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\begin{document}

\title{Establishing a Standardized Approach for Elucidating Glomeromycota
Life-History Traits: Advancing Consistency in Mycorrhizal Fungal
Research}



\author[1]{Pedro Madeira Antunes}%
\author[2]{Sidney L. Stürmer}%
\author[3]{James D. Bever}%
\author[4]{Pierre-Luc Chagnon}%
\author[5]{V. Bala Chaudhary}%
\author[6]{Coline Deveautour}%
\author[1]{Catherine Fahey}%
\author[7]{Vasilis Kokkoris}%
\author[8]{Ylva Lekberg}%
\author[9]{Jeff R. Powell}%
\author[10]{Carlos A. Aguilar-Trigueros}%
\author[9]{Haiyang ZHANG}%
\affil[1]{Biology Department, Algoma University, Sault Ste. Marie, Ontario, Canada, P6A 2G4}%
\affil[2]{Departamento de Ciências Naturais, Universidade Regional de Blumenau, Blumenau, SC, 89030-903, Brazil}%
\affil[3]{Kansas Biological Survey and Center for Ecological Research and Department of Ecology and Evolutionary Biology, University of Kansas, Lawrence, Kansas, USA}%
\affil[4]{Institut de Recherche en Biologie Vegetale, Universite de Montreal, 4101 Sherbrooke Est, Montreal, QC, H1X2B2, Canada}%
\affil[5]{Department of Environmental Studies, Dartmouth College, Hanover, New Hampshire, USA}%
\affil[6]{UniLaSalle, Rouen, France}%
\affil[7]{Amsterdam Institute for Life and Environment (A-LIFE), Faculty of Science, Section Systems Ecology, Vrije Universiteit Amsterdam, De Boelelaan 1085, 1081 HV Amsterdam, the Netherlands}%
\affil[8]{MPG Ranch \& Department of Ecosystem and Conservation Sciences, W.A. Franke College of Forestry and Conservation, University of Montana, Missoula, Montana, USA}%
\affil[9]{Hawkesbury Institute for the Environment, Western Sydney University, Penrith, New South Wales, Australia}%
\affil[10]{Institute of Biology, Freie Universität Berlin, Altensteinstrasse 6, 14195 Berlin, Germany.}%


\vspace{-1em}



  
  \date{May 26, 2024}


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\textsuperscript{1}Pedro M Antunes*, \textsuperscript{2}Sidney L.
St\selectlanguage{ngerman}ürmer*, \textsuperscript{3}James D.
Bever,\textsuperscript{4}Pierre-Luc Chagnon, \textsuperscript{5}V. Bala
Chaudhary, \textsuperscript{10}Coline
Deveautour,\textsuperscript{1}Catherine Fahey,
\textsuperscript{6}Vasilis Kokkoris,\textsuperscript{7}Ylva Lekberg,
\textsuperscript{8}Jeff R. Powell,\textsuperscript{9}Carlos A.
Aguilar-Trigueros,\textsuperscript{8}Haiyang Zhang

\textsuperscript{1}Biology Department, Algoma University, Sault Ste.
Marie, Ontario, Canada, P6A 2G4

\textsuperscript{2}Departamento de Ciências Naturais, Universidade
Regional de Blumenau, Blumenau, SC, 89030-903, Brazil

\textsuperscript{3}Kansas Biological Survey and Center for Ecological
Research and Department of Ecology and Evolutionary Biology, University
of Kansas, Lawrence, Kansas, USA

\textsuperscript{4}Institut de Recherche en Biologie Vegetale,
Universite de Montreal, 4101 Sherbrooke Est, Montreal, QC, H1X2B2,
Canada

\textsuperscript{5}Department of Environmental Studies, Dartmouth
College, Hanover, New Hampshire, USA

\textsuperscript{6}Amsterdam Institute for Life and Environment
(A-LIFE), Faculty of Science, Section~Systems Ecology, Vrije
Universiteit Amsterdam, De Boelelaan 1085, 1081 HV Amsterdam, the
Netherlands

\textsuperscript{7}MPG Ranch \& Department of Ecosystem and Conservation
Sciences, W.A. Franke College of Forestry and Conservation, University
of Montana, Missoula, Montana, USA

\textsuperscript{8}Hawkesbury Institute for the Environment, Western
Sydney University, Penrith, New South Wales, Australia

\textsuperscript{9}Institute of Biology, Freie Universität Berlin,
Altensteinstrasse 6, 14195 Berlin, Germany.

