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Research review
Blackwell Science Ltd

Community ecology of ectomycorrhizal

fungi: an advancing interdisciplinary field

Author for correspondence: Anders Dahlberg

Tel: + 46 (0) 18 67 27 94
Department of Forest Mycology and Pathology, PO Box 7026, Swedish University of Agricultural
Fax: + 46 (0) 18 30 92 45
Email: Anders.dahlberg@ mykopat.slu.se Sciences, S-750 07 Uppsala, SWEDEN

Received: 13 December 2000

Accepted: 16 February 2001

Summary

Key words: ectomycorrhizal
community ecology, mycorrhizas, plant
ecology, ecosystem processes.
A long-term goal of community ecology is to identify spatial and temporal factors
that underlie observed community structures. Ultimately, ecologists seek to relate
community patterns to ecosystem processes and functions. Since the mid 1990s,
ectomycorrhizal (ECM) research has been equipped with tools to identify and fully
quantify the taxonomic diversity in below-ground ECM fungal communities in detail
and address such questions. Many of the most important functions of terrestrial ecosystems,
as well as interactions, between plants take place below ground and mycorrhizal fungi
are among the key players in soil ecology. Here the rapidly increasing knowledge of
ECM fungal community ecology is reviewed and the prospects discussed for elucidating
processes that structure ECM fungal communities and the way in which such knowledge
might be integrated with, and advance, the understanding of plant ecology and
ecosystem processes.

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membrane interfaces, within chloroplasts mitochondria

Introduction
Mutualistic associations between organisms, which reciprocally
increase the fitness of both partners, have been a significant driving
force in the course of evolution. The importance of endosymbiotic
associations between different procaryotes in the evolution
of eucaryotic cells is now firmly established and there is also
increasing recognition that symbiosis at the level of more
complex organisms is the rule, rather than the exception (Smith
& Douglas, 1987). An ancient and ecologically important
example is the mycorrhizal symbiosis between vascular plants
and soil fungi dating back to the first land plants (Blackwell,
2000). However, despite major advances in our understanding
of mycorrhizas, the extent and significance of mycorrhizal root
symbioses remains greatly underrated in plant ecology. In fact,
vascular plants can be regarded as successful entrepreneurs
that have benefited from symbiotic association with procaryotes
and later fungi to form energy and nutrient transferring biological
and root-fungal organs, the mycorrhizas. Effective membrane
surface area is significantly increased through invagination
and external enlargement, and in the case of the mycorrhizal
interface, this has critical implications for efficient root uptake
of water and nutrients. Through these symbioses plants have
succeeded in colonizing all terrestrial biomes (Blackwell, 2000)
and consequently, plant-communities are the template for
these terrestrial communities of microorganisms. In a strikingly
similar manner to the well-being of humans and animals
that is intimately linked to community composition of gut
bacteria, so also is the functioning of plants closely dependent
upon the composition and activities of their below-ground
mycorrhizal associates.
World-wide patterns and regulation of plant species rich-
ness and their impacts on ecosystem processes have long been of
interest, but less is known about the diversity, its regulation and
functional importance of mycorrhizal fungi (Molina et al., 1992).

