Government level.
The Society is committed to the management of nonavian taxa on its
reserves, and is UK champion for nonbird species such as the medicinal
leech, Hirudo medicinalis. The Society’s interest in other biotic groups has
been somewhat biased towards invertebrates and lower plants, for the
most part to the exclusion of the fungi. This latter fact is understandable
given the difficulties with identification and the complex taxonomy within
mycology. In recent years an interest in the fungi on RSPB reserves has
been encouraged by one or two active field mycologists, and has gradually
led to the development of an in-house strategy for their conservation.
Threatened in Europe
A rapid decline of fungi across northern Europe has been reported since
the 1980s. Before discussing habitats and species on RSPB reserves the
European situation needs to be understood.
An estimated 5106 species of macromycetes are thought to occur in
144
Fungal conservation on RSPB reserves
145
northern Europe. Of these, 2658 (52%) are found on one or more
European Red Data list (Arnolds & de Vries, 1993). Proportionately more
extinct or endangered species on Red Data lists occur in highly industrialised and/or densely populated areas such as The Netherlands, Germany and Czechoslovakia. Nordic countries and Britain have a larger
proportion of vulnerable, rare or potentially threatened fungi.
No less than 55% of the European agaric fungi appear on one or more
Red Data lists (Arnolds & de Vries, 1993). Many of these species are
ectomycorrhizal and their ecological role is considered vital to the health
of the forests (Rayner, 1993). Some forests in industrialised areas of
Europe are showing sharp declines in their ectomycorrhizal species; this
often occurs before any signs of deterioration are detected in the host tree
species. Such declines particularly occur within hillside forests on poor
soils, an ecosystem commonly found in north Europe, where the habitat is
affected by acidification from air pollution (Fellner, 1993). This causes
decreases both in numbers of species of fungi and numbers of fruiting
bodies.
Of the fungal species characteristic of dry nutrient-poor grassland, 78%
are threatened in The Netherlands (Arnolds, 1989). These grassland species were once widespread, but changing agricultural methods coupled
with excessive use of fertilisers has caused a worrying decline in north
Europe generally. Mycologically rich grassland is often several centuries
old but it can be mirrored on sensitively managed old lawns.
The main external threats to European fungal communities can thus be
summarised as air pollution, change of land use and agricultural methods,
and urban encroachment. Commercial picking and overzealous clearing
up of dead wood may also seriously threaten fungi and organisms such as
fungus gnats, beetles and slugs that depend upon them. All of these threats
are relevant in the British Isles and on RSPB nature reserves.
Recording of fungi at RSPB reserves
Macrofungi fall into three broad habitat categories. The ectomycorrhizal
species associating with tree roots; the saprotrophic species of fungi living
on dead organic matter and recycling essential nutrients, and the lignicolous species inhabiting wood and thus creating the hollow trees needed
for hole nesting birds. For the purposes of describing fungi at RSPB
reserves, the species will be restricted to the agarics and boletes, the
Aphyllophorales and selected ascomycetes.
Many records on RSPB reserves originate from organised forays where
146
Martin Allison
Table 12.1. Important RSPB reserves for fungi showing totals of Red Data
list species
Reserve
Score? R
V
E
A
B
C
D
Abernethy
Tudeley Woods
Lake Vyrnwy
Geltsdale
Blean Woods
Northward Hill
Killiekrankie
Stour Wood
The Lodge
Minsmere
Leighton Moss
Wolves Wood
Loch of Strathbeg
180
109
28
24
19
18
17
16
15
14
13
12
11
38
15
6
5
1
1
3
5
1
1
7
2
2
1
8
5
1
1
1
1
21
25
6
5
5
5
5
3
6
3
6
3
3
2
4
4
1
1
1
1
2
1
1
1
1
1
1
1
British Red Data (preliminary) list Categories: R, Rare; V, Vulnerable; E,
Endangered.
European Red Data list Categories: A, widespread losses, rapidly declining
populations, many national extinctions, high-level concern; B, widespread losses,
evidence of steady decline, some national extinctions, medium-level concern; C,
widespread but scattered populations, fewer extinctions, lower-level concern; D,
local losses, some extinctions but mainly at edge of geographical range.
?Score weightings based on numerical values attributed to differing rarity
categories.
voucher specimens of the rarer species have been deposited in herbaria.
Other lists are supplied by regional amateur mycologists, or by the reserve
staff themselves. Individual mycologists tend to concentrate on certain
taxa that interest them. This is somewhat reflected in the choice of Red
Data list species.
Forty-four RSPB reserves have contributed lists totalling over 1860
species. Of these, 81 species occur on the European, and 77 on the British
provisional Red Data lists (Ing, 1992). Important reserves for threatened
fungi are shown in Table 12.1.
Some reserves have been studied in much greater detail than other
potentially rich reserves: Abernethy Forest, Highland; Tudeley Woods,
Kent; Leighton Moss, Cumbria; Nagshead, Gloucestershire, and The
Lodge, Bedfordshire, are among those that have been better studied (see
Fig. 12.1). Potentially rich sites include: Ynys hir, Powys; Wye-Elan
Valley, Powys; Geltsdale, Cumbria; Coombes Valley, Staffordshire;
Hobbister, Orkney.
Of the habitats represented on reserves, the most important mycologi-
Fungal conservation on RSPB reserves
147
Fig. 12.1. Locations of RSPB reserves that are important for fungal
conservation.
cally are Caledonian pine forest; the nutrient-poor, unimproved grasslands
of mainly northern and western Britain; and the semi-natural ancient
woodlands of southeast England. Sand dunes and western oak woodlands
also offer potential but, unfortunately, few data were available from
reserves.
Geographical distribution at reserves
Generally, fungi are widespread across the whole of Britain, and indeed
many British species are transglobal through temperate regions. Some
148
Martin Allison
species, however, show distinct geographical limitations. Of the more
interesting species at RSPB reserves, horse hoof fungus Fomes fomentarius,
Rozites caperatus (commonly found in Highland coniferous forests), and
the endangered Lactarius musteus found at Abernethy Forest, are all
predominantly fungi of northern Britain. By contrast, Russula aurata, R.
pseudointegra, Boletus pseudoregius, B. aereus and Leccinum crocipodium
are among the fungi that show a preference for warmer southern woodlands. There are a few predominantly western species, the most distinctive
of which is the delicate grey-blue fairy bonnet Mycena pseudocorticola,
recorded from Wye-Elan Valley, Powys, and West Sedgemoor, Somerset.
Important habitats for fungi at RSPB reserves
Caledonian pine forest
The Abernethy Forest reserve (Fig. 12.1) boasts one of the largest remaining areas of Caledonian pine forest. It is an important habitat for many
rare and threatened species and assemblages of fungi, both within the
forest and on the surrounding moorland. The reserve supports 33
European and 49 British Red Data list species; seven of the latter are
considered endangered.
The milk cap Lactarius musteus and the agaric Tricholoma focale are two
endangered species on the British list associated with Scots pine Pinus
sylvestris. No fewer than 10 stipitate hydnoid fungi can be found at
Abernethy; all are on the Red Data lists and two, Bankera fuligineoalba
and Phellodon confluens, are endangered in both the UK and Europe
(Table 12.2). Another endangered species is the purple cup-fungus Ombrophila violacea that grows on decaying leaves in boggy places.
Abernethy Forest also supports impressive assemblages of some important genera, notably 55 species of Cortinarius including 9 on the British Red
Data list, 40 species of Russula (7 on the Red Data list) and 14 species of
Tricholoma (3 Red Data list species).
Semi-natural ancient woodland
These predominantly southern woodlands are the strongholds of the ectomycorrhizal species in Britain and harbour important and increasingly
rare assemblages of fungi. The most representative reserve for this group is
at Tudeley Woods in Kent (Fig. 12.1), which supports 26 European and 16
British Red Data list species; two of the latter are considered endangered.
Fungal conservation on RSPB reserves
149
Table 12.2. Stipitate hydnoid species on RSPB reserves
Species
Red Data list
categories
Reserve
Bankera fulgineoalba
Hydnellum aurantiacum
H. caeruleum
H. concrescens
H. peckii
H. scrobiculatum
H. spongiosipes
Hydnum repandum
Phellodon confluens
P. melaleuceum
P. niger
P. tomentosum
Sarcodon glaucopus
S. imbricatum
S. scabrosum
A, E
B, V
B, V
C, V
B, E
B, E
C, R
widespread
A, E
C, V
B, R
B, E
B
C, V
B, E
Abernethy
Abernethy
Abernethy
Abernethy, Tudeley Woods
Abernethy
Blean Woods, Tudeley Woods
Tudeley Woods
Abernethy
Abernethy, Northward Hill
Tudeley Woods
Abernethy, Northward Hill
Abernethy
Abernethy, Tudeley Woods
Tudeley Woods
For Red Data list categories see Table 12.1.
Boletus pseudoregius is a rare and possibly endangered European fungus
associated with warm, southern oak woodland and was first noted in 1995.
Along with B. regius and B. speciosus, this taxonomically difficult group is
currently under review (Marren, 1998). Other European Red Data list
boletes at Tudeley are B. aereus, B. impolitus, B. queletii, B. radicans and B.
appendiculatus. These fungi indicate a continuity of woodland cover over
many centuries. The second endangered British species at Tudeley Woods
is the hydnoid Sarcodon scabrosus. Hedgehog fungi associated with RSPB
woodlands in Kent are also found at Blean Wood and Northward Hill
(Fig. 12.1).
Other rare European species of southern woodlands are Cortinarius
bulliardii from Fore Wood, East Sussex (Fig. 12.1), and the attractive C.
violaceus at the small Barfold Copse reserve in Surrey. Minsmere in Suffolk
holds Boletus queletii and Hericium erinaceus, whilst Boletus impolitus is
found in Blean Woods. The black-capped bolete Strobilomyces floccopus is
rare outside the Severn and Thames basins but occurs at a few outlying
stations including Tudeley Woods and Fore Wood, East Sussex. Entoloma
euchroum is a small blue fungus that is included on eight of the eleven
individual country Red Data lists across Europe, but not on the British
one. It occurs at Stour Wood and Wolves Wood in Essex as well as at
Tudeley where it grows only at the base of coppiced hazel Corylus avellana.
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Martin Allison
Unimproved nutrient-poor grassland
This habitat is mainly represented in the north and west of Britain, where it
has suffered less from intensive agriculture than in lowland Britain. Such
grassland is represented in the upland pastures of: Abernethy; Killiecrankie, Perthshire; Geltsdale, Cumbria; Lake Vyrnwy, Powys (Fig. 12.1).
Important groups of fungi associated with these pastures are the waxcaps
and the entolomas. Many of these species are declining at alarming rates in
some parts of Europe.
Of the 43 waxcap species recorded from RSPB reserves, 28 are found on
unimproved pasture at Abernethy, 20 at Lake Vyrnwy, 16 at Geltsdale, 15
each at Killiecrankie and Leighton Moss, Lancashire. Most of the 13
waxcaps that occur at Tudeley Woods are found within the coppiced
woodland, including the Red Data listed Hygrocybe calyptriformis. It is
assumed that the presence of these waxcap species reflects the fact that the
woodland was once grazed as wood pasture when extensive grassy swards
would have been present. Forty-nine species of Entoloma have been recorded at RSPB reserves: 25 occur at Lake Vyrnwy, and 24 at Abernethy.
Other declining grassland fungi are the clavarioid fungi, 16 of which occur
at RSPB reserves. Of the four grassland earth-tongues recorded at reserves, three are European Red Data species, Microglossum viride at
Killiecrankie, Geoglossum nigritum at Leighton Moss and Trichoglossum
hirsutum at Leighton Moss, Tudeley Woods and on the lawns at The
Lodge, Bedfordshire, which is headquarters to the RSPB. Such wellestablished lawns can be rich in fungi such as Hygrocybe, Entoloma,
Clavaria and Geoglossum.
Heathlands
Heathlands support a few uncommon species of fungi, but are mainly
noted for their rare bird, higher plant and reptile assemblages. Of the more
interesting ones on reserves are Clavaria argillacea (Abernethy, The
Lodge, Minsmere, Vane Farm and Loch of Strathbeg), Cantharellula
umbonata (Vane Farm and Lake Vyrnwy) and Mycena uracea (Stour
Wood, Blean Woods, Northward Hill, The Lodge and Abernethy). This
latter species is often associated with burnt ground and is found particularly around heather shoots.
Plantations
Habitats within nonindigenous woodlands should not be ignored. Of
particular worth are conifer plantations and sweet chestnut coppice Casta-
151
Fungal conservation on RSPB reserves
Table 12.3. Regional importance of selected taxa from two RSPB reserves
in southeast England shown as a percentage of the total species for
southeast England
Taxon
Tudeley (%)
Blean (%)
Total for the
two reserves (%)
Lactarius
Russula
Mycena
Pluteus
68
53
43
32
44
35
31
14
72
55
49
41
nea sativa, both occasionally supporting rare species and communities of
fungi. Sweet chestnut coppice in southern England often harbours species
found in native mixed oakwoods, particularly the boletes, russulas and red
data hydnoid species. The rare Strobilomyces floccopus mentioned earlier
has been found in chestnut coppice. A resupinate corticioid species, Byssocorticium efibulatum, was recorded from the underside of chestnut logs
at Tudeley Woods in 1994 and was a new British record (Roberts, 1994).
Introduced conifers play host to many species of important genera such
as Lepiota and Cortinarius. At The Lodge the European Red Data list
species Volvariella caesiotincta, Geastrum pectinatum, G. triplex and Hygrophorus hypothejus are found alongside several scarce lepiotas within the
softwood plantations. Accompanying these is the very rare Hygrophorus
speciosus, being newly recorded for Britain in 1984, and the only known
site in the country for this fungus.
Regional importance of two RSPB reserves in southeast England
for fungi
Comparisons with major groups of agarics in southeast England are
possible with the publication of an updated checklist for East and West
Sussex, Kent and Surrey (Dennis, 1995). For the two large RSPB woodland complexes of Tudeley Woods and Blean Woods, both in Kent,
regional and national percentages of selected groups of fungi are shown in
Table 12.3. The two reserves together support 90% of the Lactarius species
and 72% of Russula that occur in southeast England. Tudeley Woods with
its extensive derelict coppice and much fallen mature timber, including elm
and beech, is a favoured site for lignicolous Pluteus fungi.
A further comparison with a major mycological site, Bedgbury Pinetum
in Kent, shows that Tudeley Woods supports 26% of the southeast
152
Martin Allison
England agaric biota, compared with 32% at Bedgbury (Weightman,
1993). Tudeley also supports 26% of the region’s aphyllophoralean fungi.
The conservation of fungi
The conservation of many fungi at nature reserves tends to be limited by
the fact that most threats are beyond the control of the site’s management.
By far the greatest of these is acidification and eutrophication from airborne pollutants, although Britain suffers less than most mainland
European countries. This can best be tackled through international environmental policy. Some experimental work is in progress in other parts of
Europe where soil manipulation has been monitored and compared with
fungal productivity (Baar & Kuyper, 1993). Results from this work are not
yet conclusive but do suggest that a rich ectomycorrhizal biota correlates
with a thin or absent organic layer. This can be achieved artificially by the
removal of litter/mulch layers from the soil, thus helping reverse the trend
in fungal decline by removing damaging nitrogenous compounds. However, this type of management could prove more harmful than acidification
to ancient woodland soil ecosystems and their saprotrophic fungal species.
To date there are no comparisons available between managed and
nonintervention forestry in this country with regards to fungal communities (Keizer, 1993). Opportunities present themselves on extensive woodland reserves such as Tudeley, Blean or Abernethy where nonintervention
and actively managed compartments coexist. In order to make any study
feasible either large permanent quadrats or substantial transects need to be
set up and monitored for a minimum ten year period. This would ideally
merge with studies of other biotic groups such as National Vegetation
Classification (NVC) vegetation recording or Institute of Terrestrial Ecology (ITE) butterfly and other invertebrate monitoring programmes.
Wood-inhabiting fungi, along with many invertebrates, are threatened
by timber management/tidying up operations. Apart from essential safety
measures, dead wood is left in situ, standing or fallen, as important habitat
(Green & Alexander, 1993). Although few RSPB reserves hold areas of old
parkland, which are often internationally important for fungi, there are
many overmature individual trees within wooded areas that may well hold
important lignicolous species. Some uncommon bracket fungi rely on
access to the heartwood of trees to survive. A proportion of healthy mature
trees is always retained in perpetuity on reserves as potential future hosts
for these fungi. Heartwood species do not kill off the host trees as the outer
sapwood is unharmed and continues to feed the tree, thus producing the
Fungal conservation on RSPB reserves
153
hollow oaks and beeches of sites such as Windsor Great Park.
Management of the remaining unimproved nutrient-low grassland on
RSPB reserves is similar to managing for their bird or botanical interest. A
regime of low-density grazing, or mowing/forage harvesting followed by
grazing, is generally recommended. Fertilisers are avoided other than
natural droppings from livestock.
The management at Abernethy reserve is especially delicate as it is a key
site for both Caledonian pine forest species and fungi of unimproved
pasture. A balance needs to be sought here to stop overencroachment onto
the important Hygrocybe/Entoloma grassland communities, whilst ensuring the pinewoods are given full protection and allowed to perpetuate for
the future.
Large-scale collection of fungi can be indirectly damaging. The penny
bun bolete Boletus edulis is considered threatened through overpicking in
Poland. Research has shown, however, that collection may actually stimulate the growth of fruit bodies. The problems arise more from soil compaction through trampling, and the removal of habitat for the mammals and
invertebrates that feed on fungi. In Britain, commercial and private collecting of the penny bun and other species is increasing through exposure
via natural history and cookery programmes on television.
Internal threats to fungi on reserves
Potentially more threatening than human trampling is the modern practice
of using heavy timber extraction machinery on reserves, which can lead to
extensive ground disturbance and soil compaction. This can sever the
mycorrhizal links between tree and fungus, causing long-term ill health
within the forest ecosystem.
Conflicts may arise within conservation management when one group of
animals or plants is favoured above another. An example encountered on
RSPB reserves is short-term coppice and ride management to encourage
butterflies and other invertebrates. This creates large areas of dense scrub
and bramble which limits the fruiting of sometimes rare species of fungi
which once thrived in the damp and shaded uncut coppice. This particular
situation is normally rectified by allowing the coppice to regrow whilst
creating scalloping and glades away from the threatened fungus species.
At the RSPB headquarters in Bedfordshire the rare Hygrophorus
speciosus was seen to be threatened by a heathland restoration project
where trees were to be removed close by the fungus. An island of conifer
was therefore retained and later saplings were replanted on the periphery
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Martin Allison
to act as a windbreak against desiccation. Furthermore, an attempt will be
made to translocate the fungus into other areas of the reserve through
encouraging mycorrhizal links with young planted conifers that will then
be removed and transplanted elsewhere.
The RSPB has begun to address these problems through its nonavian
biodiversity monitoring programme (Cadbury & Evans, 1995). Species
currently within the programme include Boletus pseudoregius, Hygrophorus speciosus, Hygrocybe calyptriformis, Hericium erinaceum and
several hydnoid species.
Conclusions
In summary, the RSPB hopes to manage for fungi under each reserve’s
broad policy statements. To that effect, all RSPB reserve management
plans will eventually contain reference to British and European Red Data
listed species and threatened communities, and include prescriptions for
their conservation alongside those of other taxa.
It is difficult to provide precise management prescriptions for such
understudied organisms as the macrofungi. As has already been mentioned, many threats are external and habitats have already been destroyed. The rule of thumb must be that, if the fungus is present on the
reserve then the habitat is fundamentally suitable and the status quo
should be retained. Research should then be encouraged to ensure the
populations are stable or increasing and this should include the effects of
grazing, the effects of coppicing, the study of fungal communities in
nonintervention zones, etc., depending on the habitat. Guidelines should
be sought from scientists specialising in field mycology on ways to approach such research.
The important factor in fungal conservation on reserves is that the land
managers in charge of each site are aware of the presence and status of
fungi and that they retain the long-term traditional management or ‘nonmanagement’ that has allowed them to survive up to the present. To do
this successfully, regular recording is necessary, preferably by knowledgeable amateurs from outside the Society.
The RSPB has taken an essential first step in acknowledging the importance of the fungi on its nature reserves by ensuring the management
planning system is aware of their presence and current status, and by
including the rarest species and communities in its nonavian biodiversity
monitoring programme.
Fungal conservation on RSPB reserves
155
Acknowledgement
Adapted by the author from his contribution to the RSPB Conservation
Review 1997 (Allison, 1997).
References
Allison. M. (1997). The importance of RSPB reserves for fungi. RSPB
Conservation Review 11, 88–95.
Arnolds, E. (1989). A preliminary red data list of macrofungi in The Netherlands.
Persoonia 14, 77–125.
Arnolds, E. & de Vries, B. (1993). Conservation of fungi in Europe. In Fungi of
Europe: Investigation, Recording and Conservation (ed. D. N. Pegler, L.
Boddy, B. Ing & P. H. Kirk), pp. 211–230. Royal Botanic Gardens: Kew.
Baar, J. & Kuyper, T. W. (1993). Litter removal in forests and effects on
mycorrhizal fungi. In Fungi of Europe: Investigation, Recording and
Conservation (ed. D. N. Pegler, L. Boddy, B. Ing & P. H. Kirk), pp. 275–286.
Royal Botanic Gardens: Kew.
Cadbury, J. & Evans, C. (1995). The monitoring of non-avian species of
conservation importance at RSPB reserves: a strategy. Unpublished report,
RSPB: Sandy.
Dennis, R. W. G. (1995). Fungi of South East England. Royal Botanic Gardens:
Kew.
Fellner, R. (1993). Air pollution and mycorrhizal fungi in Central Europe. In
Fungi of Europe: Investigation, Recording and Conservation (ed. D. N. Pegler,
L. Boddy, B. Ing & P. H. Kirk), pp. 239–250. Royal Botanic Gardens: Kew.
Green, T. & Alexander, K. (1993). Dead wood – eyesore or ecosystem? Enact 1,
11–14.
Ing, B. (1992). A provisional Red Data list of British fungi. Mycologist 6,
124–128.
Ing, B. (1993). Towards a red list of endangered European macrofungi. In Fungi
of Europe: Investigation, Recording and Conservation (ed. D. N. Pegler, L.
Boddy, B. Ing & P. H. Kirk), pp. 231–237. Royal Botanic Gardens: Kew.
Keizer, P. J. (1993). The influence of nature management on the macromycete
flora. In Fungi of Europe: Investigation, Recording and Conservation (ed. D.
N. Pegler, L. Boddy, B. Ing & P. H. Kirk), pp. 251–269. Royal Botanic
Gardens: Kew.
Marren, P. (1998). Boletus regius Kromb., the Royal Bolete and the pretender
Boletus pseudoregius Estades. Species Recovery Report no. 111. English
Nature/Plantlife.
Rayner, A. D. M. (1993). The fundamental importance of fungi in woodlands.
British Wildlife 4, 205–215.
Roberts, P. (1994). Interesting and unusual corticioid fungi from Slapton, Devon
II. Mycologist 8, 115–118.
Weightman, J. (1993). The fungi of Bedgbury forest. Transactions of the Kent
Field Club 13, 1–28.
13
Strategies for conservation of fungi in
the Madonie Park, North Sicily
G I U S E P P E V E N T U R E L L A & SA L V A TO R E L A
R O CC A
Introduction
The Madonie Park, covering 40 000 hectares, is one of the most interesting
and floristically differentiated areas in the Mediterranean basin. Its territory shows a very high degree of diversity as instanced by about 1600 taxa
of the vascular flora with a high percentage of endemic species. The forest
vegetation, in the Mediterranean to subatlantic belt, is varied and mainly
characterised by evergreen oak (both Quercus ilex and Q. suber) and
deciduous woods with Quercus pubescens, Q.virgiliana, Castanea sativa, Q.
petraea and Fagus sylvatica. Exotic trees are widespread in all the Madonie
mountains, including Pinus halepensis, P.pinea, P. nigra, Cedrus atlantica,
C. deodara, Cupressus sempervirens, C. arizonica, C. macrocarpa, Abies
alba, A.cephalonica, Robinia pseudoacacia and Eucalyptus camaldulensis.
The landscape is also composed of rocky environments, grasslands and
wetland sites. All these environments are also very rich in cryptogams
(lower plants) but knowledge about them is still inadequate.
As regards fungi, it is noteworthy that since the second half of the
seventeenth century the local population has exploited the understorey
products as a source of food or for income. In the last decade mushroom
picking, previously limited to relatively few people, has become a widespread pastime with potentially adverse consequences to mycelium growth
and fruit body appearance. This threat, together with the lack of knowledge of Sicilian fungi, causes serious problems for any attempt to safeguard the integrity of the ecosystem. In addition, the concern for fungi by
local authorities and the scientific community is still very slight.
As a first step towards conservation of the Madonie mycota, a census
project was started in 1991 (Venturella, 1992). The aims of this programme
are to increase the knowledge on the distribution and ecology of the
156
Conservation of fungi in the Madonie Park, North Sicily
157
Sicilian macrofungi, to locate rare and endangered taxa and to propose
conservative actions.
Data recorded
At present 614 taxa included in 200 genera belonging to 79 families have
been recorded. This corresponds to 50% of the known Sicilian mycota.
Considering that in a previous paper (Venturella & Mazzola, 1991) concerning the state of the mycological exploration in Sicily, only 65 taxa
belonging to 49 genera were reported for the Madonie Mountains, it is
clear that the new data represent a major improvement in knowledge.
Among the taxa recorded three main groups were identified: taxa with
wide distribution and ecology, taxa strictly linked to some specific environments, and taxa which are rare and/or distributed only in restricted areas.
Among the taxa showing a wide distribution and ecology, Amanita
pantherina, A. phalloides and Lepista nuda are widespread. They occur,
mainly during autumn and winter, in all the woodlands and are abundantly represented in terms of their numbers of fruit bodies. Mycena pura and
Schizophyllum commune have similarly wide distribution as these taxa but
their fruiting period was extended all year long. The fruiting period of
Russula delica, R. chloroides and R. foetens is, however, restricted to
autumn. These taxa also show a wide distribution, being frequently recorded.
Agaricus silvicola, Clitocybe nebularis, C. odora, Entoloma lividum and
Lactarius chrysorrheus are widely distributed in the broad-leaved woods
and in many other localities within the area investigated. Amanita rubescens, Inocybe rimosa and Tricholoma scalpturatum are also widely distributed, but their appearance is limited as far as the number of records is
concerned. Among the boletes, Boletus aereus is the most widely distributed and recorded species. In marked contrast with its occurrence in the
Apennine woods, it is recorded not only in evergreen oak woods and
thermophilic broad-leaved woods but also in the beech woods which are
distributed in the northern and eastern parts of the island from 1200 to
2000 m.
Of taxa strictly linked to particular environments, Hebeloma cistophilum, Lactarius tesquorum and Leccinum corsicum are widespread in
the areas characterised by the presence of Cistus creticus, C. monspeliensis
and C. salvifolius. Cortinarius cedretorum, Geopora sumneriana and
Tricholoma tridentinum var. cedretorum (Venturella, 1995) are prevalent in
the woods with Cedrus atlantica and C. deodara, which were introduced
158
Giuseppe Venturella & Salvatore La Rocca
throughout the area investigated since the 1960s, after the reforestation
carried out by the local Forestry Administration.
The use of wood for charcoal caused the occurrence of fungi specifically
associated with burnt areas such as Faerberia carbonaria, Pholiota highlandensis, Hebeloma anthracophilum and Tephrocybe atrata. In areas reforested with Eucalyptus camaldulensis, the occurrence of Setchelliogaster
tenuipes is noteworthy (La Rocca & Anastase, 1997a).
Although present knowledge is incomplete, we can go some way to
evaluating the state of decline of Sicilian macrofungi by making a comparison between this survey and the provisional European Red Data list of
endangered macrofungi published by Ing (1993). As shown in Table 13.1,
Boletus rhodoxanthus, belonging to Group A, seems not to be declining
since it was recorded 23 times during the period of investigation. Of taxa
included in Group B, Boletus impolitus, B. queletii and Hygrophorus russula are quite common in the Madonie area, whilst Entoloma bloxamii,
Myriostoma coliforme (La Rocca & Anastase, 1997b) and Boletus fechtneri
also confirm their status as being infrequent in the Mediterranean area.
In Group C, Boletus radicans and B. aereus are really common, while
Omphalotus olearius and Leucopaxillus gentianeus were frequently recorded in the broad-leaved woods. Of the taxa included in Group D,
Amanita caesarea was prevalent only in the siliceous areas of the Madonie
Mountains, where fruit bodies could be abundant. Among the taxa recorded, Agaricus bernardii, A. pseudopratensis, Choiromyces meandriformis, Collybia cookei, Coprinus vosoustii, Entoloma undatum, Gyroporus
cyanescens, Helvella solitaria, Lactarius decipiens, Melanoleuca
pseudovenosa and Tricholoma bresadolanum were infrequent, while Entoloma atrocoeruleum, Macrotyphula fistulosa, Ossicaulis lignatilis and
Pleurotus nebrodensis were rare not only in the Madonie Mountains but
also in all the Sicilian territory.
These taxa are not included in Ing’s list (Ing, 1993) but they could be
regarded as worthy of protection, at least in Sicily.