\textsuperscript{10}UniLaSalle, Rouen, France

*Equal contribution

\textbf{The problem}

A trait is an attribute influencing an organism's performance within its
environment, encompassing morphological, genetic and physiological
characteristics measured at the individual or population levels
(Salguero-Gómez \emph{et al.} , 2018; Zhang \emph{et al.} , 2023a).
Understanding the ecology of species using a trait-based approach can
contribute to a mechanistic explanation of processes mediated by
microbes, including those that affect ecosystem functioning
(Romero-Olivares \emph{et al.} , 2021). This approach holds particular
significance for arbuscular mycorrhizal (AM) fungi - Phylum
Glomeromycota. As obligate symbionts of plants, where multiple species
colonize both roots and soils in a network, predicting the functional
outcomes (e.g., host growth, plant community diversity, soil
characteristics) of individual AM fungal genotypes and communities
within ecosystems remains challenging, despite major developments in
molecular methods in the last two decades (Tisserant \emph{et al.} ,
2013; Montoliu-Nerin \emph{et al.} , 2021). Indeed, establishing
connections between AM fungal taxa and/or genotypes (e.g., within
species) and their functional roles is a laborious process, which is
expected to continue in the foreseeable future (Serghi \emph{et al.} ,
2021; Manley \emph{et al.} , 2023). This is needed due to the complex
links between AM fungi and functional outcomes for both hosts
(\emph{e.g.} , plant growth and fitness, nutrient uptake and stress
tolerance) and soil functions/properties (\emph{e.g.} , carbon storage,
aggregate stability, biotic diversity), which appear to be highly
context dependent and relatively poorly predicted by taxonomy alone
(Munkvold \emph{et al.} , 2004; Koch \emph{et al.} , 2017; Yang \emph{et
al.} , 2017; Qiu \emph{et al.} , 2021). However, this effort is also
required because AM fungal traits have not been systematically assessed
alongside with hypotheses of adaptation or with specific mechanisms in
mind. For example, small-spored AM fungi may be dispersed longer
distances by wind than large-spored AM fungi. It is then a reasonable
hypothesis that small spore size is an adaptation for wind dispersal.
One could empirically observe that small-spored AM fungi are
geographically more widespread than large-spored fungi and this
potential result could be viewed as evidence in support of this
hypothesis. However, this finding would not necessarily prove that such
dispersal difference has ``functional'' or ``adaptive'' value.
Alternatively, producing small spores is a correlated response to
producing many spores quickly, which itself could be an adaptive
response to the likelihood of unpredictable soil disturbance such as
from tillage. In this scenario, the adaptation and/or function is the
production of many spores quickly to confer resistance to disturbance
and then, after soil disturbance with wind erosion, small spores may
also be blown farther (which may or may not improve fitness). Another
example, variation in rooting depth among plants in a community may
contribute to resource partitioning. But the mechanism (differential
resource depletion with depth) still needs to be demonstrated separately
from the trait evidence. AM fungi could contribute to equalize resource
partitioning if plants with short roots associate with AM fungi that
form more extensive extra-radical mycelium and vice-versa. Given these
complexities, we consider the development of a robust, universally
applicable trait-based framework for predicting key AM fungal functional
outcomes a priority. To achieve this objective, first we must identify
AM fungal traits that can be measured at morphological, physiological,
and genetic levels. Second, considering the important roles of AM fungi
in ecosystems, affecting host plants, soil, and the AM fungi themselves,
we need to discern/hypothesize how measuring AM fungal traits impacts
each of these components. For the host plant, it is crucial to consider
nutrition, biomass, fitness, and survival in face of pathogens, heavy
metals, salinity, drought, etc. (Delavaux \emph{et al.} , 2017;
Wehner\emph{et al.} ). Within the soil environment, AM fungal effects on
soil structure (Rillig \& Mummey, 2006), nutrient cycling, carbon
storage, and other members of the soil food-web are paramount (Antunes
\& Koyama, 2016; Frew \emph{et al.} , 2021; Horsch \emph{et al.} ,
2023a). Regarding the fungal organism, we should focus on key aspects of
their life-history strategies; reproduction and fitness, survival,
dispersal, competitive ability, infectivity and abundance both within
the host and soil environments (Aguilar-Trigueros \emph{et al.} , 2019;
Chaudhary\emph{et al.} , 2020; Deveautour \emph{et al.} , 2020). This
requires identifying relevant proxies (sometimes termed ``soft traits''
in the plant ecophysiology literature) to provide easy-to-measure
quantitative metrics for such complex facets of fungal life-history that
can be measured across several species. Third, we need to evaluate
existing standardized methods and experimental designs, or develop new
ones, to measure such relevant (soft) traits, as has been done in plant
ecophysiology (Pérez-Harguindeguy \emph{et al.} , 2013). Measurement
standardization and relevant metadata for hypothesis-driven analysis and
interpretation is essential if we are to aggregate trait information
from different studies into a public database, facilitating their
incorporation into earth system models (e.g., (Fry \emph{et al.} , 2019)
and enhancing the predictability of functional processes and/or
adaptations associated with AM fungi. Analogous libraries on plant
traits (Kattge \emph{et al.} , 2020) have proved useful to better
understand trait variation along global climatic gradients
(Butler\emph{et al.} , 2017). Here, we aim to:

\begin{enumerate}
\tightlist
\item
  To comprehensively catalog and define AM fungal functional traits
  (morphological, physiological/phenological, and genetic) while
  avoiding redundancy.
\item
  To elucidate the relationships between these traits and their
  functional outcomes for host plants, soil environments, and the AM
  fungi themselves.
\item
  To critically review the historical methods and experimental designs
  employed in measuring AM fungal traits, highlighting their strengths
  and limitations.
\item
  To propose standardized methodologies and protocols for measuring AM
  fungal traits.
\item
  To explore the integration of AM fungal trait information into
  ecological models to enhance ecosystem processes' predictability.
\end{enumerate}

\textbf{Contextual basis}

The scientific literature on the life-history traits of AM fungi (i.e.,
the biological characteristics and features that influence their growth,
reproduction, and survival) has predominantly centered on aspects
related to plant growth and nutrition, largely through an agronomic
lens. Although not explicitly reported as such, early studies employing
experimental approaches to assess, for example, AM fungal root
colonization, abundance of external hyphae, and spore counts for
specific species under certain experimental conditions have yielded
insights into AM fungal trait variation (Abbott, 1982; Reich, 1988;
Jakobsen \emph{et al.} , 1992a; Gazey \emph{et al.} , 1992; Bever
\emph{et al.} , 1996). Given the wide variation observed, these and
other seminal studies provided a foundation for further inquiry into the
complex dynamics of AM fungal life-history traits and their broader
implications to the AM symbiosis.

Studies of distinct traits within a taxonomic framework started with the
comparison of mycelium form and function, and root colonization
strategies among major families of the Glomeromycota. For example,
(Dodd\emph{et al.} , 2000) compared the morphology and mycelial
architecture of different AM fungal genera, discussing form and
function, (Hart \& Reader, 2002b) showed in a comparative study of 21 AM
fungal isolates (i.e., an AM fungus isolated in the laboratory into pure
culture but without genetic characterization, at which point it becomes
a certified strain with a collection number) spanning 16 species from
North America that the isolates of the Glomeraceae family, on average,
colonized roots before those of Acaulosporaceae and Gigasporaceae
families. Additionally, the proportion of fungal biomass in roots versus
soil also diverged, on average, among isolates of those families. Those
in the Glomeraceae exhibited high root colonization but low soil
colonization, Gigasporaceae tended to have low root colonization but
high soil colonization, while Acaulosporaceae displayed low colonization
in both roots and soil. These findings revealed a strong association
between AM fungal function and taxonomy for these fungi, as isolates
from the main families could be differentiated based on colonization
rate, biomass allocation, and the onset of sporulation. These
observations were corroborated by subsequent studies, albeit using AM
fungi from the same community and, possibly, the same isolations (Hart
\& Reader, 2002b, 2005; Maherali \& Klironomos, 2007; Powell \emph{et
al.} , 2009; Sikes\emph{et al.} , 2009). In fact, using the same data,
Aguilar-Trigueros et al. 2019, showed that large-spore species produced,
on average, fewer spores than small-spore species, suggesting that AM
fungi experience similar resource allocation constraints during
reproduction as plants seeds (Moles \emph{et al.} , 2005). Results from
these studies suggest differences between Glomeraceae and Gigasporaceae
concerning life-history traits and their relationship with host
benefits. However, new comparative studies that include more fungal
species isolated from other ecological contexts are necessary to confirm
these differences.