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While a relatively small number of plants, c. 8000, form ecto-
mycorrhiza (ECM), their global importance is greatly increased
by their disproportionate occupancy of terrestrial ecosystems.
Many trees, such as those in the Betulaceae, Pinaceae, Fagaceae,
and Dipterocaraceae are important and obligate ECM plants,
forming vast boreal, temperate and subtropical forests (Smith
& Read, 1997). A conservative estimate of the number of ECM
fungal species in the world is approaching 6000 (Molina et al.,
1992). Beside the diversity of known taxa, there are other ECM
fungi which do not produce conspicuous reproductive structures,
a feature that precludes their inclusion in known listings, and still
further, a huge number of fungal taxa remain to be discovered.
Until the middle of the 1990s, knowledge of ECM fungal
community structure, and the degree to which species assem-
blages show predictable colonization patterns in space and
time, was almost exclusively based on sporocarp surveys. A
major advantage of such surveys is the potential to record and
identify all fruiting ECM fungi within an area with relatively
little work load. However, because of the importance of
exogenous environmental factors in triggering fruiting, and the
consequent irregularity of reproduction, such characterization
requires long-term monitoring. Moreover, even after decades
of surveying, additional fungal species may still appear. Even
more alarmingly, recent demonstrations of the poor corres-
pondence between the species composition of sporocarps and
fungi colonizing roots raises some concern about the validity
of sporocarp survey data. It was, for example, observed that
whereas the most common sporocarps in a jack pine stand
were those of Suillus tomentosum, the mycorrhizas identified
by distinct morphology rarely constituted more than 5% of
the total mycorrhizas (Danielson, 1984).
During the past decade major advances in molecular biology
and morphotyping have made it possible to identify myco-
bionts from single mm-sized mycorrhizas, enabling monitoring
and spatiotemporal analysis of below-ground ECM fungal
communities (Horton & Bruns, 2001). Thereby, many diversity
and functional properties of ECM communities be integrated
and their key contributions related to the overall diversity and
ecosystem processes, an area of growing interest in terrestrial
ecology (Copley, 2000). Examples of such recent break-through
achievements are demonstrations of significant net transfer of
nutrients and carbon between different plant taxa through
common mycorrhizal mycelia (Simard et al., 1997) and the
implication that richness and composition of arbuscular
mycorrhizal fungi affect diversity and growth of plants (van
der Heijden et al., 1998).
Here, I briefly discuss the current potential for terrestrial
ecologists to identify and quantify ECM fungal communities.
In doing this I provide: an overview of mycorrhizal distribu-
tions in soils and methods to identify mycobionts in single
mycorrhizas; a description of recent findings in spatiotemporal
characteristics of ECM fungal communities; and a discussion
of how such an appreciation may advance the understanding
of plant ecology and ecosystem processes.
Standing mass and identification of mycorrhizas
Within days of emergence, almost every fine root of an
ectomycorrhizal plant is colonized by mycorrhizal fungi
(Smith & Read, 1997; Taylor et al., 2000). The number of the
mycorrhizal tips, commonly a few mm in length, is often very
high; in boreal forests the numbers typically range between
104 and 5 × 105 m–2 forest floor, a figure that in Norway spruce
forests roughly corresponds to the number of green needles
(Dahlberg et al., 1997). Mycorrhizas are short-lived structures,
being regenerated on a yearly basis. Regeneration may occur
throughout the growing season, but peaks at certain periods.
Mycorrhizas are vertically distributed to depths occupied by
host tree roots (i.e. down to several meters). However, the
overwhelming majority are located in surface soil layers (e.g.
at the interface of organic matter and mineral soil) where
mineralization processes are most active. In dry forest eco-
systems, mycorrhizas are located deeper in the mineral soil
than their mesic counterparts (Visser, 1995; Taylor & Bruns,
1999). Typically up to 90% of the mycorrhizas are located
within the upper 10 cm of the soil profile in boreal soils,
mainly associated with the thick organic humus or mor layer.
Fungal tissue in mycorrhizal roots have fungal biomass that
is 10–100 times higher than that in the sporocarps, even
without considering the often extensive extramatrical mycelia
(Dahlberg et al., 1997).
One well-developed approach to identify ECM fungi
colonizing roots involves detailed morphological and struct-
ural descriptions of mycorrhizas – morphotyping (Agerer,
1987–1999). In certain cases, morphology of mycorrhizas is
species-specific and allows unambiguous identification of
mycorrhizas collected in the field (e.g. Smith et al., 2000). More
generally, this approach only allows grouping into mycor-
rhizal morphotypes, which do not represent single species.
However, the most significant methodological advance has
been the application of high-resolution molecular tools that
allow identification of individual mycorrhizas (Gardes &
Bruns, 1993; Horton & Bruns, 2001). These methods offer
three advantages over morphological methods: they can be
directed at any taxonomic level or to detect genotypes; they
require less time to learn, and are clearly described; and they
are fully reproducible in time and space. Fungal, or plant
DNA can be selectively amplified from mixtures of plant and
fungal DNA from single mycorrhizas and, with subsequent
analysis, species identified (Gardes & Bruns, 1993; Horton &
Bruns, 1998). The internal transcribed spacer (ITS) region in
the nuclear ribosomal unit is frequently sequenced, digested
with restriction enzymes and resulting restriction fragment
length polymorphism analysed (Gardes & Bruns, 1996; Kårén
et al., 1997; Peter et al., 2000). ITS nucleotide databases are
also being constructed for certain ECM fungal groups for
identification and phylogenetic purposes allowing for direct
sequence comparison; particularly informative is the 5′ end of
the LSU gene (Kõljalg et al., 2000; Moncalvo et al., 2000).
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Table 1 A compilation of the three most abundant or frequent ectomycorrhizal (ECM fungal taxa, as recorded from mycorrhizas, in 49 ECM
fungal community studies published so far. Only studies where the taxonomic identity of at least one of the three most abundant taxa was
certified are included. Figures indicate at how many studies certain taxa dominated. The data from Canada includes four assessments of ECM
communities on seedlings. Besides 21 papers referred to in the present review paper, 28 additional studies are included