Conservation strategies
In parallel with the monitoring and inventory of fungal diversity in the
Madonie area, a protection plan for the mycota was proposed (Venturella,
1992). It provides for introduction of rules concerning picking and the
trade in fungi within the Madonie Park. In particular, it includes a prohibition of the use of rakes in the searching process, because they can cause
damage to the mycelium and roots, and also the use of plastic bags for
Conservation of fungi in the Madonie Park, North Sicily
Table 13.1 Taxa included in Ing’s list compared with the number of
records of the Madonie area
Taxa
Group A
Group B
Group C
Group D
?Ing (1993).
Records
Boletus regius
Boletus rhodoxanthus
Boletus satanas
Aureoboletus gentilis
Boletus fechtneri
Boletus impolitus
Boletus queletii
Cortinarius bulliardii
Cortinarius violaceus
Entoloma bloxamii
Hericium erinaceus
Hygrophorus russula
Lactarius mairei
Myriostoma coliforme
Phellodon niger
Tricholoma acerbum
Amanita franchetii
Astraeus hygrometricus
Boletus aereus
Boletus appendiculatus
Boletus fragrans
Boletus radicans
Cantharellus cinereus
Coriolopsis gallica
Cortinarius cinnabarinus
Cortinarius praestans
Ganoderma resinaceum
Geastrum rufescens
Grifola frondosa
Hygrocybe punicea
Hygrophorus arbustivus
Hygrophorus camarophyllus
Lactarius violascens
Lepiota cortinarius
Leucopaxillus gentianeus
Mutinus caninus
Omphalotus olearius
Oudemansiella pudens
Pisolithus arhizus
Pulcherricium caeruleum
Ramaria botrytis
Strobilomyces strobilaceus
Tricholoma sejunctum
Tricholoma squarrulosum
Tulostoma brumale
Amanita caesarea
Boletus torosus
Clavariadelphus truncatus
Cortinarius cedretorum
Geastrum triplex
9
23
12
3
1
28
26
3
4
1
5
18
7
1
3
12
4
8
136
4
11
98
3
2
1
9
4
1
2
3
7
1
1
8
27
1
30
10
8
8
8
3
2
8
1
39
8
2
3
8
159
160
Giuseppe Venturella & Salvatore La Rocca
storage purposes because they accelerate fruit body fermentation. Actions
promoting good forest management and restricting grazing were also
carried out.
A fundamental requirement is also to set a limit on the amount of
mushrooms that may be picked by amateur and professional collectors.
The rules also include that some species, such as Boletus edulis and related
boletes, together with Calocybe gambosa and Cantharellus cibarius should
not be collected if their caps are less than 2–3 cm in size. The picking of
Caesar’s mushroom (Amanita caesarea) in the early stages of development
will also be forbidden in order predominantly to safeguard public health
but also as a conservation measure.
Finally, a specific protection plan is proposed for Pleurotus nebrodensis,
a rare Sicilian spring taxon with isolated distribution in the Madonie
mountains of northern Sicily and the Etna mountain of eastern Sicily. The
growing sites for P. nebrodensis are in the A zone (integral reserve) of the
Madonie and Etna regional parks between 1400 and 2000 m, in mesoxerophilous pastures with Cachrys ferulacea. Since the seventeenth century, P. nebrodensis has been in great demand in the Madonie area because
of its peculiar organoleptic characters. Because of the decrease in its
natural fruiting, demand by consumers and restaurants exceeds the supply
and results in high prices that vary from ITL 80 000 to 100 000 per kg. The
increase in the number of collectors picking immature fruit bodies caused a
conspicuous decline in fruiting and raised the risk of irreparable damage to
the sustainability of this particular taxon. The protection plan for P.
nebrodensis deals with legislative actions in order strictly to forbid all
picking of fruit bodies throughout the Madonie area and to include it in
the Italian Red Data list for fungi, as previously proposed by Venturella et
al. (1997). In parallel to these safeguarding actions on this taxon, its ex situ
cultivation was started (Venturella & Ferri, 1996; Venturella, 1999) to
safeguard natural fruiting and as a sustainable alternative for the economy
of the Madonie area.
Acknowledgements
The financial support of Ente Parco delle Madonie is gratefully acknowledged.
References
Ing, B. (1993). Towards a red list of endangered European macrofungi. In Fungi
of Europe: Investigation, Recording and Conservation (ed. D. N. Pegler, L.
Conservation of fungi in the Madonie Park, North Sicily
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Boddy, B. Ing & P. M. Kirk), pp. 231–237. Royal Botanical Gardens: Kew,
UK.
La Rocca, S. & Anastase, A. (1997a). Un interessante gasteromicete agaricoide in
Sicilia: Setchelliogaster tenuipes (Setchell.) Pouzar. Atti 4e Giornate
Confederatio Europaea Micologiae Mediterraneae, Poggibonsi (Italia), pp.
21–22.
La Rocca, S. & Anastase, A. (1997b). Myriostoma coliforme in Sicilia. Micologia
e Vegetazione Mediterranea 12 (2), 114–120.
Venturella, G. (1992). Progetto per una banca dati sulla micoflora siciliana.
Quaderni di Botanica Ambientale e Applicata 2 (1991), 107–110.
Venturella, G. (1995). Tricholoma tridentinum var. cedretorum Bon
(Tricholomataceae), a misappreciated taxon from Sicily. Documents
Mycologiques 98–100, 465–467.
Venturella, G. (1999). The conservation of Pleurotus nebrodensis through its
cultivation. In Abstracts, XIII Congress of European Mycologists, p. 131.
Alcalà de Henares, Madrid, Spain, 21–25 September 1999.
Venturella, G. & Ferri, F. (1996). Preliminary results of ex situ cultivation tests
on Pleurotus nebrodensis. Quaderni di Botanica Ambientale e Applicata 5
(1994), 61–65.
Venturella, G. & Mazzola, P. (1991). Present state of the mycological exploration
in Sicily. 6th OPTIMA Meeting’s Proceedings. Botanika Chronika 10,
889–894, Delphi, 1989.
Venturella, G., Perini, C., Barluzzi, C., Pacioni, G., Bernicchia, A., Padovan, F.,
Quadraccia, L. & Onofri, S. (1997). Towards a Red Data List of fungi in
Italy. Bocconea 5, 867–872.
14
Fungal conservation in Ukraine
D. W. M INTER
Introduction
In countries that made up the former Soviet Union, however much one
wants to concentrate on fungal conservation, general infrastructure problems demand attention. Until they have been addressed, little conservation
of anything can take place. Ukraine is no exception. A number of agencies
are committed to eliminating poverty and sustaining development in the
transition countries in Central and Eastern Europe; the Department for
International Development (DFID) is the British government department
responsible for this. DFID has published a concise assessment of
Ukraine’s political, economic and environmental background which can
be found in the UK Government’s 1998 country strategy paper for
Ukraine at the web site :http://www.dfid.gov.uk/public/what/pdf/
ukraine–csp.pdf9, with the UK Government’s environmental strategy for
Ukraine being on :http://bc.kiev.ua/english/work/envstr.pdf9. The realities described in these two straight-talking documents make one admire
all the more the dedication of those Ukrainian mycologists who have
stayed in Ukraine and who somehow manage to continue their work in
surroundings that are often difficult. Problems in Ukrainian science and
education are a microcosm of those at national level. The issues that must
be faced in solving those problems are extremely difficult, and many are
contentious. It is inappropriate to discuss them here.
Ukraine does, however, have a national policy on nature conservation.
The present work describes efforts by a team of Ukrainian and British
scientists to provide infrastructure for fungal conservation within that
national policy, through improved computing and informational resources. We have been able to organise a programme for the supply of
second-hand computers. These are suitable for basic data entry of text
information, but have severe limitations in more sophisticated use. Mod162
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163
ern machines are needed for work with graphics, and for internet and
modern e-mail connections. They need to be equipped with modems and
CD-ROM drives, including at least some CD-ROM writers, and should be
accompanied by some scanners and printers of a reasonable quality.
Without at least some machines of that quality, fungal conservation in the
former Soviet Union will always be behind the times. That sort of equipment, however, generally has to be purchased new. At present two major
awards, from the UK Darwin Initiative (Biodiversity Information in the
Former Soviet Union, 1999–2002), and the European Union organisation
INTAS (Infrastructure Action for the Life Sciences, 2000–2002), and a
smaller third award from the Royal Society of London (Electronic Distribution Maps of Ukrainian Fungi and Plants, 1998–2000) are helping to
provide such equipment for critical institutions working with neglected
groups of organisms, including fungi, in Ukraine and other newly independent states. The two major awards are also helping to pay for some e-mail
connections and access to the Internet.
Mycologists in Ukraine
Ukraine has a great mycological tradition. The Flora of Fungi of Ukraine,
the Handbook of the Fungi of Ukraine and various other major works
document the mycological exploration of the country (Dudka & Burdiukova, 1996; Heluta, 1989; Kopachevskaya, 1986; Makarevich, Navrotskaya & Yudina, 1982; Merezhko, 1980; Merezhko & Smik, 1991;
Morochkovskiı̈ et al., 1967, 1969, 1971; Oxner, 1937, 1956, 1968, 1993;
Smik, 1980; Smitskaya, 1980, 1991; Smitskaya, Smik & Merezhko, 1986;
Wasser, 1980, 1992; Zerova et al., 1971; Zerova, Radzievskiı̈ & Shevchenko, 1972; Zerova, Sosin & Rozhenko, 1979).
During the Soviet period, contact between Ukrainian scientists and their
western counterparts was often difficult. There was a lack of money for
travel and collaborative work, with the limited resources being allocated to
senior scientists and people with good contacts in Moscow. Although the
political barrier has gone, some of the other factors remain, so that a
residual isolation still exists. It is timely, therefore, that the forthcoming
14th Congress of European Mycologists (14CEM) will be held in Crimea
in 2003.
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Computing resources
Conservation of biodiversity involves decisions and good decision-making
requires accurate and readily available information. Nowadays, computers service this requirement. In the former Soviet Union computing
resources available to people working with nature conservation vary enormously. The Zoology Institute of the Russian Academy of Sciences in St
Petersburg has extensive computing skills and at least some modern computers. Some other institutions, for example biological research centres
formerly with military significance, and some institutes and departments
for experimental biology also have considerable facilities and skills. On the
whole, though, the range of resources is highly skewed. Most biological
institutions, universities, polytechnics and the like are very poorly equipped. The same is largely true for scientific societies, amateur societies
(where they exist) and nongovernmental organisations, and even more so
for schools, nature reserves, national parks, national nature parks, and
private individuals with an interest in conservation. Some have no computers at all. This picture is general throughout the former Soviet Union.
Ukraine is probably no better off than many other republics and, within
Ukraine, mycology has often fared worse than other sciences.
Collecting and handling information about Ukrainian fungi
With a programme in place delivering computers to Ukraine for fungal
conservation, the details of keyboarding, storing and using the desired
information can be addressed. For fungi, biological records stem from
four main sources: published records, unpublished lists, labels of reference
collections, and field observations not backed by any specimens. To handle
this information, three main databases are needed: a Nomenclatural and
Taxonomic database, a Bibliographic database, and a database of the
Biological Records themselves.
Problems with alphabets and instability of data
The first and most obvious problem when working in the former Soviet
Union is one of alphabets. Scientific names, their authors, and places of
publication of those names are essential components of any information
system dealing with biological recording. These components, by and large,
use the Latin alphabet. In the former Soviet Union, however, the Cyrillic
alphabet is in general use for vernacular names of organisms, locations,
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names of collectors and identifiers, substrata, notes and many other aspects of biological recording. This means all keyboarding work needs
ready and convenient access to both alphabets. Indeed, one quickly learns
that the Cyrillic alphabet, like the Latin alphabet, exists in several forms:
Russian, Ukrainian or Belarussian versions may be required at different
times. Keyboarding is made much slower and more difficult by the constant need to switch between alphabets.
Anyone working with biological recording of neglected groups soon
understands how labile information can be. The information system must
accommodate, for example, the rapid changes in taxonomy which characterise contemporary mycology. This is done by keeping an unchanged
copy of the original scientific name used as an identification of the organism in a separate field from current opinion of what that organism should
be called. Biological databases that do not provide this basic service are of
very doubtful value. Even so, handling data from the former Soviet Union
can be a shock. Not only organism names but also, seemingly, every other
item of data is prone to change. Within one human life span, for example,
St Petersburg has been renamed to Petrograd, Leningrad and back again
to St Petersburg. How many other place names in the former Soviet Union
have also undergone name changes in the last ten years? New countries
have appeared which did not previously exist. Regions and cities, towns
and villages have changed their names. Vast numbers of ‘Lenin Streets’
have been swept away, together with ‘Comsomol Avenues’ and ‘October
Revolution Squares’.
The latter reminds us that the October Revolution itself occurred in
November: an overdue calendar reform in the early 1920s changed the
date, and thereby incidentally also changed the meaning of dates on every
herbarium packet already in existence in every former Soviet Union botanical collection.
The inescapable conclusion is that, for proper data recording, particularly in the former Soviet Union, an intact copy of all original data should
be stored in a suite of fields totally separate from those storing current
opinion of what that information means. Filling in fields storing current
information is, of course, a different job from data gathering. It is a large
overlay, ideally carried out by trained and experienced editors. There is
also, however, a further complication. If the only objective for using the
resulting information is domestic, records with the organism name in the
Latin alphabet, and all other information in the Cyrillic alphabet may be
adequate. But if the longer-term objective is international use of the data,
then a second copy of all of the information, all in the Latin alphabet, is
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essential. This means that, for all biological records collected in the former
Soviet Union, the database system must have, at least for the critical data
elements, three copies of all information: first, the original data; second, a
modern opinion of what that information means, stored in the language of
the owner country; third, an English language translation of that modern
opinion. In all the collaborative work done by my colleagues in Ukraine,
this daunting challenge has been faced.
Sources of information and information storage
At an early stage in collaboration, in 1994, a decision was made to use a
Nomenclatural & Taxonomic database already in existence at the former
International Mycological Institute (now CABI Bioscience). In this way, a
large amount of duplication of effort was avoided. The Nomenclatural &
Taxonomic database then stored about 30 000 different scientific names of
organisms. The number has subsequently grown to a current figure of over
450 000 names including not only fungi, but also plants, bacteria, protozoans and animals (mainly insects). The system is designed to cope easily
with all taxonomic ranks, and has a built-in warning system to alert the
user to ambiguities arising from homonyms and orthographic errors. In
most cases, the information needed to connect to the appropriate record in
this database is simply the Latin name of the organism, stripped of any
accents or hyphens.
The Nomenclature & Taxonomic database is itself linked to a Bibliographic database then containing about 3000 (now more than 40 000)
records of different published works on mycology, and it was therefore
natural to decide to use that database too. There were, however, problems.
That Bibliographic database, having been developed in western Europe,
was (and still is) rather weak in its coverage of Cyrillic alphabet publications. Furthermore, those Cyrillic language records that it holds generally
only contain either an English translation or a Latin alphabet transliteration of the original data. A small project was therefore undertaken from
1995 to 1996, generously funded by the British Council in Kiev, to establish a new Bibliographic database in Ukraine specifically for mycological
publications in the Cyrillic alphabet. That database is structurally identical
to the main Bibliographic database at CABI Bioscience, and in due course
a copy of the Cyrillic records will be shared by both systems. As a first
stage, all the bibliographic information about mycological publications in
Mikologiya i Fitopatologiya and Ukrainian Botanical Journal is being
input.
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A fully structured database for Biological Records is very complex, and
involves many different tables (Minter, 1996). Such a database is suitable
for long-term storage of data, but is rarely appropriate for the data
gathering stage. For that work, far simpler customised data-entry schemes
are needed. For collaboration in Ukraine, a feeder database intermediate
in complexity and specially designed as a general data-entry system was
adopted. Such a database permits rapid keyboarding, while ensuring that
the critical data elements are separated, thus minimising the later editorial
overhead (a range of computer programs exists at CABI Bioscience to
carry out mechanically all the commoner editorial jobs). Copies of that
feeder database were first installed in Kiev in February 1994, with the first
donated computers to reach mycologists in Ukraine. Installation of that
system marked the beginning of an intensive phase of computerising
records of fungi in Ukraine, which continues at the time of writing.
A computerised checklist of the fungi of Ukraine
The priority of the first collaborative work, a UK Darwin Initiative project
(Fungi of Ukraine, 1993–1996), was to computerise the information on
herbarium labels in fungal collections of the M. G. Kholodny Institute of
Botany in Kiev. The initial target was to digitise 24 000 records in 27
months. In the event, over 48 000 records of fungi were keyboarded, an
impressive total, given the difficulties of setting up the system, and of
abstracting and keyboarding often almost illegible data hand-written in
Cyrillic on frequently faded packets. Many of these 48 000 records were of
fungi occurring on a host plant, or some other associated organism. In the
fully structured database for Biological Records, these observations of
associated organisms are treated as separate linked records valid in their
own right. The result was that, after editing, another approximately 32 000
records of associated organisms were added to the original 48 000 fungal
records, making a total of around 80 000 biological records of all groups of
organisms from Ukraine generated by that project. These were used to
produce Ukraine’s first computerised checklist of its fungi (Minter &
Dudka, 1996).
Generation of new data
It is always important to maintain a continued flow of new observations in
addition to computerising existing records. The UK Darwin Initiative
project, the Royal Geographical Society’s first Ralph Brown Prize (for an
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expedition to the Pripyat Marshes of Ukraine) and awards from the British
Council and the UK Know How Fund have helped to keep fieldwork in
Ukraine alive. British Council funding permitted British and Ukrainian
scientists to visit Crimea in 1993. Through the Darwin Initiative project,
various collecting trips were organised for teams of Ukrainian scientists,
including the Carpathian Mountains (1994, 1995), L’viv Oblast (1995) and
Crimea (1996). The British Mycological Society paid for one Ukrainian
student to attend their Polish Foray in 1995 (incidentally providing an
opportunity to transport back-numbers of Mycological Research and The
Mycologist donated by the Society to Ukraine at the same time). Know
How Fund support enabled British and Ukrainian scientists to visit the
Ukrainian Steppe Reserve in Donetsk Oblast in 1997. The Royal Geographical Society Ralph Brown Expedition put up to fifty scientists and
students of a wide range of disciplines into the field for periods of up to two
months during summer 1998. One result was over one hundred species of
fungi new for Ukraine. Realities of life in Ukraine mean that much
fieldwork has to be done under canvas. These awards, and particularly the
Ralph Brown Prize, provided much important field equipment to Ukrainian scientists – cagoules, boots, overtrousers, tents, sleeping bags,
airbeds, cooking equipment, canoes and many other items – without which
future fieldwork would be greatly hampered.
Electronic distribution maps of Ukrainian fungi
After the 1994–1996 UK Darwin Initiative project ended, development of
the computerised databases continued. The Royal Society project (Electronic Distribution Maps of Ukrainian Fungi and Plants, 1998–2000)
enabled mycologists in Ukraine to add latitude and longitude co-ordinates
to all records for which locality information was sufficiently precise (over
35 000). For place names not included in the various available gazetteers
this was no small task. The best maps readily available in Ukraine do not
show latitude and longitude, and have place names in Russian rather than
Ukrainian. It was frequently necessary to locate a place on these maps, and
then calculate the location by reference to another more general map that
did show latitude and longitude. Under these difficult conditions more
than normal care was needed to ensure accuracy of data. In collaboration
with Dr A. H. Thomas at the NERC Institute of Terrestrial Ecology,
Bangor, simple electronic distribution maps were then produced of a
sample 75 species, and these have been made available on the Internet
(:http://www.biodiversity.ac.psiweb.com/9).
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Greatly extended computerisation of fungal records
With the new UK Darwin Initiative award (Biodiversity Information in
the Former Soviet Union, 1999–2002), it has been possible to extend the
work computerising the fungal collections in the herbarium of the M. G.
Kholodny Institute of Botany. Attention is now being directed to the
larger basidiomycetes which, for various operational reasons, could not be
prioritised during the first project. Work is also beginning to computerise
fungal records in the reference collection of the Nikita Botanic Garden
(Yalta), using this opportunity to issue proper accessions numbers for the
first time. Now that their bibliographic information has been computerised, work is also under way extracting records of fungi from
Mikologiya i Fitopatologiya and Ukrainian Botanical Journal. For this
work, a policy decision was made to extract all records, and not just those
from Ukraine. To date, about 7000 fungal records have been computerised
from these sources, but the final total is likely to run to several tens of
thousands. When this work is complete there will be an enormous resource
of information about fungi from all parts of the former Soviet Union.
The new UK Darwin Initiative project is also, incidentally, permitting
our team to extend computerised biodiversity coverage in other ways,
some reaching beyond Ukraine. The already published checklist of the
fungi of Georgia (Nakhutsrishvili, 1986) is being computerised, with nearly 17 000 records already keyboarded, and collaborative work has begun
computerising mycological herbaria in Russia and Kazakhstan. During
2000, Ukrainian mycologists expect to help Georgian and Armenian mycologists to begin computerising fungal collections in the Botanical Institutes of Tblisi and Yerevan. Finally, and important for Ukrainian fungal
conservation, this project is helping with the publication of a new edition
of the computerised checklist of Ukrainian flowering plants.
A new directory of scientists
Anyone wishing to improve infrastructure in the former Soviet Union is
spoilt for choice. The trick is to select a problem that can be resolved within
a practical time frame. At present there is no up-to-date directory listing
names and addresses of people and institutions working on biodiversity
and nature conservation in the former Soviet Union. Such a directory is
particularly necessary because the ten years since the fall of the Soviet
Union have been chaotic: new countries have come into existence, and
with them new governments, new ministries, new academies and new
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administrative systems; many cities, towns, villages and streets have
changed their names; people have emigrated or moved in response to
economic and social pressures, and sometimes even to war; new technologies, such as fax, e-mail and the Internet have become widespread. The old
Soviet directories are no longer usable. This is one of the biggest infrastructure problems for this area of science, but it is also a problem that can be
resolved. Ukrainian mycologists at the M. G. Kholodny Institute of
Botany, in collaboration with biologists at the N. K. Koltsov Institute of
Developmental Biology in Moscow, are playing a major role in the development of a new computerised directory.
This work is funded by awards from the UK Darwin Initiative (Biodiversity Information in the Former Soviet Union, 1999–2002), and the
European Union organisation INTAS (Infrastructure Action for the Life
Sciences, 2000–2002). The first version of the new directory will use the
Latin alphabet (English language), with the aim of making it easier for
outsiders interested in biodiversity and nature conservation to establish
contact with key people in former Soviet Union countries. Subsequently
one or more versions of the directory using the Cyrillic alphabet will be
distributed within the former Soviet Union, to enhance internal contacts.
A meeting will be held in 2002, probably in Ukraine, under the auspices of
BioNET INTERNATIONAL. That meeting will draw together key
people, particularly those working on neglected groups such as invertebrates, fungi and micro-organisms, with the aim of establishing a new
general infrastructure for biodiversity and nature conservation work within the former Soviet Union.
14th Congress of European Mycologists
The acceptance of Ukraine’s bid to host 14CEM in Yalta (Crimea) in 2003
is a major boost to the country’s mycologists. It will help them enormously
in developing infrastructure for mycology in Ukraine and, with the traditional emphasis of the Congress on conservation, they will be stimulated to
formulate a new national strategy for fungal conservation. The congress
will put Ukraine at the centre of European mycology for three years, at a
time when fungal mapping and fungal conservation at a European level are
getting off the ground, and when a team appointed by the 13th Congress is
looking at the possibility of a European Mycological Association. The
influx of substantial numbers of western European mycologists to the
congress will permit mycologists from Ukraine (and other former Soviet
Union countries) to establish new collaborative networks, while the or-
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ganisation of the congress itself will require development of new skills with
e-mail and the Internet, and a much greater facility with western European
languages. Furthermore, the western mycologists visiting Crimea will gain
new insights into the mycological treasures so worth conserving in
Ukraine. The biggest challenge will be to find the funding necessary to
bring in the new computers, printers, photocopiers, modems and other
Internet paraphernalia which the organisation of this congress will require.
Plans for the future
As Ukrainian mycologists adapt and survive in the new post-Soviet environment, a trend can be detected for them to become more adventurous.
The model of Spain, an isolated country on the death of Franco but now
fully integrated into Europe, and the impressive achievements of Spanish
mycologists provide them with hope that Ukraine’s larger infrastructure
problems will be resolved, and with inspiration that at the same time they
can grow. They realise that broader issues can now be identified and
tackled, because of the solid informational base which now exists, and
because they have considerably more experience of collaborating with
westerners and approaching donor organisations. This means the range of
areas in which they are prepared to work to conserve fungi is increasing,
and there is lively debate about which problems are now the highest
priority: their ideas are taking on the character of a manifesto.
As everywhere in the world, the loss of habitat is probably the most
significant threat to fungi in Ukraine, and again that means broader
infrastructure issues. At the time of writing, the team is involved with
various proposals, including: emergency work to reduce catastrophic
flooding in Volyn Oblast (based on experience and information gained
during the Ralph Brown expedition); introduction of controlled grazing to
prevent reversion of steppe to scrub (steppe is one of the most endangered
habitats in Ukraine); a publicity campaign to discourage illegal mass
gathering and selling of wild flowers. It seems likely that there will also be
future interest in at least some of the following measures: developing a
specialist industry in sustainable tourism (an award from the British
Council is currently helping part of the team to look at possibilities in this
area); protecting Crimea’s beautiful steppe meadows, monitoring the effects of new housing schemes; the damage done to fungi by pollution; the
role of fungi in pollution monitoring, and bioremediation of pollution
(particularly relevant with regard to radioactive pollution after Chernobyl).
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As more computers reach different biosphere reserves, nature reserves
and national nature parks, there will be also an increased emphasis on
helping those organisations to develop management plans that take into
account conservation of the fungi. In this work, Ukrainian mycologists
will be looking to their western colleagues from whom they will need
considerable support. They will also try to use their computerised data and
other information to identify sites of special scientific interest for mycology, and will seek with other biologists to encourage the Ukrainian military to recognise the conservation value of some of their training grounds.
Funds will be sought to help the country’s two great fungal culture
collections (in the M. G. Kholodny Institute of Botany, and the D. K.
Zabolotny Institute of Microbiology and Virology) to survive and grow.
Every effort will be made to ensure that conservation nongovernmental
organisations, like the National Ecocentre of Ukraine, are supported and
in turn support and publicise fungal conservation. In the midst of these
simultaneously daunting and exciting opportunities, two particular
courses of action are receiving detailed attention.
Red lists, grey lists, green lists
As information becomes available in computerised format, many exciting
prospects are opening up for fungal conservation in Ukraine. It is already
possible to assess Ukraine’s known fungi by taxonomy, to identify which
groups are under-recorded. It is possible to pick out all taxa with fewer
than a specified number of records, or taxa with a most recent date of
observation earlier than, say, forty years ago. Such mechanical analyses
are essential in preparing new red data lists according to IUCN criteria,
but they can also be used as a tool to show the inadequacies of those
criteria, developed for the charismatic megafauna, when applied to fungi.
There is currently no red data book in Ukraine for fungi, but a few fungi
are present in the current edition of the red data book of Ukraine for plants
(Shelyag-Sosonko, 1996), almost all larger basidiomycetes. Now, however,
mycologists in Ukraine have a real chance to produce a meaningful red
data list of their fungi for the first time.
That list is unlikely to be a book, and probably won’t be a single list
either. It is more likely to take the form of a CD-ROM and perhaps a
website, with several different lists. These will include the conventional red
list of fungi known to be endangered, but if the IUCN criteria are used,
that list may contain so many presumed extinctions – species not observed
for more than 40 years – that some additional notation would be necess-
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ary. There will also be a list of fungi recorded in Ukraine fewer than, say,
20 times, and lists of species known to be limited in distribution. The
existence of fully computerised checklists for Ukraine of both fungi and
flowering plants, together with computerised information about fungal
plant associations and substrata, however, also permits the production of
lists of Ukrainian plants on which no fungi have been observed, and lists of
Ukrainian plants for which, for example, fungi have only been observed on
leaves, but not on twigs, branches, trunks, wood, roots, flowers etc. These
lists – grey lists of the unknown rather than red lists of the known – will for
the first time allow reasonably precise indications about what remains to
be explored. Ukraine is unusual in that it already has lists (known within
Ukraine as ‘Green Lists’) of endangered plant communities. Since fungal
conservation is so tied to habitat conservation, these plant community lists
will be another important factor in future in assessing the status of the
country’s fungi.
A proposal is now in preparation to produce that CD-ROM in time for
14CEM. In addition to the various lists, the disk would also contain over
5000 distribution maps, one for every fungal species known from Ukraine,
building on the earlier Royal Society project. Although in most cases the
amount of information is insufficient for a real concept of the species’
geographical distribution, this product will provide a base line for future
recording work. Work is also beginning to develop one or more associated
websites, to raise the profile of Ukrainian fungal conservation internationally. Ukrainian mycologists are aware that ‘Red Lists’, of whatever
colour, are political documents, and care will be taken to ensure that any
resulting CD-ROMs will produce the best possible political outcomes. One
of the targets may be a revised Red Data Law for Ukraine taking more
account of the need to conserve fungi.
A mycological society for Ukraine?