The patterns described above underscore the utility of employing a
comparative framework to test hypotheses concerning AM fungal function
by examining trait expression. For instance, based on soil mycelium
production, Gigasporaceae would be expected to outperform Glomeraceae in
nutrient uptake (Maherali and Klironomos, 2007). Alternatively, if early
or extensive root colonization (with abundant coils/arbuscules) is more
important for nutrient delivery to the host, then Glomeraceae could be
more beneficial partners under nutrient limiting conditions (e.g.,
(Horsch \emph{et al.} , 2023b). Despite inconsistencies among studies,
which may to some extent be explained by variability in mycorrhizal
dependency among hosts (Pringle \& Bever, 2008; Sikes \emph{et al.} ,
2009), a meta-analysis (Yang \emph{et al.} , 2017) suggested that, on
average, Glomeraceae were better at acquiring P and reducing pathogen
growth compared to other AM fungal families. Is it also of interest,
that this family also appears to be the most abundant in many locations
(Öpik \emph{et al.} , 2010).

The preceding studies lacked a broader environmental perspective. For
instance, considering diverse environmental conditions, such as varying
soil types or climatic factors, could unveil how AM fungal traits
respond and adapt. Currently, most data reporting the impact of
different AM fungi on their host originate from short-term experiments,
using fungal taxa that readily sporulate and are easily amenable to pure
cultures (Ohsowski \emph{et al.} , 2014) which may not reflect the
reality in natural environments. The study by Sikes et al. (2009)
investigating differences in plant pathogen protection between AM fungal
taxa, as well as that of (Lerat \emph{et al.} , 2003) on C-sink strength
among different AM fungal families suggests that certain functional
outcomes resulting from the symbiosis depend on the combination of plant
and fungal traits (Johnson \emph{et al.} , 1997). As such, considering
fungal traits alone (i.e., in absence of plant and soil characteristics)
may limit predictions of functional outcomes of the symbiosis. This
brings an additional layer of complexity to the study of AM fungal
ecophysiology, or trait-based ecology, as intricate relationships
between fungal and plant traits are to be expected (Chagnon \emph{et
al.} , 2013).

\textbf{Proposed trait-based frameworks for AM fungi}

(Van Der Heijden \& Scheublin, 2007) conducted the first comprehensive
review on AM fungal traits to predict plant growth and ecosystem
functioning. The authors provided a list of 13 AM fungal functional
traits, which they categorized into morphological traits
(\emph{e.g.,}hyphal length, rate and extent of root colonization, spore
production) and physiological traits (\emph{e.g.} , fungal carbon
acquisition, host preference, nutrient uptake efficiency). Subsequently,
(Behm \& Kiers, 2014) noted substantial intraspecific trait variation
among AM fungal species (also see (Koch \emph{et al.} , 2017)),
complicating the characterization of traits and their incorporation into
functional trait models. To address this issue, they proposed a
five-part framework for characterizing intraspecific trait variation of
AM fungal species within the context of nutrient cycling, based on
experimental design and trait measurement considerations. According to
Behm and Kiers (2014), AM fungal genetic units should be subjected to
diverse environmental conditions (e.g., host plants, soil nutrient
concentrations). Subsequent measurements would encompass the degree of
variation, trait reversibility, relationships among traits, the adaptive
nature of variation, and the potential for variation to evolve. Through
these five dimensions, researchers could map traits onto an evolutionary
tree and incorporate them into functional models for predicting nutrient
cycling dynamics.

Chaudhary et al., (2022) highlighted the challenges in defining traits
for organismal networks such as the mycorrhizal symbiosis. They proposed
a unified trait framework, complemented by a standardized vocabulary,
with the objective of establishing a clear connection between
trait-based mycorrhizal ecology, AM fungal niches and community assembly
rules. The authors categorized traits into three main groups:
morphological, physiological, and phenological. Within each of these
categories, they pinpointed distinctive mycorrhizal traits specific to
both the host plant (such as root:shoot ratio, growth form,
photosynthetic pathways) and the fungal partner (e.g., spore size,
hyphal length, and melanin content). Beyond these bifurcated traits by
plant or fungal traits, (Chaudhary \emph{et al.} , 2022) introduced the
concept of mycorrhizal traits. These are unique attributes that emerge
during symbiosis and are co-dependent on both partners. They encompass
aspects such as root colonization-induced structures, plant mycorrhizal
response, and resource exchange rates. This novel framework provides an
enriched understanding of mycorrhizal ecology and serves as a basis for
the empirical framework proposed here.

(Chagnon \emph{et al.} , 2013) put forth an AM fungal trait-based
framework building on Grime's CSR (competitive, stress-tolerant,
ruderal) framework - which identifies stress, disturbance and
competition as the major filters driving trait selection and evolution
in natural communities. By allowing speculative connections to be made
regarding potential linkages between fungal traits (e.g., hyphal fusion,
sporulation phenology, carbon sink strength, growth rates, etc.) and
environmental filters (e.g., soil disturbances, scarce C transfer from
host, low soil pH), this framework could tentatively identify priority
traits for measurement, and combinations of host and fungal traits that
may lead to the highest mutual benefits. Building on the apparent
family-level conservatism of many traits or responses to environmental
filters, parallels were drawn between AM fungal major families and C, S
and R strategies. However, as stressed by Chagnon et al. (2013), this
family-to-strategy association is simplistic and unlikely to stand the
test of time. In addition, it several AM fungal families (e.g.,
Pacisporaceae, Entrophosporaceae, Diversisporaceae, or more basal
lineages in the Paraglomerales or the Archaeosporales). It also fails to
consider the relative distribution of different AM fungal families in
certain biomes or at certain latitudes. For example,
\emph{Acaulospora}is a common genus in the tropics, where it can be
dominant both in natural forests and under intensive land-use where
ruderal traits are crucial (e.g., (González-Cortés \emph{et al.} ,
2012). The primary significance of the CSR framework in AM fungal
trait-based ecology should not be considered as merely a framework for
associating families with strategies. Instead, it should be recognized
as a tool for leveraging well-established life-history trade-offs in
plant ecology to pinpoint pertinent fungal traits that should be
incorporated into our research agenda.