Number of studies with dominance of the following taxa

Sporocarps lacking or Sporocarps lacking

nonconspiceous Conspicous sporocarps or nonconspiceous

Total no Cenococcum Corticea- Thelephor- Suilloid (incl Cortinaria- MRA

Forest type Country of studies geophilum ceae aceae Rhizopogon) Russulales ceae E-strain Risseia

Coniferous forest
Boreal Canada 15 10 7 3 4 4 1 9
Sweden 12 6 12 3 2 2 1 1

Temperate Switzerland 3 1 3 3 1
& Germany
USA 15 4 7 8 7

Deciduous forests
USA 1 1
Germany 1 1
Netherlands 1 1

Rainforest
Indonesia 1 1

In total: 49 22 22 17 14 14 3 10 1

Sequence data in the ML5/ML6 region of the mitochondrial
LrDNA representing all major ECM basidiomycete genera is
established (Bruns et al., 1998). Thus, any single mycorrhiza
can potentially be identified to species either by ITS-RFLP
comparison with reference material or by sequence comparison
narrowed down to smaller subsets of putative species affinity
that ultimately, with supplementation of appropriate references,
will be identified.
Slightly more than 50 such ECM community studies have
been published in the period between 1996 and late 2000, few
before (cf. Horton & Bruns, 2001; cf. Table 1). Two thirds of
these have been based on morphotyping and one third on
molecular methods, sometimes with morphotyping-based
stratification as a powerful approach (Jonsson et al., 1999a).
Characteristics of ECM communities
Studies from North America and Europe unanimously confirm
that fungi that do not form obvious fruiting structures form the
major mycorrhizal abundance. Fruiting species merely constitute
20–30% of the mycorrhizas (e.g. Gardes & Bruns, 1996; Jonsson
et al., 1999a; Fig. 1). Apparently, carbon budget or resource alloca-
tion to production of sporocarps vs mycorrhizas and mycelial
growth vary among species, as discussed in Gardes & Bruns
(1996) seminal community paper. Typically, certain species
that commonly form mycorrhizas are poorly or nonrepresented
in the above-ground fruiting record, and commonly fruiting spe-
cies may conversely have only a limited representation in below-
ground mycorrhizal networks. Thelephoraceae, Russulaceae and,
at least in post fire settings, also suilloid fungi repeatedly emerge
as the most frequent or abundant mycorrhizas in forests of
western USA, whereas Corticeaceous fungi and Cenococcum
geophilum, followed by Russulaceae and Thelephoraceae appear
as the most common in boreal forests (Table 1).
The general rule is that a few fungal ECM taxa account for the
most, (> 50%), of the mycorrhizal abundance and are widely
spread whereas the majority of species are only rarely encoun-
tered (Gehring et al., 1998; Jonsson et al., 1999a,b; Peter et al.,
2000; Stendell et al., 1999; Grogan et al., 2000). In two
stands of Douglas-fir, 69 morphotypes were recognized from
17 500 assessed mycorrhizas; most types were rare, 19 types
colonized > 1% of the mycorrhizas and 9 types colonized
together 67% (Goodman & Trofymow, 1998a). This pattern
of dominance of a few species and rarity of most others is a com-
mon community pattern of any taxonomic group of organisms.
Major determinants of the composition of ECM fungal
community are host plant composition ( Molina et al., 1992)
and edaphic factors (Gehring et al., 1998; Kernaghan & Harper,
2001). Within local forests, the spatial variation of ECM fungi
is very high and most species show aggregated distributions
(Gardes & Bruns, 1996; Kranabetter & Wylie, 1998; Bidartondo
et al., 2000). A study of pinyon trees showed one or a few ECM
fungal taxa to dominate single trees, and the dominant fungi
varied between trees (Gehring et al., 1998). Spatial variation
was concluded to have a larger effect on ECM fungal species
composition, than that caused by low intensity wildfire, the