There are various scientific societies in Ukraine dealing with different
groups of organisms. Comparison of these societies with analogous bodies
in, for example, the European Union is instructive. Scientists from outside
Ukraine might find the practice in certain societies of having the same
person president for many years with no bar on re-election a little unusual.
In the west a regular change at the top is considered desirable to prevent
stagnation and the development of undesirable patronage. There is no
scientific society in Ukraine specifically for the study of fungi. Mycologists
(including lichenologists) tend to be members of the Ukrainian Botanical
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Society, which has a section devoted to Plant Pathology. With the fungi
now universally accepted as having at least one kingdom of their own,
perhaps the time is coming to establish a new and separate mycological
society in Ukraine, run on modern lines, and free from the historical
constraints which continue to affect other already existing scientific societies in Ukraine.
Such a society could play an important role in fungal conservation, even
with limited funds. This could be realised through the development of field
meetings, workshops, lectures, displays, fostering of local groups, recording and mapping schemes, an educational programme (including pressure
to raise the profile of mycology in schools and tertiary education), posters,
publicity, modernised publications, organised fund-raising (perhaps
through an encouraged amateur membership), websites and all the other
tools so successfully used by other learned societies outside Ukraine. A
new mycological society for Ukraine could also act as a unified voice for
Ukrainian mycology within the European and international scientific communities, and would probably be able to attract significant grants at
establishment stage from European Union bodies fostering the development of democratic institutions within former Soviet Union countries.
Acknowledgements
None of the work for fungal conservation in Ukraine described in this
chapter would have been possible without generous support from a range
of people and organisations. The National Academy of Sciences of
Ukraine has always supported our work directly or indirectly, particularly
through the M. G. Kholodny Institute of Botany, the country’s principal
institution for work on fungal biodiversity and conservation and its Director, Academician K. M. Sytnik. Awards from the UK Darwin Initiative,
the European Union organisation INTAS, the Royal Geographical Society’s Ralph Brown Award, the British Council in Kiev, the Royal Society
of London, the UK Know How Fund and the British Mycological Society
have provided the financial foundations for the work. Donations of computing and other equipment, particularly from CAB International, the
NERC Institute of Terrestrial Ecology Bangor, Richard Ellis & Co. and
the National Trust, and assistance in transporting that equipment from
SAS, British Airways, Ukrainian International Airlines and Zoloti Kolesa
have made the seemingly impossible seem easy. The staff of the Embassy of
Ukraine in London is thanked for their kind co-operation. The enthusiasm
of all members of the ‘Darwin Team’ in Kiev, and their families, is
Conservation in Ukraine
175
gratefully acknowledged. Lastly the author expresses his gratitude to his
employers for their support, and to his family for their toleration of the
piles of second-hand computers which have periodically invaded the
house.
References
Dudka, I. A. & Burdiukova, L. I. (1996). Oomitsety: Fitoftorovye i albugovye
griby [Oomycetes: Phytophthoraceae Pethybr. and Albuginaceae Schröt.].
Flora Gribov Ukrainy [Flora of the Fungi of Ukraine], 208 pp. Naukova
Dumka: Kiev.
Heluta, V. P. (1989). Muchnistorotsyanye griby [Powdery mildew fungi]. Flora
Gribov Ukrainy [Flora of the Fungi of Ukraine], 256 pp. Naukova Dumka:
Kiev.
Kopachevskaya, E. H. (1986). Likhenoflora Kryma i yeye Analiz [The Lichen
Flora of Crimea and its Evaluation], 296 pp. Naukova Dumka: Kiev.
Makarevich, M. F., Navrotskaya, I. L. & Yudina, I. V. (1982). Atlas
Geograficheskogo Rasprostraneniya Lishainikov v Ukrainskikh Karpatakh
[Geographical Atlas of the Distribution of Lichens of the Ukrainian
Carpathians], 403 pp. Naukova Dumka: Kiev.
Merezhko, T. A. (1980). Sferopsidal’ny e griby [Sphaeropsidalean fungi]. Flora
Gribov Ukrainy [Flora of the Fungi of Ukraine], 208 pp. Naukova Dumka:
Kiev.
Merezhko, T. A. & Smik, L. V. (1991). Diaportal’nye griby [Diaporthalean
fungi]. Flora Gribov Ukrainy [Flora of the Fungi of Ukraine], 216 pp.
Naukova Dumka: Kiev.
Minter, D. W. (1996). Recording and mapping fungi. In A Century of Mycology
(ed. B. C. Sutton), pp. 321–382. Cambridge University Press: Cambridge.
Minter, D. W. & Dudka, I. O. (1996). Fungi of Ukraine, a Preliminary Checklist,
361 pp. International Mycological Institute: Egham, & M. G. Kholodny
Institute of Botany: Kiev.
Morochkovskiı̈, S. F., Radzievskiı̈, G. G., Zerova, M. Ya., Dudka, I. O.,
Smits’ka, M. F. & Rozhenko, G. L. (1971). Nezaversheny gribi [Imperfect
fungi]. Visnachnik Hribyv Ukraini [Handbook of the Fungi of Ukraine] 3,
1–696. Naukova Dumka: Kiev.
Morochkovskiı̈, S. F., Zerova, M. Ya., Dudka, I. O., Radzievskiı̈, G. G. &
Smits’ka, M. F. (1967). Slizoviki (Myxophyta), Hribi (Mycophita):
Arkhymytseti, Fykomytseti [Slime moulds (Myxophyta); Fungi
(Mycophyta): Archymycetes, Phycomycetes]. Visnachnik Hribyv Ukraini
[Handbook of the Fungi of Ukraine] 1, 1–255. Naukova Dumka: Kiev.
Morochkovskiı̈, S. F.; Zerova, M. Ya.; Lavitskaya, Z. G.; Smitskaya, M. F.
(1969). Askomytseti [Ascomycetes]. Visnachnik Hribyv Ukraini [Handbook of
the Fungi of Ukraine] 2, 1–518. Naukova Dumka: Kiev.
Nakhutsrishvili, I. G. (1986). Flora Sporovykh Rasteniy Gruzii [Flora of
Sporulating Plants of Georgia], 888 pp. Metsniereba: Tblisi.
Oxner, A. M. (1937). Viznachnik Lishainikyv URSR [Handbook of Lichens of the
Ukrainian SSR], 341 pp. Institute of Botany, Academy of Sciences of the
Ukrainian SSR: Kiev.
Oxner, A. M. (1956). Flora Lishainikyv Ukraini [Flora of the Lichens of Ukraine]
1, 1–495. Naukova Dumka: Kiev.
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Oxner, A. M. (1968). Flora Lishainikyv Ukraini [Flora of the Lichens of Ukraine] 2
(1), 1–500. Naukova Dumka: Kiev.
Oxner, A. M. (1993). Flora Lishainikyv Ukraini [Flora of the Lichens of Ukraine] 2
(2), 1–544. Naukova Dumka: Kiev.
Shelyag-Sosonko, Y. R. (1996). Chervona Kniga Ukraini: Roslinniy Svit [Red
Book of Ukraine: Plant Kingdom]. Edition 2, 668 pp. Vidavnitvo ‘Ukrains’ka
Entsiklopediya’ im. M.P. Bazhana: Kiev.
Smik, L. V. (1980). Sferial’nye griby [Sphaerialean fungi]. Flora Gribov Ukrainy
[Flora of the Fungi of Ukraine], 184 pp. Naukova Dumka: Kiev.
Smitskaya, M. F. (1980). Operkulyatnye diskomitsety [Operculate discomycetes].
Flora Gribov Ukrainy [Flora of the Fungi of Ukraine], 224 pp. Naukova
Dumka: Kiev.
Smitskaya, M. F. (1991). Gipokreal’nye griby [Hypocrealean fungi]. Flora Gribov
Ukrainy [Flora of the Fungi of Ukraine], 89 pp. Naukova Dumka: Kiev.
Smitskaya, M. F., Smik, L. V. & Merezhko, T. A. (1986). Opredelitel’
pirenomitsetov USSR [Handbook of the Pyrenomycetes of the Ukrainian
SSR], 364 pp. Naukova Dumka: Kiev.
Wasser, S. P. (1980). Basidiomitsety. Agarikovye griby [Basidiomycetes:
Agaricalean fungi]. Flora Gribov Ukrainy [Flora of the Fungi of Ukraine], 326
pp. Naukova Dumka: Kiev.
Wasser, S. P. (1992). Basidiomitsety. Amanital’nye gryby [Basidiomycetes:
Amanitalean fungi]. Flora Gribov Ukrainy [Flora of the Fungi of Ukraine],
186 pp. Naukova Dumka: Kiev.
Zerova, M. Y., Morochkovskiı̈, S. F., Radzievskiı̈, G. G. & Smits’ka, M. F.
(1971). Basidyomytseti: Dakrymitsetal’ni, Tremelal’ni, Aurykularyal’ni,
Sazhkovn’dni, Irzhasti [Basidiomycetes: Dacrymycetales, Tremelales,
Auriculariales, Ustilaginales, Uredinales]. Visnachnik Hribyv Ukraini
[Handbook of the Fungi of Ukraine] 4, 1–316. Naukova Dumka: Kiev.
Zerova, M. Y., Radzievskiı̈, G. G. & Shevchenko, S. V. (1972). Basidyomytseti:
Eksobasydyal’ni, Afyloforal’ni, Kanteral’ni [Basidiomycetes: Exobasidiales,
Aphyllophorales, Cantharellales]. Visnachnik Hribyv Ukraini [Handbook of
the Fungi of Ukraine] 5 (1), 1–240. Naukova Dumka: Kiev.
Zerova, M. Y., Sosin, P. E. & Rozhenko, G. L. (1979). Basidyomytseti:
Boletal’ni, Strobilomitsetal’ni, Trikholomatal’ni, Entolomatal’ni,
Russulal’ni, Agarykal’ni, Gasteromitseti [Basidiomycetes: Boletales,
Strobilomycetales, Tricholomatales, Entolomatales, Russulales, Agaricales,
Gasteromycetes]. Visnachnik Hribyv Ukraini [Handbook of the Fungi of
Ukraine] 5 (2), 1–566. Naukova Dumka: Kiev.
15
The threatened and near-threatened
Aphyllophorales of Finland
H. K O T I R A N T A
Introduction
The total area of Finland is 337 000 km. The land area is 305 000 km and
65.8% (about 200 000 km) of this is wooded. Most of the aphyllophoroid
fungi are wood-decayers, saprotrophs or parasites, and only a handful can
form mycorrhizas.
Altogether 848 aphyllophoroid taxa were evaluated according to the
‘IUCN Red List Categories’ (Anon., 1995), with a national application. It
means that the numbers used in different criteria (see below) on the
national level are smaller than in the original IUCN criteria.
Categories
The following categories were used: EX (Extinct), CR (Critically Endangered), EN (Endangered), VU (Vulnerable), NT (Near Threatened), DD
(Data Deficient), LC (Least Concern) and NE (Not Evaluated). In the
tables that follow, the species which are near threatened (NT) are considered also, even though they are not threatened (yet?) in the strict sense of the
IUCN criteria.
Criteria
The criteria (which are fully described in Anon., 1995) that have been used
in different threat classes are briefly as follows; they have been applied as
recommended by the 40th meeting of the IUCN Council, except in class
NT, which is a IUCN category for a population that has diminished by
more than 10% during ten years or three generations:
177
178
H. Kotiranta
Table 15.1. Aphyllophoroid taxa in different threat categories
Categories
Aphyllophoroid taxa
Percentage value
EX
CR
EN
VU
NT
DD
LC
NE
5
0.6
22
2.6
31
3.7
62
7.3
75
8.8
171
20.1
478
56.4
4
0.5
A, reduction of population (observed, estimated, inferred or
suspected);
B, distribution area (small, strongly fragmented, etc.);
C, small and continuously declining population;
D, very small and restricted population, which is then
subdivided into subcategories on the basis of population size
or distribution area thus: D1 comprises CR, less than 50
individuals; EN, less than 250 individuals; VU, less than 1000
individuals; and NT, less than 5000 individuals; whilst in D2
the distribution area of the population is not more than CR,
:5 km or one area of occupancy only; EN, :50 km or two
areas of occupancy only, VU, :100 km or 3–5 areas of
occupancy only; and NT, :500 km or 25 areas of occupancy
only.
Results
Table 15.1 shows that the threatened categories, EX, CR, EN and VU,
together comprised 120 species, which represented 14% of the total taxa.
The few species that were ‘Not Evaluated’ (category NE) all grow inside
some sort of building, such as Serpula lacrymans (in dwellings) or
Gloeophyllum trabeum (found in saunas), and are not found in natural
habitats in Finland. The group described as Data Deficient (DD) is relatively large. There are tens of species which have been found once or only a
few times. Their right place, at least for the moment, is in class DD rather
than in one of the threat classes, such as CR. Some newly described species
are classified in DD also. In the ‘Near Threatened’ class (NT) are species
which do not fulfil the usual criteria of vulnerable species. Many of the taxa
in class NT are old-growth forest species.
Of the criteria used in defining the different classes (Table 15.2), criterion
A (reduction of population) is often used in the less-threatened classes
(VU, NT). These species have their optimum in virgin forests. The reduc-
179
Aphyllophorales of Finland
Table 15.2. The criteria used in different classes (note: all the criteria are
tested, and several criteria per species are used if they fulfil them)
% in each threat category
IUCN criterion
CR
EN
VU
NT
A (reduction of population)
B (distribution area)
C (small and continuously declining population)
D (very small and restricted population)
22.7
40.9
13.6
100
38.7
25.8
35.5
87.0
45.2
46.8
12.9
74.2
53.3
5.3
4.0
72.0
Table 15.3. Frequency of instances in which only one criterion is fulfilled in
different classes
Proportion in each threat category
IUCN criterion
A (reduction of population)
B (distribution area)
C (small and continuously declining
population)
D (very small and restricted population)
CR
EN
VU
NT
0/22
0/22
0/22
0/31
0/31
0/31
11/62
1/62
0/62
19/75
0/75
0/75
11/22
11/31
10/62
29/75
tion of the old-growth forests over three generations has greatly increased.
In Finland, the age of a generation of most wood-rotting fungi that infest
large trunks only is concluded to be 25 years, even longer in many cases.
Criterion D (very small and restricted population) is used in all of the
Critically Endangered species (Table 15.2). The high percentage even in
class NT shows that the listed species have small populations and the
number of individuals is not very high. In this study one tree trunk which is
occupied by a mycelium and bears fruit bodies of one species is considered
to be one individual, regardless of how many fruit bodies there are on the
trunk. This is, of course, not the true situation for every species (e.g.
Trametes spp.), but it is the only practical way to solve the problem of
defining an individual at the moment without extensive in vitro studies.
The frequency with which only one criterion is fulfilled in the different
classes is shown in Table 15.3.
Table 15.4 shows the primary threats in different classes. The greatest
threats for the Finnish Aphyllophorales are the changes in age structure of
the forests (reduction of virgin forests) and the reduction of decaying
wood. It is noteworthy that none of the species is threatened because of
180
H. Kotiranta
Table 15.4. The primary threats in different classes
% in each threat category
Primary threat
Changes in age structure of forests/forest
stands (reduction of primeval forests, giant
trees and burnt areas)
Reduction of decaying wood (e.g. hollow or
decayed trees)
Plantation and forest management activities
Changes in tree species composition
(domination by spruce and reduction in
deciduous tree stand)
Construction: towns, rural areas, shores, road
building etc.
Construction of waterways
Peatland drainage for forestry and peat
harvesting
Reason/cause unknown
EX
CR
EN
VU
NT
0
41.0
48.4
46.8
52.0
60
27.3
29.0
24.2
26.6
0
40
13.6
13.6
6.5
0
11.3
8.1
8.0
8.0
0
4.5
12.9
6.4
2.7
0
0
0
0
3.2
0
0
1.6
0
0
0
0
0
1.6
2.7
Note: In very small populations the small size is already a serious threat, but it is
used only if no other threats are known.
Table 15.5. The primary habitats in different classes
% in each threat category
Primary habitat
EX
CR
EN
VU
NT
Old heath forests
Heath forests
Old herb-rich forests
Herb-rich forests
Lake shores and river banks
Parks, yards and gardens
0
100
0
0
0
0
40.9
13.6
4.5
36.4
0
4.5
45.2
6.5
3.2
32.2
9.7
0
46.8
8.1
11.3
29.0
1.7
1.7
48.0
29.3
5.3
13.3
0
1.3
Note: Only the most common habitats are mentioned, habitats of one species
only are excluded.
picking or collecting or by chemical disturbance like atmospheric pollution.
The overwhelming majority of threatened species (also those categorised
as near threatened) are forest species and the most common habitat is
old-growth heath forest (Table 15.5) with Norwegian spruce (Picea abies),
pine (Pinus sylvestris), birches (Betula pendula, B. pubescens) and aspen
trees (Populus tremula). The most important host is spruce, followed by
pine and aspen.
Aphyllophorales of Finland
181
Future
During the last ten years an intensive survey of old-growth forests took
place in Finland. In the survey the most valuable areas were investigated.
To evaluate the conservation value of forests several criteria were used,
including the amount, quality and continuity of dead wood, and some
polypores were used as indicators of primeval forests.
The survey led to protection of large areas of old-growth forests. Moreover, the National Parks, Strict Nature Reserves and areas otherwise
protected by law nowadays form a good network of old-growth forests.
The already existing protected areas and those that are in different nature
conservation programmes (e.g. Natura 2000) now cover 714 300 ha. If less
productive forests are also included (mires with pines etc.) the protected
area is over 1.5 million ha. However, most of the protected areas are
situated in the northern part of the country, and the area protected in
southern Finland is still too small. The situation of the most threatened
Aphyllophorales is now good in northern Finland, but needs further
action in southern Finland.
Some of the species (mostly in the threat classes CR and EN) are classed
as Specially Protected. If some of these species are found outside a protected area the management (forest cutting, road building etc.) of the area
is interrupted until estimations of the management needed to protect the
species is established.
The new forest cutting recommendations, that some amount of wood
(often old, large or already dead trees) is always left in the managed area,
may help some species to survive even in commercially managed forests.
Reference
Anon. (1995). IUCN Red List Categories. IUCN: Gland, Switzerland.
16
Fungal conservation in Cuba
D. W. M INTER
Introduction
Cuba, the largest island in the Caribbean, is in many ways a remarkable
place. Its location gives it this special character. The country lies just inside
the tropics, in the Greater Antilles archipelago, between the Dominican
Republic and the Yucatan Peninsula of Mexico, with Florida to the north.
The island’s main crops are sugar cane and tobacco, and much of the land
is devoted to agriculture. There are also, however, many areas of unspoilt
and outstanding natural beauty. These include the mogotes (limestone
hills) in the west, the huge expanses of the Ciénaga de Zapata mangrove
swamps in the south, the Sierra Maestra range in the east, and numerous
coral reefs and small islands around the main island. At its western end, the
island’s flora and fauna contain elements from North America, for
example native oaks. Further east the mix of organisms gradually becomes
truly Antillean. Cuba is, furthermore, sufficiently isolated from other
landmasses to ensure that about 50% of its approximately 6500 native
plant species are endemics, making it the region’s principal centre of
speciation.
The human population of about 11.2 million is of mixed racial origin,
Spanish speaking and partly urbanised. The country’s natural trading
partner is the USA and, before 1959, Cuba was for many years virtually a
colony of the US. Since the revolution, however, Fidel Castro’s government has maintained a policy of genuine independence from the USA, with
strong backing from the Soviet Union: his famous foreign minister, Che
Guevara, organised the equally famous barter of Cuban sugar for Soviet
oil. The USA responded with an economic blockade, which has lasted to
today, and is still being tightened. The blockade is unilateral, and regularly
opposed by most members of the United Nations, including the UK and
other countries of the European Union. The revolution, while criticised in
182
Conservation in Cuba
183
some quarters, resulted in phenomenal improvements to education and
public health. Cuba now has a highly educated workforce, with low levels
of infant mortality, excellent life expectancy statistics, and an infrastructure unparalleled in any other country with a similar level of GNP.
The collapse of the Soviet Union, however, removed an important
economic prop and, combined with the blockade, resulted in significant
hardship for the civilian population over the last decade. In response to
that hardship, which was known in Cuba as ‘the special period’, various
initiatives were launched, particularly in biotechnology and tourism. Certain economic adjustments were also introduced which removed some
restrictions on the handling of foreign currency. These have resulted in
considerable trade growth in recent years with Canada, Mexico and the
European Union, and some easing of the economic hardship. They have
also made some international scientific collaboration easier.
In terms of mycology Cuba is no less remarkable. The earliest fungal
explorations were made in the mid nineteenth century by the British and
French, followed by considerable work at the end of the nineteenth century and the beginning of the twentieth century by mycologists based in the
USA. Large numbers of early and significant Cuban fungal specimens
therefore reside in reference collections outside the country, particularly in
North America. Over the last 40 years, the island has produced a string of
world-class scientists, trained largely in the former East Germany and
former Czechoslovakia but also in Canada and Russia. Their publications,
and those of their mentors, particularly Kreisel (1971a,b) and Arnold
(1986) have contributed greatly to our knowledge of Caribbean fungi.
These mycologists have also organised impressive, large and fully functional herbaria and culture collections within Cuba, and it can be reasonably argued that no other country in Latin America is better equipped to
carry out serious work on fungal biodiversity.
Under the very difficult conditions of the US blockade, the same scientists were instrumental in organising the first two Latin-American Congresses of Mycology, thereby effectively starting the Asociación
Latinoamericana de Micologı́a. My participation in the second of those
congresses, in Havana in 1996, gave me the stimulus to propose a project
through the UK Darwin Initiative with the objectives of preparing a
computerised checklist of the fungi of the Caribbean, putting into place the
infrastructure for a regional fungal identification service, and drawing up
proposals for national fungal conservation strategies for two Caribbean
countries, one of them Cuba. This three-year project began in April 1997.
Proposals for a Cuban national fungal conservation strategy are being
184
D. W. Minter
produced largely in collaboration with mycologists at the Instituto de
Ecologı́a y Sistemática (IES) in Havana, with additional input, particularly from the Jardı́n Botánico Nacional (HAJB) in Havana. The document
draws heavily on the computerised data collected and keyboarded for the
Caribbean fungal checklist by these two teams and other Cuban mycologists from the Instituto de Investigaciones Fundamentales en Agricultura
Tropicál (INIFAT) and the Instituto de Investigaciones de Sanidad
Vegetál (INISAV), both also in Havana. The result is already a bulky and
complex document which currently exists only in an unpublished Spanish
version (Mena Portales et al., 2000). Until adopted by the appropriate
authorities, this is not a strategy, but a series of scientific proposals. This
chapter provides a short and annotated English summary of its first part:
an assessment of the current state of mycology in Cuba. The following
specific topics are examined: threats to fungal biodiversity, current state of
knowledge by taxonomic group, habitat and substratum, fungi in protected areas, public awareness of mycology, the lack of environmental
legislation for fungi, and resources for mycology. The chapter ends with a
few notes on how fungal conservation in Cuba might develop in the next
few years.
Threats to fungal biodiversity in Cuba
Most mycologists believe that the best way to conserve fungal biodiversity
is in situ, through habitat conservation. Cuban mycologists support this
general view. In Cuba, most threats to natural habitats are much the same
as in many other parts of the world. Deforestation for agriculture, poor
farming methods, civil engineering projects such as dams, roads and urban
spread, mining and other industrial activities, environmental pollution
from fertilisers, pest control, nitrification of water and, simply, rubbish all
contribute to the destruction of habitats. Other familiar threats include
overexploitation of habitats, and illegal hunting and fishing.
A further factor in recent years has been the rapid growth of tourism in
Cuba. This has resulted in increased human pressure on key sites, such as
beaches and coral reefs, and picturesque mountain areas. Developments in
some of these areas are already causing alarm, since tourism also results in
a demand for new hotels, bigger roads, extended airports, and many
activities detrimental or potentially detrimental to ecosystems. Even activities billed as ‘ecotourism’ are often not truly sustainable. At the root of all
of these threats, arguably, is an ignorance of the true value of biodiversity.
There are, however, additional problems that may particularly affect
Conservation in Cuba
185
Cuban biodiversity. These relate to quarantine and biosecurity, particularly the wrongful introduction of exotic organisms. Apart from casual
damage done by ill-informed visitors, there is a controversial political
element to these issues. The Cuban government has repeatedly claimed
that in addition to waging economic war through the blockade, the USA
has introduced various pests and diseases as a form of biological warfare.
The most detailed allegations claim that the US government introduced
the exotic insect Thrips palmi, said to be a vector of plant viruses and an
important crop pest in its own right, by aerial spraying over Matanzas
province on 21 October 1996. Other allegations concern human dengue
fever, swine fever, rusts and smuts of sugar cane, and other exotic diseases
which were also said to have been introduced as biological warfare
(Castro, 1998). This problem is taken seriously by reputable Cuban scientists, and has been judged sufficiently important to merit inclusion in the
Cuban National Study on Biodiversity as a significant threat to the country’s biodiversity.
Current knowledge of Cuban fungi by taxonomic group
The Darwin Initiative project’s computerisation of fungal specimens in the
herbaria of IES and HAJB, together with further information derived
from the literature, culture collections and various other sources, has
resulted in a database of many thousands of records. At the time of
writing, these records are still being edited, but for the Caribbean in
general they are likely to exceed 140 000. Of these, perhaps 70 000 will
represent Cuban fungi and their associated organisms. Preliminary analysis of the records indicates that over 3870 fungal species are known from
Cuba, in nearly 2750 genera, of almost 600 families and 106 orders. These
3870 species, which include lichenised fungi, represent an impressive 5.3%
of the species already discovered world-wide, but less than 0.3% of the 1.5
million total number of species estimated by some to exist world-wide
(Hawksworth, 1991).
Within these records, all fungal phyla are represented in Cuba with the
exception of the Labyrinthulomycota and Hyphochytriomycota. It is
possible to compare actual species numbers for each group with world
estimates of species numbers for the same groups as provided by Hawksworth et al. (1995). These comparisons provide a rough estimate of how
well each group known from Cuba has been studied in Cuba. As a
percentage of known world numbers, the phyla best represented in Cuba
are the Ascomycota (9.4%), Acrasiomycota (8.3%), Zygomycota (7.5%)
186
D. W. Minter
Table 16.1. Orders of fungi that are not represented or poorly represented
in records from Cuba
Not recorded?
Poorly recorded
Ceratobasidiales
Cryptobasidiales
Cryptomycocolacales
Dimargaritales
Echinosteliopsidales
Elaphomycetales
Lahmiales
Medeolariales
Melanogastrales
Myzocytiopsidales
Neocallismastigales
Neolectales
Pneumocystidales
Protomycetales
Rhipidiales
Salilagenidales
Schizosaccharomycetales
Sclerosporales
Septobasidiales
Spathulosporales
Taphrinales
Triblidiales
Cantharellales
Cortinariales
Dacrymycetales
Diatrypales
Dothideales
Entomophthorales
Erysiphales
Exobasidiales
Gomphales
Graphiolales
Laboulbeniales
Lachnocladiales
Leptomitales
Lichiniales
Phallales
Phyllachorales
Platygloeales
Rhytismatales
Saprolegniales
Teloschistales
Thelephorales
Trichosphaeriales
Uredinales
?One or two other orders, such as the Cyttariales, with specialised distributions or
host ranges not expected to occur in Cuba, have not been listed.
and Dictyosteliomycota (6.5%), and the phyla most poorly represented are
the Basidiomycota (2.8%) and Plasmodiophoromycota (2.2%). At class
level there is no information about Protosteliomycetes and Trichomycetes,
and information about Coelomycetes remains poor. These statistics are,
however, to be used with care. For example, within the Basidiomycota
there are some relatively well-studied groups, like the Hymenochaetales
(15.5%) and Poriales (12.5%), while others are poorly represented, like the
Uredinales (0.6%). Orders of fungi that are not represented or poorly
represented in records from Cuba are listed in Table 16.1.
Current knowledge of Cuban fungi by habitat and substratum
Purely in terms of numbers of records, plant parts are the best studied
fungal substrata in Cuba, and the fungi studied on those plant parts tend to
Conservation in Cuba
187
be saprotrophs. Numbers of records can be misleading, however. In a large
number of these records, as in other countries, the identification of the
plant is inadequate: if the herbarium packet reads, for example, ‘on wood’
or ‘on bark’ or ‘on dead fallen leaves’, with no further information, then all
one can say about the associated organism is that it is a plant. By implication, the identification is only to Kingdom level. Even for many large,
common and abundant vascular plants, therefore, if the substratum is
difficult to identify at the time of collection, the level of knowledge remains
generally poor.
There are, furthermore, biases in the records of fungi on plant parts.
Most information is about fungi on leaves, twigs, stems, bark and wood,
with virtually no information about fungi on roots, buds, flowers, fruits or
seeds. Furthermore, most information about fungi on cultivated plants,
and other plants of economic importance, such as forest trees, relates to
parasitic species: saprotrophic fungi on these plants are almost unstudied.
For other forest and scrub plants information about saprotrophic fungi
tends to be better, but there is less information about parasitic fungi. Many
wayside and ruderal plants, or plants of special ecosystems, are not studied
at all. In a country where 50% of the native flora is endemic, the biodiversity implication is clear: the groups of plants which are least studied, and
usually totally unstudied, are the endemics and those which are threatened.
