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impacted on mycology. There are many reasons for this, but the diversity of fungi* is so

great, and so much of it is as yet unknown, that a habitat or community approach has to

be a surrogate for species conservation in fungi. Yet we remain far from the point where

a community- or ecosystem-based system for fungi can be recommended for general use

in conservation criteria or legislation at international, regional, and national levels.

The issue requires a consensual and transparent approach that could in time be

adopted or endorsed by bodies such as IUCN–The World Conservation Union and the

Subsidiary Body for Scientific, Technical and Technological Advice (SBSTTA) of the

Convention on Biological Diversity. This vision is probably at least a decade to realization.

Here we can but point to constraints, provide an eclectic view of fungal diversity across

biomes and ecological niches, discuss correlations with plant and animal diversity and

forces driving and influencing fungal diversity, and point to ways we might proceed.



2.2



CONSTRAINTS



2.2.1

Circumscription

A key problem in a community approach to fungal diversity and distribution is their

interdependence with other organisms. With the exception of fungi that form lichens,

fungi are not primary producers; in consequence, they cannot form separate self-sustaining communities, and their occurrence is irrevocably linked with that of organisms on

which they depend for their nutrition. A further complication is that many fungi are found

outside natural vegetation systems, occurring on particular cultivated crops or garden

plants or as biodeteriogens and contaminants of foodstuffs and manufactured goods.

Consequently, any attempt to circumscribe fungal communities in an independent manner,

parallel to that used in botany, and without regard to the organisms on which they depend,

is surely unrealistic.

2.2.2

Mycosociology

Mycosociology, the study and classification of fungal communities in their own right, has

had few advocates. Höfler (1938) and Hueck (1953) considered that fungal communities

could be named independently because they were dependent on factors other than those

that controlled the plant communities within which they occurred, but few have endeavored

to take this further. The problems have been aired by Apinis (1972) and Barkman (1973)

in particular and need not be repeated here.

In the case of macromycetes, the only author to assiduously endeavor to introduce

a formal system for naming fungal communities independently of the plants with which

they were associated was Darimont (1975), who attempted to lay down a formal system

of mycosociological nomenclature to apply to data he had collated from woodlands in

Belgium, for example, using the suffixes -ecea (class; e.g., Cortinario-Boletacea), -ecia

(order; e.g., Boleto-Amanitecia), -ecion (alliance; e.g., Boletacion scabri), and -ecium

(sociomycie, equivalent to the association of phytosociologists; e.g., Amanitecium muscariae). He recognized 24 sociomycies in the Belgian woodlands, 18 alliances, 8 orders, and

4 classes. The approach does not appear to have been taken up subsequently.



* The term fungi is used throughout this chapter to embrace all organisms that belong to the kingdom

Fungi together with others traditionally studied by mycologists, i.e., lichens, slime molds, straminiples (oomycetes), and yeasts, as well as mushrooms and molds.



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Some European lichenologists, however, enthusiastically adopted phytosociological

principles and developed systems for the formal naming of communities regardless of the

plant communities in which they occur (e.g., Barkman, 1958; Wirth, 1972). Consequently,

a staggering number of Latinized community names for lichen assemblages have been

proposed (Delzenne-van Haluwyn, 1976). This is not so surprising, as lichens are the

predominant biomass in some ecological situations, such as on rock surfaces and on the

ground in boreal and arctic-alpine situations, and those on bark can be more related to its

chemical characteristics and the ambient environment (e.g., relative humidity, temperature)

than the trees involved. Provided that a broad-brush approach is adopted, the use of a

hierarchy of Latinized names can provide both a useful framework for the description and

a shorthand method for communicating complex concepts of assemblages for lichenologists (James et al., 1977). Community names such as Lobarion and Xanthorion are

consequently in widespread use in the 21st century, even though few workers now devote

time to characterizing and describing lichen communities in the formal manner required

by the International Code of Phytosociological Nomenclature (Barkman et al., 1986).*