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major natural disturbance in Swedish boreal forests (Jonsson
et al., 1999a). One exception is reported in a study of 1000-
yr-old bristlecone pine stands at high elevation in California;
here one ECM Pyronemataceae sp. was consistently present as
inoculum in soil at all forest sites (Bidartondo et al., 2001).
Commonly, small volumes of soil contain several ECM species,
for example single four year old seedlings were colonized by
up to 11 ECM fungi (Kranabetter & Wylie, 1998). Certain
taxa also appear to grow better in either the organic or mineral
layer (cf. Goodman & Trofymow, 1998).
A significant step forward in understanding ECM fungal
community structure is to establish studies that allow estimation
of the relative importance of different mechanisms that create
this diversity, and take into consideration spatial and temporal
scales. Hypotheses to explore can be placed in two categories:
those based on niche differentiation between coexisting species;
and, paradoxically, those based on similarities in competitive
ability, that is nonequilibrium models. An example of the
former is the intermediate disturbance hypothesis stating that
a disturbance leads to a predictable sequence of species repla-
cing each other. An intermediate range of disturbance would
allow a range of species traits, early to late successional species,
to coexist and lead to a high species richness. The latter assumes
elements of stochasticity which enhance coexistence: so called
lottery models state that due to poor dispersal ability, low
local abundance and chance events, species may be absent in
a neighbourhood and their abundance recruitment may thus
be limited. Therefore the best competitor that happens to
colonize a particular local site may well be inferior on a larger
scale. Potentially, this can lead to unlimited species richness.
Temporal dynamics of ECM communities
Successional changes in primary vegetation successions, and
thus of primary ECM fungal colonization, are described from
glacial retreats and farmlands. Such questions have been studied
in the course of natural secondary forest succession in a few
cases. It must be remembered that forest management procedures,
such as clearcutting, impose novel selection pressure upon ECM
fungi. Marked changes in ECM community composition occur
after intense, stand replacing fire that totally consumes the
organic layer, a natural occurrence in certain forest ecosystems
( Visser, 1995; Grogan et al., 2000). However, in low intensity
fires, where part of the organic layer remains unburnt and a
large proportion of trees survive the fire, the legacy of ECM
fungi appears large (Jonsson et al., 1999a). Short-term variation
in abundance of certain taxa of mycorrhizas may occur, but as
yet has been poorly investigated, as weather conditions affect
photosynthesis and thereby allocation of assimilates to the roots,
Fig. 1 (opposite) Late successional Scots pine stands, in northern Sweden, where both ectomycorrhizal fungal community structure from
sporocarps and mycorrhizas, and fungal population structure of Suillus variegatus have been analysed. The last major disturbance was a forest
fire in 1647. The relative proportion of mass in sporocarps (open red bars, 34 taxa) and of mycorrhizal abundance (closed blue bars, 24 taxa,
distinguished by ITS-RFLP) is shown (Jonsson et al., 1999b). At the same site the spatial distribution Suillus variegatus genets was monitored
from sporocarps analysed by somatic incompatibility reactions (from Dahlberg (1997), with permission).
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Research review Review 559
and the mycorrhizas, and the temperature and humidity of
the soil. This certainly affects the patterns of fruiting. However,
a detailed molecular study of mycorrhizas in a Swiss Norway
spruce forest conducted over three years showed a striking
similarity in abundance of the 10 most abundant taxa (Peter
et al., 2001).
Processes in ECM fungal populations underlie ECM fungal
community patterns and thus, a knowledge of the ability of
individual ECM fungal species to bridge distances in time and
space is central to the understanding of ECM community
structure. However, little is known about the relative impor-
tance of mycelial spread and longevity vs novel colonization
from spores for any ECM fungal species. From the nearly 20
