fungal biology reviews 44 (2023) 100294

journal homepage: www.elsevier.com/locate/fbr

Review

Reviewing the contributions of macrofungi to forest

ecosystem processes and services

Allen Grace T. NIEGOa,b,c, Sylvie RAPIORd, Naritsada THONGKLANGa,b,**,

b, Kevin D. HYDEa,b,e,f, Peter MORTIMERg,*
Olivier RASPE
a
Center of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai 57100, Thailand
b
School of Science, Mae Fah Luang University, Chiang Rai 57100, Thailand
c
Iloilo Science and Technology University, La Paz, Iloilo, 5000, Philippines
d
CEFE, CNRS, Univ Montpellier, EPHE, IRD, Laboratory of Botany, Phytochemistry and Mycology, Faculty of
Pharmacy, CS 14491, 15 Avenue Charles Flahault, 34093 Montpellier Cedex 5, France
e
Department of Biology, Faculty of Science, Chiang Mai University, Chiang Mai 50200, Thailand
f
Institute for Plant Health, Zhongkai University of Agriculture and Engineering, Haizhu District, Guangzhou 510225,
PR China
g
Center for Mountain Futures, Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650201,
Yunnan, PR China

article info abstract

Article history: Macrofungi are vital components of any forest ecosystem, performing different roles
Received 4 July 2022 crucial to ecosystem functioning. Macrofungi play effective roles in ecosystem processes
Received in revised form such as nutrient cycling. Ectomycorrhizal fungi and plant symbionts work together to
24 October 2022 accumulate, use and transfer essential nutrients, especially in nitrogen/phosphorus-
Accepted 10 November 2022 limited environments. Pathogenic and predatory macrofungi exploit other resources
such as plants and animals to obtain nutrients needed for growth. These groups of macro-
Keywords: fungi also contribute to the species diversity of forest ecosystems. Saprotrophic macro-
Biodiversity fungi degrade available organic matter from dead plants and soil organic matter.
Conservation Macrofungi also play an important role in carbon sequestration in the forest underground
Mycorrhiza as well as in soil formation. Macrofungi remediate pollutants in the environment via extra-
Nutrient cycling cellular enzymes. Mycelial networks that connect macrofungi to their symbiotic hosts and
substrates enable most of these functions. Mycelial networks facilitate the absorption and
Abbreviations: transport of nutrients as well as the secretion of enzymes and other organic substances.
(AM) Spore-producing bodies of macrofungi serve as a food source for wildlife. Fungal spores
Arbuscular mycorrhiza can act as aerosols in rain formation. All of these functions of macrofungi are necessary
(C) for maintaining biodiversity and healthy forest ecosystems. However, the contributions
Carbon of macrofungi to ecosystem processes are often taken for granted or not fully recognized,
(CO2) offering key services that are easily overlooked in planning processes and policymaking.
Carbon dioxide The present review summarizes the major roles of macrofungi in ecosystem functioning

* Corresponding author.
** Corresponding author. Center of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai 57100. Thailand.
E-mail addresses: naritsada.t@gmail.com (N. Thongklang), peter@mail.kib.ac.cn (P. Mortimer).
https://doi.org/10.1016/j.fbr.2022.11.002
1749-4613/ª 2022 British Mycological Society. Published by Elsevier Ltd. All rights reserved.
2 A. G. T. Niego et al.

(EcMF) and services rendered, and the interrelationships between these functions and services in
Ectomycorrhizal fungi the forest ecosystems.
(ErM) ª 2022 British Mycological Society. Published by Elsevier Ltd. All rights reserved.
Ericoid mycorrhiza
(LiP)
Lignin peroxidase
(MnP)
Manganese-dependent peroxidase
(N)
Nitrogen
(P)
Phosphorus
(PAHs)
Polycyclic aromatic hydrocarbons
(SOM)
Soil organic matter

1. Introduction Ecosystem services refer to the benefits humans get from

Macrofungi are a group of fungi with large, easily observed
spore-bearing structures. They are heterotrophic organisms
belonging to the phyla Basidiomycota and Ascomycota in
the kingdom of Fungi (Kinge et al., 2020). Macrofungi are
morphologically diverse with either epigeous or hypogeous
spore-producing bodies. There are around 14,000 identified
species of macrofungi out of which over 2000 species are
edible and can be consumed safely (Dupont et al., 2017; H. Li
et al., 2021), some with great gastronomic value (Kalac, 2013;
Meenu and Xu, 2019; Mortimer et al., 2021), serving as a food
source for humans (Ache et al., 2021; Mortimer et al., 2021)
and forest-dwelling animals (Katarz yte_ and Kutorga, 2011;
Nuske et al., 2019; Ori et al., 2021).
the processes arising from healthy ecosystems (Fu et al.,
2013). Macrofungi are an integral part of forests and provide
a diverse range of ecosystem services, including innovative
biotechnological, medicinal, and ecological applications
(Hyde et al., 2019; Kinge et al., 2020; Niego et al., 2021).
. Through their role in various ecosystem processes, mac-
rofungi contribute to regulate and improve ecosystem func-
tioning. Consequently, macrofungi also indirectly contribute
to the forest ecosystem services. Thus, contributions of mac-
rofungi to forest ecosystem services can be classified either
as direct contributions or indirect contributions (Fig. 1). Direct
contributions to ecosystem services such as the provision of
food sources, improvement of livelihoods and recreation
contribute on human welfare, while other functions of

Fig. 1 e Direct and indirect contributions of macrofungi to forest ecosystem services in relation to humans.
Contributions of macrofungi to forest ecosystem
macrofungi in ecosystem processes have indirect effects on
humans. For instance, the role played by macrofungi in
nutrient cycling can be considered as an indirect contribution
to the ecosystem services rendered by forest ecosystems.