The approach is unlikely to ever be fully implemented for nonlichenized fungi in

view of the problems in thoroughly recording what species may be present in a particular

site. However, as most other fungi are an integral functional part of plant-dominated

communities (Dighton, 2003), this is surely not an inappropriate outcome of the debate

as to whether fungal communities in general should be named in a formal manner. Because

of issues of differing spatial scale and processes of structuring species assemblages

between plants and fungi, invoking the concept of synusia (a grouping, within one layer

of a community, of species characterized by similar life-forms and similar ecological

requirements), or assemblage, may be of use in describing fungal species assemblages,

e.g., epiphyllous ascomycetes of oak leaves, soil microfungi in particular microhabitats,

and gallery beetle associate fungi.

2.2.3

Recording

The first issue to be considered in endeavoring to survey the fungi in a particular site is

the number of different ecological niches in which they can occur and that have to be

searched if a total inventory is to be attained. This is a major constraint in view of both

the number of niches meriting scrutiny (Table 2.1) and the different expertise and methodologies required to examine the species present in many of those niches (Rossman et

al., 1998; Mueller et al., 2004). The problem is then compounded by seasonality or

periodicity in the production of visible fruiting bodies. In some cases fruit bodies are

ephemeral and may last for only a few hours, while in others the same species may not

produce fruit bodies every year, or even decade. For example, in a 21-year study of forest

plots in Switzerland, Straatsma et al. (2001) encountered fruit bodies of 408 species.

However, the number recorded in each year, even after repeated visits, ranged from 18 to

194, with 19 species not previously encountered at all found in the last year of study.

The time necessary to produce a definitive list requires long-term commitment and

investment. The two most intensively studied sites to date are Esher Common (Surrey,

U.K.) and the Slapton Ley Nature Reserve (Devon, U.K.), with 2900 and 2500 species

recorded, respectively (Cannon et al., 2001). Yet while both sites have been studied for



* For a brief introduction to the complex procedures and history of the formal naming of communities

in a mycological (and lichenological) context and further early references on this topic, see

Hawksworth (1974).



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Table 2.1 Principle Niches and Microhabitats Occupied by Fungi

Living Vascular Plants

Biotrophs and necrotrophs of leaves, stems, flowers, fruits, seeds, roots, etc.

Commensals on bark and leaves (especially lichen-forming fungi)

Endophytes of leaves, stems, bark, and roots

Secondary colonizers of dead attached tissues and leaf spots, etc.

Mycorrhizas (endo-, ecto-, ericoid, orchid, etc.)

Leaf surfaces

Nectar

Resin

Dead Vascular Plants

Saprobes on wood, bark, and litter

Burnt plant tissues

Saprobes on submerged and inundated plants

Pollen in water samples

Nonvascular Plants

Algae (marine, terrestrial, and freshwater)

Bryophytes

Fungi

Biotrophs, necrotrophs, and saprobes of other fungi

Lichenicolous fungi

Myxomyceticolous fungi

Vertebrates

Skin, feathers, hair, bone, etc.

Dung

Nests, lairs, etc.

Ruminant guts

Fish scales and guts

Invertebrates

Biotrophs and nectrotrophs

Arthropod exoskeletons

Arthropod and annelid guts

Nematodes

Insect nests

Rock

Lichens

Epilithic fungi

Endolithic fungi

Soil

Surface

Soil cores

Water

Foam

Streams, permanent and temporary ponds

Litter and wood immersed in sea- and freshwater

Plants (e.g., bromeliads)

Adapted from Hawksworth et al. (1997) and Hyde and Hawksworth (1997).



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Fungal Communities: Their Diversity and Distribution



31



over 25 years by numerous mycologists to reach these figures, only 40% of the species

are in common despite many similarities between the localities, and additional species

are added each year. The actual number of species of fungi present in these sites could

easily be around 4000, but both are much disturbed and neither has a long history of

ecological continuity.