studies of ECM fungal population structures conducted so far
(cf Redecker et al., 2001), it is apparent that genets in some
species are numerous, short-lived (1–2 yr) and small (< few
m, i.e. Hebeloma cylindrosporum, Laccaria amethystina, Amanita
franchetii, Russula cremoricolor). Other species may have few
and due to persistence (decades to centuries) and mycelial
spread, relatively large genets (20–40 m in diameter, i.e.
Suillus spp. and Cortinarius. Small genets may suggest colon-
ization from recent meiospores, whereas fewer, larger genets
indicate old mycelial structures, that have grown from a point
source over decades. The latter are often found in old forests
(Fig. 1). Some studies suggest mycelial colonization as the
primary avenue of ECM fungal colonization in mature un-
disturbed forests, i.e. that clonal traits are of higher fitness
than sexual traits under undisturbed conditions (e.g. Jonsson
et al., 1999b; Kranabetter et al., 1999). Resistant ECM fungal
propagules also persist, probably in the mineral soil, after major
disturbances such as stand replacing wildfires (Baar et al., 1999).
Exceptions do occur, for example L. amethystina appear to
be short-lived in old beech forests (Gherbi et al., 1999).
Demographic patterns in ECM fungi, or any soil dwelling
fungus, have still not been explored.
There is a need to launch studies of regional scale dynamics
of ECM fungal populations, and communities, to elucidate
important population processes. There is also a need to deter-
mine which models are most applicable for different ECM fungi:
theories of remnant population dynamics with discontinuous,
nonequalibrium conditions and a pervasive impact of ‘accidents
of history’ or, on the other hand, meta-population models
with continuous, equilibrium conditions and without historic
dimensions (cf. Eriksson, 1996).
Interactions with plant communities
Most ECM fungal species typically have broad, but differing,
plant host ranges. They may form mycorrhizas with any ECM
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angiosperm or gymnosperm or only with angiosperms or gym-
nosperms, or a single family of hosts. This potentially extensive,
but importantly sometimes also limited, linking of ECM plants
via intra- and interspecific ECM mycelia, within and between
species, may have major consequences for our understanding
of plant community characteristics, plant competitive interac-
tions and performance ( Molina et al., 1992). These linkages
potentially allow for nutrient and carbon transfer between
tree species, but their significance in nature remains to be
settled. Evidence for the potential of such mycelial linkages is
accumulating. Field and laboratory soil bioassays showed
Douglas fir to share ECM fungi and potentially form hyphal
links with several other plant species (Massicotte et al., 1999).
Jones et al. (1997) found that 91% of paper birch and 56%
of Douglas fir intermingled mycorrhizal roots examined were
colonized by the same mycorrhizal fungi. In these tree species,
a 4–7% net transfer of isotopically labelled carbon from
birch to Douglas fir has been demonstrated, which may be
ecologically significant (Simard et al., 1997). If net transfer is
predominantly from pioneer plant species to late successional
species, ECM fungi may be an important contributing factor
determining plant community development.
In a mixed stand of bishop pine and Douglas fir the majority
of identified mycobionts were associated with both hosts, even
within small soil volumes, suggesting that individual fungal
genotypes linked the putatively competing tree hosts (Horton
& Bruns, 1998). One study in the Canadian Rockies recorded
all present ECM fungi to form mycorrhizas with both Dryas and
Salix (Kernaghan & Harper, 2001). In a comparison between
seedlings of lodgepole pine, white spruce and subalpine fir
seedlings, 43 of 74 mycorrhizal morphotypes were shared
by all conifers, constituting 52% of the total mycorrhizal
abundance (Kranabetter et al., 1999). However, it is also
important to appreciate certain ECM fungi, for example
suilloid fungi, as rigidly restricted to certain plant taxa and
thus only potentially linking intraspecific plants.
Ericoid and ectomycorrhizal plants often coexist in boreal and
Mediterranean forests and are normally considered as separate
mycorrhizal systems with mycobionts of different taxonomic