A further bias can be detected in the groups of fungi recorded from
Cuban plants. Most are Hyphomycetes, or from the families Coriolaceae,
Meliolaceae, Hymenochaetaceae and Lentinaceae. That bias extends to
known information of fungi on endemic and threatened plants, where
most of the little available information relates to Hyphomycetes and the
Coriolaceae. This probably only means that mycologists with particular
interests have been rather active. The splendid work carried out in recent
years on fungi of the royal palm, Roystonea regia, shows how many
unusual fungi potentially could be found on such endemics (e.g. Mercado
Sierra, 1983).
Some mutualistic fungi, such as VA mycorrhizal fungi, are relatively
well studied, but there is much less information about others, such as
ectomycorrhizal fungi, and virtually nothing is known about endophytic
fungi in Cuba. The other great group of mutualistic fungi is those forming
lichens. In general, lichens remain inadequately studied in Cuba, particularly those inhabiting rocks, and there is a need for many more collections
throughout the country, from different habitats. With the exception of a
very few records of species colonising basidiomycetes, fungi parasitic on
lichens or on other fungi are almost unrecorded from Cuba.
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D. W. Minter
Given Cuba’s excellent recent record in medicine, fungi of clinical
importance are well studied, with about 85 species of 36 genera recorded
on Homo sapiens in the country. Much work is being carried out, not least
with opportunistic species associated with AIDS, and particularly at the
Pedro Kouri Institute in Havana. Some information is available about
fungi allergenic to humans (and other animals), particularly through studies of airborne fungi, but this ecological group needs further study in
Cuba, particularly in the interior of the country. Little remains known
about many mycoses associated with work conditions, or of environmental origin, or about mycotoxicoses, and Cuban scientists feel it is necessary
to intensify techniques for rapid detection and control of opportunistic
mycoses.
Knowledge of fungi that develop on other vertebrates is insufficient, and
research has so far been directed mainly towards fungi pathogenic on
domestic animals and other animals in captivity. Almost nothing is known
about fungi associated with but not pathogenic on these animals, or about
fungi with any kind of association with wild vertebrates (these include, of
course, not only mammals, but birds, fish, reptiles and amphibians). Fungi
on dung, similarly, remain practically unstudied in Cuba. There has been
some work on fungi associated with insects, particularly bees, but information about fungi on most other insects, and on all other groups of invertebrates is more or less absent.
About 60 species of fungi have been reported from Cuba as growing on
non-naturally occurring materials, as agents of biodeterioration or biodegradation. Particular studies include biodegradation of fats, degradation
of cellulose, biodeterioration of wood in roofs and of monuments, contamination of microfiches, biodeterioration of museum exhibits (particularly of skulls and other bones), and of books, journals and other documents, and corrosion of plastics, textiles and glass. These investigations are
being carried out at a range of centres including the Facultad de Biologı́a
Universidad de Havana, the Instituto de Historı́a de Cuba, the Centro
Nacional de Investigaciones Cientı́ficas and the Centro de Investigaciones
Quı́micas.
By and large, fungi growing on naturally occurring materials, or in
unusual ecosystems, remain almost or completely unstudied in Cuba.
About 50 species of phycomycetes were recorded from freshwater in the
early 1950s, and there is one work about Ingoldian fungi in foam from
Cuban waterfalls. Marine fungi are similarly poorly known, with 43 species recorded in collections of the Instituto de Oceanologı́a in Havana.
Nothing is known about fungi inhabiting rocks (‘cryptoendolithic fungi’)
Conservation in Cuba
189
in Cuba. Given their potential importance, soil-inhabiting fungi remain
remarkably little studied in Cuba. There are collections of soil fungi,
including a few thermophiles, in INIFAT. The only information about
fungi from Cuban caves, another interesting but specialised habitat, relates
to the human pathogen Histoplasma capsulatum.
Fungi in protected areas
Cuba has a national system of nature reserves and protected areas which
covers about 22% of the country’s area, and all major vegetation types.
The level of protection provided, however, varies with different categories
of reserve, and in reality, only in about 6% of the country can nature be
described as having strict protection. These areas include Natural Reserves, National Parks, Ecological Reserves, certain Natural Monuments,
Floristic Reserves, Animal Refuges, and Nature Parks. Although no information is currently available, areas held by the Cuban military, such as
training areas or other restricted areas are also likely to be of value in
nature conservation. Fungi found in these reserves and other areas, or
known to be associated with ecosystems covered by these reserves, are
likely to enjoy some sort of protection. Cuba’s superb infrastructure, in
this sense, however, is a two-edged sword: the tight legislation protecting
these ecosystems has meant that getting permission to collect and study
fungi in protected areas is often very difficult because of the sheer number
of administrative, not to say bureaucratic, hurdles which must be overcome.
Four of the reserves have been accorded ‘UNESCO Biosphere’ status.
In these reserves 950 fungal species, about 25% of all fungi known from
Cuba, have been recorded. The biosphere reserve where most fungi have
been collected is Cuchillas del Toa, while that with the smallest number of
recorded species is Peninsula de Guahanacabibes. In terms of area occupied, the best studied is Sierra del Rosario, followed by Peninsula de
Guahanacabibes, Cuchillas del Toa and Baconao, in that order. In biosphere reserves, the best-recorded fungal groups are conidial fungi (250
species), and the families Meliolaceae (63 species), Coriolaceae (55 species)
and Hymenochaetaceae (38 species). With some notable exceptions, such
as the Alturas de Banao region, information about the status of fungi in
other protected areas is almost totally lacking. The lack of material resources in many reserves frequently means that adequate local conservation strategies simply do not exist, even for vascular plants.
Although Cuba’s botanic gardens contain no areas set aside specifically
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D. W. Minter
for fungi, they do have a role to play in fungal conservation and, when
mycologists are present on the garden staff, often quite good information
is available about their fungi. Of those which have recorded fungi, the
botanic garden with most is HAJB with 171 species, mainly
basidiomycetes, and the one with fewest is the Jardı́n Botanico de
Cupeinicú, with 72 species. Some specialist botanic gardens, however, have
no information at all about their fungi, but could be interesting and
significant. These include the Orchid Garden in Soroa and the Fern
Garden in Santiago de Cuba. The fungal group best documented for
occurrence in botanical gardens nation-wide is the conidial fungi.
Public awareness
Cuba has a high general level of education. Even so public awareness of the
fungi is very low. Partly this is cultural, the Cuban population being
markedly different in this respect from those of neighbouring Guatemala
and Mexico, with their long traditions of use of fungi in religious rites. In
one recent unpublished study of 900 people within a single borough of
Havana, significant differences in public perception of the fungi were
revealed. The over 40s generally considered them to belong in the plant
kingdom, to be a nuisance, and to be associated mainly with plants. The
under 40s in comparison tended to understand that they belonged in their
own kingdom, could be beneficial, and believed their main significance was
medical. The study related these different views directly to different treatments of the fungi in schools and higher education during different periods, suggesting that for mycology education really does make a difference.
In Cuba, at present, the national environmental strategy does not adequately cover the country’s fungi. As a result, treatment of the fungi in
programmes of basic education is generally insufficient. Only in higher
education, where it meets the requirements of different specialisms, is there
proper coverage of mycology. Very few of the 174 generalist museums in
Cuba provide educational coverage of the fungi. Other organisations
combining public recreation and scientific education, such as aquaria,
botanic gardens, zoos and cultural centres, rarely or never hold exhibits
about fungi. Conservation of fungi gets little direct coverage on radio and
television in Cuba, although environmental and nature programmes make
occasional references to fungi.
Unlike in Britain, the formation of amateur and other societies is strictly
controlled by legislation in Cuba. There is therefore no well-developed
network of amateur nature clubs in Cuba, even for bird-watchers, and
Conservation in Cuba
191
effective nongovernmental organisations for nature conservation as understood in Britain are also not well developed. Amateur groups for the study
of fungi are, a fortiori, nonexistent. Scientific societies exist, which to some
extent cover mycology. These include the Sociedad Cubana de Botánica,
the Sociedad Cubana de Microbiologı́a and the Sociedad Cubana de
Fitopatologı́a. These are, however, purely for professional scientists, with
no amateur membership, and the need for societies that unite amateurs
and professionals is still not fully recognised. Furthermore, although over
140 mycologists are known to be active in Cuba, indicating a clear need,
there is still no Cuban Mycological Society.
Environmental legislation
Environmental legislation of one sort or other has existed in Cuba from the
pre-revolutionary period, and even from Spanish colonial days. There was
considerable legislative activity on this topic during the 1980s,
strengthened in the 1990s particularly after the Rio Summit. The various
laws, however, while indicating a positive attitude to nature conservation,
are rather complex and, at times, conflicting. Cuban environmental experts recognise that their implementation contains many practical problems, and there is concern, for example, that fines are generally derisory.
As far as fungi are concerned, current environmental legislation in Cuba
provides no specific coverage of their use or conservation, and no specific
legal status seems to exist for this group of organisms.
Resources
Cuba’s major mycological resource is, of course, its mycologists, but even
apart from these, the country has the best infrastructure for work in
mycology in the Caribbean and, arguably, the whole of Latin America.
There are at least 40 institutions in the country which include at least some
research on fungi in their remit (coverage includes systematics, ecology,
medical and veterinary mycology, industrial mycology, biotechnology,
plant pathology, quarantine, biodeterioration and biodegradation). Some
of these institutions have adequate material resources. In addition there is
a good network of regional laboratories and extension scientists and
technicians.
There are 17 culture collections for fungi in Cuba, including several with
specialist interests. The most important is in INIFAT, with about 8000
isolates. This remarkable culture collection, undoubtedly the largest in
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D. W. Minter
Central America, and one of the largest in the tropics, has survived and
even grown despite inadequate equipment and frequent power cuts. There
are rather more herbaria than culture collections in Cuba. The three main
collections are in IES (an estimated 20 000 specimens), HAJB (an estimated 7500 specimens) and INIFAT (an estimated 3000 specimens).
Among smaller herbaria, the collections of hyphomycetes in the Centro de
Evaluación y Conservación de Ecosistemas Terrestres de Camagüey, and
of pathogenic forest basidiomycetes in the Instituto de Investigaciones
Forestales in Havana, deserve particular mention.
The United States’ blockade, combined with the collapse of the former
Soviet Union, has resulted in significant shortfalls in material resources for
most if not all of these institutions. There are problems with obsolete
scientific equipment (particularly poor-quality microscopes), and difficulties in obtaining reagents and consumables for research. Culture collections lack equipment that, outside Cuba, would be considered essential:
freeze-drying techniques are unavailable, and cultures have to be maintained in oil, or regularly subcultured on agar, both highly labour-intensive techniques. Chemicals for fumigation of herbaria cannot be obtained
from the cheapest source (the USA) and are therefore more expensive,
more difficult, and sometimes even impossible to obtain. The result has
been the loss, over the last twenty or so years, of significant numbers of
type specimens to the depredations of insects. Expedition vehicles tend to
be old fuel-inefficient Soviet-built jeeps and trucks for which spare parts
are now largely unavailable. These, combined with a general shortage of
fuel, make access to distant places and places rich in biodiversity very
difficult. For the present Darwin Initiative project, much of the day-to-day
travel between institutes in and around Havana has used an ancient
Russian-built ‘Jupiter’ motorbike and sidecar – pleasant enough travelling
in the tropics, but disconcerting when being overtaken by energetic cyclists.
There are also severe informational problems. Institutional libraries
have great problems in maintaining up-to-date runs of even major scientific journals. Essential literature, for example catalogues like the Bibliography of Systematic Mycology, and the Index of Fungi, are incomplete,
even in the country’s main systematic institutions. Specialist literature on
particular taxonomic groups is also generally lacking. The scientific visitor
to Cuba soon encounters the resulting thirst for new literature, and rapidly
learns to pack his or her bag with books on the outward journey, usually
coming home with piles of mail which Cuban colleagues are unwilling to
entrust to their own postal services. Cuban scientists face additional prob-
Conservation in Cuba
193
lems in publishing their work. Until very recently, one distinguished
Cuban mycologist wrote all his papers using a typewriter made in 1909.
When a manuscript is successfully sent to a journal, perhaps by handdelivered mail through a returning foreign visitor, all too frequently an
editor may return the manuscript for corrections, entrusting it to conventional post without imagining the delays, difficulties, frustrations and
dismay this can cause. One superb and major mycological manuscript by
Cuban and Czech authors waited nine years before it could be published in
1998.
Computers and computing resources remain another very important
limiting factor. With the exception of certain flagship reserves, computers
are almost entirely lacking from most nature reserve offices, making it
difficult for staff there to organise their reserve records efficiently, or to
produce machine-readable reserve management plans. Access to computers is also severely limited in many research institutions, and enormous
frustrations can result from loss of computing time because of power cuts
and other shortages. Another computing and informational problem is
e-mail and Internet access. Various difficulties continue to hinder easy
access for researchers and technicians to these services.
Some possible future developments
This chapter has briefly reviewed the current state of mycology and fungal
conservation in Cuba. Although it would be premature to comment on the
proposed strategy itself, a few remarks about future directions may be
appropriate. Fungal conservation is only likely to happen if there are
people who care enough to make it happen. Those people are mycologists.
In many parts of the world mycologists are an endangered species. It
follows that fungal conservation can only occur if mycologists are conserved. That means that Cuba with its wealth of expertise is likely to become
increasingly important. There is a very good case for directing resources to
its mycologists. This is something the British, Mexicans and Spanish in
particular are already doing, but not in sufficient quantities. Cuba, nevertheless, can do much to help itself here, through the formation of a Cuban
Mycological Society. This single act would, at present, do more than any
other to forward fungal conservation in that country, and the Cuban
authorities are urged to facilitate establishment of that society.
Furthermore, assistance to Cuban mycologists is a matter that
should transcend the political rancour which has existed between the
USA and Cuba. The restrictions imposed on US scientists by their own
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D. W. Minter
administration through the blockade are having a deleterious effect on
their own important scientific projects. There is, for example, a large
US-funded project on the flora of the Greater Antilles (and this flora
includes fungi and lichens). But it is almost completely impossible for the
organisers of this project to access and include recent information about
the biggest and, in biological terms, most important island in the archipelago. The efforts being made by some scientists in the USA to repatriate
botanical data to Cuba (:http://www.ascoll.org/cuban3.html9) are to be
applauded. It is a pity that this has not yet extended to fungal records.
Although much has been done for Cuban mycology over the last three
years, an enormous amount remains to be done. Computerising the floristic records and producing a checklist is only a start. Subsequent stages
should include the production of electronic distribution maps of fungi,
Red Data lists of species known to be endangered or of limited distribution, grey lists setting out the plants, plant parts and other substrata for
which few or no fungi have been recorded, and many other uses of that
information. Further in the future, but not too distant, should be the
large-scale production of electronic keys, descriptions and expert systems,
all fully illustrated with digitised photographs. There is furthermore a clear
need for a simple guide to the identification of dead fallen leaves by shape
and size, and of dead fallen twigs and wood by anatomy observable with a
hand-lens and other field characteristics. Such a guide would surely improve the proportion of future records with an adequate identification of
the associated plant. Even if a guide like that were available, however, field
identification of dead parts of grasses, sedges and similar plants would still
usually be impractical.
The growth of tourism in Cuba also provides an urgent stimulus to
survey and assess particular areas rich in biodiversity and of outstanding
natural beauty. British mycologists are justifiably proud of a small number
of sites where fungi have been intensively recorded, such as Esher
Common or Slapton Ley. No equivalent sites exist anywhere in the tropics.
Given the splendid human resources Cuba has in terms of high calibre
systematists, not only in mycology, but also in many other areas of
biology, a good case can be made for combining such tourism impact
surveys with the establishment of a small number of such sites in the
tropics, at least for the fungi, perhaps also for one or two other groups.
This would, of course, fall far short of an all taxon biodiversity inventory,
but it could mark the first steps in that direction. Localities that spring to
mind as suitable include the mogotes hills near Viñales, the swamps of
Ciénaga de Zapata, and the reserve of Alturas de Banao, all mentioned
Conservation in Cuba
195
earlier in this chapter. There are, doubtless, many other appropriate
sites.
Conclusion
Any scientist visiting Cuba is likely to come away with a whole spectrum of
strong impressions. Among these, the high quality of Cuban science, and
the calibre, enthusiasm and dedication of its scientists who achieve wonders with minimal resources, and the well-developed infrastructure are
likely to be prominent. The Cuban National Study on Biodiversity, produced before this Darwin Initiative project began, and without any external assistance, is a profoundly impressive work. It is hard to imagine any
other country with so low a per capita GNP being able to produce such a
document unaided. The apparent ease with which Cuban colleagues were
able to pull together huge amounts of information for the much more
detailed proposed Cuban national conservation strategy for fungi was,
frankly, amazing. It must be evident that, while Cuban mycologists need
material resources, they do not need aid. They are seeking collaboration
from a strong position, and have much to contribute. It means that, in
Cuba, perhaps more than in any other tropical country in the world, there
is a real chance to carry out real fungal conservation over the next few
years.
Acknowledgements
The UK Darwin Initiative is warmly thanked for support through its
project Fungi of the Caribbean. Part of this chapter constitutes an English
language summary of parts of the proposed Cuban national conservation
strategy for fungi (Mena Portales et al., 2000), and is thus derived from the
work, thoughts and ideas of my Cuban colleagues and friends. Because I
have annotated that summary with my own comments and interpretations
(including some with which they may not agree), it has seemed better to
offer the present chapter under my sole authorship. It is therefore only
appropriate to acknowledge with thanks their contribution to this work.
References
Arnold, G. R. W. (1986). Lista de Hongos Fitopatógenos de Cuba. 206 pp. Edit.
Cientı́fico Técnica: Ciudad de La Habana, Cuba.
Castro, F. (1998). Informe central al V Congreso del Partido Comunista de
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Cuba. Granma Internacional 32 (47), 3–14.
Hawksworth, D. L. (1991). The fungal dimension of biodiversity: magnitude,
significance, and conservation. Mycological Research 95, 641–655.
Hawksworth, D. L., Kirk, P. M., Sutton, B. C. & Pegler, D. N. (1995). Ainsworth
& Bisby’s Dictionary of the Fungi. Eighth edn, 616 pp. CAB International:
Wallingford, Oxon.
Kreisel, H. (1971a). Clave para la identificación de los macromicetos de Cuba.
Ciencias Serie Ciencias Biológicas 4 (16), 1–101.
Kreisel, H. (1971b). Clave y catálogo de los hongos fitopatógenos de Cuba.
Ciencias Serie Ciencias Biológicas 4 (20), 1–104.
Mena Portales, J., Herrera Figueroa, S., Mercado Sierra, Á. & Minter, D. W.
(2000). Estrategia para la Conservación de la Diversidad Fúngica en Cuba.
Estado de Conocimiento, Estrategia y Plan de Acción. Unpublished report,
154 pp. Instituto de Ecologı́a y Sistemática: La Habana, Cuba and CABI
Bioscience: Egham, Surrey, UK.
Mercado Sierra, Á. (1983). La palma real (Roystonea regia): un sustrato idóneo
para el desarrollo de hifomicetes demaciáceos. Acta Botánica Cubana 15,
1–13.
17
Microfungus diversity and the
conservation agenda in Kenya
P. F . C AN N O N , R . K. M I B EY & G . M. S I BOE
Introduction
Microfungi are rarely considered within conservation policies for a number of reasons: they are small, poorly known (especially in the tropics),
extremely diverse, and their fruit bodies are often ephemeral. They are
frequently perceived at best as not charismatic, and at worst as threats to
other species. However, it is probably true to say that the majority of fungi
would be describable as ‘microfungi’ and many are likely to be rare and
threatened. Nevertheless, they may play important roles in the ecosystem
through positive interactions with other organisms, and they represent an
enormous range of genetic and metabolic resources. Thus, consideration
of microfungi in relation to the issues that are addressed in conservation
programmes for African animals and plants is appropriate, and represents
a valuable model for many other organism groups.
One of the principal barriers to the inclusion of microfungi as targets for
conservation is knowing whether species are genuinely rare, or simply
rarely recorded. With the assistance of the UK Government’s Darwin
Initiative, we have addressed this problem in Kenya by collecting and
studying fungal species which are associated with rare and endangered
plants, and which are likely to be host limited. We can then be confident
that the fungi are at least narrowly distributed, and are threatened to at
least the same degree as their plant hosts. We have identified a range of
biotrophic species associated with rare Kenyan plants, many of which are
being published as new taxa, and are now monitoring populations and
establishing host ranges by searching for them on related host plants. We
hope then to be in a position to introduce microfungi to the conservation
agenda, by demonstrating that the loss of particular plant species from the
Kenyan flora will result in the loss also of associated fungi. Plants are
197
198
P. F. Cannon, R. K. Mibey & G. M. Siboe
actually community microcosms on which many other organisms depend.
We hope that this realisation will lead to enhanced consideration by
conservation decision-makers in Kenya of the small-bodied but speciesrich groups such as fungi and insects.
The fungi comprise one of the most species-rich and ecologically important organism groups on Earth. Despite this, until recently they have rarely
been considered seriously in the course of in situ conservation decisionmaking, especially in tropical regions, and funding for ex situ conservation
in the form of culture collections remains inadequate (Kirsop & Canhos,
1998). The reasons for this are not difficult to discover. Firstly, the conservation movement, largely funded through voluntary donations, has historically concentrated upon large charismatic organisms with which the
general public empathises. Popular perceptions of fungi tend to revolve
around toxicity and disease, with few species considered in a positive light.
Secondly, there are significant problems in analysis and monitoring of
fungal populations, and in the sheer number of potential candidate species
for conservation. Nevertheless, there are compelling reasons to include
fungi in conservation strategy, and the project outlined here aims to
provide basic data to overcome one major obstacle, that of deciding
whether fungi which are rarely recorded are genuinely rare or simply rarely
detected.
Why are fungi difficult to conserve?
There are very large numbers of fungal species, making prioritisation
complex and time-consuming. Estimates have been the subject of vigorous
debate over the last ten years, but most agree that at least 1 million and
perhaps as many as 1.5 million species exist world-wide (Hawksworth,
1991; Rossman, 1994). Predictions of the number of species occurring in
particular localities or habitats are also difficult as fungal distribution
patterns do not necessarily mirror those of better known organism groups.
However, two protected sites in southern Britain, Esher Common and
Slapton Ley National Nature Reserve, both harbour between 2500 and
3000 recorded species with only about 40% overlap (D. L. Hawksworth, J.
A. Cooper, pers. comm.), suggesting that substantial numbers remain to
be discovered at both sites. Even at the individual sample level, species
numbers can be overwhelming, with 50–100 commonly detected from
individual substratum collections, even where the techniques used do not
result in a complete survey (e.g. Bills & Polishook, 1994; Lodge & Cantrell,
1995). There is very little reliable information which can be used to
Conservation in Kenya
199
contrast fungal diversity in tropical and temperate regions (Lodge et al.,
1995), but as a large proportion of species is host specific to some degree, it
is likely that fungal diversity shows a similar bias towards the tropics as
does the diversity of many other, better studied organisms (Hawksworth &
Kalin-Arroyo, 1995).
Fungi are also poorly known. Only an estimated 5% have received even
a rudimentary description (Hawksworth, 1991; Cannon & Hawksworth,
1995). Even of these, the vast majority has not been treated in a modern
monograph, and information that is available is inaccurate, sketchy and
often buried in inaccessible journals. Fungal classification has undergone a
series of major changes in recent years with the incorporation of lichens
into mainstream fungal systems, the continuing integration of anamorph
and teleomorph classifications, and now the molecular revolution which is
providing dramatic new evidence of relationships. These are all crucially
important advances, but in the short term the instability makes life even
more difficult for conservationists and other applied biologists. Species
concepts are poorly understood, with traditional morphology-based systems being replaced by nucleic acid-based phylogenies and biological
species concepts (Brasier, 1997). This has particular relevance for
necrotrophic plant pathogens, with great uncertainty over the extent of
host specificity in many genera (Rehner & Uecker, 1994; Cannon et al.,
2000).
Almost all fungi exist primarily as mycelium, often immersed within
host tissue, with fruit body production (currently essential for both detection and identification) ephemeral and greatly influenced by external factors. This means that it is often difficult to establish whether a given species
is genuinely absent from a location, or is simply behaving cryptically. Both
fruit bodies and colonies are extremely small in many cases, and most are
not conspicuously pigmented. Detection and subsequent monitoring is
therefore sometimes a painstaking process, involving careful microscopic
observation over an extended period.
Why are fungi important?
Fungi and bacteria constitute the dominant organism groups involved in
C/N cycling, primarily in the degradation of organic matter (Christensen,
1989). Critical stages in lignin decomposition are almost exclusively carried out by fungi, and the breakdown of complex biomolecules such as
cellulose and tannins in soils is due mostly to fungal enzymatic activity
(Cooke & Whipps, 1993). Hyphal length has frequently been estimated at
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P. F. Cannon, R. K. Mibey & G. M. Siboe
over 1 km g\ in grassland soil (Kjøller & Struwe, 1982), and fungi
account for up to 90% of total living biomass in forest soils (Frankland,
1982).
Fungi mediate plant and animal health and growth via symbiotic associations. These may be mutualistic, for example mycorrhizas, endophytes
and ant-nest symbionts (Read et al., 1992; Redlin & Carris, 1996; Chapela
et al., 1994), or antagonistic with a plethora of well-known plant and
animal diseases. They provide a food source for a wide range of vertebrates
and invertebrates including insects, mites, molluscs and nematodes (Shaw,
1992), and gain nutrition in turn from living or dead animals as well as
plants.
Large numbers of fungal species, especially saprotrophs and necrotrophic taxa, can be isolated easily into pure culture. This has led to
extensive screening for chemicals with useful pharmaceutical and industrial properties. Some of the most valuable and widely used pharmaceuticals from natural products originate from fungi, including the penicillin
antibiotics, the antifungal cyclosporins used in transplant surgery, and
recent ‘cholesterol-busting’ drugs based on zaragozic acids (Dreyfuss &
Chapela, 1994; Bergstrom et al., 1995).
How can we include fungi in conservation strategies?
Most fungal species are poorly known, if they are described at all. Even
when species are known, the difficulties of sampling and recognition mean
that it is problematical to assess whether species that have been detected
only a small number of times are really rare, or merely under-recorded.
In Kenya, we have been addressing this issue with the assistance of a
major grant from the UK Department of the Environment’s Darwin
Initiative. The approach taken is to target fungi which are likely to be host
specific, and which are associated with plant species which are themselves
rare or endangered. We can then be confident that these fungal species are
at least as rare as their plant partners are. The aim is to provide information that can be used by policy-makers in prioritising conservation activities; the project leaders have no direct influence on national and regional
environmental protection strategies. However, we are in a position to tell
those who do make the decisions that if particular plant species become
extinct in Kenya, fungal species will also be lost.
The information may also be used to prioritise protection for particular
populations of plants. The distribution of fungi associated with endangered plants may not be coincident with that of their hosts; some popula-
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201
tions may lack the associated fungi. In Kenya as in many tropical countries
pressure on native vegetation is intense due to a rapidly increasing population with burgeoning needs for farmland, timber for construction, firewood etc. Hard decisions sometimes have to be taken to sacrifice stands of
natural vegetation to meet the needs of the people at large. In those
circumstances, decisions may be taken to promote conservation of rare
plant populations which harbour endangered fungal taxa.
Most host-specific species of fungi are biotrophic or necrotrophic, gaining their nutrition from living plant tissue. The relationship with the plant
is by no means always deleterious. The fungus may compensate the plant
for providing nutrients and shelter by secreting toxic metabolites that
discourage herbivory by vertebrate or invertebrate animals; this is particularly well documented for endophytic fungi associated with grasses (Redlin
& Carris, 1996), but may be a widespread phenomenon. It is rare in natural
environments for an indigenous fungus to have a serious deleterious effect
on a native plant species (Allen et al., 1999); good parasites don’t kill their
hosts. Where significant damage does occur, it is often through the introduction of alien species, or modifications of the environment that upset the
balance between fungus and plant. Positive contributions to ecosystem
function by parasitic fungi are increasingly being considered by plant
pathologists (Browning, 1974; Helfer, 1993; Ingram, 1999), leading to calls
for their conservation. Coevolution of pathogens with wild crop hosts has
resulted in a wide range of disease resistance mechanisms, which have not
always been utilised effectively in crop breeding (Wood & Lenné, 1997;
Allen et al., 1999). Wild relatives of crop plants are therefore acquiring new
significance in world economics, and genetic manipulation systems already
allow the transfer of pathogen resistance between unrelated plants, perhaps providing new roles for genes from native plants in food production.
The need for integrated conservation action plans
Species cannot be preserved in situ in isolation. If rare or endangered
plants are to be protected, an array of associated species must also be taken
into account. A wide variety of organisms may be dependent for survival
on preservation of the plant species. These may include the following.
Biotrophic and necrotrophic fungi, bacteria, other plants etc.
directly associated with the plant.
Organisms using the plant as a platform, e.g. foliar epiphytes
and lichenised fungi on bark.
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P. F. Cannon, R. K. Mibey & G. M. Siboe
Endophytes, e.g. fungi, bacteria and viruses within plant tissues.