Without macrofungi, the services rendered by forests, like
timber production or carbon stocking, would definitely be
reduced. Consequently, those indirect contributions should
not be ignored in policy making or when evaluating the contri-
bution of fungi to forest ecosystem services.
Some of the functions of macrofungi to the ecosystem pro-
cesses include macrofungi acting as decomposers and mutu-
alistic symbionts (Lutzoni et al., 2018; Richard et al., 2004; Ye
et al., 2019). Macrofungi are crucial in recycling organic matter
needed for the growth and survival of living organisms
including humans, and thus are important for maintaining
biodiversity (de Mattos-Shipley et al., 2016). They play an inte-
gral role in the transport, storage, and release of nutrients
such as carbon (C), phosphorus (P), and nitrogen (N) (Bortier
et al., 2018; Ekblad et al., 2013; Varma et al., 2017). Some macro-
fungi form ectomycorrhizal associations with terrestrial
plants thereby supporting the growth and survival of the plant
(Sharma, 2017; Taschen et al., 2020). Macrofungi can be patho-
genic and regulate dominating plant species, creating space
and opportunities for other plant species to thrive, thereby
maintaining, or even increasing, the biodiversity of a forest
(Angelini et al., 2015; Hennon, 1995; Mujica et al., 2020).
Macrofungi also play a key role in C sequestration, with
mycorrhizal symbioses functioning as C sinks, enhancing
the use of atmospheric carbon dioxide (CO2) for both plant
and fungal growth to a greater extent than non-mycorrhizal
plants (Wang et al., 2016). Saprotrophic fungi are responsible
for the degradation of organic matter and assimilation of C-
rich degradation products, thereby also acting as a C sink, by
retaining C in the soil (He et al., 2017). Macrofungi also have
the capability to remediate environmental pollutants by pro-
ducing composite enzymes capable of degrading these pollut-
ants (Ali et al., 2017; Uddin et al., 2020; Yadav et al., 2021). They
also bolster soil formation by producing organic compounds
that bind soil particles, thus improving soil structure and
porosity (Oregon State University Extension Service, 2021).
Despite the services provided by macrofungi and their
importance to humankind and ecosystems, they remain
underrecognized and undervalued, and the economic value
of these services remains largely unknown. Macrofungi are
underappreciated in the value of the contributions made to-
wards forest ecosystems, many only view macrofungi as a
source of mushrooms and little else. We thus hope to bridge
this knowledge gap by providing insight into the range of
functions performed by macrofungi in forests. In addition,
and partially as a result of the mentioned knowledge gap,
macrofungi are seldom considered in policymaking decisions
relating to forest conservation (Zotti et al., 2013). Therefore, we
need to increase recognition of ecosystem services to help pol-
icymakers and resource managers consider the conservation
of macrofungi and recognize the value of their contribution
to forest ecosystems and humans alike. This review compre-
hensively summarizes the major roles of macrofungi in for-
ests as sentinel organisms, and to promote a better
understanding of the ecosystem processes and services deliv-
ered in forests.
3
2. Types of macrofungi in forest ecosystems
2.1. Ectomycorrhizal fungi
Ectomycorrhizal fungi (EcMF) are the only mycorrhizal fungi
considered to be macrofungi (Policelli et al., 2020). EcMF
comprise >5000 spp. and >250 genera, mostly Basidiomycota
but also some Ascomycota, and form mutualistic relation-
ships with the fine roots of various plant species, primarily
comprising woody plants from both gymnosperms and angio-
sperms (Geml, 2017; Sharma, 2017). Of the 1928 genera of mac-
rofungi belonging to Basidiomycota (He et al., 2019), around
100 are presumed to have ectomycorrhizal associations with
approximately 2% of vascular plants (mainly including Dipter-
ocarpaceae, Fagaceae, Myrtaceae, and Pinaceae) (Brundrett
and Tedersoo, 2018; Corrales et al., 2018; T. Li et al., 2021).
EcMF have the capacity to mine soils more effectively than
tree roots, transporting nutrients to the host plant for which
the host provides sugars in exchange (Nehls et al., 2010). It is
thus an association between the fine mycelia of fungi and
roots of plants whereby nutrients are exchanged, and forest
ecosystem functions are strengthened via enhanced host
growth and performance (Bonfante and Genre, 2010). EcMF
also improve host plant resistance to diseases and environ-
mental stresses (Liu et al., 2020). For instance, in soil contam-
inated with high concentrations of heavy metals, EcMF have
been shown to improve the host plants’ tolerance of heavy
metal stress, and thus improve host growth and performance
in contaminated soils (Chot and Reddy, 2022).
Environmental and biological factors greatly affect the di-
versity of EcMF (Peay et al., 2010), however, host specificity
also greatly influences EcMF diversity (Kernaghan et al.,
2003). Furthermore, soil chemistry and temperature have a
direct effect on the community composition of EcMF and the
related production of spore-producing bodies (Ruhling and
Tyler, 1990). In the tropics, for instance, high soil temperatures
and rapid decomposition rates lower the availability of
organic matter as substrates (Tedersoo et al., 2014). Soil forma-
tion also has indirect effects on the diversity of EcMF since the
soil can influence plant community composition, thereby
influencing the EcMF community (Kernaghan et al., 2003).
Other factors affecting EcMF diversity are climate and topog-
raphy. Around 80%e90% of all temperate and boreal forest
trees form a beneficial symbiotic relationship with EcMF
(Brundrett, 2004; Read, 1991; Wilson et al., 2017) in which spe-
cies richness of EcMF has been found to be higher in mature
forests (Dejene et al., 2021).