We build upon prior frameworks, emphasizing that the discrepancies
observed among studies, stemming from non-standardized experimental
approaches and the absence of a comprehensive database on AM fungal
traits, represent significant barriers to achieving a more predictive
understanding of AM fungal ecology. Moreover, the validity and relevance
of the isolates and species employed in these studies are reliant on the
taxa available in culture collections or from a few natural communities.
A deliberate inclusion of numerous uncultured taxa, or other taxa
hitherto overlooked fungal mutualisms in conjunction with AM
fungi---such as Mucoromycotina, as suggested by (Hoysted \emph{et al.} ,
2023) ---remains an important task. Given the existing data showing
large variability in plant and soil responses to the AM symbiosis both
among and within AM fungal species, we must address these issues to
assess if, and to what extent, AM fungal traits determine growth
responses or effects on ecosystems.

\textbf{Traits and Function}

\textbf{Morphological and physiological traits}

Arbuscular mycorrhizal fungal traits, including for example hyphal
length, arbuscule morphology, or the robustness of spore walls, can
modulate key functions/processes with ramifications not only to the
health of the fungus itself but also the associated plant and the soil
environment (see Table 1 for detailed descriptions of key traits, their
hypothesized function, and methods for trait measurement). Here we
define AM fungal traits primarily as ``functional markers'', which are
indicators of mycorrhizal function and are dependent on the
morphological, physiological, or phenological characteristics of the
fungal partner (Chaudhary \emph{et al.} , 2022). In this context, AM
fungal traits are most likely instrumental in defining ecosystem
resilience and adaptability to environmental fluctuations, as certain
fungal isolates with specific traits may demonstrate superior robustness
or flexibility under changing conditions.

Conceptualizing the form and function of AM fungal traits becomes
clearer when contextualized within the lifecycle of the fungal organism.
We can broadly categorize the lifecycle of an AM fungus into two phases:
(1) the asymbiotic phase, in which the dispersed spores are activated,
germinate and explore the soil for a compatible host, and (2) the
symbiotic phase, which includes the four following stages: a) the
initiation of root colonization; b) the formation of structures within
the root cortex; 3) the extension of mycelium into the soil matrix and
possibly other hosts; and (4) spore production and dispersal. Briefly,
spores, hyphal networks, and colonized root fragments, identified as the
three principal types of propagules, remain dormant until the proper
abiotic/biotic conditions emerge (Lanfranco \emph{et al.} , 2018).
Hyphae emerging from these propagules identify a host root, adhere to
its surface, and commence root colonization. A swollen hyphopodium forms
subsequently, from which a single hypha penetrates the root epidermis to
access the root cortex. A series of morphogenetic and molecular
processes come into play at these initial stages, enabling the plant to
identify the presence of the fungus (as reviewed by (Bonfante \&
Perotto, 1995; Gianinazzi-Pearson \emph{et al.} , 2007; Bonfante \&
Genre, 2010; Luginbuehl \& Oldroyd, 2017). Upon reaching the root
cortex, the fungus colonizes intercellular spaces, forming the
intraradical mycelium (IRM). This mycelium then differentiates into
structures such as arbuscules or coils, and, in some taxa, vesicles/
spores and the extraradical mycelium (ERM) which consists of runner
hyphae, branched absorbing structures (BAS), spore associated BAS, and
''asexual'' spores. Upon attaining a certain threshold of root
colonization, hyphae extend beyond the root system into the soil matrix,
forming the extraradical mycelium (ERM) and generating asexual spores.
The expansive hyphal network, comprising IRM and ERM and spores,
embodies the traits that underpin several ecosystem-level processes
attributed to AM fungi. These traits impact not just the host plants and
soil environment, but also the fungal organism itself.

AM fungal spores

Arbuscular mycorrhizal fungal spores are among the largest
(Aguilar-Trigueros \emph{et al.} , 2023) and most multinucleated spores
(Kokkoris et al., 2020) known in the kingdom Fungi and exhibit the
phenotypic characteristics that enable species identification. Three
types of spore formation are recognized (Walker \emph{et al.} , 2018).
Glomoid spores are formed blastically at the tip of a hyphae or by
intercalary inflation of a hypha. Acaulosporoid spores involve the
blastic formation of a sporiferous saccule with a neck, followed by the
differentiation of spores laterally or inside the neck, or inside the
sporiferous saccule. Gigasporoid spores are differentiated at the tip of
a small bulb or suspensor cell.

Spores range widely in their traits including size, shape, color, and
wall thickness (see (Morton, 1988) for a review) across and within
species. In fact, single isolates of some species are known to produce
more than one type of spores (even the model fungus, \emph{Rhizophagus
irregularis} (Kokkoris \emph{et al.} , 2023)). It has been reported that
spores can be produced singly in the soil, in loose aggregates and in
small or large sporocarps. The cytoplasm of the spores contains not only
the nuclei (ranging from hundreds to thousands) but also lipid reserves
that assist the germination and early colonization. Based on spore
ontogeny, three main phenotypic characters are observed in AM fungal
spores: spore wall, germinal walls, and germination structure - with the
latter two not present in many species. In addition, the spore walls of
many species show appendages and indentations. Some AM fungi produce
sporocarps (i.e., aggregations of spores), which are called
``glomerocarps'' (Jobim \emph{et al.} , 2019). Sporocarps have functions
in reproduction (Yamato \emph{et al.} , 2022) and dispersal, including
dispersal by mammals (Mangan \& Adler, 2002). Overall, variation in
spore traits across species are hypothesized to reflect differences in
reproduction (investing in fewer larger spores or many small spores),
dispersal (long or short, different dispersal vector), survival in the
absence of the host (e.g., resistance to desiccation and pathogenesis)
and early colonization strategies (Chaudhary \emph{et al.} , 2018).