Root-inhabiting nematodes, protozoans, etc.
Bark and stem-boring insects, leaf-cutting ants and wasps,
leaf-miners, gall-formers, etc.
Pollinators (insects, birds, bats, etc.).
Yeasts and other organisms associated with nectar.
Fruit-eating/ingesting organisms, from mammals to bacteria.
Organisms eating, degrading or taking shelter in dead plant
material.
In some circumstances, for example mycorrhizas or pollinators, the loss
of the associated organism will cause death or at least have a serious
deleterious effect on survival of the plant. In other cases, the dependence is
only one-way, but loss of the plant will result in multiple extinctions.
Health audits of endangered plants
Many organisms do not have a beneficial effect on plants. In most cases
their prominence in natural ecosystems may be due to human intervention
in the past or present (Allen et al., 1999), such as encroaching agriculture,
logging or firewood collection, or industrial pollution. Major environmental modifications such as those associated with global warming are
likely to result in more extreme perturbations of the natural balance
between plants and other organisms.
It is valuable to perform health audits on rare or endangered plant
species, identifying species alien to their community such as plant
pathogenic fungi and bacteria spreading from neighbouring agricultural
areas, immigrating root-, stem- and bark-boring insects, or invading
weeds. It is not always easy to tackle the sources of these threats to plant
health and existence (and indirectly to the conservation of all dependent
organisms), but their identification may enable protective measures to be
taken. It is also difficult in some circumstances to decide whether fungi are
genuinely native to the region or have been introduced: the distinctions
between indigenous and alien populations may be evident only at the
genetic level. This is particularly the case with necrotrophic fungi such as
Colletotrichum, Guignardia and Phomopsis, all of which are commonly
encountered in surveys of natural habitats.
Darwin Project activities
The first step was to establish an initial list of target plants. Plant conservation research in Kenya has been centred on the National Museums in
Conservation in Kenya
203
Nairobi, and several documents are available which detail threatened taxa.
Prominent amongst these are accounts by Gillett (1979) and Luke (1991),
and much of the information was usefully summarised by Beentje (1994),
who included IUCN threat categories in his treatment of the woody plants
of Kenya, and also by Walter & Gillett (1997). Available fascicles of the
Flora of East Tropical Africa were also surveyed for endemic and/or
threatened plants. A second category of plants was also included in response to requests from the Kenyan Forestry Department, species restricted to the few remaining remnants of Guinea–Congolian rainforest in
western Kenya, centred on Kakamega Forest. This vegetation has links
with West rather than East Africa. While many of the species in the
Kakamega region have widespread distributions, their presence in Kenya
is threatened and extensive fragmentation of the forested regions west of
Lake Victoria means that populations are genetically isolated. Forests in
the Kakamega region are seriously threatened due to pressure from the
rapidly increasing surrounding population, and have been the subject of
planning for sustainable exploitation in the past (Wass, 1995). Plants from
the coastal Kaya forests of Kenya were also particularly considered, to
build on previous work by the National Museums of Kenya and the World
Wide Fund for Nature (Robertson & Luke, 1995). Kayas are forests or
woodland fragments of religious significance to local inhabitants, which
have historically been well preserved but are now in some instances being
threatened as traditional values are eroded. Finally, a small list of charismatic plant species was included, which, while not necessarily endangered,
are well known to many nonspecialists. This included the arborescent
Lobelia and Senecio species of Mount Kenya and other highland regions,
and also species of Protea with their spectacular flowers. Target species
were prioritised using the following criteria: conservation status (e.g. restricted distribution, known threats); woody habit (provides more microhabitats for fungal colonisation); systematic isolation (plants more likely
to harbour host-specific fungi); ease of recognition in the field; distinctiveness of habitat; ease of access.
The research programme has surveyed forests in a number of regions of
Kenya, including Kakamega Forest, the Cherangani Hills, Mount Elgon,
the Aberdares, Mount Kenya, the Taita and Shimba Hills, and ArabukoSokoke Forest. More information can be found in the project website,
:http://www.cabi.org/bioscience/darwin.htm9. Approximately 110
plant species have been surveyed, and each harboured at least one
identifiable fungus, with many of them being host to many species of
fungi. Surveys concentrated on living leaves to detect biotrophic
and necrotrophic Ascomycota, including the mitotic forms traditionally
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P. F. Cannon, R. K. Mibey & G. M. Siboe
referred to as hyphomycetes and coelomycetes. Some studies were also
made of fallen leaves, many fungi from which would previously have been
present in living tissues as endophytes, and of dead twigs attached to living
plants, often dying back through fungal activity.
The fungi of particular note belong to the following assemblages:
Meliolaceae (Ascomycota: Meliolales). The sooty moulds have
been the subject of a previous major study of fungi from
natural habitats in the Shimba Hills (Mibey & Hawksworth,
1997), and it is notable that a number of further new taxa
have been identified from the same region during the current
project, even after the intensive sampling which preceded the
first report. Twelve new species have been described as part of
the current project (Mibey & Kokwaro, 1999; Mibey &
Cannon, 1999), eleven of which belong to the genus Meliola
and one to Cryptomeliola (the latter on the same plant as one
of the Meliola species). There is some controversy
surrounding the host specificity of the Meliolaceae, but it
seems likely that coevolution has resulted in at least a degree
of specialisation (Mibey & Cannon, 1999).
Phyllachoraceae (Ascomycota: Phyllachorales). This family has
not been studied in detail from Kenya, but a survey during
the current project has so far identified 40 taxa, including six
new species (Cannon, in prep.). The new genus Fremitomyces
has also been described from Kenyan material (Cannon &
Evans, 1999). Most species of the family are biotrophic, and
thus generally considered to be host specific at least at the
genus level. In addition, colonies of Colletotrichum species
(sometimes with their Glomerella teleomorphs) have been
found associated with lesions on many of the rare indigenous
plants surveyed. Most are referable to the species aggregate C.
gloeosporioides, which is extremely widespread and a common
disease-causing organism of crop plants. However, there are
well-established genetic clusters within the aggregate as a
whole, some of which are clearly host specific (Cannon et al.,
2000). Regrettably, the project resources did not allow
culture and molecular characterisation of the strains from
indigenous plants, so we are unsure whether they constitute
part of the natural environment or have invaded from
elsewhere.
Conservation in Kenya
205
Asterinaceae (Ascomycota: Dothideales). Also monographed by
Mibey & Hawksworth (1997), we have amassed around 40
further collections, again including further taxa from the
Shimba Hills as well as many from the Kakamega region. A
number include only the anamorph, a part of the life cycle
which has not been studied intensively, so species concepts are
difficult to establish.
Cercospora and relatives (Ascomycota: Dothideales). Around 20
species have been recognised as undescribed, with several
already prepared for publication (Siboe, Kirk & Cannon,
1999; Siboe et al., 2000). Most are necrotrophic pathogens
found associated with leaf spots.
A very wide range of other fungi has also been studied. Some of the
better known groups of biotrophic fungi such as the Erysiphales (powdery
mildews) are not typical of forested habitats where most of the research has
been carried out. Rusts have been rarely recorded for the same reason,
though one species of Aecidium from Diospyros greenwayi and D. shimbaensis from the Shimba Hills is of particular note as it appears not to
parasitise D. kabuyeana. The first of these plant species is in the IUCN
‘rare’ conservation category, and the other two are classed as ‘vulnerable’
(Beentje, 1994). The fungus is similar but not identical to an Indian species
on other species of Diospyros. Research continues.
Conclusions
This is a distinctive research programme, which is breaking new ground in
some respects. So far as we are aware, fungi have not previously been
integrated into plant conservation strategies in this way before, either in
the tropics or in temperate regions. While we do not pretend to be able to
provide unequivocal answers during the programme, we will be able to
provide strong pointers that will facilitate the inclusion of fungal species in
protection strategies, and explore the complex relationships between fungi
and plants.
The programme charts a middle course between conservation of individual species and protection of environments. The first approach has many
drawbacks due to the complex interrelationships between target species
and other organisms, and the second provides little information about
individual species.
The project represents a small beginning, in a research area that is
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P. F. Cannon, R. K. Mibey & G. M. Siboe
potentially far-reaching. If this approach proves fruitful there is considerable potential to expand the programme to cover other plant species in
Kenya and beyond. It would also be most valuable to integrate these
studies with parallel research into other species-rich organism groups
associated with rare plants, particularly the insects. This would result in
developing knowledge of the complex food webs that underpin biodiversity, and gain much information about the component parts of threatened
ecosystems.
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18
Fungi and the UK Biodiversity Action
Plan: the process explained
L. V . F LE M I NG
Introduction
Lead partners, champions, priority species, SAPs, HAPs, and Country
Groups are just some of a bewildering array of terms, jargon and acronyms
that confront a novice to the UK Biodiversity Action Plan (BAP) (see
Table 18.1). This chapter aims to give the background to the BAP, to
explain in simple terms what is actually happening (focusing on the production of species and habitat action plans), and will consider the implications of this process for the conservation of fungi in the United Kingdom
(UK).
Background
The United Nations Conference on the Environment and Development
(the ‘Earth Summit’) in Rio de Janeiro, 1992, resulted in the widely
publicised Convention on Biological Diversity, to which the UK is a
signatory. One of the many requirements of the Convention was that
individual states produce their own action plans or programmes for the
conservation of biodiversity. Accordingly, in 1994, the UK Government
published Biodiversity: the UK Action Plan (Anon., 1994). This document
provided a summary of the state of UK biodiversity and the actions being
taken to conserve it. Most importantly, it contained in a final section a set
of ‘59 steps’ to be implemented by Government and its agencies. These 59
steps encompassed a range of activities, from the need to continue with the
programme of site designations required by European Community Directives, to education and awareness programmes, to actions for UK overseas
territories and for the limitation of UK fishing activities. Whilst these
actions were a mixture of the ongoing and the new, one step in particular
was to lead to a flurry of new activity. This step (33) reads:
209
210
L. V. Fleming
Table 18.1. Glossary of terms used in the Biodiversity Action Plan
Agenda 21
A programme for action for the 21st century, arising from the
Rio conference, to have sustainable development at its core, and
to be taken forward by Local Governments, amongst others, in
consultation with their citizens.
BIG
Biodiversity Information Group.
Champion
Any organisation or individual who wishes to support the work
of an action plan by providing resources, in cash or in kind.
Contact point Government agency, responsible for acting as initial contact for
an action plan, for stimulating action and maintaining standards
with regard to the requirements of a published plan.
Country Group The biodiversity steering group for each country in the UK.
HAP
Habitat Action Plan.
Key habitats
The name, now superseded, for those habitats for which action
plans will be/have been prepared.
LBAP
Local Biodiversity Action Plan.
Lead partner
The organisation responsible for leading and co-ordinating the
implementation of an action plan.
Long list
The name, now superseded, for the list of species from which the
short and middle lists was derived.
Middle list
The name, now superseded, for the list of species in the second
tranche of action plans.
Priority
The current nomenclature for those species or habitats for which
species or
action plans have been prepared.
habitat
SAP
Species Action Plan.
Short list
The name, now superseded, for the first tranche of species action
plans.
The name now used for the database (formerly the long list) from
Species of
which priority species are derived.
conservation
concern
Species
A statement of action (typically search only) prepared for those
statement
priority species not seen within the last 10 years.
UK BG
UK Biodiversity Group.
Prepare action plans for threatened species in priority order: globally
threatened; threatened endemics; other threatened species listed in the
relevant schedules or annexes to UK and EC legislation and international
agreements to which the UK is a party; endangered and vulnerable
species listed in Red Data Books, aiming to complete and put into
implementation plans for at least 90% of the presently known globally
threatened and threatened endemics within the next ten years.
UK Biodiversity Action Plan
211
This action may have been stimulated to a degree by the publication of
two editions of Biodiversity Challenge: an Agenda for Conservation in the
UK (Wynne et al., 1993, 1995), produced by a consortium of nongovernment organisations (NGOs). These documents contained a series of targets
and summary action plans for a range of species considered to be priorities
for action.
The Steering Group report
Subsequently, the Government established a Steering Committee to take
forward the work of the Biodiversity Action Plan. This committee consisted of representatives of Government, of statutory conservation agencies
and of NGOs. Their work culminated in the publication of Biodiversity:
the Steering Group Report (Volumes 1 & 2) (Anon., 1995a,b). This report
set the cornerstone for much of the work that has subsequently developed.
In 1996, the recommendations of this report were broadly accepted by
Government (Anon., 1996).
This response also initiated the creation of a network and hierarchy of
groups to oversee the implementation of the action plans and the recommendations of the steering group report. The UK Biodiversity Group now
takes responsibility for overseeing the development and implementation of
the BAP. Four groups, one for each country in the UK, have been
established to take the lead on matters relating to their geographic areas of
responsibility. Additional groups lead on targets, on data (the Biodiversity
Information Group), and on local issues and links to the Agenda 21
process (see Table 18.1), also arising out of Rio. Some of these groups have
created their own subgroups so that the Scottish Biodiversity Group, for
example, now has subsidiary groups dealing with costed action plans,
public awareness, and local biodiversity action plans.
Species and habitat action plans
One of the most significant developments in the steering group report was
the publication, in Volume 2, of action plans for 116 species (including
four fungi), 14 action plans for key habitats, and statements for 37 broad
habitats. The report also recommended that a further 290 species plans
and 24 habitat plans should be produced within two years. As importantly,
from a mycological point of view, it established the criteria for the further
selection of species (and habitats) for action plan production. They are:
∑ that a species is globally threatened, or is possibly so; or
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L. V. Fleming
∑ that a species has declined by over 50% or more in the last 25
years.
Both the first 116 species (which became known as the ‘short list’) and the
second tranche of species plans (known as the ‘middle list’) were selected
from a list (the ‘long list’) of about 1200 species which were considered to be
worthy of conservation attention. Subsequently, it was recognised that the
long list, whilst lengthy, was by no means comprehensive and was an
inadequate basis from which to select priorities for action. For instance,
some species such as blue tits Parus caeruleus were included on the published long list simply because they appeared in an Appendix to an international convention, whilst many red list invertebrates and plants were
excluded. Accordingly, the Joint Nature Conservation Committee (JNCC)
was asked to organise a time-limited review of the middle list in late 1996.
However, this review was to be based on the agreed criteria without
alteration, despite concerns expressed by some specialists on invertebrates
and cryptogams that data were simply not available to enable these criteria
to be satisfied. Although the revised middle list was subsequently larger
than before, additional criteria were applied to limit the number of species
subsequently to receive action plans: namely that any selected species
should be taxonomically sound, that it had been recorded within the last 10
years, and that it would actually benefit from conservation action.
Since that review, it has been determined that all the species on the
revised (and extended) middle list should be subject to some conservation
action. Those species that met the subsidiary criteria above would still
receive a full species action plan. The remainder would either be lumped,
where possible, with another species or habitat action plan (if their requirements were the same) or, if they had not been recorded for over 10 years,
the only action would be for targeted search. The latter, and any remaining
species, would be the subject of a ‘species statement’. The plans and
statements were commissioned from a variety of sources including, for
fungi and lichens, the Royal Botanic Garden Edinburgh and Plantlife, the
UK wild plant conservation charity. Editing was overseen by a group of
cryptogamic specialists from each of the Country Agencies and JNCC.
These subsequent tranches of action plans and statements are mostly now
published (Anon., 1999).
It has also been decided that the nomenclature of short, middle and long
lists should be superseded. In future, any species that is the subject of an
action plan will be referred to as a ‘priority species’. The list from which
these priorities have been determined will be referred to as ‘species of
conservation concern’. Indeed, this is now the title of a database, derived
UK Biodiversity Action Plan
213
from the original long list, holding summary data on threatened species,
which is maintained by the JNCC and managed by a consortium of
country agency and NGO representatives. In due course, this database will
be accessible via the JNCC web page (:http://www.jncc.gov.uk9), where
copies of the species action plans can also be obtained from the UK
Biodiversity website.
Implementing the action plans
In taking forward these action plans, a number of roles have been established. These include the following.
∑ Contact point. Always a Government agency, responsible for
acting as initial contact for queries, stimulating action and
maintaining standards, reporting progress, and confirming that
the work programme meets the requirements of a published plan.
∑ Lead partner. Any organisation of conservation competence,
responsible for co-ordinating the work programme with other
partners, directing resources and stimulating publicity,
establishing a partnership network to ensure that the work
programme can be delivered, and reporting on progress.
∑ Champions. Any organisation or individual that wishes to
support the work of an action plan by providing resources, in
cash or in kind.
So far, no champions have been recruited to support the work of a
fungus or lichen action plan. Plantlife have accepted the role of lead
partner for many of the fungi and will be consulting with the mycological
community. English Nature and Scottish Natural Heritage will lead on
others. Habitat action plans are also assigned a lead agency that is always a
part of Government or one of its agencies. The lead partners or agencies
for these plans may establish a steering group to oversee and guide the
implementation of a plan and to co-ordinate the, often high, number of
contributors.
Local biodiversity action plans
The BAP steering group report also set in train the development of Local
Biodiversity Action Plans (LBAPs). Typically led by Local Authorities,
through the development of partnerships of relevant organisations and
individuals, they aim to ensure that national priorities are taken into
account at local level. As importantly, they seek to ensure that local
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L. V. Fleming
Table 18.2. Fungi for which species action plans have been published
Published 1995
Battarea phalloides
Boletus satanus
Poronia punctata
Tulostoma niveum
Published 1999
Armillaria ectypa
Boletus regius
Buglossoporus pulvinus
Hericium erinaceum
Hygrocybe calyptraeformis
Hygrocybe spadicea
Hypocreopsis rhododendri
Microglossum olivaceum
Stipitate hydnoid fungi@
Species statements (in preparation)
Boletopsis leucomelaena?
?Search only – not recorded since 1963; @a combined plan for all threatened
species in the genera Bankera, Hydnellum, Phellodon and Sarcodon. Lists of all
action plan species can be found at :http://www.jncc.gov.uk9.
priorities and concerns are also addressed. A key element of this process is
that there is full involvement of local people. An audit of the biodiversity
of the area under consideration is usually the first step in the process before
going on to define priorities and to implement action. Following a number
of pilot projects, there are now a significant number of LBAPs under way
in the UK.
Discussion
How have fungi fared in all this?
Of the 116 original action plans, only 4 were for nonlichenised fungi (Table
18.2). A further 9 taxa have subsequently had action plans produced (with
an additional single ‘species statement’). Lichenised fungi have fared better
with 7 species in the first tranche and a further 29 species in the second
tranche. Indeed, of all the taxonomic groups, fungi have fared least well in
terms of the proportion of the group receiving action plans (0.1% for fungi,
2.1% for lichens and 6.7% for birds) (Anon., 1998). Only ‘arthropods other
than insects’ fare almost as badly. However, one may ask whether even this
UK Biodiversity Action Plan
215
limited number of fungi deserve to be included based on a rigid application
of the criteria. Often there is little evidence of decline in fungi due to a
paucity of earlier records. Current or increased recording effort may
paradoxically, even inevitably, provide evidence of an apparent increase in
range or abundance (e.g. Lawson, 1999) even though the underlying trend
may be one of genuine decline. For some of these species, especially the
grassland Hygrocybe and Microglossum species, decline was inferred based
on known losses of grassland habitat. Evidence of global threat or decline
is generally still more poorly known unless a species is confined to Europe
or it is a putative UK endemic. Indeed, the criteria as a whole may have
placed an undue emphasis on the endemic biota of the UK rather than
dealing with threatened species per se.
On the current criteria those fungi receiving action plans are, at best,
going to be a token, and perhaps ad hoc, sample of priorities for conservation. It is likely that the selection of these fungi, from a broad range of
possible candidates, will be the subject of debate for some time. Intuitively,
mycologists may view the rarest fungi, or those with the most restricted
distribution, as being most worthy of attention. Yet, typically, these species may already occur in protected areas, such as Sites of Special Scientific
Interest. By contrast, action plans for species, such as Hygrocybe calyptraeformis, that are more widespread (but which were perhaps once even
more common) will require the needs of fungi to be taken into account in
management decisions for conservation over a much broader area. Consequently, they may have a greater impact on raising the profile of fungus
conservation overall. Compare, for example, the distributions of the H.
calyptraeformis and Armillaria ectypa (Anon., 1999), the latter being
known from only a single site whilst the former is widespread though
uncommon throughout the UK. Many of the vertebrates targeted in the
BAP for action are also ‘common’ and widespread farmland, or even
garden, species such as song thrush Turdus philomelos and skylark Alauda
arvensis.
Regardless of the selection process, there is now a group of fungi, from a
variety of habitats and of a variety of life strategies, which are firmly on the
conservation agenda and which have started to receive targeted action by
the conservation agencies and others (Table 18.3). For example, Tulostoma
niveum has been subject to detailed studies of phenology and population at
its, then, single known site in Scotland (Fleming, Ing & Scouller, 1998); a
second site has since been discovered (E. Holden, pers. comm.). The
Countryside Council for Wales (CCW) has commissioned studies of waxcap grasslands (Rotheroe, 1999; and see Chapter 10) and Scottish Natural
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L. V. Fleming
Table 18.3. Examples of projects on fungi undertaken or commissioned by
the statutory conservation agencies in the UK since the initiation of the
BAP
Agency
Project
Countryside Council Waxcap grassland survey of Camarthen; identification of
for Wales
950 sites for Hygrocybe calyptraeformis.
Training on fungi, their identification and conservation.
Production of leaflets and articles in media on fungus
conservation.
Database established on the rare fungi of Wales.
English Nature
Grant aid for survey of Boletus regius by Plantlife.
Leaflet on collecting of fungi (1998).
Contribution to waxcap survey.
Environment &
National checklist of fungi of the British Isles (a
Heritage Service
partnership of all the Country Agencies and Royal Botanic
(Northern Ireland) Gardens Kew).
Survey of Boletaceae and Amanitacaea of County Down.
In-house surveys for waxcaps and Microglossum olivaceum.
Comparative survey of waxcap grassland fungi in Britain
Joint Nature
and Ireland.
Conservation
Fungi of National Vegetation Classification grassland
Committee
communities.
Collation and digitisation of foray records for the BMS
database.
Scottish Natural
Survey of tooth (stipitate hydnoid) fungi in native
Heritage
pinewoods (1998–2000); including search for Boletopsis
leucomelaena.
Technical information on the occurrence of cryptogams in
Scotland to assist the implementation of LBAPS.
Phenology and monitoring of Tulostoma niveum at
Inchnadamph.
Waxcap grassland survey of Scotland 1999–2001.
Inventory and database of fungi in the Cairngorms.
Heritage (SNH) is about to follow suit. SNH has also commissioned a
study of tooth (stipitate hydnelloid) fungi in native pinewoods, English
Nature have grant-aided Plantlife to undertake work on Boletus regius
(Marren, 1998) and further work is planned. SNH and CCW have provided technical information on the occurrence of fungi by Local Authority
boundaries and, in Scotland, by natural areas (Ward, 1999). It is no
exaggeration to say that without the BAP process, the claim of fungi on the
resources of the statutory conservation agencies, at least, would have been
much smaller (even though the resources being committed are currently
much less than those for other taxa).
UK Biodiversity Action Plan
217
What is the role of the British Mycological Society in
implementing these plans?
Already, the British Mycological Society (BMS), through its waxcap grassland survey (Rotheroe et al., 1996) is taking forward action that is fundamental to implementing these plans. Indeed, the BMS survey is recognised
in the action plans for the two Hygrocybe species. All the plans will require
good data if they are to be implemented effectively and the BMS database,
and records collected by BMS members, will be vital in assisting those
charged with taking plans forward. It is also recognised in this process that
the selection of priorities is not set in stone. Better information in time may
indicate that further species need to be considered as priorities for action.
Alternatively, conservation or recording effort may indicate that a species
no longer deserves the action being committed to it – a conclusion reached
by Marren (1998) in his study of Boletus regius. Such reporting and
feedback on progress with plan implementation are an integral part of the
BAP.
Although this account has focused on the species action plans, there will
also need to be an input by mycologists to the implementation of the Local
and Habitat Action Plans if they are to achieve their potential to contribute to the conservation of fungi. Indeed, the maximum gains for the
conservation of fungi, and of all fungi not just those selected for action
plan production, may best be achieved by this approach. Many of the
fungus action plans, and indeed those of other taxa, have been grouped by
habitat preference to achieve synergy of effort (Simonson & Thomas,
1999). For example, implementation of the habitat action plan for wood
pasture should benefit Boletus regius, Buglossoporus pulvinus, Hericium
erinaceum, and some stipitate hydnoid fungi as well as other species such as
the lichen Enterographa sorediata or the stag beetle Lucanus cervus. Other
Habitat Action Plans cover ecosystems, such as sand dunes, native pinewoods, calcareous grasslands and wet woodlands, which are also rich in
fungi yet the fungus interest will only be fully recognised with the active
involvement of mycologists. Many Local Biodiversity Action Plans already include some fungi in their list of priorities.
Regardless of any faults in the process, the Biodiversity Action Plan has
given a priority, momentum and resource commitment to the conservation
of fungi in the UK that was inconceivable in its absence; it is now up to
mycologists to make the most of the opportunities that have been presented to us.
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L. V. Fleming
Acknowledgements
I thank Maurice Rotheroe for comments on drafts of this article. I am also
grateful to Stephen Ward (SNH), Nick Hodgetts (JNCC), Ray Woods
(CCW), Dave Stone and Will Simonson (English Nature), and David
Mitchel (Environment & Heritage Service) for the provision of information used in this chapter. The views expressed here are those of the author
and do not necessarily reflect those of the Joint Nature Conservation
Committee.
References
Anon. (1994). Biodiversity: the UK Action Plan. Cm2428. HMSO: London.
Anon. (1995a). Biodiversity: the UK Steering Group Report. Vol. 1, Meeting the
Rio Challenge. HMSO: London.
Anon. (1995b). Biodiversity: the UK Steering Group Report. Vol. 2, Action Plans.
HMSO: London.
Anon. (1996). Government Response to the UK Steering Group Report. Cm3260.
HMSO: London.
Anon. (1998). UK Biodiversity Group. Tranche 2 Action Plans. Volume I –
Vertebrates and Vascular Plants. English Nature: Peterborough.
Anon. (1999). UK Biodiversity Group. Tranche 2 Action Plans. Volume III –
Plants and Fungi. English Nature: Peterborough.
Fleming, L. V., Ing, B. & Scouller, C. E. K. (1998). Current status and phenology
of fruiting in Scotland of the endangered fungus Tulostoma niveum.
Mycologist 12, 126–131.
Lawson, P. (1999). Sandy stilt puffball. British Wildlife 10, 361.
Marren, P. (1998). Boletus regius Kromb., the royal bolete and the pretender
Boletus pseudoregius Estades. Plantlife Report No. 111. Plantlife: London.
Rotheroe, M. (1999). Mycological survey of selected semi-natural grasslands in
Carmarthenshire. CCW Contract Science Report no. 340. Countryside
Council for Wales: Bangor.
Rotheroe, M., Newton, A., Evans, S. & Feehan, J. (1996). Waxcap-grassland
survey. Mycologist 10, 23–25.
Simonson, W. & Thomas, R. (1999). Biodiversity: Making the Links. English
Nature: Peterborough.
Ward, S. D. (1999). Local Biodiversity Action Plans – Technical Information on
Species: I. Cryptogamic Plants and Fungi. Scottish Natural Heritage Review
No. 70. Scottish Natural Heritage: Battleby, Perth.
Wynne, G., Avery, M., Campbell, L., Gubbay, S., Hawkswell, S., Juniper, T.,
King, M., Newbery, P., Smart, J., Steel, C., Stones, C., Stubbs, A., Taylor,
J., Tydeman, C. & Wynde, R. (1993). Biodiversity Challenge: an Agenda for
Conservation in the UK, 1st edn. Royal Society for the Protection of Birds:
Sandy.
Wynne, G., Avery, M., Campbell, L., Gubbay, S., Hawkswell, S., Juniper, T.,
King, M., Newbery, P., Smart, J., Steel, C., Stones, C., Stubbs, A., Taylor,
J., Tydeman, C. & Wynde, R. (1995). Biodiversity Challenge: an Agenda for
Conservation in the UK, 2nd edn. Royal Society for the Protection of Birds:
Sandy.
19
The Scottish Wild Mushroom Forum
ALIS ON DYKE
Introduction
The Scottish Wild Mushroom Forum was developed in response to the
rapid growth of interest in harvesting wild mushrooms. Both picking for
the pot and commercial harvesting are popular and bring people back into
the countryside in an enjoyable and sometimes profitable way. The commercial industry, which has developed in the last fifteen years, involves five
principal companies and up to 400 casual harvesters. Commercial harvesting is fairly important as it provides an additional source of income for
rural communities.
Until recently there has been little mushroom harvesting in Scotland,
and the traditions and widespread knowledge that exist on the Continent
have not had time to develop. The Forum sees the enjoyment that people
gain from harvesting mushrooms as an opportunity for positive action,
and seeks to involve harvesters in the creation of a voluntary Code of
Practice. As well as producing a Code, the Forum will seek to implement it
in ways that will reach the largest audience. By involving people in the
management of the resource that they use, the Forum intends to build a
culture of sustainable harvesting.
Grants from Scottish Natural Heritage and Moray Badenoch and
Strathspey Enterprise fund the Forum, together with a Millennium Award
from the Millennium Forest for Scotland Trust.