2.2. Saprotrophs
Saprotrophic macrofungi are the major decomposers of a for-
est, growing on different substrates such as leaf litter, fallen
debris, bark and wood of standing trees, and animal waste
(Ye et al., 2019). Saprotrophic fungi secrete enzymes to break
down organic materials, releasing mineral nutrients into the
surrounding soil, thereby improving soil fertility (Kubicek
and Druzhinina, 2007). Wood- and litter-degrading fungi play
an important role in degrading lignocellulose, the most abun-
dant terrestrial organic material, and are critical for
4 A. G. T. Niego et al.
facilitating C cycling in nature (Vos et al., 2021). These groups
of macrofungi are able to produce lignocellulolytic enzymes
to access organic nutrients stored within dead wood as well
as other plant materials (Crowther et al., 2012b), especially
lignocellulose. Lignocellulose is a complex substrate primarily
consisting of cellulose, hemicellulose, and lignin but also con-
taining traces of pectin, N compounds, and mineral residues
(Andlar et al., 2018). It is among the most recalcitrant com-
pounds in nature. Around 30% of the observed macrofungi
in the Greater Mekong Sub-Region were reported to be sapro-
trophs living on decomposing plant litter, with Marasmius spe-
cies featuring as the most abundant (Ye et al., 2019). He et al.
(2019) and Salo et al. (2019) listed the saprotrophic macrofun-
gal genera belonging to Basidiomycota and Ascomycota.
2.3. Pathogenic macrofungi
Fungal pathogens are an important component of healthy for-
est ecosystems, fulfilling the role of eliminating weaker plants
(Burgess and Wingfield, 2006) as well as highly abundant spe-
cies, thus affecting species occurrence and distribution
(Castello et al., 1995). Although pathogenic basidiomycetes
can be considered invasive in plantations and reforested
areas, often resulting in high losses of individual trees, in their
native habitats they exist in balance with their hosts, insti-
gating the death of only scattered, usually old or weak, trees
(Old et al., 2000).
About 10% of wood-decaying fungi mostly attack soft-
woods and are known as brown-rot fungi (Sigoillot et al.,
2012). Brown-rot fungi, which constitute a physiological group
of fungi belonging to Basidiomycota and to some lesser extent
Ascomycota, are capable of rapidly degrading cellulose and
hemicellulose (Arantes and Goodell, 2014); however, they do
not produce lignin-degrading enzymes (Langer et al., 2021).
Brown-rot fungi accomplish decomposition by degrading car-
bohydrates and extensive demethylation of lignin resulting in
the brownish color of the wood residue (Langer et al., 2021).
Brown-rot fungi preferentially attack coniferous woods and
have been the most destructive in decaying structural wood
products (Goodell et al., 2003).
White-rot fungi, on the other hand, are the most effective
degraders in forests (Rodr
ıguez-Couto, 2017). White-rot fungi
mineralize lignocellulosic biomass, including cellulose, hemi-
celluloses, and lignins (Suryadi et al., 2022). The extensive
degradation of lignin, using heme-peroxidases and laccases,
results in the bleaching of decayed wood (Xie et al., 2021).
This class of fungi are restricted to Agaricomycetes within
the Basidiomycota, and are mostly dominated by Polyporales
(Go et al., 2021; He et al., 2019; Riley et al., 2014).
2.4. Predatory/parasitic macrofungi
Some macrofungi have evolved complex mechanisms for
trapping food sources such as amoebas, nematodes, and roti-
fers, through an array of trapping devices to catch, kill and
digest their prey (Barron, 2003; Bava et al., 2022). Hohenbuehelia
sp., a wood rotting-fungus, was the first documented macro-
fungus found growing in a parasitized nematode (Barron
and Dierkes, 1977). Some Pleurotus species become carnivo-
rous to supplement their N intake under nutrient-limiting
conditions (Lee et al., 2020). Pleurotus ostreatus can cause rapid
nematode paralysis in which they secrete adhesive sub-
stances around their hyphae to trap and catch nematodes
(Lee et al., 2020). Pleurotus eryngii has been shown to be preda-
tory on Ancylostoma caninum (Braga, 2015) and Panagrellus sp.
(Sufiate et al., 2017). Coprinus comatus preys on Panagrellus redi-
vivus and Meloidogyne arenaria, nematodes that attack plants
(Luo et al., 2004). This macrofungus has toxic, spiny ball struc-
tures at the end of the hyphal branches that pierce the body of
the prey, killing it in minutes (Luo et al., 2004). Colonizing hy-
phae then penetrate the prey’s body through the wounds to
digest and assimilate its contents (Mora et al., 2018). Entomo-
pathogenic fungi are fungal pathogens that often kill or seri-
ously incapacitate a variety of insect species (Mantzoukas
et al., 2022). For example, the entomopathogenic fungus Cordy-
ceps militaris is well documented as parasite of insects or other
arthropods (Shrestha et al., 2012). Ophiocordyceps species are
also well-known in infecting insects (Coleoptera, Elateridae)
(Arau
jo et al., 2021; Xiao et al., 2019; Yang et al., 2021).
3. Active roles of macrofungi in ecosystem
processes
Indirect contributions to ecosystem services rendered by for-
est ecosystems.
3.1. Nutrient cycling
It can be safely assumed that all life relies on the presence of
fungi to recycle the basic building blocks needed for growth
and survival (de Mattos-Shipley et al., 2016). Fungi, along
with other microbes, play a crucial role in the transport, stor-
age, release, and cycling of C and nutrients such as N and P
(Rashid et al., 2016). The distinctive functions of macrofungi
such as ectomycorrhizal symbionts, decomposers, pathogens,
and predators all contribute toward nutrient cycling in a forest
ecosystem (Fig. 2). EcMF assist in the assimilation and trans-
portation of nutrients from the soil to their host plants, and
the movement of photosynthetically assimilated C from the
host plant into the soil (Lakhanpal, 2000). Alternatively, sapro-
trophic fungi degrade these host plants, releasing the assimi-
lated nutrients back into the environment, further driving the
nutrient cycle (Crowther et al., 2012a). Pathogens and preda-
tors ultimately contribute to the death of plants and animals
and the release of organic matter into the ecosystem,
providing substrates to sustain saprotrophic fungi (Boddy,
2016).