\textbf{Intraradical mycelium (IRM)}

The AM fungal mycelial system colonizes two distinct environments: the
IRM grows inside the roots in a very constant environment while the ERM
grows in the soil under highly variable environmental conditions (Smith
\& Read, 2008). Two broad anatomical groups of IRM can be recognized in
mycorrhizal roots , the \emph{Arum} -type, dominated by arbuscules, and
the \emph{Paris-} type, dominated by coils, although evidence suggests a
continuum between these types, depending on the host plant and the
fungus (Dickson, 2004). Presence of 'H' branches in the IRM is more
common in Glomeraceae taxa compared to Acaulosporaceae, while looping
hyphae and hyphae with small bumped projections are prevalent in species
forming gigasporoid spores (see (Dodd \emph{et al.} , 2000)for a
review). Arbuscules are highly branched structures with a turnover rate
ranging from 7 to 16 days (Alexander \emph{et al.} , 1989) or longer in
woody plants (Brundrett \& Kendrick, 1990), and they serve as the
primary site of nutrient exchange between the fungus and the host. The
main differences observed in arbuscule architecture relate to branching
patterns. In Gigasporaceae, the trunk is wide and branching is abrupt,
whereas the trunk is narrow and branching is gradual in Acaulosporaceae
and Glomeraceae. Vesicles are thick-walled, globose to lobed structures
that store lipids and contain many nuclei (Smith \& Read, 2008). They
are not formed by members of Gigasporaceae, and there is some evidence
the same is true for basal families such as Ambisporaceae,
Archaeosporaceae, and Paraglomeraceae.

\textbf{Extraradical mycelium (ERM)}

The extraradical mycelium (ERM) is composed of two types of hyphae:
unbranched runner hyphae, which run parallel to the root length to
initiate secondary colonization, and highly branched absorptive hyphae
responsible for nutrient uptake from the soil and subsequent
translocation to the host (Friese \& Allen, 1991). (Bago \emph{et al.} ,
1998a) and Dodd et al. (2000) observed the formation of 'branched
absorbing structures' - small groups of dichotomous hyphae - within the
ERM in species of Glomeraceae. The ERM is also accountable for the
formation of spores and auxiliary cells in the soil. Phenotypic
variables associated with the ERM, such as hyphal length and density,
interconnectedness, and hyphal diameter, have been studied in some AM
fungal species (Dodd \emph{et al.} , 2000; Avio \emph{et al.} , 2006).
The extraradical mycelium can form Common Mycorrhizal Networks (CMNs),
where a single AM fungus associates with multiple plant hosts,
interconnecting them within a shared mycelial network. These networks
may serve as biological bridges, facilitating resource exchange and,
possibly, communication among different plant hosts (Barto \emph{et al.}
, 2011; Babikova \emph{et al.} , 2013). However, more research is needed
to determine the role of CMNs in natural environments and to what extent
the capacity to form CMNs can be considered a fungal trait
(Karst\emph{et al.} , 2023).

\textbf{Genetic traits}

In this section we define genetic measurements that have been proven or
have the potential to reflect differences in life history strategies as
``genetic traits'' and we discuss how they can fit in a trait-based
framework. Among these traits are the genetic organization of AM fungal
strains, the spore nuclear content, the genome size, and the GC content
of the genome. Recent findings demonstrated that AM fungal strains
belonging to one species carry thousands of nuclei in their coenocytic
mycelia that either belong to one (i.e., homokaryotic strains) or two
nuclear genotypes (i.e., dikaryotic strains (Ropars \emph{et al.} ,
2016) with each of these genotypes having unique structure, genetic
content and epigenetics (Sperschneider \emph{et al.} , 2023).
Interestingly, the relative abundance of the coexisting genotypes in the
dikaryotic strains appears to be deterministic and their regulation to
be responsive to biotic (e.g., plant host identity) (Kokkoris \emph{et
al.} , 2021) and abiotic factors (e.g., pH, temperature, nutrient
content) (Cornell\emph{et al.} , 2022). Carrying two genomes instead of
one may reflect differences in life histories strategies if the same is
shown in multiple species (Serghi \emph{et al.} , 2021). Particularly,
the homokaryotic strains have higher germination rates and faster
germination compared to the low germination rate of the dikaryotic
strains. In contrast, dikaryotic strains grow faster and produce larger
and more interconnected ERM compared to the homokaryotic ones. The
nuclear organization can affect the mycorrhizal response of their plant
hosts. Specifically, and in contrast to expectations that two genomes
might result in more mutualistic interactions, dikaryotic strains were
inferior mutualists compared to the homokaryons when interacting with
multiple potato cultivars (a highly mycorrhizal dependent crop) in
greenhouse conditions (Terry \emph{et al.} , 2023). While we recognize
that nuclear organization may be an important function trait, until mono
vs dikaryons are found in more AM fungal species it might be premature
to suggest this trait should be included in a program for
standardization of trait measurement across all AM fungal taxa.

The nuclear content of the spores also seems to be associated with
particular life history traits although further experimental evidence is
needed. The range of nuclei present in spores correlates with spore
size, ranging from 35000 nuclei for spores of \emph{Gigaspora
decipiens}which have an average diameter of 400um, to 130 nuclei for the
spores of\emph{Glomus cerebriforme} with an average diameter of 80um
(Kokkoris\emph{et al.} , 2020). These huge differences in nuclear
content could be associated to spore viability and germination, and
overall colonization ability after dispersal. For example, multiple
re-germination events have been observed for Gigasporaceae spores when
no host is encountered initially (Sward, 1981), a trait that does not
appear in \emph{Glomus}sp. spores which usually demonstrate a more
ruderal behavior. It has been hypothesized that the numerous nuclei,
could serve as resource reserve via nucleophagy when facing starvation,
a phenomenon previously observed in fungi (Shoji \emph{et al.} , 2010;
Kokkoris \emph{et al.} , 2020). Despite the number of genotypes and the
number of nuclei present in the AM fungal networks and spores, the
overall genome size might influence the reproductive rate, the
environmental adaptability and in the overall resource economy of a
species/strain. Although not very common for fungi, linkage of genome
size to life history affiliations is not a novel concept. (Grime \&
Mowforth, 1982), linked plant genome size to climate growth conditions.
(Veselý \emph{et al.} , 2012) linked larger plant genome sizes to early
flowering events and preference for humid conditions, and (Bhadra
\emph{et al.} , 2023) linked genome size to multiple functional traits
related to plant morphology, physiology, performance and survival. Our
knowledge on the variation of genome size in AM fungi is limited due to
the low number of sequenced genomes. Regardless, we know that the
variation is extreme, with larger species (Gigasporaceae) having genomes
that reach 740 Mb and smaller species (e.g., \emph{Rhizophagus clarus} )
116Mb (Kokkoris \emph{et al.} , 2020). It is important to keep expanding
our datasets with such information to identify links between genome size
and the morphological, physiological, and phenological traits of AM
fungi. Finally, a particular genetic trait, the guanine plus cytosine
(G+C) content of genomic DNA which have the potential to reflect
ecological niche or pathogenicity in fungi (Yoder \& Turgeon, 2001).
Once again with limited data due only few complete genomes available
substantial variation in G+C content exists in AM fungi (range from 25
to 36 (mol\%) (Malar C \emph{et al.} , 2022). These differences could
potentially reflect differences observed in mycorrhizal response and
host preference.