The process
The groups involved in the harvest of wild fungi were identified during an
earlier study (Dyke & Newton, 1999) and are shown in Table 19.1. Contact
was made with these groups. In the case of mushroom harvesters announcements were made in the local press and at points of sale in three
219
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Alison Dyke
Table 19.1. Groups involved in harvesting
Forum members
Landowners’ representatives
Mushroom Buyers
Mushroom Pickers
The Association of British Fungus Groups
The British Mycological Society
Cairngorm Partnership
Forest Enterprise
Grampian Fungus Group
The Joint Committee for the Conservation of
British Insects
The National Trust for Scotland
Scottish Native Woodlands
Scottish Natural Heritage
Nonparticipating members Plantlife
The Royal Society for the Protection of Birds
World Wide Fund for Nature
Scottish Wildlife Trust
Highland Birchwoods
Table 19.2. Some of the issues surrounding wild mushroom harvesting
Beneficial aspects of
harvesting
Conservation aspects
Exercise and access to the countryside
Money – the commercial aspect
Waste – picking and dumping of inedible or unwanted
species
Effects on colonisation
Habitat disturbance – plants and animals
Designated sites: Sites of Special Scientific Interest,
Nature Reserves, etc.
Effects on estate business Damage (deer fences cut, litter dropped)
Stalking and cattle disturbance
Permission
Effects of land management on fungi
Competition between
pickers
areas where mushroom harvesting is concentrated (Deeside, Speyside and
the Black Isle). Regional meetings were held where issues were discussed
and representatives nominated. The issues the Forum aims to address are
shown in Table 19.2.
Having established an agenda, Forum members were provided with
background information to ensure that they were all aware of the concerns
of other groups. A meeting of all representatives was held and issues that
Scottish Wild Mushroom Forum
221
Table 19.3. The Scottish Wild Mushroom Code
The countryside is a working landscape. Please be aware of safety and follow the
Countryside and Access Codes. As a matter of courtesy you are advised to ask
for permission before you pick mushrooms.
By respecting the natural environment you can help to manage and conserve the
countryside. When picking mushrooms for any purpose, please consider the
following points:
∑ Wildlife, especially insects, need mushrooms too, so only pick what you
will use.
∑ Do not pick mushrooms until the cap has opened out and leave those
that are past their best.
∑ The main part of the mushroom is below the surface, take care not to
damage or trample it, and not to disturb its surroundings.
∑ Scatter trimmings discreetly in the same area as the mushrooms came
from.
∑ Some mushrooms are poisonous and others rare and should not be
picked – only pick what you know and take a field guide with you to
identify mushrooms where you find them.
∑ Before you collect mushrooms at a nature reserve please always seek
advice from the manager, as special conditions may apply.
If you own or manage land:
∑ Be aware that your management activities may affect mushrooms.
If you wish to run a foray or collect for scientific purposes:
∑ In order to ensure the safety of your party obtain permission in writing.
∑ Give a record of what you have found to the landowner or manager and
explain the significance of your findings.
This Code was created by the Scottish Wild Mushroom Forum, a group
consisting of representatives of: conservation organisations, landowners, public
landowning bodies, mushroom buyers and mushroom pickers. The Forum was
funded by Scottish Natural Heritage, The Millennium Forest for Scotland Trust
and Moray, Badenoch and Strathspey Enterprise.
could be addressed by a Code were identified. In order to reach a variety of
audiences the Code uses direct and positive language and will be produced
in a number of formats.
The draft text of the Code is shown in Table 19.3 and will be supported
by an informational leaflet. The first five points could be used as a set of
‘golden rules’ on signage, as a credit card sized handout, or even on wild
mushroom packaging. It is hoped that educational materials for use by
rangers and people wishing to run forays will also be available.
The Code will be endorsed by a number of organisations and will be
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Alison Dyke
distributed through a range of outlets in order to reach the widest possible
audience. Plans are also being made for a pilot consent scheme, involving
the issue of consents to mushroom harvesters. This would allow landowners to limit the number of commercial harvesters, to identify cars
parked on their land and could also allow mushroom harvesters access to
areas they would not normally be able to reach. A consent could also be
used as a vehicle to introduce the Code to commercial harvesters.
It is intended that the process should remain open, with ongoing participation to review the Code and its implementation.
Reference
Dyke, A. J. & Newton, A. C. (1999). Commercial harvesting of wild mushrooms
in Scottish forests: is it sustainable? Scottish Forestry 53, 77–85.
20
The contribution of national
mycological societies: establishing a
British Mycological Society policy
DA VID MOORE
Introduction
The British Mycological Society (BMS) was founded in 1896 and today
has about 2000 members who are located all over the world. The constitutional objective of the Society is to promote mycology in all its aspects by
publications, meetings and such other means as it shall deem appropriate.
The Council of the BMS is the executive that implements on a day-to-day
basis the activity of the Society that is decided by the members of the
Society at the Annual General Meeting, usually held in early December of
each year.
Field mycology and an awareness and appreciation for the natural
world were at the heart of the business of the Society from its foundation
and these concerns continue as one of the Society’s major activities today.
The origins of the BMS trace back to the mid-nineteenth century (Webster,
1997). The Woolhope Field Naturalists’ Club was based on the Hereford
Museum, though members of the club dined at the Green Dragon Hotel in
Hereford. In 1867 the Curator of the museum, Dr H. G. Bull, encouraged
the club to take a special interest in fungi. He invited them to join him in ‘a
foray among the funguses’ and this became an annual event, traditionally
held in Hereford during the first week of October. The Woolhope Club
meetings became a focus for all with an interest in fungi and attracted
mycologists both from Britain and abroad. These forays lost their popularity when Dr Bull died in 1885 and stopped in 1892. By that time, though,
the Yorkshire Naturalists’ Union (YNU) was organising regular forays in
different parts of Yorkshire and a Mycological Committee was formed in
1892. The stated aim of the YNU Mycological Committee was that their
annual forays would take the place of the Hereford Foray and ‘by avoiding
the weak points of its predecessor, which were mainly confined to an excess
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David Moore
of hospitality − prove at least equally attractive and instructive to mycologists’ (Massee & Crossland, 1893, quoted by Ramsbottom, 1948). A need
was also felt to provide an outlet for the publication of scientific articles on
fungi. The idea of forming a ‘National Mycological Union’ emerged at the
YNU meeting in Huddersfield in 1895 and the decision to set up the British
Mycological Society was taken on 19 September 1896 at a meeting of the
YNU Mycological Committee at the Londesborough Arms in Selby. The
first officers were G. E. Massee as President (then Mycologist at the Royal
Botanic Gardens, Kew), Charles Crossland as Treasurer (a Halifax
butcher by trade, Crossland compiled, with Massee, The Fungus-flora of
Yorkshire in 1902–1905) and Carleton Rea as Secretary. Rea was a barrister by profession but he gave this up in 1907 and wrote British
Basidiomycetae (published 1922) which was the standard work on identification of the group for many years.
Autumn forays were the main activity of the Society initially and their
arrangement was the chief responsibility of the Secretary. From 1919 a
Foray Secretary took on this task. The first was A. A. Pearson (1919–24),
who was a gifted amateur and an authority on the identification of agarics
and boletes. He wrote popular keys to Russula, Lactarius, Boletus, Inocybe
and Mycena, and in 1948, with R. W. G. Dennis, A Revised List of British
Agarics and Boleti. The Annual General Meeting in 1942 established a
Foray Committee. Later the committee was renamed the Foray and
Conservation Committee.
Autumn, Spring and Day forays became a regular feature of the Society’s annual programme. Regular specialist forays were inaugurated in
1982 with an Upland foray to Wester Ross. The first ‘official’ Truffle Hunt
took place in the Cotswolds in 1984. They were enthusiastically continued
for several years and included a truffle meeting in Italy in 1987. Regular
annual hunts were discontinued because of the possible threat to rare
fungi. Overseas Forays have been held in Northern France (1984), Northern Greece (1988), Southern Denmark (1991), Norway (1994) and are now
also a regular feature of the programme. The first Tropical Expedition
organised by the Society was an excursion to Cuyabeno, Ecuador, in 1992
(Hedger et al., 1995). Collecting trips have more recently been made to
Thailand and it is expected that tropical expeditions will be organised
every three to four years. In 1998 the BMS held a joint meeting in Chiba,
Japan, with the Mycological Society of Japan. Although not a field meeting, this sealed close contacts between the two Societies and will lead to
further and closer intercontinental collaborations in the future.
Collections made during BMS forays and expeditions provided valuable
British Mycological Society conservation policy
225
information on numbers of species and species distributions: for example,
of British truffles (Pegler, Spooner & Young, 1993). Lists of fungi collected
on forays formed a regular part of Foray Reports in the Transactions of the
British Mycological Society and the earlier volumes of the Society’s Bulletin. With the establishment of a BMS Fungal Records Database in 1986,
computerisation of records began (Minter 1986a,b) and that database is
now becoming a major resource.
In the present committee structure of the Society there are several
Special Interest Committees (SICs): Biodiversity; Ecology and Environmental Mycology; Foray; Fungus–Invertebrate Interactions; Conservation; Genetics, Molecular Biology and Evolution; Pathogenic and Mutualistic Interactions; Physiology; Systematics and Structure. The SICs for
Conservation and for Biodiversity were set up in 1996 in recognition of the
need for better focus on these particular aspects of mycology.
Developing the conservation agenda within BMS
Mounting concern over adverse effects of environmental and atmospheric
pollution was reflected in data showing similar decline in fungal populations published from the 1970s onwards. This is not the place for a
comprehensive set of references, but it is interesting to note the time span
represented in the following few papers: Wilkins & Patrick, 1940; Wilkins
& Harris, 1946; Richardson, 1970; Arnolds, 1988a,b; Eveling et al., 1990.
In a report to Council dated 9 December 1986, the 1987 BMS President,
Professor Roy Watling, reminded Council that ‘In 1976 a list of rare larger
fungi was discussed with BRC [Biological Records Centre] (Greenhalgh,
Whalley and Watling representing BMS). It is possible to produce a list for
Britain and a list of indicator species . . . ’.
Subsequently, in April 1987, Roy Watling was even more instrumental
in placing conservation firmly on BMS Council’s agenda, in a perceptive
document that stated: ‘The British Mycological Society is committed to
the conservation of our national heritage and to playing some defined and
active role to meet these aims. The Society is in a good position to offer
expert advice on individual sites and expert opinions on specific fungal
records and this will improve in the future as more records are keyed into
the Society’s computer. To fulfil the above role, which is really one of
communication and co-operation, the Society has already appointed a
representative on the Conservation Committee of the European Mycological Congress, and its own Conservation Advisory Officer. It is hoped that
as time goes on the names of these Officers will be more widely known, and
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David Moore
their expertise called upon more frequently . . . Council should think in
terms of setting up a small working party probably best linked to the
Systematics, Structure and Foray SIC to consider how the proposed
mapping of British Fungi can be used to monitor the decrease of particular
species etc., and so find out the impact that foraying has on a selected
fungus flora, or whether such a factor can even be measured.’
On 22 February 1988, Roy Watling made the specific suggestion that
Council should finance some urgent research ‘To circulate and collate
information concerning fungal protection in European countries and to
relate this to the situation in the British Isles.’ The supporting documentation for this proposal clearly identifies the major concern being expressed
at that time about ‘the effects of collecting fungi, either for recreational or
scientific purposes, on their productivity and, therefore, their long term
continuance in nature. The Plant Protection Act has made a major contribution to the safeguarding of higher plants, but sadly the fungi were not
included therein owing to lack of knowledge. Unfortunately, this lack of
knowledge continues but there is an ever-increasing pressure on our fungi,
including demands from abroad for edible species. Certain European
countries already have a ‘picking policy’ and the British Mycological
Society needs urgently to make a preliminary study of (a) the information
which has brought this about, (b) the desirability of similar policies in the
British Isles with all its attendant problems, and (c) the effects of introduction of such policies if considered necessary.’
Council agreed to support the project on ‘Effects of picking fleshy fungi
on the Countryside’s resources’ and the recently appointed Conservation
Officer, Bruce Ing, was able to report to Council in July, 1988, details of the
programme of research and that the work would be undertaken by Dr
Thomas Læssøe, who then reported in November of the same year.
As far as I am aware the Læssøe report Conservation of Fungi remained a
Council discussion paper which was never published, but as its findings
formed much of the foundation for subsequent BMS activity it is worth
giving some extensive quotations from it. The first few paragraphs established the history and background of the study.
Since the invitation to European mycologists by the Dutch group to form
a committee on fungal conservation [at] the European Mycological Congress in 1985 the subject has become topical. Many journals including the
Mycologist and The Transactions of the British Mycological Society have
published papers on the protection of fungi and have pointed to alarming
decline in some fungal groups in specific areas. Acid rain, over-picking,
bad management, etc., have been put forward as reasons for the alleged
British Mycological Society conservation policy
227
decline. The Council of the British Mycological Society also felt that
something should be done and appointed an officer to the European
Committee (Roy Watling) and an officer to deal with the local matters
(Bruce Ing). These two officers together with the General Secretary (Tony
Whalley) became the board for my project: To write a summary of the
situation and give some suggestions of how to proceed.
History of mycological conservation
For obvious reasons the direct conservation of fungus species have been a
very insignificant part of general conservation schemes and only recently
have fungi been taken into consideration by conservationists.
In the USSR all fungi included on the Red Data List are automatically
protected. In Poland a special list of protected fungi have been prepared
and e.g. advertised to the public by issuing a stamp series with pictures of
the different species. In some countries (e.g. the German Federal Republic) it is prohibited to market certain edible but rare species.
A more important thing for a long-term conservation purpose is of
course to protect sites rather than species. But again very few sites have
been protected because of a known important mycoflora. There is only
one nature reserve in the UK created to protect the mycoflora (a ‘hedgerow-locality’ with Battarrea phalloides). One or two other sites have been
listed as sites of special scientific interest because of an interesting and
well-documented mycoflora. Protection of sites because of mycological
interests is equally rare in other parts of the world. . . .
Possible causes of fungal decline (or increase): Picking . . . Although it
is commonly believed that mushrooms are threatened by overpicking, no
scientific data support this view (Jansen & van Dobben, 1987, Arnolds
1988[b], etc.). Fruiting of some species seems to be enhanced by cropping
and it has been postulated that hypogeous fungi are favoured by soil
disturbance. Nitare (1988) warns against continued picking of fungi
already recorded in an area although the taxonomic difficulties often
make it necessary to collect specimens. In Oregon a project [has] recently
been set up to study the cropping of Cantharellus cibarius over a 10 year
period. The aim is also to measure the effect of trampling (mechanical
damage to the soil and supporting vegetation). This might be a more
severe threat than the actual picking. Arnolds (1988b) made a seven year
study where fungal fruit bodies were removed from plots. No decrease in
fruiting was observed.
. . . The British Mycological Society policy concerning picking is
somewhat unclear, since on [the] one hand R. Watling in CAB News and
in the Mycologist advises the membership to collect only what is necessary for a determination while in each issue of the Mycologist you will find
instructions of how to cook (etc.) your mushrooms. In most European
countries, with the exception of The Netherlands, it would be considered
completely non-desirable to condemn picking for culinary purposes.
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Land management
‘‘The principal reasons for the decline of fungi are forest management
practices, including the reduction in the numbers of rotting trees in
commercial forests, the artificial drainage of wetlands, the encroachment
of forest and especially spruce trees on what was originally meadowland,
and constructions.’’ (from the Finnish Red Data Book).
This statement is almost universal in the different conservation papers.
Another general factor is the agreement on the negative effect of commercial fertilisers both in grasslands and, in forests. Even coprophilous
species such as some Coprinus species and Poronia punctata are negatively
affected by adding commercial fertilisers (Arnolds, 1988b).
The removal of dead trunks and larger fallen branches is also considered a general problem.
In a table [unpublished] communicated by Bruce Ing the loss of major
habitats in the UK since 1949 is listed. Only 3% of ‘lowland neutral
grasslands’ remain undamaged and only 15% of ‘ancient lowland woods
of native broad leafed trees’ remain undamaged . . .
Pollution
Both Arnolds (1988b) and Nitare (1988) (and various Dutch and German
papers) stress the importance of airborne pollutants in fungal decline. In
all probability airborne pollutants are responsible for the decline of
ectomycorrhizal fungi in The Netherlands, exemplified by detailed studies on the Chanterelle. Jansen & van Dobben (1987) stated about the
decline of Cantharellus cibarius ‘‘although our data indicate an effect of
acidification or eutrophication, effects caused by heavy metals are also
possible. Heavy-metal pollution may also lead to an increase in the
accumulation of organic matter. In one case there is evidence of sensitivity of C. cibarius to heavy metals (a Swedish study)’’. In the Dutch
material levels of heavy metals were lower than in the proven cases of
toxicity, and thus heavy metals were less likely to be the cause of the
decline.
Judged by the communications in Poland also the situation in Czechoslovakia and in parts of Germany is very grave concerning the ectomycorrhizal fungi, while the situation in Scandinavia is either not documented
or much less severe. The same seems to apply for the UK.
Climatic changes
Many fungi considered rare in various countries can probably best be
regarded as being outposts from their natural distribution and they are
thus sensible to even small changes in the climate. No doubt some of the
disappearing species can be regarded as ‘threatened’ by climatic changes
(e.g. several elements in the south eastern British mycoflora). The annual
variation in climate no doubt also is responsible for many records of so
called rare fungi, e.g. in this summer in Denmark where the unusually hot
and humid July has resulted in a number of records of ‘‘rare, southern’’
species.
British Mycological Society conservation policy
229
In reaching his conclusions, Læssøe referred to a paper by Kirby (1988)
from the Chief Scientific Directorate of the Nature Conservancy, who
raised three questions about fungal conservation: (A) are special conservation measures for fungi needed or justified? (B) How should we judge
which sites should be conserved for their fungi? (C) How should these sites
be managed to maintain their fungal value?
Læssøe concluded that special conservation measures for fungi are
justified, that Red Data lists can provide the basis for deciding which sites
are most important, and recommended that site management should
ensure, for example, that nutrient poor grasslands should be kept nutrient
poor (by avoiding fertilisers) and should be grazed or cut; old wood should
be left in forest reserves, and drying of the topsoil should be avoided.
A number of specific projects for BMS to undertake were proposed in
the Læssøe report. These were, in the order of preference quoted by
Læssøe: production of a booklet on conservation of fungi; production of a
code of conduct leaflet for forays and mushroom pickers; updating of the
BMS database so it can help in preparing Red Data lists; protection of
valuable grasslands (including dunes); study of airborne pollution and its
effect on the mycota; and issuing questionnaires to the membership to
assess change in distribution of indicator species.
Also in November 1988, Roy Watling reported to Council a successful
meeting on Fungal Conservation held on 12 November which was organised for the BMS by David Minter. Delegates at the meeting included
Dr A.-E. Jansen, the Secretary of the European Conservation Committee,
Dr N. Stewart from the Conservation Association of Botanical Societies,
and Dr N. Hodgetts and Dr Keith Kirby, of the Chief Scientific Directorate, of the Nature Conservancy Council. This was the first in a number of
scientific meetings on this topic area sponsored by the Society. Later ones
included a Symposium on Fungi and Environmental Change held in 1994
(Frankland, Magan & Gadd, 1996), and the Symposium at the Royal
Botanic Gardens, Kew (13 November 1999) entitled Fungal Conservation
in the 21st Century, which was the origin of this book.
All the above shows the careful scientific approach expected of a learned
society. There is a need for cogent logical arguments if we hope to change
what people are doing, but above all there is a need for relevant scientific
knowledge.
In the course of the few years immediately following the Læssøe report
the Society’s then Conservation Officer, Bruce Ing, published a provisional
Red Data list of British fungi (Ing, 1992); he has revised and re-evaluated
these data on a regular basis since then. However, the first formal outcome
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of discussions prompted by the Læssøe report was published as the ten
point BMS policy on conservation (Anon., 1990), stating:
1. The British Mycological Society is committed to the
conservation of non-harmful fungi and their habitats. To this
end it will foster and support those activities which will ensure
the survival of fungal populations.
2. The Society will compile a Red List, using strict criteria and
related to habitat type, of rare and endangered fungal taxa,
which may be helpful in evaluating sites and in drafting any
future legislation on species protection.
3. The Society will, where possible, seek to provide information
concerning the mycoflora of sites which may be threatened, so
that management appropriate to fungal conservation may be
planned. The Society will provide advice to landowners,
conservation bodies and local authorities on the mycological
importance of their land and of suitable conservation measures
that may be adopted.
4. The Society will encourage research into the decline, or
otherwise, of fungal populations and will help to make available
the results of such research. The Society’s database of fungal
observations will assist in this motoring activity.
5. The Society maintains that there is no evidence that the
responsible collection of fungi for scientific purposes presents
any threat to populations and, moreover, that it is essential for
the accurate identification of species and the compilation of site
inventories.
6. The Society will provide a Code of Conduct for its members,
other conservation bodies and individuals who wish to collect
fungi.
7. The Society does not condone commercial collection of wild
mushrooms, particularly as there is uncertainty as to the effects
of such large-scale collection on fungal populations.
8. The Society will strengthen its links with those organisations
concerned with forest ecosystems, especially where such systems
are seen to be under stress, so that the vital role that fungi play
in such ecosystems becomes more widely appreciated.
9. The Society will extend its relationship with its European and
other overseas counterparts to exchange information concerning
continental and global changes in mycoflora, and will encourage
British Mycological Society conservation policy
231
appropriate research programmes. It will review and update the
policy regularly so as to take into account new research findings
and policy decisions in other countries.
10. The Society will actively promote a wider understanding of the
importance of fungi and their biology, and in particular their
significance in the conservation of natural communities, as a
contribution to environmental education.
During the next few years the Society concentrated on codifying its
conservation policies, with particular emphasis on collaboration with
other bodies, both national and international, in an effort to increase and
widen the effectiveness of those policies.
Codifying policy
At a meeting of Foray Group Leaders convened by the Society at Littledean in May 1995 Maurice Rotheroe, then speaking as Deputy Conservation Officer (and Conservation Officer-elect) expressed the view that the
Society could make a much greater contribution to local and national
conservation issues and there was a need for conservation to have a much
higher profile within the BMS. He outlined the role that the BMS could
play in liaison with English Nature, the County Trusts and other wildlife
and conservation bodies. The BMS could offer specialist knowledge of the
relationship between fungi and other organisms not available elsewhere.
There was much work to do in publicising fungi, in educating reserve
managers and others in mycology and management for conservation of
fungi. He also issued a questionnaire to Group Leaders to enable him to
collect the first-hand experience of these groups and to establish links with
local County Trusts and English Nature representatives in each area.
Minutes of the BMS Council meeting of 6 December 1996 featured the
annual report to Council of the BMS Conservation Officer, Maurice
Rotheroe. This was especially notable because in the course of 1996 a
Conservation Special Interest Committee had been established. The SIC’s
first meeting lasted four hours during which its members took on responsibility for gathering more data on endangered species to contribute to
future Red Data lists; initiated work on the production of a draft guide to
site management techniques which are sensitive to the requirements of
fungi; and identified commercial collecting of wild fungi as the most
prominent current issue following intense publicity about the topic earlier
in the year.
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On behalf of the Conservation SIC, Patrick Leonard compiled a consultation document entitled Outline Policy on Commercial Collecting of Wild
Fungi which was presented to Council at this December 1996 meeting. This
report notes that the BMS had set out its policy on commercial collecting
of macrofungi as part of its policy on conservation, published in 1990
(Anon., 1990; and see above). Public concern had mounted as commercial
collecting had expanded in the early 1990s and the policy needed amendment. The vehemence of the concern among some members of the public
had been brought home to the Society in a face-to-face confrontation
several weeks before this Council meeting. 1996 was the BMS Centenary
year and at the Society’s ‘Fungus 100’ exhibition in London in 1996, a
wild-mushroom cooking demonstration by one of the world’s great chefs,
Antonio Carluccio, was noisily interrupted by ‘green activists’ accusing
him of pillaging the environment!
Increased collecting of fungi gave rise to concerns about: (1) the effect on
the long-term survival of the fungus, and the indirect effect on organisms
further down the food chain, that is, the overall wildlife conservation issue;
(2) potential distortion of scientific records; (3) the effect on local people
who thought they had de facto collecting rights in public forests; (4) effects
on landowners, and particularly woodland owners, of the nuisance of ever
more determined pickers; (5) adverse aesthetic effects on the natural environment caused by indiscriminate collecting in autumn woods.
Patrick Leonard argued that the then published position of the BMS
appeared weak and difficult to explain to concerned but nonexpert members of the public. It also failed to address the question of the cause of
observed declines in fungal populations because point 7 in the 1990 policy
statement (see above) simply refers to ‘uncertainty as to the effects of such
large-scale collection’. Inevitably, also, the 1990 policy had no reference to
the International Convention on Biological Diversity that the UK signed
in 1992.
Leonard suggested that ‘A much more viable policy position would be
to acknowledge that there are declines in the populations of common
edible fungi. To point clearly to habitat losses and the effects of pollution
and make clear the BMS stance on these. To adopt the precautionary
principle in relation to commercially collected species where numbers are
declining rapidly and propose to Government that action plans should be
drawn up for two or three of these. To protect habitats where rare and
threatened fungi are present by halting commercial collecting. To make
common cause with the conservation agencies, the Forestry Authority,
landowners and with the major conservation voluntary bodies such as the
British Mycological Society conservation policy
233
National Trust and the Wildlife Trusts . . . In the light of increasing
concern about the rapid expansion of commercial collecting of fungi in
Britain it would seem desirable for the BMS to re-examine and strengthen
its conservation policy. Such a change would be seen as a positive response
to the concerns and a timely updating of the policy following new national
and international policies on conservation . . . ’.
Council agreed on the need for a revised conservation policy and that
there should be a statement which presented the Society’s views on the
commercial collecting of wild fungi. The President (John Webster) asked
Maurice Rotheroe to receive and collate contributions to a revised policy
document which would be brought back to Council before being published
in the Mycologist.
Consultations continued through 1997 and the final policy document
was approved at the December 1997 meeting of Council, with David
Moore as President. The text, published two months later (Anon., 1998a),
was as follows.
British Mycological Society Policy on Conservation
1. General Statement:The British Mycological Society is committed to the
conservation of fungal populations and communities. To this end it will
foster and promote those activities that contribute to survival of viable
fungal populations and communities.
2. Threatened Fungi: The Society will compile and publish a Red List that
conforms to international standards and will press for positive
conservation of threatened fungi through national and international
measures.
3. Habitat Conservation: The Society will seek protection for important
mycological sites against loss, deterioration or fragmentation,
howsoever caused.
4. Edible Fungi: The Society acknowledges the importance of edible wild
fungi as a resource to be utilised, but accepts harvesting of such fungi
only where it is non-threatening to the viability of fungal populations,
and their associated organisms and habitats.
5. Code of Conduct: The Society will publish a Code of Conduct for the
responsible collecting of fungi.
6. Research: The Society’s constitutional object ‘to promote mycology’
encompasses encouragement of research on the biology, including
taxonomy and ecology, of fungi; on the causes of decline of fungal
populations; and on the cultivation of edible fungi.
7. Information: The Society will monitor and record the occurrence of
fungi and make its Database available to its members and to outside
organisations and individuals.
8. Education: The Society will promote a wider understanding of the
importance of the conservation of fungi.
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David Moore
9. Collaboration: The Society will enhance its links with organisations
concerned with conservation and the protection of the environment at
local, national and international levels.
10. Review: The Society will review and update its conservation policy, as
required, to take account of new research findings and changes in
relevant legislation and environmental policy.
The published policy was accompanied by a paragraph describing how
practical implementation of the BMS Conservation Policy would be incorporated into a Five-year Plan which the Conservation SIC had developed
and had been approved in principle at Council. As recorded in Council
Minutes, the consultation document described the provisional five-year
plan in the following terms.
1. General: The British Mycological Society will disseminate its policy on
the conservation of fungi both inside and outside the scientific
community, using appropriate media including publication via the
World Wide Web.
2. Threatened Fungi: The Society will review and update the Provisional
Red Data List of British Fungi and will press for national and
European legislation to protect threatened fungi where appropriate. It
will urge the Joint Nature Conservation Committee to publish a Red
Data Book for fungi and will assist in its production. The Society will
actively participate in the implementation of action plans for fungi
published under the UK Biodiversity Action Plan.
3. Habitat Conservation: The Society will co-operate with statutory and
voluntary conservation organisations to identify mycologically
important sites and habitats. It will compile a management handbook,
covering all ecosystems, to assist landowners and managers.
4. Edible Fungi: The Society will monitor the impact of commercial
collecting and will press for regulation where such collection is shown to
threaten the viability of fungal populations and their associated
organisms. It will actively discourage the collecting of species that are
identified as being under threat. The Society will co-operate with land
managers and other bodies to promote the sustainable harvesting of
edible fungi. It will initiate action through the European Council for the
Conservation of Fungi to urge governments of importing and exporting
countries to curb excesses of commercial collecting through legislation.
5. Code of Conduct: The Society will seek sponsorship for the production,
publication and promotion of its Code of Conduct.