In the case of the Guanacaste Conservation Area in Costa Rica, an international

working group estimated that the number of fungi to be found would be around 50,000

species and that an inventory would cost U.S.$10 to 30 million over 7 years, depending

on the level to which identifications were made (Rossman et al., 1998). However, even if

ample funds were available, the shortage of available specialists in many groups of fungi

would pose a major hurdle to such an intensive recording project.

Data quality of records is also a problem, as a significant proportion of reports of

species from a particular community made by nonspecialists may be unreliable due to

insufficiently critical determinations being made. Further, in many cases dried or cultured

voucher material is not preserved in institutions such as herbaria, museums, and collections

of fungus cultures, rendering many records of questionable long-term value (Agerer et al.,

2000). Additionally, many countries do not yet have fungus recording schemes in place,

and those that do are generally underfunded.

Finally, until the recent publication of recommended standardized sampling methodologies (Mueller et al., 2004), there had been little progress toward the adoption and

universal use of recommended and internationally accepted standard protocols for sampling fungi in particular ecological niches, thus making comparisons of data sets from,

for example, soil and leaf isolations, difficult. Similarly, suggestions to focus on target

groups of fungi as surrogates for overall species richness (Hawksworth et al., 1997) have

hardly progressed.

2.2.4

Diversity

The suggestion that fungi are an exceptionally diverse and poorly known group of organisms with around 1.5 million fungi on Earth, of which only 74,000 to 120,000 have so

far been identified (Hawksworth, 2001), continues to be supported by fresh analyses

(Heykoop et al., 2003; Mueller et al., in press). Schmit and Mueller (in press) conservatively estimate a minimum of 600,000 species worldwide based on the ratios of fungalto-plant species in well-studied regions and taking into account data on endemism. This

conservative figure was calculated to establish a lower boundary for the number of fungal

species, which will be revised upward as more information becomes available. Whatever

the final figure may prove to be, there is no doubt that the magnitude of the species numbers

present in any detailed community study poses special problems, in that species that are

as yet unnamed are likely to be encountered, especially when working in hitherto littleexplored ecological niches or geographical locations.

Further, there is a lack of modern monographic revisions and keys for many groups

of fungi, which makes the identifications necessary for community characterization particularly time-consuming.

2.2.5

Species Concepts

The species concepts traditionally used in different fungal groups vary, depending on which

characteristics are considered important. However, incompatibility studies and the advent

of molecular phylogenetic approaches are increasingly showing that in many cases, what

has traditionally been interpreted as a single species on morphological criteria alone is in

reality a complex of biologically and evolutionary distinct species. This is as true for lichenforming fungi (Grube and Kroken, 2000) as it is for macromycetes (Petersen and Hughes,



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Hawksworth and Mueller



1999) and plant pathogens (O’Donnell et al., 2000). At the extreme end of changes in a

group’s species richness based on molecular data are suggestions that the diversity of

arbuscular mycorrhizal fungi may be closer to hundreds of thousands of species rather than

the traditional view of hundreds of species (Fitter, 2003). It is clear that the formal recognition of such species will increasingly become the norm in mycology (Taylor et al., 2000).

While these so-called cryptic species are evidently a major component of the estimated

unrecognized global species numbers of fungi, their existence poses major problems for

the documentation of fungal communities. The need to distinguish species that are so similar

morphologically that they cannot be conclusively identified without cultural or molecular

studies will inevitably hinder critical survey work of all kinds in mycology.

2.2.6

Fallacies

Several fallacies impinge on endeavors to characterize fungal communities, three of which

are deeply embedded in the minds of many biologists:

1.



2.



3.



“Everything is everywhere and the environment selects” (Baas-Becking, 1934).

While this may be true for some saprobic, soil, and opportunistic spoilage microfungi with enormous potential for dispersal (Gams, 1992), it clearly does not

apply to the huge numbers of host-specific fungi, many macrofungi (see below),

or lichen-forming species. This is especially so when species are studied at the

cryptic species and population levels, where species thought to have widespread

distributions prove to be complexes of two or more distinct taxa (see above).