\textbf{The way forward on AM fungal traits research}

Considering the above discussion and to overcome challenges related to
measurement of AM fungal traits, and to obtain a more accurate
understanding of AM fungal-plant interactions, we suggest the following
points for future research.

1) \emph{Create and maintain a centralized database of AM fungal traits}
. This is a keystone task to integrate data and analyses to use
appropriate AM fungal traits to predict plant responses and ecosystem
processes. Databases are available on AM fungal traits but they are
limited to mycorrhizal type and intensity of root colonization primarily
in the early stages of plant development (Soudzilovskaia \emph{et al.} ,
2020). We propose the development of a generic structure for the AM
fungal database for individual taxa stemming from the traits described
in Table 1 and that is consistent with the principles of the Observation
and Measurement Ontogeny (Madin \emph{et al.} , 2007). The central
tabulation in this database is the taxonomy at the species and strain
level, and accession codes if available. Ancillary data are related to
a) the site of origin such as latitude and longitude, along with the
date and the observer b) its source either from field observations, a
culture collection or literature data, and c) metadata and information
about experimental treatments used to measure traits including, but not
limited to, the levels of replication and variation associated with each
estimate. A trait and measurement component integrates all information
related to a specific trait (spore, mycelium, arbuscules, vesicles) and
its measurement values and units. A trait database could facilitate the
synthesis of research findings and improve our understanding of the
functional diversity and ecological roles of AMF taxa.

2) \emph{Expand the scope of research to include a broader range of AM
fungi, with a particular focus on uncultured and underrepresented taxa}
. Traits have been studied focusing on Glomeraceae, Acaulosporaceae, and
Gigasporaceae, all common components of AM fungal communities and
harboring 76\% of the total number of species in the Glomeromycota.
Other families such as Diversisporaceae, Claroideoglomeraceae, and basal
families such as Paraglomeraceae and Archaeosporaceae are rarely
included in experiments and there is very little information on their
traits, despite being common components of AM fungal communities.
Glomeromycota comprise 13 families and 337 species but likely contain
many more (Öpik \emph{et al.} , 2010). However, we estimate that
only\emph{ca.} 88 species are represented in culture collections
worldwide. To ensure a more comprehensive understanding of AM fungal
ecological roles and interactions, it is essential to record traits from
taxa in these poorly studied families and attempt to increase taxonomic
diversity in culture collections.

3) \emph{Determine traits at fine levels of taxonomic resolution} .
Results from distinct experimental approaches indicate that AM fungal
traits related with internal and external mycelium and sporulation time
exhibit some conservation at the family level, although variation within
these clades was also observed (Hart \& Reader, 2002a; Maherali \&
Klironomos, 2012). Seemingly inconsistent with this finding, high
intraspecific variability in root colonization, external mycelium length
and plant responses has been demonstrated for several species
(Mensah\emph{et al.} , 2015; Schoen \emph{et al.} , 2021; Stahlhut
\emph{et al.} , 2023). As argued above, there is a need to conduct
additional comparative studies using different species within the same
genus to investigate trait conservatism.

4) \emph{Measure and report AM fungal traits using standardized
experimental approaches} . AM fungal traits reported in the literature
have been measured using different experimental approaches, which makes
comparisons difficult. We propose herein (Table 2) a set of minimum
parameters that can be used when studying AM fungal traits that can
minimize biases and account for the complex interplay between fungal and
plant traits. We expect that by adopting these standard approaches
database data supplied by different laboratories are more directly
comparable. An obvious research imperative will be to validate the
reproducibility of AM fungal trait measurements by different
teams/infrastructures but using the same starting inoculum material.
This reinforces the value of culture collections.

5) \emph{Determine the variability (plasticity) of AM fungal trait
expression} . A gap in our understanding of AM fungal traits, partially
responsible for our inability to predict symbiotic function under field
conditions, lies in our knowledge of how consistent traits are under
varying environmental conditions. Experiments designed to test how
specific environmental factors impact AM fungal traits are needed.
Additionally, the scope of trait variations across different traits and
taxa could be studied to determine if some traits are more conserved
than others and/or vary more significantly in one taxon compared to
others.

6) \emph{Embrace AM fungal community diversity} : A basic premise of
ecophysiology is that environmental filters will select for specific
traits/adaptations (Lambers \emph{et al.} , 2008). Given that some
traits can be measured at the community level (e.g., hyphal nutrient
stoichiometry (Zhang \emph{et al.} , 2023b)), it can be envisaged to
conduct experiments (physical disturbance, nutrient additions, drought
etc.) on whole natural AM fungal communities and look at correlations
between environmental filters and traits (Chagnon, 2023). Coupled with
rotating and static cores (Johnson et al. 2001), these experiments could
also assess AM fungal growth and mycorrhizal function. With synthesis
studies identifying major drivers of AM fungal community structure at
global scales (Davison et al., 2021), the next frontier is to move
beyond taxonomy and assess the functional biogeography (e.g.,
(Violle\emph{et al.} , 2014) of AM fungi.

7) \emph{Utilize AM fungal isolates deposited in culture
collections.}Culture collections worldwide uphold a considerable variety
of AM fungal isolates in terms of physiology, genetics, taxonomy, and
geographic origin. These isolates, which are cultivated either in a
mineral substrate or in conjunction with root organ cultures, are well
characterized taxonomically, thereby representing important resources in
comparative studies of traits. These centers are instrumental in
training personnel, and specific workshops can be developed to provide
hands-on experience and theoretical knowledge about measurement of AM
fungal traits.