6. Research: The Society will press for research relevant to conservation
of fungi and will compile a list of key areas for investigation, to be
distributed to grant-awarding and research organisations in the public
and private sectors. Specific areas for priority investigation will include:
the impact of commercial collecting on fungal populations; the
cultivation of mycorrhizal species of edible fungi; and the effect of
British Mycological Society conservation policy
7.
8.
9.
10.
235
government agricultural and land-use policies on fungal populations
and communities.
Information: The Society will ensure that its database is efficiently
managed and updated and the design modified, where necessary, to
make it compatible with similar databases held by other conservation
agencies. It will place a high priority on participation in the National
Biodiversity Network. The Society will encourage the exchange of data
on fungi with European mycological societies.
Education: Officers of the Society will publicise the conservation
activities of the Society, will promote educational training in fungal
conservation and encourage other bodies to include consideration of
fungi in their conservation and management activities. The Society will
press for a programme of training to be set up for commercial
collectors, restaurateurs and other traders in wild fungi in order to
encourage responsible collecting and consideration of the importance of
the long-term sustainability of the resource.
Collaboration: The Society will work with the European Council for the
Conservation of Fungi, the International Union for Conservation of
Nature and Natural Resources and the Berne Convention to seek
protection for mycological habitats of European importance through
the implementation of appropriate government land-use policies.
Opportunities will be investigated for the financing of international
fungus conservation research programmes. The Society will organise an
international symposium in 1999 in order to review the future of fungal
conservation in the 21st century.
Review: The Society will encourage its members to inform the
Conservation Special Interest Committee of publication of new research
relating to fungal conservation, of impending changes in legislation on
environmental matters and of any modifications of policy by statutory
or voluntary conservation agencies.
Consultations continued through 1997 and 1998 with English Nature,
The Woodland Trust, The National Trust and the Forestry Commission,
leading towards the publication of two leaflets entitled The Wild Mushroom Pickers Code of Conduct and The Conservation of Wild Mushrooms
(Anon., 1998b,c).
The BMS input to these leaflets, which was crystallised over many years
from the contributions of numerous individuals and committees can be
paraphrased as follows.
British Mycological Society: collecting fungi from the wild, a code of
conduct
Beauty, intrigue and value. All these can be found amongst the fungi.
Exotic displays of mushrooms and toadstools in our woodlands and
pastures, particularly during the autumn, are as much a part of our
natural heritage as the more commonly appreciated displays of wild
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flowers. And the flushes of fungi can provide as much aesthetic appeal
as their cousins, the flowers.
But fungi are also important for their medical and industrial value.
They are sources of life-saving drugs and other products in daily use by
us all. But more than this: fungi are essential components of natural
habitats like woodlands, meadows and pastures and without them the
ecology of these places could not function.
Fungi of all sizes are used as food by a range of animals. Mammals
like squirrels, voles, deer, and we, of course, take the larger fruit bodies.
Indeed, these provide homes as well as nutrient to many invertebrates
(worms, insects, slugs), some of which may be rare or endangered
species in their own right.
We collect fungi too. Some for scientific study and identification;
some for screening for new drugs; but most, perhaps, for food use by
the collectors themselves or for sale to groceries and restaurants.
We treat wild fungi as a natural harvest, the hunter-gatherer’s
perquisite! Which is all well and good, but humans don’t have a
particularly good track record for reasonable exploitation of such
natural resources. The intention of this Code is to minimise the adverse
impacts of our collecting activities. We want to maximise enjoyment of
fungi whilst minimising the damage we might cause to other wildlife and
wild places, to fungal populations in nature and, indeed, to landowners,
site managers and other collectors.
Need for a code
Mycologists (people who study fungi) have known for many years that
populations of wild fungi are subjected to the same adverse pressures as
are other wildlife by our modern life style. This has led to the
compilation of Red Data Lists of threatened species in the hope that
recognising the threat will assist in their protection.
Right across Europe there has been a decline in the fruiting of some
wild fungi. A particularly worrying aspect of this is that it is especially
true for mycorrhizal mushrooms − which form an intimate symbiotic
association with living tree roots − and which cannot yet be cultivated.
Two popular edible mushrooms belong to this category: the Cep,
Boletus edulis, and the Chanterelle, Cantharellus cibarius. The main
causes of the decline appear to be loss of habitat and air pollution, and
edible and inedible species of mushroom are equally affected.
But there are other pressures. Increasing popularity of ‘field
mushrooms’ and ‘wild fungi’ on the menus of fashionable restaurants
and in recipes of fashionable cookery books has increased the
profitability of collecting from native populations. Collecting for
commercial gain has become more common and is often concentrated
on a few areas known to yield good harvests and has led to inevitable
resentment and conflict with local residents and landowners.
The aim of this Code is to recommend good practice for all those who
collecting fungi from the wild.
British Mycological Society conservation policy
237
Principles
A number of common-sense principles underlie this Code:
∑ Scientific research has not detected any ecological damage arising
directly from harvesting fungi, even on a large scale for commercial
purposes. BUT, such studies are in their early stages. What IS clear is
that the act of collecting can cause collateral damage by the effects of
trampling, disturbance, removal and even destruction of natural areas.
∑ Mushrooms, toadstools and other fruiting structures are only the visible
spore-producing bodies of the fungus, like the fruits of our orchard
trees. The bulk of the fungus exists below ground as the fungal
mycelium. Removing a mushroom may do no more damage than taking
an apple from the tree (even though dispersal of spores, like discharge
of plant seeds, is necessary for sexual reproduction). But trampling the
mycelium to death can cause untold damage to the existing population.
∑ While we are so ignorant of the biological requirements of many fungi
are not, it would be prudent to be cautious in our exploitation of
natural resources. We must not risk long-term damage by ignorant
exercise of our ‘right to collect’.
∑ Nevertheless, wild fungi must be seen as a legitimate resource to be
harvested. But we must ensure that harvesting is sustainable and does
no damage either to the fungal population or to populations of
organisms associated with the fungi.
Scientific study of fungi, such as recording their occurrence,
sometimes requires removal of fruit bodies, even when the study is
aimed at conservation. The reason is that proper identification may
need microscope observations, or even molecular analyses.
Fungi and the law
Two main laws may protect fungi.
The Theft Act (1968), which applies only to England and Wales,
makes it an offence to:
∑ Dig up and take any plant, tree or shrub or any soil etc. which is part of
the land, being the property of somebody, so digging up fungi could
constitute theft unless the landowner has given permission.
∑ Take the property of somebody and sell it for gain. The emphasis here is
on selling for gain. The custom of taking wild fruit and flowers,
including fungi, is permitted by the Act so long as there is no personal
financial gain. Sale of collected fungi without the landowner’s
permission may be an offence.
The Wildlife and Countryside Act (1981), and a similar law in Northern
Ireland, makes it an offence to:
∑ Pick, uproot, destroy or sell, and/or collect and cut any plant listed on
Schedule 8 attached to the Act − and that schedule now includes some
species of fungi.
∑ Uproot any wild plant, unless the person is authorised.
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David Moore
Remember also that special protection restricting collection of fungi
may apply to Sites of Special Scientific Interest (SSSIs), designated
under the Wildlife & Countryside Act, and National Nature Reserves
(NNRs).
Local Bylaws forbidding the picking of fungi and plants on National
Trust property, Crown Estates, Local Nature Reserves, Forestry
Commission and Local Authority land may also apply.
The fungus pickers’ Code
General guidelines
Many of the recommendations below are simply common courtesy or
countryside etiquette. Remember that we share the environment with
many other organisms and interests and so are the custodians of our
natural heritage.
Before entering any land, get the landowner’s or site manager’s
permission, explaining the purpose of your visit. Check for Bylaws
concerning picking of fungi. Collecting is not allowed on some SSSIs,
Nature reserves and other protected areas.
Follow the Country Code
Be sensitive to the structure of the natural habitat. Do not damage
vegetation or soil, nor disturb unnecessarily leaf litter or other features.
Do not move or remove dead wood unless essential to identify a fungus.
Collecting for food
Be VERY aware that some fungi contain deadly toxins and many more
may make you unwell. Some people suffer allergic reactions after eating
particular fungi, even though they may be well known as ‘edible’.
Many mushrooms can concentrate heavy metals so do not collect in
heavily polluted areas. This warning applies to the verges of busy roads
(lead pollution) and to reclaimed land sited on old landfill waste dumps
(where the danger is of industrial heavy metals like cadmium and
mercury).
Do not collect large numbers of specimens that you don’t recognise
on the off chance that some might be edible.
Some edible species have poisonous look-alikes. Beginners should not
eat anything that has not been checked by an experienced field
mycologist. And remember that mainland Europe, where amateur
mushroom hunting is a popular pastime, has the highest reported
incidence of mushroom poisoning in the world!
Respect, and protect, all species, including poisonous ones.
Never collect rare or Red Data list species − and certainly not the
species that are scheduled in the Wildlife and Countryside Act (listed
below).
Even if the population is plentiful, take no more than you need for
your personal consumption or use.
Collecting with a view to selling for profit or other commercial use
British Mycological Society conservation policy
239
must be agreed beforehand with the landowner.
Generally, never take more than half of the fruit bodies you can see.
Generally, never collect unopened or ‘button’ mushrooms. Giving the
fruit bodies time to expand will allow spores to be discharged and result
in bigger, perhaps tastier, mushrooms to be picked later.
Scientific collecting
Collect the minimum amount of material or number of specimens
required for proper description and reliable identification.
Minimise the disruptive effects of taking samples. For example,
replace rolled-over logs and avoid the unsightly damage caused by
cutting chunks out of long-lived bracket fungi.
Always offer the results of your fieldwork to the landowner or site
manager, with explanation of the significance of your findings.
Record localities and habitat data for rare species accurately and
retain dried ‘voucher specimens’ for deposit in herbarium collections.
Supply information to local and national databases. Remember that
science needs to be communicated!
If you have permission to collect for scientific purposes do not abuse
it by collecting edible fungi for eating ‘on the side’!
The following fungi are protected under the Amendment to Schedule
8 of the Wildlife & Countryside Act, 1981:
Sandy Stilt Puffball, Battarrea phalloides − inedible.
Royal Bolete, Boletus regius − edible.
Oak Polypore, Buglossoporus pulvinus − inedible.
Hedgehog Fungus or Lion’s Mane, Hericium erinaceum − edible.
. . . but you can’t please all of the people all of the time
One of the most remarkable, and unexpected, aspects of the reaction to
publication of the Code of Conduct was a thunderous editorial in the Daily
Telegraph of 31 August 1998 headlined ‘Mushrooming bureaucracy’. An
accompanying article by the newspaper’s Environment Editor, Charles
Clover, gave a fairly measured account, though it bore the challenging
headline ‘Country code to curb the mushroom picker’. However, the
editorial started off ‘After Greenpeace, green police? This week will see the
publication of one of those increasingly familiar ‘codes of conduct’ which
tell you what to do in areas where we have managed well enough without
rules. This one is to control mushroom picking, for heaven’s sake, and it
promises to provide splendid new opportunities for minor-league jobsworths whose hobby is bossing us about.’ This gem of the journalist’s
trade ended ‘Leave us to pick our mushrooms in peace, and if we wish to
sell them, that is our own affair.’
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In some quarters, then, the Code of Conduct might well be seen as overly
challenging, but this could reflect the attitudes of at least a few of those
involved in bringing together the contributions of different organisations.
Certainly, during the discussion phase a representative of one of the
partner organisations with which we were trying to establish a joint
compromise approach e-mailed the BMS Conservation Officer stating ‘I
would remind you that much of your text . . . is the intellectual property of
K--------, M----- and I, and you do not have our consent to use it . . . ’ −
not a very helpful stance for one’s co-authors! Nevertheless, careful political navigation around the jealously self-protective (the ‘jobsworths’ in the
eyes of the Daily Telegraph’s Editor) can, and in this case did, lead to a
satisfactory conclusion − a document agreed by all parties concerned
which could actually help the situation.
Dissemination of the advice and continued education is the key to
further progress. It is clearly not good enough to limit our targets to the
mycological community. The message must be spread to other concerned
naturalists and then to the wider lay audience, especially managing and
legislating authorities. This is something a national Society can, and
should, take into its responsibility.
References
Anon. (1990). British Mycological Society policy on conservation. Mycologist 4,
52.
Anon. (1998a). British Mycological Society policy on conservation. Mycologist
12, 35.
Anon. (1998b). The Wild Mushroom Pickers Code of Conduct. English Nature:
Peterborough, UK.
Anon. (1998c). The Conservation of Wild Mushrooms. English Nature:
Peterborough, UK.
Arnolds, E. (1988a). Dynamics of macrofungi in two moist heathlands in
Drenthe, The Netherlands. Acta Botanica Neerlandica 37, 291–305.
Arnolds, E. (1988b). The changing macromycete flora in The Netherlands.
Transactions of the British Mycological Society 90, 391–406.
Eveling, D. W., Wilson, R. N., Gillespie, E. S. & Bataillé, A. (1990).
Environmental effects on sporocarp counts over fourteen years in a forest
area. Mycological Research 94, 998–1002.
Frankland, J. C., Magan, N. & Gadd, G. M. (1996). Fungi and Environmental
Change. Cambridge University Press: Cambridge, UK.
Hedger, J., Lodge, D. J., Dickson, G., Gitay, H., Læssøe, T & Watling, R.
(1995). The BMS expedition to Cuyabeno, Ecuador: and introduction.
Mycologist 9, 146–148.
Ing, B. (1992). A provisional Red Data list of British Fungi. Mycologist 6,
124–128.
Jansen, E. & van Dobben, H. F. (1987). Is decline of Cantharellus cibarius in The
British Mycological Society conservation policy
241
Netherlands due to air pollution? Ambio 16, 211–213.
Kirby, K. J. (1988). The conservation of fungi in Britain. Mycologist 2, 5–7.
Minter, D. W. (1986a). Fungus recording. Computerization of foray records.
Bulletin of the British Mycological Society 20, 34–38.
Minter, D. W. (1986b). Fungus recording. Foray records database. Further
information for users. Bulletin of the British Mycological Society 20,
101–105.
Nitare, J. (1988). Skydd av hotade svampar, svensk och internationellt arbete
[Protection of endangered fungi, Swedish and international work].
Jordstjärnan 9, 25–33.
Pegler, D. N., Spooner, B. M. & Young, T. W. K. (1993). A Revision of British
Hypogeous Fungi. Royal Botanic Gardens: Kew.
Ramsbottom, J. (1948). The British Mycological Society. Transactions of the
British Mycological Society 30, 1–12.
Richardson, M. J. (1970). Studies of Russula emetica and other agarics in a Scots
pine plantation. Transactions of the British Mycological Society 55, 217–229.
Rotheroe, M. (1998). Wild fungi and the controversy over collecting for the pot.
British Wildlife 9 (6), 349–356.
Webster, J. (1997). The British Mycological Society, 1896–1996. Mycological
Research 101, 1153–1178.
Wilkins, W. H. & Harris, G. C. M. (1946). The ecology of the larger fungi. V. An
investigation into the influence of rainfall and temperature on the seasonal
production of fungi in a beechwood and a pinewood. Annals of Applied
Biology 33, 179–188.
Wilkins, W. H. & Patrick, S. H. M. (1940). The ecology of the larger fungi. IV.
The seasonal frequency of grassland fungi with special reference to the
influence of environmental factors. Annals of Applied Biology 27, 17–34.
21
The contribution of national
mycological societies: The Dutch
Mycological Society and its Committee
for Fungi and Nature Conservation
M A R I J K E M . N A U T A & L EO M . J A L I N K
The Dutch Mycological Society
The Dutch Mycological Society was set up in 1908 and the object of the
Society is to promote the study and understanding of fungi. This is done by
organising meetings and field excursions, and by publication of the Society’s journal, Coolia. The Society has a number of Committees, of which
the Committee for Fungi and Nature Conservation deals with all aspects
of conservation of fungi and nature management.
Apart from the yearly meeting in which all matters concerning the
organisation of the Society are discussed, two general days with lectures
are organised each year: one with a special theme organised by the Scientific Committee and a more general one organised by the Society itself.
These meetings are very well attended, and at the general meeting in
January it is often the case that 170 of the approximately 500 members
attend.
In the mushroom season at least every weekend and often also during
the week field excursions are organised for members. Every year two
week-long forays are organised, one inland and one abroad. A good policy
concerning conservation of fungi is impossible without knowledge of their
distribution. Therefore, we will first briefly discuss the activities of the
Dutch Mycological Society in the study of fungal distribution.
Distribution of fungi: recording
The Society is very active in gathering information about the distribution
of fungi in The Netherlands. From its early days, the Dutch mycologist
Frencken kept lists of records of field excursions. In the 1960s mycologists
became aware of a change in mycota in The Netherlands (Bas, 1959, 1978),
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Dutch Mycological Society Conservation Committee
243
and assessment of this was one of the motives behind the conception of a
national recording project in 1980. This project aimed at improving knowledge of the distribution and ecology of (macro)fungi. Because of organisational and financial obstacles it was not until 1984 that the project ran well.
Since a recording project cannot work sufficiently without a checklist of
the species to be recorded, the Standaardlijst van Nederlandse macrofungi
was published in 1984 (Arnolds, 1984). This checklist contained an annotated list of the macrofungi to be expected in The Netherlands, and was
based on literature, data from private and institutional herbaria and expert
judgement. In relation to the publication of this preliminary Red Data list
the Dutch Mycological Society organised a meeting in 1984 on the changes
in the mycota of The Netherlands. At this meeting a great deal of scientific
and statistical evidence for the decline of macrofungi was presented (Arnolds, 1985).
The recording project was a collaboration of the Dutch Mycological
Society, the Institute of Forestry and Nature management, the Biological
Station Wijster and the IKC Naturemanagement, and was financially
supported by the Ministry of Agriculture, Nature Management and Fisheries. At present only the Dutch Mycological Society runs the project. The
database now contains over a million records. Besides records from the
recording project, volunteers have entered many data of older date into the
database, derived from literature and herbaria. Many members of the
Society, as well as naturalists from outside the Society, contribute to the
database and field excursions are often planned in areas of The Netherlands which are under-recorded.
Recently, a monitoring project has been started, organised by the Dutch
Mycological Society and the Central Bureau for Statistics, also financed by
the Information and Knowledge Centre Naturemanagement (IKCN).
This is aiming to monitor a limited number of rare and threatened species
in woodlands on sandy soil (Arnolds & Veerkamp, 1999). This project is
based on yearly counts in permanent plots of 1000 m and hopefully will
permit an evaluation of fungi in woodlands that is statistically more
reliable than before.
Important publications on distribution of fungi
The mapping project has enabled publication of a new checklist (Arnolds,
Kuyper & Noordeloos, 1995), which was written by 24 authors, including
both amateur and professional mycologists. This new checklist contained
data on nomenclature, distribution, rarity and ecology of the recorded
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Marijke M. Nauta & Leo M. Jalink
macrofungi. Also, after publication of the preliminary Red Data list in
1989 (Arnolds, 1989), it has been possible to compile a new Red Data list
of threatened fungi applying the official guidelines of the International
Union for Conservation of Nature and natural resources (IUCN), by the
direction of the Ministry of Agriculture, Nature Management and Fisheries (Arnolds & Kuyper, 1996; Arnolds & van Ommering, 1996). A digital
version of the new checklist with addition of the most recent Red Data list
data is included in BioBase (Anon., 1997), a CD-ROM with coded data on
many groups of organisms.
Another important publication was the distribution atlas containing
detailed information of a selected number of 375 taxa on distribution,
ecology and phenology and their changes (Nauta & Vellinga, 1995). The
data allowed statistical analysis and showed once more the serious decline
in macrofungi in The Netherlands. The project was made possible by a
grant from the Ministry of Agriculture, Nature Management and Fisheries
which was awarded to the Dutch Mycological Society.
Conservation of fungi
The Committee for Fungi and Nature Conservation gives advice and
answers questions concerning conservation of fungi. When Environmental
Impact Studies are initiated for projects in which nature is affected, the
Committee verifies whether data are available concerning fungi and tries to
ensure that no important habitats are destroyed. Popularising fungi and
emphasising the important role fungi play in nature is one of the ways to
achieve a better understanding and protection for fungi. For this purpose a
popular booklet of 20 pages was published in 1989 (Anon., 1989).
At present the most important project of the Committee for Fungi and
Nature Conservation is the project ‘Mycological Reserves’, in which a
better protection for mycologically valuable sites is one of the goals. The
procedure and background of this project are discussed in Chapter 6. The
major tool for conservation of fungi in The Netherlands is the Red list. By
counting the number of species occurring in a site or using a weightingfactor according to the amount of threat experienced by a species, the
mycological value of a site can be established. By means of giving specific
management advice to people or councils dealing with management it aims
to protect and improve the mycota.
In contrast to other countries (see, among others, Chapter 18) protection of fungi is solely tackled through protection of habitats and sites, since
it is believed that fungi themselves can hardly be protected independently
Dutch Mycological Society Conservation Committee
245
of their habitat. An exception is the case where larger woodland fungi grow
on trees that are threatened with removal but, then, even this is habitat
destruction. No biodiversity action plans for fungi exist in The Netherlands. Instead, protection of fungi is pursued by promoting understanding
of fungi at all management levels and by indicating threatened habitats
and sites.
Collecting fungi
In The Netherlands collecting fungi has also become an issue for discussion. In some nature reserves commercial picking is experienced as a
problem (Anon., 2000). In principle, the Society wants to stimulate study
of fungi but is not encouraging people to pick large quantities of mushrooms. Direct damaging effects to the mycelia by picking the fruit bodies
has never been proven, but in an overcrowded country like The Netherlands the visual damage and the indirect damage resulting from trampling
or disturbance of the forest floor is too great to encourage harvesting of
fungi. The Society takes the position that fungi should only be collected in
limited quantities when it is necessary for scientific, identification or educational purposes. In most nature reserves, strict rules already prevent the
picking of mushrooms, and field excursions aiming at recording of fungi
need special permission to collect the necessary specimens for identification.
References
Anon. (1989). Paddestoelen. Commissie Paddestoelen en Natuurbeheer,
Nederlandse Mycologische Vereniging: Wijster.
Anon. (1997). BioBase 1997, Register Biodiversiteit. Centraal Bureau voor de
Statistiek: Voorburg/Heerlen.
Anon. (2000). Grootschalige paddestoelenpluk, een toenemend probleem? Coolia
43, 45–46.
Arnolds, E. (1984). Standaardlijst van Nederlandse macrofungi. Supplement,
Coolia 26.
Arnolds, E. (1985). Veranderingen in de paddestoelenflora (Mycoflora).
Wetenschappelijke Mededelingen van de Koninklijke Nederlandse
Natuurhistorische Vereniging 167, 1–101.
Arnolds, E. (1989). A preliminary Red Data List of macrofungi in The
Netherlands. Persoonia 14, 77–125.
Arnolds, E., Kuyper, T. W. & Noordeloos, M. E. (1995). Overzicht van de
paddestoelen in Nederland. Nederlandse Mycologische Vereniging: Wijster.
Arnolds, E. & Kuyper, T. W. (1996). Bedreigde en kwetsbare paddestoelen in
Nederland. Basisrapport met voorstel voor de rode lijst. Nederlandse
Mycologische Vereniging/Biologisch Station LUW: Wijster.
246
Marijke M. Nauta & Leo M. Jalink
Arnolds, E. & van Ommering, G. (1996). Bedreigde en kwetsbare paddestoelen in
Nederland, Toelichting op de Rode lijst. IKCN, Wageningen.
Arnolds, E. & Veerkamp, M. T. (1999). Handleiding Paddestoelenmonitoring.
Nederlandse Mycologische Vereniging, Baarn.
Bas, C. (1959). Verdwijnende landschappen en hun fungi. Coolia 6, 7–8.
Bas, C. (1978). Veranderingen in de paddestoelenflora. Coolia 21, 98–104.
Nauta, M. M. & Vellinga, E. C. (1995). Atlas van Nederlandse paddestoelen. A. A.
Balkema Uitgevers B.V.: Rotterdam.
22
Fungal conservation in the 21st century:
optimism and pessimism for the future
DA V I D MO O R E , M A R I JK E M. N A U T A , S H E LL E Y
E. E V A N S & M A U R I C E R O TH ER O E
This book has examined a variety of problems associated with fungal
conservation. We have tried to go beyond mere debate by including
constructive guidance for management of nature in ways favourable to
fungi. The geographical range of the examples presented is from Finland in
the North to Kenya in the South, and from Washington State, USA, in the
West to Fujian Province, China, in the East. Our authors suggest solutions
that are equally wide ranging: from voluntary agreements, through ‘fungus-favourable’ land management techniques and on to primary legislation. Taken together, the book offers practical advice on how to include
fungi in conservation projects in a range of circumstances.
The book has its origin in a British Mycological Society Symposium at
the Royal Botanic Gardens in Kew, UK, in November 1999, combined
with a selection of papers delivered at the XIIIth Congress of European
Mycologists, held in Alcala de Henares, Spain, in September 1999. There
were lively discussions at both meetings but verbal discussion is difficult to
represent in writing. We felt it was important, however, to attempt to
convey some feeling of the discussions that occurred during the meetings,
so we asked contributors to respond, briefly, to the question ‘Are you
optimistic or pessimistic about fungal conservation in the 21st century?’
Our contributors replied from their different standpoints and their
replies are given here without editorial interference from us. The replies are
arranged in the same order as the authors’ chapters in this book and the
opinions expressed are those of the individual authors.
247
248
D. Moore, M. M. Nauta, S. E. Evans & M. Rotheroe
‘Are you optimistic or pessimistic about fungal conservation in the
21st century?’
From Régis Courtecuisse, Lille, France
I can say that I am reasonably optimistic, in the sense that more and more
people, especially those involved in nature management (foresters, nature
managers, officials from government or nongovernment organisations)
turn out to be interested and concerned with the role of fungi in ecosystems. Some examples of positive impact of fungal conservation (in The
Netherlands for example) are encouraging to go further in this research
and activity. Of course, this good impression on the future of fungal
conservation is counterbalanced by the global threat to the environment,
especially to the forests world-wide. These global problems (forest cutting
as well as global pollution on the other hand) must be also considered and
addressed, even if it is evidently more difficult to hope for a serious
improvement in the situation from this point of view. Local (I mean at
nonglobal scale) actions for fungal conservation and for the conservation
of their natural habitats may be successful, especially in the perspective of
future improved collaborations with nature management professionals of
any kind. That is encouraging, but the global environmental problems will
probably go further in degradation and threat. What makes me still
optimistic, despite this globally pessimistic remark, is the actual possibility
to improve the collective and federated actions and efforts, among mycologists but also among naturalists and nature managers. That’s what I’ll try
to do and organise in the near future, as chair of the ECCF and the
Specialist Group for Fungi (SSC/IUCN).
From Randy Molina, Corvallis, Oregon, USA
Overall, I am optimistic regarding fungal conservation, but mycologists
have a lot of work to do in educating the public, government management
agencies, and other biologists as to why and how. One pitfall to avoid is
advocating ‘single species management’ because there are simply too many
fungal species for this approach. Instead, our focus should be on maintaining or building habitat at the appropriate scale to provide for the development and normal function of fungal guilds and communities. To do so we
must aim our field research at questions and scales relevant to conservation issues. These include successional development of fungal communities, disturbance thresholds and adaptations, habitat preferences, and
ecological amplitude. Fundamental work is also needed at the population
Optimism and pessimism for the future
249
level to understand how we can incorporate genetic data into our concepts
of fungal species viability. Mycologists must begin to think more broadly
in scale and scope in regards to fungal conservation. We must become
better versed in the quickly growing field of conservation biology, publish
our work in international conservation journals, and look for opportunities to work with other conservation biologists and field ecologists in
integrated studies with common goals. Mycology cannot stand alone in its
pursuit of species conservation. Success will only come about when we
demonstrate and communicate the great importance of maintaining fungal
populations to the sustainability of our world’s diverse ecosystems.
From Eef Arnolds, Beilen, The Netherlands
I am rather optimistic about the fungi in forest communities in Europe.
There is an international trend to maintain remnants of old-growth forests
and to create new strict reserves. The exploitation of forests tends to
become less rigorous and more often based on ecological principles and
durability. On the other hand, continued economic growth may have a
negative impact on environmental quality that is harmful for many fungi
by expansion of polluted areas. I am pessimistic about the future of fungi
in other habitats. Intensification of agriculture will have a strong negative
impact on the environmental quality and the existence of semi-natural
habitats. Expansion of the EU will lead to great losses in the rural habitats
in eastern Europe. These developments may be compensated by increased
efforts for conservation of valuable habitats, but only in part. However,
the greatest losses will take place in the tropics during an ever increasing
process of habitat destruction, stimulated by the modern religion of the
free market, invented in the rich countries. I have great concern about the
future of field-mycology as a scientific discipline in Europe. Without
experts we shall simply not know what happens to the mycota.
From Erast Parmasto, Tartu, Estonia
Very optimistic. Fungal species will be included in the European Red Data
list soon – maybe some few species in the beginning, a few more later.
Fungi will be mentioned in international conventions. It will take much
more time (maybe, a dozen of years), but hopefully European mycologists
will agree which species will be there in a European Red List (for ‘internal
use’); 25, 50 or 100 species will be selected then for distribution mapping.