Most fungi occur in damp places (countries). May (1994) cautioned against

extrapolations based on data from countries such as the U.K., which were “damp

and fungal place[s]” from an Australian viewpoint. This may apply to some

groups of fungi, but cannot be supported as a general rule as the most speciesrich habitats may vary from region to region. For example, in Australia huge

numbers of microfungi are found associated with native species (e.g., Eucalyptus; Sankaran et al., 1995), macrofungal diversity is high (T.W. May, personal

communication), and the continent appears to be a gold mine for undescribed

hypogeous macromycetes (Claridge et al., 2000; J. Trappe, personal communication) and certain groups of rock-inhabiting lichens (e.g., Xanthoparmelia; Elix

et al., 1986; Elix, 2003).

Yeasts and lichens are not fungi. Old concepts die hard, and even to this day it

is not uncommon to see phrases such as “yeasts and fungi” and “lichens and

fungi,” which are oxymorons.* Yeasts are firmly part of the kingdom Fungi,

and strictly lichens have no names; the names used are those ruled as applying

to the fungal partner (the photosynthetic symbionts maintaining independent

names; Hawksworth, 1997a). It is especially surprisingly to find lichens treated

in a series of floras (e.g., Flora of Australia), while the “other” fungi have an

independent sister series (e.g., Fungi of Australia).



In addition, the historical inclusion of mycology within botany, and also the treatment

of fungi as a kind of cryptogam or lower plant, has been deeply damaging to the perception,

organization, and development of the subject (Hawksworth, 1997b).



* Oxymoron: “A figure of speech by means of which contradictory terms are combined,” literally

“pointedly foolish” (Kirkpatrick, 1983).



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2.3



33



DIVERSITY ACROSS BIOMES



Fungi are so diverse that it is difficult for any single person to address the issue of diversity

across biomes. However, Lodge et al. (1995) approached this by soliciting the opinions

of a range of mycologists with experience in more than one continent or hemisphere. Most

of those consulted who worked on basidiomycetes felt diversity was more correlated with

host and habitat than resource abundance, while those whose expertise was with ascomycetes (and their hyphomycete anamorphs) considered all three factors important. The

areas most frequently mentioned as sources of novel unknown taxa were humid forests

on islands, tropical mountaintops, and large tropical river basins. However, some hostrestricted heterobasidiomycetes occurred in all areas and habitats with the hosts, while

ranges in many agarics appeared to be limited regardless of the region. Overall, diversity

in most groups, except rusts and smuts, was judged to be greatest in the tropics and

subtropics and most strongly related to habitat and host diversity. While this was a

refreshing way to visit the overall issue, it was essentially qualitative rather than quantitative, a valuable source of hypotheses based on impressions that merit testing by substantial data sets. Mueller et al. (in press) pooled data on diversity and distribution patterns

for macrofungi from different geographical regions. They compiled 21,679 names during

this study and found that the percentage of unique names varied from 37% for temperate

Asia to 72% for Australasia. No comparable data set for other fungal groups has been

compiled and analyzed, and fresh and broader studies on the lines of those undertaken by

Pirozynski and Weresub (1979) are to be commended.

A different approach was taken by Schmit et al. (2005). They undertook a metadata

analysis of published and available unpublished point diversity studies of macrofungi that

included species lists of the plants in the sampled plots. Macrofungal species included in

the analyzed studies displayed neither larger nor smaller species ranges than the plants in

the data set, and not surprisingly, the diversity of macrofungi in each site was high. While

this study documented that tree diversity proved to be a good predictor of macrofungal

diversity at each site, plant community data could not be used as a surrogate to predict

macrofungal community composition.

Nevertheless, it appears from casual studies of the available information that some

generalizations as to the major differences in the diversity of fungi and the communities

developed in different biomes can be made (Table 2.2). Estimating species numbers has

been attempted by extrapolation from the numbers of vascular plant species present

(Rodríguez, 2000). While such approximations are open to debate, they do suggest that

much of the species diversity in fungi is in the tropics and remains to be discovered

(Table 2.3).