3.1.1. Role of EcMF in nutrient cycling
EcMF contribute towards nutrient cycling in two ways: 1) via
the increased rate of photosynthesis in their host plants,
and the subsequent movement of that C belowground; and
2) via the assimilation and transportation of soil nutrients
(Heinonsalo et al., 2015; Liu et al., 2020; Read and Perez-
Moreno, 2003). These combined processes are crucial to
ecosystem functions within any forest; however, they are
most pronounced in the northern latitudes, such as in boreal
forests, where ectomycorrhizal trees are the dominant plant
type (Qu et al., 2010).
Contributions of macrofungi to forest ecosystem
Fig. 2 e Roles of macrofungi in nutrient cycling.
EcMF are dependent on host plants for meeting organic C
requirements necessary for growth, respiration, and other
biological maintenance activities. Some environmental condi-
tions affect the need of EcM for C, such as the availability of P,
availability of CO2, light quality, and plant density (Jansa et al.,
2011). Studies have shown that between 5% and 50% of a
plant’s photo-assimilates can be transferred to EcM
(Deckmyn et al., 2014; Simard et al., 2003). Less than 2% of
the C in the biomass of EcMF originates from decomposing
litter, providing evidence that EcMF acquire C mostly from
their plant symbionts, via direct transfer from the host roots
(Treseder et al., 2006).
EcMF, similarly to other mycorrhizal fungi, are involved in
the nutrient cycling of the forest ecosystems by promoting the
uptake of growth-limiting nutrients such as N and P as well as
water by host plants (Liu et al., 2020). EcMF form a more bene-
ficial relationship for the host trees in nutrient-limited envi-
ronments, especially for receiving N, which is the main
growth-limiting factor in many forest ecosystems (Vitousek
et al., 2002). EcMF also facilitates the P nutrition of forest trees
(Cairney, 2011). Among the different P forms present in the
soil, only orthophosphate is taken up in large amounts by
plant symbionts and EcMF (Jansa et al., 2011). Apart from C,
N, and P, EcMF also help promote the acquisition of other ele-
ments such as calcium, magnesium, and potassium to the
host plant (Neba et al., 2016; Schmalenberger et al., 2015)
through the process of weathering minerals and rocks (Liu
et al., 2020). Weathering involves the acidification, complexa-
tion, and physical fragmentation delivering metal ions into
the soil (Wei and Jia, 2014).
EcMF use different mechanisms in nutrient regulations
such as the enzymatic and non-enzymatic (Fenton chemistry)
degradation of organic matter (Liu et al., 2020). In the case of
enzymatic reactions, EcMF produce extracellular enzymes
involved in organic matter degradation (Lu et al., 2020;
5
Tedersoo et al., 2012). EcMF, however, have lost the genetic ca-
pacity to produce lignocellulose-degrading enzymes acquired
by their saprotrophic lineages, and have accordingly devel-
oped an oxidative mechanism (Fenton reaction) to facilitate
organic N acquisition (Op De Beeck et al., 2018). This mecha-
nism is also used by brown-rot fungi in decomposing lignocel-
lulosic material in soil organic matter SOM (Rineau et al., 2012).
The interaction between EcMF and the plant host also pro-
motes the priming effects in which simple organic com-
pounds are released from the fine roots of the plant
symbionts, stimulating the growth of other microorganisms
and thereby promoting the decomposition of old organic mat-
ter needed in both C and N cycles (Lindahl and Tunlid, 2015;
Liu et al., 2020). Moreover, EcMF can compete with free-living
saprotrophs for N, thus delaying the decomposition of C and
increasing soil C accumulation, which is known as the Gadgil
effect (Sterkenburg et al., 2018). Any increases in C sequestra-
tion will ultimately reduce the amount of CO2 going into the
atmosphere, thus aiding in the mitigation of global warming
(BBC, 2022).
The role of EcMF in acquiring organic N and P from SOM
and transferring this to their plant symbiont is critical in the
understanding of plant-fungal interactions as well as the
cycling and sequestering of C and N in forest ecosystems
(Zak et al., 2019). However, there are still no comprehensive
studies quantifying the accumulative effects of EcMF in
nutrient cycling. Moreover, there is no generally accepted uni-
versal mechanism in the transfer of nutrients between fungal
and plant symbionts. The contribution of EcMF to these pro-
cesses remains poorly quantified. Evidence of the involve-
ment of EcMF in P cycling remains inconsistent and not
properly understood (Jansa et al., 2011). More work must be
done to better understand the impact on C sequestration,
the value of sequestered C, or amounts of N and P that are
extracted from the soils.
6 A. G. T. Niego et al.
3.1.2. Role of fungal decomposers in nutrient cycling
Saprotrophic fungi are heterotrophic organisms that can be
considered primary decomposers and are key drivers of
ecosystem function. They are the primary agents of plant
litter decomposition via hyphal networks that grow
throughout the soil-litter interface through which nutrients
are readily distributed (Crowther et al., 2012a). Although the
vast majority of saprotrophic fungal species are classified as
microfungi, a number of macrofungal species occur in forests
that can be found growing on fallen logs, dead or dying parts
of still-standing trees, and dead remains of other organisms
(Ferraro et al., 2022; Ye et al., 2019). Those saprotrophic macro-
fungi are distributed throughout Basidiomycota (He et al.,
2019) and Ascomycota (Salo et al., 2019). Some studies have
shown that factors affecting the diversity of wood-inhabiting
fungal communities are climate, availability of dead wood in
the surrounding landscape, and characteristics of the colo-
nized dead-wood material. However, Krah et al. (2018) demon-
strated that host tree species is also an important factor in the
diversity of wood-inhabiting decomposers. During the decom-
position process, a succession in the fungal decomposer com-
munity takes place within woody substrates (Hyde and Jones,
2002). Accordingly, the fungal decomposers are grouped into
three categories: primary, secondary, and tertiary decom-
posers, depending on the stage of wood decomposition in
which they are active (Hiscox et al., 2015).