affinity. However, recently an ericoid mycorrhizal fungus was
shown to associate with Quercus roots (Bergero et al., 2000). The
possibility that some sharing may occur between ericaceous
plants and ECM trees has been discussed, but questions concerning
whether it occurs and what possible ecological significance it
may have still need to be answered in appropriate, controlled
studies. The results of such experiments may have important
implications for our understanding of interactions between
trees and the understorey.
Effects on ecosystem function
To date, ECM fungal community studies have been mainly of
a descriptive or correlative nature and few experimental studies
have been performed. Causal mechanisms underlying the
Fig. 2 Diversity and ecosystem function have no direct relationships
to each other, but are both functions of the presence and activities
of individual species or functional groups and their interactions. The
environment affects species, and the outcome of their interactions,
but not diversity or ecosystem processes directly. The question mark
indicates the question as to whether diversity and ecosystem
processes are related (redrawn from Bengtsson, 1998).
dynamics, and implications for ecological functions, have
rarely been addressed under controlled experimental conditions
(Lilleskov & Bruns, 2001). It is likely that many ECM fungi
may fulfil similar ecological functions and that a degree of
functional redundancy exists in ECM fungal communities. In
an experiment on the effect of plant litter composition, there
was no evidence that the composition of the present ECM fungi
affected the performance of seedlings (Nilsson et al., 1999).
Given their taxonomic diversity, communities are still likely
to retain vast amounts of functional heterogeneity. Yet it is
difficult to generalize about the ecological functionality of ECM
communities because relatively few investigations of ECM
fungal physiology have been conducted with more than a few
isolates, and those typically include only a handful of taxa that
typically occur in low abundance (Cairney, 1999). The potential
relationship between ECM fungal species richness and ecosystem
productivity has once been explored without finding any
correlation (Gehring et al., 1998). As debated at length, there
is no mechanistic relationship between species richness, or any
diversity measure, and ecosystem function (Bengtsson, 1998).
Probably, it is more vital to understand linkages between the
actions of key species or functional groups and ecological
functions, as diversity is an abstract aggregated property of
species in the context of communities or ecosystems (Fig. 2).
The functioning of terrestrial ecosystems seems to depend heavily
on soil biodiversity. We now have the potential to address and
experimentally test questions of ecological significance, as
abundant ECM fungi, potentially being key species, can be
identified. One approach could be to use A set of ECM fungi
from selected environmental settings, and to investigate whether
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the abilities of these ECM fungal communities differ for
various functions, such as carbon cost, rate of nutrient-cycling
or response to perturbations.
One promising approach is to explore ECM fungal community
functioning under simulated forest conditions using micro-
cosm systems (Tammi et al., 2001). In these systems, intact
mycorrhizal root systems comprising individual species, for
example abundant taxa likely to be functionally important, or
natural communities, can be manipulated and analysed to
determine carbon and nitrogen relations and host root-fungal
metabolic activities that contribute to plant growth or plant
community productivity. The application of molecular methods
is already providing detailed insights into the complexity of
ECM fungal communities and offers exciting prospects to
elucidate processes structuring ECM fungal communities
and advance our understanding of plant ecology, such as plant
interactions, and ecosystem processes.
Acknowledgements
I thank Robin Sen, Marie-Charlotte Nilsson, Tom Bruns,
Åke Olsson, Roger Finlay and Jan Stenlid for critical reading
of earlier drafts of this paper. Major support over the years
for this research came from the Swedish Council for Forestry
and Agricultural Research.
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