8) The use of microphotography and machine learning: The integration of
microphotography and machine learning algorithms could not only help
standardize and accelerate trait quantification but also eliminate the
subjectivity of the observer, a common issue entangled with our current
quantification approaches. Successful integration of the two can create
automations that will allow for large dataset acquisition, no longer
limited by space and time (e.g., continuous growth measurements of ERM
and its traits, or continuous progression of root colonization with the
help of fluorescent markers). These approaches can help reveal
behavioral patterns that have so far remained undetected due to
technical limitations.

As mentioned earlier, some ecologically significant functions of the AM
fungal symbiosis, such as the ability to promote host plant growth,
depend not only on the properties of an individual AM fungal isolate but
also on its interaction with the host. While we recognize that this may
limit the scope of some trait measurements, we consider that even though
a hypothesis-driven approach as outlined in this study for effectively
assessing AM fungal traits may not address all questions about the role
and impact of AM fungi in ecology, it represents a significant step
towards that goal.

\textbf{Experimental approaches BOX 1}

Various experimental approaches can be employed to investigate
morphological and physiological AM fungal traits in semi-realistic
conditions including soil or substrate and plant host(s). We identified
five main approaches in the literature. Gazey et al. (1992) utilized
a\textbf{sterile mesh bag} approach to examine the association between
the initiation of sporulation and external hyphae production in
two\emph{Acaulospora} species. This technique involves the use of 25 \selectlanguage{greek}µ\selectlanguage{english}m
mesh bags (2 cm wide, 10 cm across, and 10 cm long) containing 200 g of
uninoculated, steamed soil. The mesh bags can be placed in pots anywhere
along the soil profile. Data are gathered at weekly intervals from 3 to
8 weeks post-plant host germination within the ''sterile mesh bag,''
encompassing external hyphae length, spore numbers. Outside the
''sterile mesh bag'' host biomass and root colonization, among other
variables, can be measured. This approach allows measuring mycorrhizal
traits within a controlled sterile soil environment without the
interference of propagules present in the original inoculum. The sterile
soil should be free from AM fungal propagules and researchers should
have controls with non-inoculated mesh bags. Furthermore, as samples are
collected from a small soil volume inside the mesh bag, correlations
between root colonization, spores, and external hyphae are more likely
to represent realistic relationships among these traits. The temporal
approach (weekly sampling) permits close monitoring of the entire AM
fungal and plant life-cycles. Nonetheless, there are potential
limitations to this approach, such as the requirement for propagules to
infiltrate the mesh bag, which may be limiting for some taxa (e.g.,
(Thonar \emph{et al.} , 2011)). Additionally, if many isolates or
species are being analyzed concurrently, the entire experiment may be
time-consuming and labor-intensive.

A second approach similar to the sterile mesh bag was used by
(Jakobsen\emph{et al.} , 1992a) to study the abundance of AM fungal
hyphae in soil and P uptake into roots. \textbf{Root compartment bags}
consisted of cylindrical (60 mm diameter) bags constructed using 25 \selectlanguage{greek}µ\selectlanguage{english}m
nylon mesh and filled with AM fungal inoculum. These bags are placed in
the center of 1.5 L pots, surrounded by steamed sterilized soil and
pre-germinated seeds are transplanted into each bag. Thus, the host
generated roots are confined within the compartment bag, while AM fungal
hyphae are unrestricted to extend into the surrounding soil. After 25
days, root compartment bags can be transplanted into rectangular PVC
boxes (L x W x H = 300 x 185 x 130 mm) surrounded by 7 kg of steamed dry
soil. To measure hyphal growth, 10 mm diameter soil cores are obtained
on five sampling dates at different distances from the root compartment.
Similar to the sterile mesh bag method, the root compartment approach
enables the measurement of hyphal length in an environment devoid of
pre-existing mycorrhizal inoculum. By transplanting the root compartment
into large rectangular boxes, this approach is very suitable to study
distance and rate of spread of AM fungal external hyphae, allowing a
direct comparison of these traits among fungal taxa. However, the
experimental design requires a substantial quantity of soil for the
setup. A similar approach uses a trap plant on the other end, which
allows measurements of resource movement between hosts in a community
(Mikkelsen \emph{et al.} , 2008).

A third approach, named \textbf{inoculated containers,} was used by
(Hart \& Reader, 2002a) to establish the taxonomic basis for the
observed variation in root colonization strategies among AMF families.
Fungal biomass is initially measured based on ergosterol concentration
to equalize the amount of inoculum at the onset of the experiment.
However, we recommend using a different approach (e.g., fatty acids) as
has been shown that AM fungi do not produce ergosterol (Olsson \emph{et
al.} , 2003; Olsson \& Lekberg, 2022). Containers (4 cm diameter x 20.5
cm deep) are  2/3  filled with soil, then inoculated with soil inoculum
containing spores, hyphae and colonized root fragments and sown with
leek as a surrogate host. Following a 30 d period, the shoots are
harvested, and the soil subjected to experimental treatments, including
different hosts. Containers are harvested six times over a 12 w period
to measure the extent of root and soil colonization. The inoculated
container approach, using small-volume containers, allows the concurrent
study of many isolates over time. The standardization of fungal biomass
among species allows a direct final comparison among taxa. Given that
the abundance of AM fungal external hyphae is assessed in the same
container in which the inoculum was introduced, differentiating between
newly produced hyphae and those present in the original inoculum is not
feasible.

Direct measurements of physiological/chemical traits can require
efficient separation of fungal hyphae from the growth substrate.
(Zhang\emph{et al.} , 2023b) employed \textbf{hyphal in-growth bags} ,
consisting of glass beads and fine silt and clay particles (as a
nutrient source) and surrounded by fine nylon mesh (pore size = 38 um)
based on the original design by (Neumann \& George, 2005), to harvest
enough mycelium so that carbon and nitrogen concentrations could be
assessed following combustion, and phosphorus concentration following
wet digestion, after eight weeks of incubation. In-growth bags were 2 cm
wide and 10 cm long and filled with 40 g of the soil-glass bead mixture
and buried lengthwise in the top 10 cm of each pot. The mesh size was
sufficient to prevent in-growth of \emph{Medicago sativa} roots but not
root hairs of \emph{Festuca arundinacea} , which requires a mesh size of
10 um to exclude (Zhang \emph{et al.} , 2021). Soil particles in the bag
can still stick to hyphae and need to be carefully removed before
analysis. This method, while similar to the sterile mesh bag, is more
suitable to study physiology.