Actual or ‘true’ conservation is mainly possible in nature reserves. It
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D. Moore, M. M. Nauta, S. E. Evans & M. Rotheroe
depends on mycologists themselves: are they interested to take part in
all-species inventories of protected areas? Do they care to spend some time
for discussions with managers of reserves? Have they something to propose to them for better protection of habitats rich in fungi? Do they
quarrel with ornitholocracy- or ornithologarchy-minded persons on equal
rights of all endangered living beings to be protected? Do they dare to send
articles on fungi to newspapers and popular scientific journals, to participate in TV shows in a studio or even in a forest stand? Are they ready to
lobby MPs for selecting a State Mushroom (as we have State Flowers in
many countries), using for this action of the legislature? Fate of fungal
conservation is in our own hands.
From Leo Jalink & Marijke Nauta, National Herbarium of The
Netherlands
We are moderately optimistic for the situation in The Netherlands. There
is more and more attention for fungi in management plans now, and the
general (changed) approach to nature conservation is usually beneficial for
fungi as well. Of course, on the other hand there is an ongoing habitat
destruction, and some habitats are still severely threatened (nutrient-poor
grasslands, for example) but, at least, more attention is being paid to fungi.
From David Arora, Santa Cruz, California, USA
Since I didn’t hear the discussion, and am about to leave for the airport, I
will just have to leave it at, ‘Optimistic, providing habitat is conserved . . . ’.
From Siu Wai Chiu, Chinese University of Hong Kong, China
China is one of the megadiversity countries and is also one of the most
highly densely populated countries by man. Another unique feature of
China is its diverse geography and thus the fauna, flora and mycota are
very rich. However, there is always a conflict between human interests and
the protection of natural resources especially when the present status of
China is still that of a developing and poor country. However, the Government plays an active role in policy making, practising and running the
conservation plans. With the continual and expanding international effort
in conserving the natural habitats in China, the international demand for
natural products from sustainable industry, the open door policy in improving communication and education with the outside world, the open-
Optimism and pessimism for the future
251
ness in accepting and assimilating favourable culture by the Chinese,
China is expected to actively participate in environmental protection. As
the Chinese have the oldest history of mushroom cultivation, with over 30
fungal species being used in traditional herbal medicines, and the country
is now the champion mushroom producer and exporter, the Chinese
people will definitely guard, protect and explore the native fungal diversity.
For fungal diversity in this biosphere, although habitat loss is still a major
problem for species extinction, fungi being mainly saprotrophs, could
survive in various artificial habitats. With the increasing interest and
recognition of the potential for exploitation of fungi, and a global view
emphasising sustainable development, humans will definitely make greater
efforts to conserve biodiversity, including fungal biodiversity, into the
twenty-first century.
From Maurice Rotheroe, Lampeter, South Wales
Like some other contributors to this chapter, I am ambivalent. The ‘cons’
seem to be just as compelling as the ‘pros’. I share many of my colleagues’
misgivings about the global future of fungal conservation, and any possible stemming of the tide of ‘progress’. But if I confine my remarks to the
British Isles, then the outlook is considerably more optimistic. Conservation is now established as a priority on the agenda of the British Mycological Society. Since 1996, the Society has had a Conservation Special Interest
Committee. The Conservation SIC is now one of the two most active
committees of the Society. Closer co-operation with CABI Bioscience and
with the Mycological Herbarium of the Royal Botanic Gardens, Kew, has
played a part in the establishment of a BMS database which is rapidly
increasing in size and is in constant use as a resource. Similarly, the Society
has given its backing to the ambitious Kew project to produce a new
Checklist of British fungi. Both of these products are essential tools to the
fungal conservationist. Concurrently with these developments, the Society
has forged a strong link with the new National Botanic Garden of Wales.
It has given £5000 of its Conservation Fund to the creation and curation of
a mycological herbarium at the new garden and the funding of work which
will eventually lead to publication of a Mycota Cymru (Fungus Flora of
Wales). Discussions are proceeding on the setting up of a so-called
‘Mycodome’ on the NBG Wales site, to interpret and celebrate the concept
‘Fungi Mould Our Lives’. The BMS is well represented on the steering
committee of this unique £2 million international initiative. If it becomes a
reality, fungal conservation will be a major element in its brief. When I
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D. Moore, M. M. Nauta, S. E. Evans & M. Rotheroe
became BMS Conservation Officer in 1996, my ‘manifesto’, was strongly
based on the idea of communication – of greater collaboration and
contact with other biological disciplines and with governmental and nongovernmental environmental agencies, as well as between mycologists,
within and outside the BMS. All of us appeared to be working towards a
common goal but, with a few notable exceptions, there was little communication or co-operation between those involved, while routine information
exchange was almost nonexistent. This situation has changed considerably
in the past five years. Government agencies and NGOs are involved in a
wide variety of different initiatives aimed at improving the representation
of fungi in the biodiversity picture. There are now some 30 local recording
groups well dispersed over the British Isles, who are not only recording and
mapping but also liaising with local environmental agencies. In more
recent times a new effective and extremely businesslike player has moved
confidently into the fungus conservation arena. Plantlife, the wild-plant
conservation society, set up Plantlife Link in 1992 in order to ‘advance
plant conservation by facilitating the exchange of information between
groups involved in plant conservation and to provide the context for
strategic planning and co-operative venture’. Fungi were regarded as
‘honorary plants’, so the forum included representatives from the BMS,
the Association of British Fungus Groups and several individual mycologists. Because fungi began to feature more and more on agendas, an
offshoot, the Fungus Conservation Forum was formed in 1999. This
committee already has a large and active agenda and fulfils my ‘manifesto’
ambitions admirably because it puts fungal conservation at the heart of the
thinking of so many different environmental agencies. I see Plantlife and its
dynamic links as being a very important catalyst in advancing the cause of
fungal conservation in Britain, certainly in the early years of the twentyfirst century.
From Martin Allison, Bedfordshire, UK
Most fungal population declines and extinctions across Northern Europe
are accounted for by problems deep-rooted in the fabric of modem society.
The insatiable demand for fossil-fuelled energy produces unacceptable
levels of atmospheric pollutants. The ever-increasing human population
creates new housing schemes spilling onto green-field sites. Intensive farming regimes controlled by inappropriate agricultural policies cause massive
declines of once-common farmland birds, insects, flowers and fungi. The
logical conclusion might well be that the future looks dismal, for wildlife
Optimism and pessimism for the future
253
and for us. Is there room for optimism within such a bleak landscape? I
believe there is. Coming from a background of bird conservation I have
heard many success stories over the years. The osprey’s return to the
Scottish Highlands, the avocets to Minsmere, red kites back in England.
Extensive tracts of heathland and reedbed restored, increasing populations
of rare species such as woodlark, Dartford warbler, Cetti’s warbler, bittern
and bearded tit. All this has been achieved through dedicated and enthusiastic conservationists, backed by sound scientific research and increasingly
more meaningful legislation. Fungal conservation has progressed in leaps
and bounds over a very short period of time. There are now Red Data lists
for most European countries, the most-threatened fungi are appearing
within biodiversity action plans, and in the UK the habitat and survival
requirements of rare fungi are being addressed by bodies such as Plantlife,
English Nature, the RSPB and of course the British Mycological Society.
To cap this, early in the New Year the UK Government announced a
long-needed new Bill to secure better legal protection for wild plants and
animals. The mere fact that this volume on Fungal Conservation exists
gives some hope for the future. Through research, education, lobbying and
that all-important ‘in-the-field’ enthusiasm many of the mycological goals
should eventually be within reach.
From Giuseppe Venturella, Palermo, Italy
I am really optimistic about the possibility that the safeguarding actions
for the mycota that need to be done in many European countries will be
achieved in the twenty-first century. If this desirable goal is to be achieved,
though, I think it is necessary to co-ordinate planning among the different
Associations and Institutions working towards the conservation of fungi,
such as BMS, ECCF, OPTIMA, CEMM, Planta Europa, Amateur Mycological Associations, etc.
From Heikki Kotiranta, Finnish Environment Institute, Finland
The intensive survey of forests done in Finland over the last ten years led to
protection of large areas of old-growth forests. Old-growth forests in
National Parks, nature reserves and some other areas are now protected by
law and incorporated in nature conservation programmes. The new forest
cutting recommendations, that always some amount of wood (often old,
large or already dead trees) is left to the managed area, may help some
species to survive even in commercially managed forests. However, most
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D. Moore, M. M. Nauta, S. E. Evans & M. Rotheroe
of the protected areas are situated in the northern part of the country, and
the area protected in southern Finland is still too small, so further actions
in southern Finland are required.
From Paul Cannon, CABI Bioscience, Surrey, UK
There are opportunities for both optimism and pessimism. On the positive
side, conservation policy is changing dramatically from individual speciesoriented approaches to ecosystem preservation, resulting in the conservation of many fungi by default. The value of fungi to the environment and
ecological processes is being more widely appreciated, and the role of
pathogenic species in the natural environment is being reassessed. The
potential is bright for sensitive and efficient detection and monitoring of
individual species during the next century, with ever more sophisticated
molecular tools being developed. Pessimists are likely to be rewarded also.
The dramatically increasing pressure on land, especially on natural habitats in the tropics, is probably causing fungal extinctions on a daily basis.
Sustainable exploitation of natural resources will continue to be aspiration
rather than reality as economic pressures force short-term decision-making. Global warming will pose added challenges for species survival. More
specifically, funding for systematics shows little sign of reversing its downward trend, despite its critical role in identification and increased emphasis
on the environment following the Rio conference. The continuing lack of
interaction between fungal systematics experts and ecologists is also depressing, especially in areas such as sampling and monitoring procedures.
Most fungal survey work is still carried out on an unscientific serendipitous
basis, which makes comparison of results almost impossible. The key to
ensuring a prominent role for fungi in conservation is to promote interdisciplinary links between scientists, and a holistic vision of fungi and their
myriad roles in the ecosystem.
From Vincent Fleming, Peterborough, UK
We should be optimistic, for the UK anyhow. There is now increasing
recognition that fungi are subject to the same pressures as, and are of equal
worth to, other wildlife. This change in attitude can be attributed to the
recent emphasis on biodiversity and it marks a radical shift from earlier
assumptions that the conservation of fungi would, at best, have to depend
on their chance representation in sites specially protected for other, more
familiar, interests. The financial resources directed to fungi by conserva-
Optimism and pessimism for the future
255
tion agencies have also increased, yet fungi still lag behind many other
taxonomic groups. Realistically, they are likely to continue to do so: we
have a shortage of specialists, a paucity of time-series data from which to
estimate trends, and little knowledge of how to manipulate populations.
To achieve maximum gain, we should seek to identify and highlight those
circumstances where the needs of fungi differ from a ‘normal’ conservation
approach. However, we should not expect to achieve total knowledge nor
to delay action until we do so. We must use the best available information
to guide and influence positive action and to stimulate public interest. For
parallels and parity, we should look less to vascular plants and more to the
conservation of invertebrates, with which fungi share many similarities:
namely, the large numbers of taxa, difficulties of identification, and the
transient appearance of observable life-stages. We have a considerable
task ahead and optimism does not guarantee success, but to be pessimistic
is not to try.
From Alison Dyke, Edinburgh, Scotland
From a Scottish perspective, and particularly on the subject of harvesting
wild mushrooms, the future of fungal conservation seems fairly rosy.
There is real interest in fungal conservation among all those involved in
harvesting wild mushrooms – after all for some it is their livelihood and
they do have a long term perspective. Effort is being made to survey
important habitats and species groups, e.g. waxcap grasslands and tooth
fungi. There are obviously some areas of concern, where important habitats are being lost, where the funds to do research are hard to come by, etc.
It is these areas where it is important that positive action is taken and it is
everyone’s responsibility to ensure that this happens.
Index
Abies 19
abundance 37, 105
Acanthophysium 36
acid rain 226
action plans 201, 245
Aecidium 205
Agaricus 157, 158
Agenda 21 210
agriculture 182
airborne fungi 188
Albatrellus 36
Aleurodiscus 86
allergen 188
Alnus 19
Alpova 36, 97, 100
Amanita 157–160
amenities 21
Amylocorticium 86
anamorph 199, 205
ancient woodland 148
ant-nest symbionts 200
Aphyllophorales 177–181
Arbutus 19
Arcangeliella 36, 97, 100
Arctostaphylos 19
Armillaria 214, 215
artificial logs 117
Asterodon 86
Asterophora 36
Astraeus 159
Aureoboletus 159
Baeospora 36
Balsamia 36, 97, 98
Bankera 148, 149
Barssia 97–99, 102
basidiospore dispersal 117
Battarrea 214, 227
BIG (Biodiversity Information Group)
210
256
biodeterioration 188, 191
Biodiversity Action Plan 209–218
bioindicators 14
biological records 166, 167
biotechnology 191
biotrophic 201, 203, 204
birds of prey 24
BMS (British Mycological Society) 121,
217, 223, 251, 253
boletes 33
Boletopsis 214
Boletus 36, 47, 86, 108, 148, 149, 153, 154,
157–160, 214, 216, 217, 236
Bondarzewia 36, 76
bracket fungi 152
Bridgeoporus 36
British Council 168, 171
Bryoglossum 36
Buglossoporus 214, 217
Byssocorticium 151
C/N cycling 199
calcicolous grassland 121
calcifugous grassland 121
Calocybe 160
Calvatia 109
Cantharellula 150
Cantharellus 33, 36, 47, 107, 108, 159, 160,
236
Castinopsis 19
Catathelasma 36
CCW (Countryside Council for Wales) 120
cep 236
Cercospora 205
Ceriporia 86
Chalciporus 36
Chamaecyparis 40
Chamonixia 36, 97, 100
champions 210, 213
change in land use 70
257
Index
chanterelle 50, 107, 236
charcoal 158
checklist 10, 67, 90, 167, 184, 194, 243
CHEG profile 125
China 106, 111–118
Chinese forests 112
Choiromyces 36, 97, 98, 158
Chromosera 36
Chroogomphus 36
Chrysomphalina 36
Clavaria 74, 121, 129, 138, 150
Clavariadelphus 36, 86, 159
clavarioid 120, 138–140
Clavicorona 36
Clavulina 36
Clavulinopsis 121, 129, 138, 139
climate 45, 128, 228
Clitocybe 36, 157
coastal dunes 136
coastal forest 19
code of conduct 219, 235, 238, 239
coelomycetes 204
Colletotrichum 202, 204
Collybia 36, 158
commercial harvesting 9, 21, 22, 26,
46–49, 52, 53, 105–110, 145, 153, 156,
158, 160, 219–222, 226, 231, 232, 236, 245
communities 25–28, 30, 68, 213
computer records 169
computing resources 164
conservation efforts 54
conservation policy 223–241
conservation value 123, 125, 127
contact point 210, 213
Convention on Biological Diversity 209
coppice 153
Coprinus 158
Cordyceps 36
Coriolopsis 159
Cortinarius 36, 148, 149, 151, 157, 159
Corylus 19, 149
Country Group 210
Crataegus 140
Craterellus 36, 108
cryptic species 21
Cryptomeliola 204
Cuba 182–196
Cudonia 36
cultivated strains 113, 114
cultivation method 111
culture collections 13, 160, 172, 183, 192,
198
CWD (coarse woody debris) 27
Cyphellostereum 36
Darwin Initiative 185, 192, 200, 202
dead wood 152
deforestation 112, 184
degradation 188, 191, 199
Dentipellis 86
Dermocybe 36
Dermoloma 129
Destuntzia 36
DFID (Department for International
Development) 162
Dichostereum 36
Diospyros 205
Diplomitoporus 86
directory of scientists 169
distribution 67, 68, 72, 92, 147, 168, 200,
215, 225, 242–244, 249
disturbance 27, 99, 102, 227
diversity 22, 29, 30, 32, 35, 37, 69, 73, 77,
85, 109, 114, 118, 134, 142, 154, 156, 163,
184, 185, 187, 197, 199, 206, 209, 211,
213, 245, 251, 252
dune slacks 137
dunes 147
Dutch Mycological Society 89, 242–246
Earth Summit 209
earth-tongues 120, 136–143, 150
ECCF (European Council for the
Conservation of Fungi) 7, 64, 68, 75,
95
ecological corridors 12
ecology 68, 99
ecosystem functions 23
ecosystem management 22
ectomycorrhiza 19, 23, 33, 45, 49, 51, 52,
69, 95, 145, 148, 152
education 118, 190, 240, 248, 253
Elaphomyces 36, 95, 97, 98
Elasmomyces 97, 100
endangered species 81, 128, 130, 201, 202
Endangered Species Act 20
Endogone 36
endophyte 187, 200–202
enoki 109
Enterographa 217
Entoloma 36, 74, 120–143, 149, 150, 153,
157–159
environmental quality 14
Eocronartium 86
ephemeral fruit bodies 42
epigeous fungus 33
epiphytes 201
Estonia 81
Eucalyptus 158
ex situ conservation 160, 198
extinct hypogeous species 102
exudates 23
Faerberia 158
258
Index
fairy-club 120, 121
Fayodia 36
Fevansia 36
fieldwork 168
fire 27, 45, 48
fire suppression 29, 30
Fischerula 97, 98
Fomes 148
Fomitopsis 86
food webs 21, 24, 29, 200
foray 223, 224
forest disease 21
forest felling 8
forest fungi 19, 22, 23, 26
forest gaps 24
forest management 20, 25, 29
forest products 21
forest soil 200
Fremitomyces 204
Fujian Province 114, 115
fungal habitat 42
fungal records 187, 225
fungivores 24
fungus linkages 25
Galerina 36
Ganoderma 86, 159
Gastroboletus 36
Gastrosporium 97, 100
Gastrosuillus 36
Gautieria 36, 97, 100
Geastrum 86, 151, 159
Gelatinodiscus 36, 40
gene flow 31, 32
gene pool 114, 118
Genea 97, 98
genet 32, 116, 117
genetic drift 31
genetic homogeneity 114
genetic isolation 34
genetic variation 32, 33
Geoglossum 74, 138, 150
geographic scales 32
Geopora 97, 98, 157
global trade 106
Gloeophyllum 178
Glomerella 204
Glomus 36
Gomphus 36, 86
grassland 74, 120–143, 145, 147, 150, 215,
217, 228, 250
grassland soil 200
grassland types 121, 127, 129, 133
grazing 140–142, 153
grazing, cattle 136
grazing, rabbit 134
grazing, sheep 134, 136
green lists 172
greenhouse effect 8
grey lists 172
Grifola 109, 159
Guangdong Province 114, 115
guidelines 54, 93, 103, 133, 134, 141, 142,
181, 221, 229, 244
Guignardia 202
Gymnomyces 36
Gymnopilus 36
Gyromitra 36
Gyroporus 158
habitat 75, 86, 95, 99, 148
habitat action plans 210, 217
habitat conservation 11, 103, 184, 250
habitat destruction 8, 69, 249, 250
habitat indices 42
habitat loss 232
habitat management 38, 103
habitat manipulation 49
habitat modelling 42–45
habitat plans 211
habitat protection 244
habitat scale 248
HAP (Habitat Action Plan) 210
Hapalopilus 86
health audits 202
heartwood 152
heathland 150, 253
Hebeloma 36, 66, 157, 158
hedgehog mushroom 108
Helvella 36, 158
hen of the woods 109
Hericium 86, 149, 154, 159, 214, 217
heritage value 10
Hippophae 140
Histoplasma 189
historic sites 40
Hubei Province 114, 115
Hydnangium 97, 100
hydnelloid 216
Hydnellum 86, 149
Hydnobolites 97, 98
hydnoid species 149
Hydnotrya 36, 97, 98
Hydnum 36, 47, 108, 149
Hydrobotrya 98
Hydropus 36
Hygrocybe 66, 74, 120–143, 150, 153, 154,
159, 214, 215
Hygrophorus 36, 151, 153, 154, 158, 159
Hymenogaster 97, 100
hyphal length in nature 199, 200
hyphomycetes 204
Hypocreopsis 214
hypogeous fungi 24, 33, 44, 95–103, 227
Index
Hypomyces 36, 108
Hysterangium 97, 100
Idaho 19
indicator species 14, 84–86, 127, 128, 132,
225
indicator-species profile 125
industrial mycology 191
information storage 166
Inocybe 157
interior forest 19
inventory 10, 35, 38, 42, 82, 83
ITE (Institute of Terrestrial Ecology) 152
IUCN specialist group 13
IUCN threat category criteria 11, 70, 72,
75, 177, 179, 203, 244
Jiangxi Province 114, 115
Kenya 197–208
key habitats 210
king bolete 108
labile information 165
Lactarius 108, 140, 148, 151, 157–159
land management 48, 54, 154, 228, 247
land use 145
land use change 9
land-owners 222
landscape scales 31, 34
Larix 19
laying yard 112
LBAP (Local Biodiversity Action Plan)
210
lead partner 210, 213
leaf spots 205
Leccinum 148, 157
legislation 8, 20, 21, 160, 173, 189, 191,
237, 239, 247, 253
Lentinula 111–118
Lepiota 151, 159
Lepista 157
Leptoporus 86
Leucangium 47, 109
Leucogaster 36, 97, 100
Leucopaxillus 158, 159
lichen 187, 199
lignin decomposition 199
limestone hills 182
Lindtneria 86
Lithocarpus 19
Lobelia 203
lobster mushroom 108
long list 210
Lucanus 217
machinery 153
259
Macowanites 36
Macrotyphula 158
management 21, 30, 31, 36, 47, 48, 93,
122, 142, 152, 153, 160, 226, 229, 247,
250
management guidelines 54, 93, 103, 133,
134, 141, 142, 181, 221, 229, 244
management history 124
management plans 89, 90
mangrove swamps 182
mapping 10, 67, 68, 168, 226, 243, 249
Marasmius 36
Martellia 36, 97, 100
matsutake 50, 106, 107
medical mycology 191
Melanogaster 97, 100
Melanoleuca 158
Meliola 204
mesotrophic grassland 121
microarthropods 24
microfungi 197–208
Microglossum 129, 131, 138, 150, 214,
215
middle list 210
migrations 8
mire 121
modelling habitat 42
molecular techniques 28
monitoring 35, 103
Morchella 47, 107
morel 50, 107
mowing 136, 141, 142, 153
Multiclavula 86
multispore spawn 117
mushroom 24, 29, 33, 47
Mushroom Code 221
Mutinus 159
mutualisms 21
mycelium 199
Mycena 36, 148, 150, 151, 157
Mycodome 251
mycological reserves 12, 90, 244
mycological sites, The Netherlands 92
mycophagy 33
mycorrhiza 25–28, 30, 73, 154, 187, 200,
202, 236
mycoses 188
mycotoxicoses 188
Myriostoma 158, 159
Mythicomyces 36
natural products 200
natural selection 31
nature management 136, 248
nature managers 93
nature reserves 89, 99, 144–155, 151, 181,
189, 198, 227, 245, 249
260
Index
NBGW (National Botanic Garden of
Wales) 123, 251
necrotrophic 201–203
Neolentinus 36
Neournula 36
Nivatogastrium 36
nomenclature 126
northern spotted owl 20, 29
Northwest Forest Plan 20, 21
nutrient cycling 21, 23
nutrient retention 23
NVC (National Vegetation Classification)
120, 121, 127, 152
Octavianina 36, 97, 100
old-growth forests 20, 22, 29, 35, 69, 73,
75, 81, 82, 84, 180, 181, 249, 253
old-growth grassland 74
old-growth woodland 92
Oligoporus 86
Ombrophila 148
Omphalina 36
Omphalotus 158, 159
optimism 247–255
Oregon 19
organic carbon sinks 23
Ossicaulus 158
Otidea 36
Oudemansiella 159
over-picking 226
Overseas Forays 224
Oxyporus 76
oyster mushroom 109
Pachyphloeus 97, 98
Panaeolus 140, 141
parasites 201
pasture 153
PCR–RFLP 28, 30
peasant-farmers 111
Perenniporia 86
pessimism 247–255
Peziza 141
Phaeocollybia 36
Phaeolus 86
pharmaceuticals 200
Phellinus 86
Phellodon 36, 86, 148, 149, 159
Phlebia 86
Pholiota 36, 158
Phomopsis 202
Physisporinus 86
Picea 19, 99
picking wild mushrooms 9, 21, 22, 26,
46–49, 52, 53, 105–110, 145, 153, 156,
158, 160, 219–222, 226, 231, 232, 236,
245
Picoa 97, 98
pine forest 147, 148, 153
Pinus 19
Pisolithus 159
Pithya 36
plant pathogens 24
plantations 150
Plectania 36
Pleurotus 158, 160
Pluteus 151
Podostroma 36
pollution 8, 9, 52, 69, 70, 145, 152, 184,
225, 228, 232, 252
Polyozellus 36
polysaccharide exudates 23
ponderosa pine 30
population 31–33, 68, 252, 255
population biology 22, 30, 113
population structure 116
Populus 19
Poronia 214
portabello 109
powdery mildews 205
primary threats 180
priority species 212
priority species or habitat 210
productivity 50, 51
Protea 203
protected areas 189
protection 103, 158, 160
Pseudaleuria 36
Pseudomerulius 86
Pseudotsuga 19
public awareness 190
puffball 109
Pulcherricium 159
Pycnoporellus 86
Quercus 19
raising yard 112
rakes 158
Ramaria 36, 159
Ramariopsis 129
ramet 32
random mating 34
rare species 30, 35, 38–43, 69, 81, 84, 85,
89, 91, 128, 130, 145, 158, 197, 200, 205,
224, 225, 228
recommendations 181, 229
red (rarity, endangerment and distribution)
data lists 11, 35, 37, 64–67, 70–72, 74,
84, 121, 128, 130, 145, 146, 148, 158, 160,
172, 177, 227, 229, 231, 249, 253
red data lists, European 71, 96–103
red data list, Idaho 37
red data list, Estonia 86
Index
261
red data list, Oregon 37
red data list, The Netherlands 90–92, 243,
244
reproduction strategies 69
rhizomorphs 23
Rhizopogon 33, 36, 97, 100
Rhodocybe 36
Rickenella 36
Rigidoporus 86
risk status 37
Rozites 148
RSPB (Royal Society for the Protection of
Birds) 144–155
RSPB reserves (map) 147
rural development 106
rural economy 105
Russula 36, 108, 148, 151, 157
Sphaerosoma 97, 98
spore dispersal 33
SSC (Species Survival Commission) 7
SSSI (Site of Special Scientific Interest)
120
Stagnicola 36
Steccherinum 86
Stephanospora 97
Stephensia 97, 98
Strobilomyces 149, 151, 159
succession 26
Suillus 33
survey 21, 28, 36, 39–41, 120, 134, 203,
216, 253
sustainable management 26, 48, 50, 52,
109, 118, 219, 250
symbionts 27
Salix 19
sampling 31, 41, 42
SAP (Species Action Plan) 210
Sarcodon 36, 86, 149
Sarcosoma 36, 86
Sarcosphaera 36
Schizophyllum 157
Sclerogaster 97
Scotland 219–222
second-hand computers 162
Sedecula 36
Senecio 203
Serpula 86, 178
Shaanxi Province 114, 115
shiang-gu 109, 111–118
shiitake 109, 111–118
short list 210
Sichuan Province 106
Sicily 156–161
silent protection 103
silviculture 29, 30, 51, 53
Sistotrema 86
site monitoring 103
Skeletocutis 86
small mammal 24, 29, 44
snowbanks 40
soil fungi 189
soil structure 23
somatic compatibility 117
sooty moulds 204
Sowerbyella 36, 86
Sparassis 36, 108
Spathularia 36
spatial scales 32, 34
spawn production 113
species action plans 211, 214
species concepts 66
species of conservation concern 210
species statement 210, 212
taxonomists 10, 39
taxonomy 38, 66
teleomorph 199, 204
Tephrocybe 158
Thaxterogaster 36
The Netherlands 242–246
threat categories 11, 98, 100, 178
threatened habitat 72
threatened species 70, 96, 97, 128, 130,144,
145, 180
threats 184
Tibet 106
timber harvest 20, 27, 45, 47, 107
timber management 152
Tomentella 86
tourism 184, 194
traditional technology 111
training 39
Trametes 179
trampling 227
Tremiscus 36, 86
Trichoglossum 138, 150
Tricholoma 36, 47, 107, 148, 157–159
Tricholomopsis 36
tropical expeditions 224
truffle 24, 29, 35, 95, 103, 107
Truffle Hunt 224
Tsuga 19
Tuber 36, 47, 97, 98, 109
Tulostoma 159, 214, 215
Tylopilus 36
UK BG (UK Biodiversity Group) 210
UK Darwin Initiative 163, 167, 168, 170
Ukraine 162–176
UNESCO biosphere reserve 189
United Kingdom 209–218
USDA (US Department of Agriculture)
20, 22
262
Index
USDI (US Department of the Interior)
20
voluntary agreements 247
voluntary code 219
Volvariella 151
Wakefieldia 97
Wales 120–135
Washington 19
waxcaps 120–143, 150, 215, 217
wild mushroom harvesting, issues 220
wild mushrooms 9, 21, 22, 26, 46–49, 52,
53, 105–110, 145, 153, 156, 158, 160,
219–222, 226, 231, 232, 236, 245
woodlands 147
woody debris 27, 30, 35, 45, 83
Xylaria 86
Xylobolus 86
Yunnan Province 106, 114, 115