Primary decomposers are fast-growing species that are
usually the first to grow on available organic matter, prefer-
ring an environment with little or no competition. These
fungi initiate the decomposition process, breaking down
the initial compounds within the substrate, and thus open-
ing it up for the secondary decomposers (Junior Letti et al.,
2018). The primary decomposers produce enzymes that can
digest mechanically robust and complex materials such as
carbohydrate, lipidic and proteic polymers (Junior Letti
et al., 2018), and can degrade starch, cellulose, hemicellulose,
lignin, and other components of plant material (Grimm and
Wo € sten, 2018). Some species (e.g. Pleurotus ostreatus) can
even digest recalcitrant and toxic substances such as oxo-
biodegradable plastic bags and green polyethylene (Luz et
al., 2020; Temporiti et al., 2022). Some primary decomposers
include Grifola frondosa, Lentinula edodes, and Pleurotus spp.
(Junior Letti et al., 2018; Luz et al., 2020; Zied et al., 2020).
Secondary decomposers generally degrade the remainder
of the cellulose and/or lignin within in a substrate complex,
but are also able to decompose other compounds present.
They prefer partially decomposed organic matter so they
can effectively digest and absorb required nutrients (Junior
Letti et al., 2018). Secondary decomposers often work together
with other microbes such as bacteria and other fungi to
further break down partially degraded substates (Wendiro
et al., 2019). They not only tolerate the presence of bacteria
but are able to extract nutrients from bacterial biomasses.
This group of decomposers prefers more biologically complex
environments (Wendiro et al., 2019). Termitomyces species are
an example of a secondary decomposer growing on naturally
composted plant materials collected by termites, with which
it has a symbiotic relationship (da Costa et al., 2019; Schmidt
et al., 2022). Other examples of well-known secondary decom-
poser macrofungi are Agaricus bisporus in temperate areas and
Volvariella volvacea in the tropics (Chang and Miles, 2004;
Zhang et al., 2019). C. comatus, and Macrolepiota bonaerensis
are also good examples of secondary decomposers usually
found in grassland and meadows (Junior Letti et al., 2018). Sis-
totremastrum niveocremeum is frequently found growing on rot-
ten wood (Floudas et al., 2020).
Tertiary decomposers are fungi that draw their nutrients
from the tail-end of the decomposition process, usually
growing under rotten tree stumps or logs, or can be found as
soil dwellers (Arora, 2019). These fungi gain their nutrients
from materials left over after the primary and secondary de-
composers have completed their degradation cycles (Grimm
and Wo € sten, 2018). Examples of tertiary macrofungal decom-
posers are Agrocybe, Conocybe, Mycena, and Pluteus (Stamets,
2005).
3.1.3. Role of macrofungal pathogens in nutrient cycle and
diversity
Pathogenic fungi can influence patterns of tree mortality in a
forest ecosystem, thereby influencing tree growth and
nutrient cycling (Castello et al., 1995). The Janzen-Connel hy-
pothesis is applicable in the role of pathogenic fungi. It pro-
poses that host-specific pathogens and herbivores can
maintain plant diversity in the forest ecosystem by attacking
conspecific plants (Comita et al., 2014). Eliminating plant spe-
cies will cause gaps in the forest canopy, thus creating more
opportunities for other species to thrive, increasing plant spe-
cies diversity (Hennon, 1995), and providing additional sub-
strates for the decomposers to further contribute to nutrient
turnover. Moreover, these pathogenic fungi are also brown-
rot or white-rot fungi, thereby contributing to the nutrient cy-
cle and other functions in the forest ecosystem (Arantes and
Goodell, 2014; Nagy et al., 2012).
3.2. Carbon stocking
Macrofungi are suggested as a key player in C sequestration
by helping the host plant assimilate atmospheric CO2 and
store it in the forest (either as woody material or as assim-
ilates in the forest floor litter and in the soil (Treseder and
Allen, 2000; Wang et al., 2016). The boreal forest biome
covers 11% of the total land surface and contains 16% of
the C stock sequestered in soils (Post et al., 1982), wherein
about 50e70% of sequestrated C is derived from roots and
root-associated microorganisms (Clemmensen et al., 2013).
The work of Clemmensen et al. (2013) highlights the role
that EcMF play in transporting and storing C in deeper soil
layers, thereby effectively storing this C into the soil system.
Root-associated fungi are important regulators of ecosystem
C dynamics, they not only compete with other microorgan-
isms for nutrients and C, but also decompose organic mat-
ter, transport C to deeper soil layers, and promote plant
growth, thereby increasing the rates of photosynthesis
(Macia
-Vicente and Popa, 2022; Schneider-Maunoury et al.,
2020).
Contributions of macrofungi to forest ecosystem
Fig. 3 e Mycelial growth in soil Agaricus subrufescens (A) and Stropharia rugosoannulata (B) used in soil amendments in field
conditions.
Under nutrient-limiting conditions there is a greater effect
of EcMF on soil C dynamics, since the host plants rely more
heavily on the fungal symbionts for the supply of nutrients,
which are exchanged with photosynthetic C (Orwin et al.,
2011). Evidence of this is provided by the recent study of
Sulman et al. (2019) in which the N-acquiring mycorrhizal
symbiosis increased terrestrial C sequestration when
compared to static N acquisition pathways. The C sequestered
in ecosystems dominated by EcMF is much higher, 70% more C
per unit N, than in those dominated by arbuscular mycor-
rhizal AM fungi (Averill et al., 2014).
3.3. Soil formation
Weathering is the process of chemical decomposition and
physical disintegration of rocks, substantially contributing
to soil fertility and ecosystem productivity (Hack, 2020).
