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Editorial
Fungal ecology beyond boundaries: From
communities to the globe
This special issue is based upon an eponymous sympo-
sium, “Fungal ecology beyond boundaries: from communities
to the globe”, that took place at the 2013 annual Mycological
Society of America meeting in Austin, Texas. My motivation
for organizing this symposium was to bring together fungal
ecologists working at different biological and spatial scales
in the hope of increasing communication and continuity
across the field. Equally important, I wanted to bring more
awareness to the way in which scale affects the type informa-
tion we can infer about the ecology of fungi. This special issue
highlights the complementary (and sometimes contradictory)
ecological information that can be obtained across spatial
scales spanning communities of fungi carried on the beaks
of hummingbirds (Belisle et al., 2014), to communities of
arbuscular mycorrhizal fungi across continents (Egan et al.,
2014), and biological scales from the detection of an individual
and rare fungus in Alaskan boreal forests (Glass et al., 2014), to
the recovery of thousands of airborne fungi in coastal south-
ern California (Kivlin et al., 2014). While the outcomes of
the symposium and this special issue highlight new and
intriguing patterns of fungal biogeography and ecology
(Hayward and Hynson 2014; Huggins et al., 2014), they also
call for further integration, and perhaps standardization
among researchers working across levels of biological organi-
zation, space, and time (Chagnon et al., 2014; Chaudhary et al.,
2014). Issues of scale dependency are not unique to the study
of fungal ecology (Chase and Knight, 2013), but this field
currently stands at a crossroads. We have the opportunity to
overcome the disjoint among researchers working at different
scales by standardizing approaches from next generation mo-
lecular methods and combining these efforts with intensive
studies of fungal autecology (Peay, 2014). However, accom-
plishing this task means marrying the skills of alpha taxono-
mists, ecologists and computational biologists to revise and
in some cases, reinvent the characterization of fungi.
While our basic understanding of the interactions of fungi
with their environments have grown substantially, especially
over the last 20 years (Peay et al., 2008), fundamental questions
regarding the ecology of fungi remain. Some of these ques-
tions are addressed throughout this issue. For instance, the ar-
ticles by Egan et al. and Kivlin et al. examine the dispersal
potentials of arbuscular mycorrhizal (AM) fungal spores over
1 yr, and total fungal propagules over 16 months, respectively.
These studies are among the first to explicitly test the dispersal
limitations of fungi by air in natural environments, relate
fungal aerobiota to neighboring soil fungal communities and,
in the case of Kivlin et al., link soil fungal community structure
to specific abiotic conditions. While the studies of Egan et al.
and Kivlin et al. assess the diversity of soil fungi across conti-
nental and regional scales, the study by Chaudhary et al. ex-
amines the diversity of AM fungi in arid soils at relatively
smaller spatial scales (0.0001e5 000 ha). Together, these three
papers demonstrate how different spatial and temporal scales
and the choice of diversity assessment tools can result in
mixed messages about the ecology of a given group of fungi.
Egan et al.’s paper examines AM fungal dispersal by air
across six biomes stretching across North America and finds
that, except for Glomus spp. in arid regions, AM fungi generally
do not aerially disperse. In soils across all ecoregions, Glomus
spp. were the most abundant, but there was no correlation be-
tween their abundance in the air and in the soil of a given
habitat. Similarly, Chaudhary et al. found that Rhizophagus
irregularis (Glomus intraradices) was the most common AM fun-
gus isolated from soils of arid habitats. Together, these studies
indicate that Glomus spp. may have the widest geographical
ranges and ecological niches of any of the AM fungi. In terms
of changes in fungal diversity across space, the findings of Kiv-
lin et al. dovetail those of Chaudhary et al.’s where both
studies found significant differences in soil fungal commu-
nities at the site scale (0.01e1 ha). However, the AM fungal
communities from Chaudhary et al.’s study were also distinct
at the regional scale (5 000 ha) where Kivlin et al. found no sig-
nificant differences in fungal diversity for all groups of soil
fungi across an area of 300 000 ha. The discrepancies between
these three studies are likely due to: (1) the more general
assessment of all fungal groups executed by Kivlin et al.; (2)
the size of the sampling area; (3) the timing of soil sampling;
and (4) the methods used for assessing fungal diversity in
Kivlin et al. versus Chaudhary et al. and Egan et al. However,
all three studies concur that AM fungi are relatively dispersal
limited compared to other groups of fungi, which the authors
attribute to their specific life history strategy.