Such broad-brush approaches clearly mask differences in the diversity of fungal

communities on a niche-by-niche (cf. Table 2.1) basis. For example, despite the considerable uniformity reported among soil microfungi (Gams, 1992), studies of communities

in soils subject to different degrees of climatic stress suggest that the proportion of sexually

reproducing species is positively correlated with increasing stress (Grishkan et al., 2003).



2.4



HOW TO PROCEED



The study of fungi as an integral part of plant communities and ecosystems, or as associates

of particular animal or plant hosts and providers of essential ecological services, rather

than in isolation, should be the underlying feature of future studies in fungal ecology; the

approach of Dighton (2003) is commendable in this respect. The issues of diversity and



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Hawksworth and Mueller



Table 2.2 Examples of Ecological Fungal Groups Predominating in Major Biomes

Biome



Fungal Groups



Arctic, Antarctic, arctic-alpine,

montane, northern tundra

Boreal

Temperate

Mediterranean and hot desert

Tropical



Lichen-forming fungi

Lichen-forming and ectomycorrhizal fungi

Plant-specific (especially rusts and smuts) and

ectomycorrhizal fungi; also slime molds

Lichen-forming and soil fungi (including hypogeous

macrofungi)

Foliicolous fungi (including sooty molds, asterines,

meliolines, fungicolous spp., foliicolous lichens), ostropalean

lichen-forming fungi (including graphids and thelotremes),

entomogenous fungi, endomycorrhizal fungi, and endophytic

fungi



Table 2.3 Approximate Numbers of Fungi (including Slime Molds, Lichen-Forming

Fungi, Straminipilous Fungi, and Yeasts) and Plantsa (Seed Plants and Ferns) Known

from Different Regions of the World



Region

Asia

Europe

Africa

North America

Central and South America

Oceania

Antarctica

Global

a



Described

Species

20,000

25,000

10,000

21,000

10,000

6,000

750

72,000



(70,000)

(12,000)

(60,000)

(18,000)

(85,000)

(17,000)

(2)



Estimated Total

Species

600,000 (77,000)

65,000 (12,000)

450,000 (67,000)

250,000 (18,000)

500,000 (100,000)

250,000 (21,000)

1,750 (2)

1,470,000



Percentage

Unknown

>95 (10)

60 (>1)

>95 (10)

>90 (1)

>95 (15)

>95 (20)

55 (0)

95



Plant figures are in parentheses.



Adapted from Rodríguez (2000).



complexity in communities are so immense, however, that small-scale case studies

designed to test particular hypotheses are likely to be keys to the understanding of patterns

and interrelationships on a global scale. Ideally, such studies should employ similar

protocols in different biomes and communities across wide geographical regions, and also

be integrated with more holistic ecosystem studies by broad-based plant and animal

ecologists. Since the mid-1990s there has been a heightened awareness of the need for

multidisciplinary approaches to ecosystem functioning, and fungal data have been taken

note of in key debates and syntheses (Schulze and Mooney, 1994; Brussaard et al., 1997;

Freckman et al., 1997). Regrettably, the funds to conduct large-scale new studies to address



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Fungal Communities: Their Diversity and Distribution



35



key issues of the structure and maintenance of ecosystem function at the international

level have remained elusive. In the interim, mycologists with an interest in the diversity

of fungi and their roles in communities should endeavor to collaborate with one another

wherever possible, in order to maximize the potential inputs that can be made to the

elucidation of our understanding of fungal community ecology. The synthesis of issues

and protocols by Mueller et al. (2004) should be a major stimulus to such an approach.



ACKNOWLEDGMENT

Support to David L. Hawksworth from the Programa Ramón y Cajal of the Ministerio de

Ciencia y Tecnología through the Facultad de Farmacia of the Universidad Complutense

de Madrid is gratefully acknowledged. Gregory M. Mueller acknowledges the financial

support of the U.S. National Science Foundation.



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