This is the primary source of all the primary minerals
required for the growth of soil dwelling organisms (Banfield
et al., 1999). Fungal weathering has become an increasingly
important focus of biogeochemical research (Hoffland et al.,
2004). The spatial organization of the soil is important in
belowground functions, affecting the delivery of ecosystem
services (Ritz and Young, 2004). Macrofungi are involved in
the soil structural dynamics in the forest by mediating the
formation of soil structure via different mechanisms such
as charge, adhesive, and enmeshment (Ritz and Young,
2004). Fungi produce organic anions and cations that help
break down weak spots in solid rock substrates (Gadd,
1999). They also produce hydrophobic compounds affecting
the water infiltration properties of the soil (Ritz and Young,
2004). Mineral grains from soils are often completely covered
by fungal hyphae, thus binding the soil, as they grow over the
grains (Hoffland et al., 2004) (Fig. 2). The fungal hyphae also
7
produce organic materials that bind soil particles together,
improving their structure and porosity, and are thus an
important component of soil structural dynamics (Muneer
et al., 2020). Some species of macrofungi such as Agaricus sub-
rufescens and Stropharia rugosoannulata (Fig. 3) have been used
in the application of soil amendments promoting the degra-
dation of pollutants and formation of soil aggregates (Hu
et al., 2021).
3.4. Mycoremediation
Macrofungi are able to remediate different types of environ-
mental pollutants, through a variety of processes collectively
referred to as mycoremediation (Uddin et al., 2020; Yadav
et al., 2021). They possess the biochemical and ecological capa-
bilities to degrade recalcitrant environmental contaminants
(Arora, 2019) and convert them into useful forms (Uddin
et al., 2020). Mycoremediation relies on efficient enzymes pro-
duced by macrofungi to degrade the wide range of xenobiotics
and other various types of pollutants (Kulshreshtha et al.,
2014). The enzymes involved in the degradation of contami-
nants include lignin peroxidase (LiP), versatile peroxidase,
Mn-peroxidase, and laccase (Pozdnyakova, 2012). Discovery
of other peroxidases such as dye peroxidase also reveal the
opportunity for other industrial applications (Sigoillot et al.,
2012).
The process by which macrofungi remediate contaminated
soils can be classified in three categories: bioaccumulation, bio-
sorption, and bioconversion (Yadav et al., 2021). Toxicity levels
of contaminants such as heavy metals in macrofungi increase
due to biosorption and biodegradation, especially if they grow
in highly toxic or polluted substrates. However, toxicity reduc-
tion depends on their ability to produce different enzymes to
degrade the pollutants (Uddin et al., 2020). Macrofungi can
8 A. G. T. Niego et al.
degrade pollutants and transform them into simpler mineral
constituents (Yadav et al., 2021).
Macrofungi can degrade recalcitrant hydrocarbon contam-
inants which can turn hydrocarbon pollutants present in the
environment into an energy source, increasing the selection
of hydrocarbon-metabolizing fungal populations (Hyde et al.,
2019). For instance, the oyster mushroom, P. ostreatus, can
degrade hydrocarbons present in the soil (Steffen et al.,
2007). Pozdnyakova et al. (2019) demonstrated the ability of
10 strains of Basidiomycota belonging to genera Agaricus, Bjer-
kandera, Pleurotus, Schizophyllum, Stropharia, and Trametes that
can degrade oil hydrocarbons, with P. ostreatus, S. commune,
and T. versicolor featuring as the most active degraders.
Several macrofungi have also been demonstrated to reme-
diate heavy metals including Agaricus bisporus, Boletus edulis,
Calocybe indica, Calvatia excipuliformis, Hygrophorus virgineus,
Lepiota rhacodes, Lepista nuda, Pleurotus platypus, Pleurotus
sajor-caju, P. ostreatus, Psalliota campestris, Russula delica and
Suillus spp. (Arora, 2019; Das, 2005; Das et al., 2008). Some mac-
rofungal species are efficient degraders of high molecular
weight PAHs. Examples of such fungi include: Agaricus (Li
et al., 2010), Armillaria (Hadibarata and Kristanti, 2013), Gano-
derma (Agrawal et al., 2018), Marasmiellus (Vieira et al., 2018),
Phanerochaete (Wang et al., 2009), Pleurotus (Hadibarata and
Teh, 2014; Li et al., 2010), and Trametes (as Coriolus) (Jang
et al., 2009). They outperform bacteria in hydrocarbon degra-
dation, given bacterial specialization in degrading smaller
molecules (Peng et al., 2008).
Heavy metals found in soils can be toxic, even at low con-
centrations, to both plants and soil organisms, and can accu-
mulate in the food chain (Singh et al., 2008). However,
macrofungi have been shown to help mitigate the negative ef-
fects of soil heavy metals, and improve plant growth as a
result. Macrofungi can absorb heavy metals from substrates
or soils, which can be an effective method of bioremediation,
and a mechanism to reclaim polluted lands (Uddin et al., 2020).
The absorption of heavy metals by fungi can result in the
accumulation of these heavy metals in their network of hy-
phae, where the metals are bound and often stored. As a
result, the fruiting bodies produced by these fungi often
possess high levels of heavy metals and thus caution is
advised when eating mushrooms collected from areas known
to have heavy metal contamination of the soils (Ab Rhaman
et al., 2021; Elekes et al., 2010). Macrofungal species such as
A. bisporus, Amanita spissa, Fomes fomentarius, Lactifluus pipera-
tus (as Lactarius piperatus), Phellinus badius, Suillus grevillei and
Tricholoma terreum have been shown to effectively extract
metals such as cadmium (Cd), chromium (Cr), copper (Cu),
lead (Pb), mercury (Hg), iron (Fe), and zinc (Zn) (Ali et al., 2017).