2 Editorial
The life history strategies and adaptations of mycorrhizal
fungi are further highlighted in the contribution by Huggins
et al. who found that some of the ectomycorrhizal (EM) fungi
associated with Alnus host species are far more tolerant of
high nitrate and acidic growth conditions than a suite of
non-Alnus-associated EM fungi. Also, the response of Alnus
and non-Alnus EM fungi to either high nitrate or acidity
tended to be species specific. However, the overall greater
tolerance of Alnus EM fungi to high nitrate and acidity indi-
cates that some fungi can adapt to the environments shaped
by their host, and in this case, their hosts’ interactions with
other symbionts such as nitrogen fixing bacteria. The work
of Belisle et al. also touches upon multi-partner direct and in-
direct symbiotic interactions by examining how environ-
ment and time affect the community composition of
microfungi associated with bird and bat pollinators. While
previous research had established that the identity of these
microfungi significantly affects plantepollinator interactions
(Vannette et al., 2013; Good et al., 2014), this study was one of
the first to examine how these communities vary between
pollinators and over time. In contrast to the findings of
Egan et al. on AM fungi, Belisle et al. found that pollinator-
dispersed microfungal communities varied more with time
rather than habitat and found no indication of dispersal lim-
itation among this group of fungi across an area of 4,300 Ha.
These conclusions stand in contrast to other recent studies of
microbial community assembly processes (reviewed in
Hanson et al., 2012) and stress the need for consistency in
methodological approaches and data interpretation among
researchers, as well as additional studies of long distance
fungal dispersal.
One prime example of historical long distance dispersal by
fungi is included in this issue: Hayward & Hynson report on
their recent discovery of EM fungi associated with a host
tree endemic to the Hawaiian Islands. Due to these islands
extreme isolation, Hawaii was previously thought to lack
any native or naturalized EM symbioses. However, based on
the authors’ findings, we now know that Pisonia sandwicensis
(Nyctaginaceae) hosts a handful of taxonomically and ecolog-
ically diverse EM fungal species. In fact, this study may be the
first to fully characterize a native EM fungal community from
an entire region, providing an exciting new study system to
examine fungal biogeography. The article by Glass et al. also
features new information on the ecology of a novel fungal
taxon (NS1) associated with white spruce in Alaskan boreal
forests, but known only from environmental DNA sequencing.
Based on repeated, spatially-explicit field sampling and DNA
sequence analysis the authors found that this fungus has a
clumped distribution and can persist for at least 6 yr. They
go on to argue that “patterns of occurrence of DNA sequences
in environmental samples can reveal valuable information
regarding the ecologies of both well-known and unknown or
novel species”.
Building upon the arguments of Glass et al., this issue in-
cludes a compelling commentary on the future of modern
fungal ecology by Kabir Peay. In this essay Peay evaluates the
best way to move the field forward given the unlimited
potential for data generation and the increasing participation
of scientists from across disciplines in fungal ecological
research. In answering this question he echoes the recent
sentiments of others in the scientific community who are call-
ing for a resurgence of the practice of natural history among
biologists and ecologists (Tewksbury et al., 2014). However,
Peay argues that in the specific context of fungal ecology this
practice must be integrated with the tools of molecular bio-
logyetwo fields that have been slow (or even resistant at times)
to unite. Along with Peay, the commentary by Chagnon et al.
describes clear and necessary steps to achieve synergy be-
tween the complementary approaches of extensive data gen-
eration and an intensive understanding of the fundamental
properties of organisms in their environments. These recom-
mendations include more investment in determining the biotic
and abiotic determinants of community patterns and less
emphasis on describing community membership (i.e., opera-
tional taxonomic units). While the returns on this type of in-
vestment for the individual researcher are uncertain, they
will certainly propel the field as a whole forward in a more uni-
fied direction. The collective results of such efforts may in-turn
ask that current ecological theory based primarily on the study
of macroorganisms also move beyond its current bounds.
Acknowledgments
I would like to thank Elsevier Publishing and the New Phytol-
ogist Trust, and Lynne Boddy for their support of the original
symposium upon which this issue is based. Also, I would
like to thank the speakers and authors who have contributed
to these efforts. Finally, I would like to thank ASA and ARA for
helpful comments on a previous version of this editorial.
references
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Editorial
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Nicole A. Hynson
University of Hawaii Manoa, Department of Botany,
3190 Maile Way, Room 101, Honolulu, HI 96822, USA
E-mail address: nhynson@hawaii.edu
http://dx.doi.org/10.1016/j.funeco.2014.09.005
1754-5048/ª 2014 Elsevier Ltd and The British Mycological
Society. All rights reserved.