White-rot fungi are also effective in the transformation of
pesticides and degradation of wastes containing cellulose in
pulp and paper industries (Malik et al., 2021). Pleurotus was
one of the most promising macrofungi for the biodegradation
of agro-pesticides because of its ligninolytic capabilities
(Camacho-Morales and Sa
nchez, 2016; Du et al., 2018). G. fron-
dosa has been shown to be able to selectively degrade a variety
of agroforestry residues (Galic et al., 2021). Trametes versicolor
was also able to degrade medium to highly polar pesticides
such as the imidacloprid, neonicotinoids acetamiprid and
organophosphate malathion (Hu et al., 2022).
3.5. Role of fungal spores in rain formation
Millions of tons of fungal spores are released into the atmo-
sphere annually, forming important components of biogenic
aerosols, specifically the hydrophilic spores discharged by
Ascomycota and Basidiomycota (Elbert et al., 2007). Around
30,000 basidiospores can be released by a gilled macrofungus
in 1 s, and it is estimated that billions of aerosols are released
per day by a single basidiome (Dressaire et al., 2016; Hassett
et al., 2015). In a further effort to quantify the amount of
spores released, and thus emphasize the impact of fungal
spores in the atmosphere, Elbert et al. (2007) estimated that
approximately 50 million tons of fungal spores are dispersed
annually. These spores can be released at specific times of
day or at irregular intervals (Lagomarsino Oneto et al., 2020).
The spores have been shown to be hygroscopic and serve as
cloud condensation nuclei, critical to initiating precipitation
that results in rain formation (Rosenfeld et al., 2008). Mannitol
and other hygroscopic sugars are secreted from the spore sur-
face, resulting in the condensation of water at the spore sur-
face, ultimately leading to an increase in localized rain
events (Hassett et al., 2015).
3.6. Food source for wildlife
Many EcMF rely on animal consumption, mainly by mam-
mals and insects, for their spore dispersal (Blaschke and
Ba€ umler, 1989; Nuske et al., 2019). Changes in mammalian
communities directly affect the diversity of the truffle-like
EcMF as shown in the study of Nuske et al. (2019). It was
assumed that mammal-mediated dispersion is the main
mechanism for the long-distance dispersal of EcMF spores
(Policelli et al., 2022). This dependence links animal
mycophagy to both EcMF diversity and ecosystem health.
Macrofungi are an important source of nutrition for wild an-
imals. Many deciduous forest animals such as squirrels, ro-
dents, deer, boars, slugs, and insects consume mushrooms
(Ori et al., 2021; Wayking, 2021). Specifically, small mammals
such as northern flying squirrels (Glaucomys sabrinus) and
red-backed voles (Myodes gapperi) have diverse fungal food
preferences, including both mycorrhizal and non-
mycorrhizal fungi in Ascomycota, Basidiomycota, Zygomy-
cota, Chytridiomycota, Glomeromycota, and other unidenti-
fied fungal taxa (Cloutier et al., 2019). Other small mammals
such as mice (Apodemus spp.), bank voles (Myodes glareolus),
and shrews (Sorex araneus and S. minutus) prefer hypogeous
fungi (Katarz yte_ and Kutorga, 2011). Approximately 6.1 kg/an-
imal/year of fresh weight fungi is consumed by an animal
with a weight of 500 g (Hanson et al., 2003). Furthermore,
many primates eat mushrooms, the Goeldi’s monkey (Calli-
mico goeldii), for example, feeds on Auricularia auricula, A. mes-
enterica, Ascopolyporus polyporoides and A. polychrous (Hanson
et al., 2003). Natural disturbances or human interventions
have brought about changes in forest composition, impacting
fungal sporocarp productivity and diversity, which has
downstream effects on food availability for mycophagous
wildlife (Dejene et al., 2021; North et al., 1997).
Mycophagus insects, especially within Coleoptera, and
macrofungi have long been documented to engage in mutual-
istic symbioses, with the insects gaining nutrition in exchange
Contributions of macrofungi to forest ecosystem
for spore dispersal and protection (Biedermann and Vega,
2020). Past research has suggested that there is a strong link
between the fungal community composition and the diversity
of mycophagus insects within a forest ecosystem (Yamashita
and Hijii, 2007). Numerous species of ants, termites and bee-
tles can cultivate macrofungi, in fungal gardens, which serve
as a source of nutrition for the insects (Epps and Arnold,
2018; Nicoletti and Becchimanzi, 2022). For example, species
of fungi from Termitomyces are cultivated in the nests of Mac-
rotermitinae termites (Ono et al., 2017), whereby the termites
collect plant material which is deposited in fungal gardens
within the termite nest. The Termitomyces fungi then digest
this material, which, in turn, acts as a food source for the ter-
mites (Ono et al., 2017; Vesala et al., 2019).
Direct contributions to ecosystem services rendered by for-
est ecosystems.
3.7. Food, income source and other uses of macrofungi to
humans
Wild mushrooms are important non-timber forest products
that provide food and income for local communities from
many parts of the world (Cai et al., 2011; Milenge Kamalebo
et al., 2018). Different studies show that people use wild mac-
rofungi either for food, medicine, recreation, or in relation to
certain beliefs or myths (Boa, 2008; Milenge Kamalebo et al.,
2018; Mortimer et al., 2021; Osarenkhoe et al., 2014). Approxi-
mately 2200 species of wild fungi are edible, of which more
than 1000 species are used as food (H. Li et al., 2021). Macrofun-
gal foraging is becoming an increasingly popular leisure activ-
ity in many countries (Egli et al., 2006; Marini Govigli et al.,
2019; Schulp et al., 2014). The collection and sales of wild edible
macrofungi account for 8% of the global macrofungal produc-
tion, which equated to approximately USD 5 billion in 2013
(Royse et al., 2017).
In addition to food and cuisine, macrofungi offer numerous
health benefits, including anticancer, antidiabetic, antimicro-
bial, antiviral, anti-inflammatory, anti-oxidant, cholesterol-
lowering and neuroprotective properties (Hyde et al., 2019;
Niego et al., 2021a, 2021b). About 270 species of macrofungi
have been reported to have medicinal value (Shamtsyan,
2010). Some of the most explored species for their medicinal
properties are Antrodia cinnamomea, Ophiocordyceps sinensis,
Ganoderma lucidum, Phellinus linteus, and Xylaria nigripes (Hyde
et al., 2019; Niego et al., 2021a). Some of the more common
bioactive compounds found in fungi include polysaccharides
(Cai et al., 2012; Liang et al., 2019; Veena and Janardhanan,
2022), proteins (El Enshasy and Hatti-Kaul, 2013), glycopro-
teins (Cohen et al., 2014), phenolic compounds (Gupta et al.,
2018), tocopherols, ergosterol (Diallo et al., 2020), unsaturated
fatty acids, lectins (Ismaya et al., 2020).
3.8. Interrelationship between the functions of macro-
fungi in forest’ ecosystem processes
The active roles of macrofungi in the forest’s ecosystem pro-
cesses, whether with direct or indirect effects to humans,
are all important to healthy and functional forests (Zotti
et al., 2013). These contributions are manifold and well estab-
lished. EcMF form beneficial symbiotic relationships with the
9
roots of 80%e90% of all temperate and boreal forest trees and
are therefore essential contributors to forest ecosystem func-
tioning (Stuart and Plett, 2019). The services rendered by mac-
rofungi are interrelated and somewhat overlapping. The
different roles macrofungi perform in nutrient cycling are of
utmost importance. Macrofungi ensure that nutrients are
cycled and made available for plant growth-enhancing forest
health and productivity. Healthy forests can support a greater
number of living organisms, thus enhancing species diversity.
The capability of macrofungi to cycle nutrients also influences
C sequestration as well as soil formation. Giving the growing
urgency of climate change mitigation, C sequestration capa-
bilities of forests with the help of macrofungi can play a vital
role. Spores of macrofungi can also initiate rain formation as
mentioned above, affecting the water cycle in forest
ecosystem. These services have indirect effects to humans
and wildlife.
Functions of macrofungi in the forest ecosystem processes
can also have direct benefits to humans (Sharma et al., 2022).
Macrofungi are beneficial not only for the vital roles they
play in the ecosystem functions but also for their influence
on human and human-related activities (Zotti et al., 2013).
Just like wildlife that consume the spore-producing bodies of
macrofungi, humans also consume many edible macrofungi
from forests. Edible macrofungi can support livelihoods in ru-
ral communities via wild mushroom foraging and sale or via
enriching household nutrition (Chauhan, 2021). These pro-
mote sustainable economic growth with minimal impact to
the forest (Mart
ınez de Arago
n et al., 2011). In return, human
and wildlife mycophagy have become important primary
mechanisms for macrofungal dispersal, thus increasing their
diversity and survival. The dispersal of the spores of EcMF
through mycophagy also actively shapes plant communities.
3.9. Future perspectives
Forest macrofungi are non-timber forest products with
diverse genetic resources. They contain immense value
because of their ecological, economic, medicinal, socio-
cultural, and biotechnological roles and applications. Their
importance in forest sustainability is well-established. The
functions of macrofungi in forest ecosystems are often
undervalued since they are considered “free” services. Few
existing policies recognize their contribution to ecosystem
functioning.
Since people depend on the multiple benefits provided by
forest ecosystems, it is essential to understand ecosystem ser-
vices and properties as well as their responses to drivers of
change (e.g., climate change, deforestation, urbanization).
Habitat transformation can drastically alter macrofungal
community structures and diversity, which could diminish
their ecosystem functioning (Go
mez-Herna
ndez et al., 2021).
It is important to understand and apply functional traits
studies to macrofungal communities integrated with ecolog-
ical and evolutionary perspectives to determine their impact
on the ecosystem and tolerance to environmental change
(D
ıaz et al., 2013). Standardized methodologies for collecting
data on traits across the taxonomic group and spatial distribu-
tion are also necessary for the precise understanding of mac-
rofungal communities in forest ecosystems (Bazzicalupo et al.,
10
2022; Dawson et al., 2019; Runnel et al., 2021). Proper identifica-
tion, characterization and conservation of macrofungal
diversity are necessary to understanding and forecasting
ecosystem cycling and preserving fungal bioresources
(Lofgren and Stajich, 2021).
In sustainable development, it is necessary to achieve a
balance between socio-economic development, exploration
of natural resources, and conservation of ecosystems. It is
important to understand the functions of macrofungi in the
maintenance of healthy forests as well as their benefits to hu-
mankind. By recognizing the ecosystem services and func-
tions of macrofungi, we hope to increase awareness of their
importance and of the necessity to consider them in environ-
mental policymaking, leading to their inclusion in conserva-
tion efforts. Furthermore, the role that macrofungi play in
carbon capture and the potential for mitigating the effects of
climate change needs to be further explored and discussed
at a higher level, as this topic needs to be considered in the
context of environmental policies in the future.
Declaration of competing interest
The authors declare that this manuscript has not been pub-
lished previously and have no conflict of interest to disclose.
Acknowledgements
This work was in part supported by the Thailand Science
Research and Innovation grant “Macrofungi diversity research
from the Lancang-Mekong Watershed and surrounding areas”
(Grant No. DBG6280009), the National Research Council of
Thailand (NRCT) grant “Total fungal diversity in a given forest
area with implications towards species numbers, chemical di-
versity and biotechnology” (Grant No. N42A650547) and Mae
Fah Luang University Thesis/Dissertation Support Grant
(Reference No. Oh 7702(6)/49). We would like to acknowledge
Austin G. Smith for language editing the manuscript. We
also would like to thank Yuwei Hu for providing photos of A.
subrufescens and S. rugosoannulata mycelial growth in the soil
under field conditions.
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