Another approach to study traits associated with common mycorrhizal
networks (CMNs) was proposed by (Johnson \emph{et al.} , 2001)
using\textbf{rotative cores} . In this method, a slot (2 cm wide and 5
cm long) is made on the side of a conical container (270 ml) and covered
with a 40 \selectlanguage{greek}µ\selectlanguage{english}m nylon mesh or hydrophobic membrane. Containers are fitted
into polystyrene foam and assembled in pots or microcosms, depending on
the study's objective. The containers are then filled with soil and sand
mixtures, inoculated with AM fungi and seeded with a host plant to
establish the CMNs for 2-3 months. Following this period, CMN treatments
can be established by either maintaining the containers' positions
throughout the experiment (undisturbed CMNs) or rotating them (one full
rotation) periodically to physically sever the AM fungal hyphal network.
This method is highly suitable for understanding the role of external
mycelium and the impact of AM fungal mycelium network disruption on
various soil (e.g., bacterial community structure, soil aggregation) and
plant (e.g., biomass production, nutrient allocation) parameters
(Babikova \emph{et al.} , 2013). Although the preparation of modified
containers is somewhat laborious, this approach enables the
investigation of the role of CMNs in plant interactions across multiple
species simultaneously.

The methods described above are appropriate for assessing fungal traits
and their interrelationships; however, these are not the sole approaches
available. The study of genetic traits typically requires the use of
isolation techniques into \emph{in vitro} culture see (Declerck \emph{et
al.} , 2010). For example, \emph{in vitro} root-organ cultures methods
have enabled major breakthroughs in the understanding of genetic and
physiological traits such as nutrient exchange ratios
(Cranenbrouck\emph{et al.} , 2005; Kiers \emph{et al.} , 2011) and
patterns of hyphal anastomosis between isolates in the same species
(Giovannetti \emph{et al.} , 1999). In addition, \emph{in vitro} systems
may be instrumental in investigating trait interactions between AM fungi
and other microorganisms (Faghihinia \emph{et al.} , 2023). Considering
the vast amount of literature on mycorrhizal research and the diverse
factors involved in experiments (e.g., plant hosts, soil type,
fertilization regime, environmental conditions), a comprehensive
understanding of traits for different AM fungal taxa has not been
achieved. Given that AM fungi are influenced by soil parameters, plant
hosts, and environmental conditions, we propose a set of standard items
that should be considered in experiments aiming to measure AM fungal
traits. We acknowledge that certain experimental items could be
standardized, while others that are more challenging to standardize
(e.g., soil type, lighting conditions) could be collected as metadata.
By adopting this approach, experiments could be conducted in different
laboratories using the same AM fungal taxa while modulating different
environmental conditions (\emph{e.g.,}soil disturbance, salinity,
drought, CO\textsubscript{2} concentration, temperature, light
intensity) to examine the degree of conservation of AM fungal traits
across systems and the validity of predictions based on taxonomic
identity.

\textbf{Toward a standardized approach to measure AM fungal traits BOX2}

Standardizing the quantification of mycorrhizal fungal traits can be
useful to optimize data quality, reduce bias, facilitate comparability,
and reproducibility among studies. This methodological uniformity can
enhance the robustness of meta-analyses and promote collaboration among
researchers, thereby advancing our ecological understanding of AM fungi.
What follows is a proposal, based on the authors' expert opinions, for
the standardization of trait-measurement experiments. Researchers may
have very good reasons to not comply with some or all of these
recommendations in their work, and we want to make clear that this would
not mean that the data lack value in the context of understanding AM
fungal life histories. As long as deviations from proposed methods are
documented along with the data, they would certainly still hold value.

a) Pot Size and Type: Measuring AM fungal traits might result in
experiments with many experimental units. Therefore, pot size and
substrate (see below) are two items that should be considered to make
the experiment feasible. We propose the use of pots with volumes greater
than 2 kg , when mesh bags are used. If inoculated containers or
rotative cores are chosen, then containers (4 cm diameter x 20.5 cm
deep) with conical open bottoms are recommended (Weremijewicz \& Janos,
2019).

b) Soil texture: Production of AM fungal external mycelium and
sporulation are certainly affected by soil texture, as it determines the
number of pores and available free space for fungi to grow. Inert media
like sand: expanded clay could be used to standardize the substrate.
These media have the advantage of not containing AM fungal propagules.
However, they hardly represent the common habitat of AM fungi.
Therefore, extrapolating AM fungal trait measurements obtained using
these media to field soil conditions might be challenging. We propose
using a sterilized loam soil in standard experiments to measure AM
fungal traits. If a loam soil is not available, adding quartzite sand or
coarse river sand to the soil is suggested to bring the texture closer
to a loam soil.

c) Soil sterilization: Autoclave 121\textsuperscript{o}C (for as long as
needed; adjust by placing a rod with autoclave tape to the center)
repeated twice with a 24h interval. However, if budget and access to
infrastructures allow, gamma radiation is a useful alternative that
limits chemical alteration of organic matter and downstream consequences
on dissolved organic carbon and aggregate stability (Berns \emph{et al.}
, 2008).

d) Nutrient solution: Plant nutrition is an important aspect to be
considered as it impacts the establishment of the mycorrhizal symbiosis.
We suggest the use of a modified Hoagland's solution for monocots to
provide the minimum amount of macro and micronutrients for the host
plant (\emph{SupplementaryMaterial}).

e) Typically a microbial wash to introduce and standardize
non-mycorrhizal soil organisms is prepared by mixing all the tested AM
fungal inocula with water and passing the resulting slurry through a 20
\selectlanguage{greek}µ\selectlanguage{english}m sieve to be added to the test pots (Ames \emph{et al.} , 1987). This
procedure is arguably the hardest to standardize. We suggest including a
control treatment, without a microbial wash to ascertain the role of
microorganisms on AM fungal traits.

f) Host plant: Mycorrhizal host plants vary widely in their growth
habits (e.g., grasses, trees, forbs), growth rates, and root
architecture, which affect the amount of root colonization and spread of
hyphae in the soil. Host preference is also another factor to be
considered, as it impacts AMF sporulation (Bever \emph{et al.} , 1996).
We suggest the use of \selectlanguage{ngerman}\emph{Sorghum × drummondii} (Sudan grass) as a
standard host because a) it has been widely used to grow and maintain a
vast array of AMF germplasm in culture collections (Morton \emph{et al.}
, 1993), b) it has a fasciculated root system that provides space for
root colonization, and c) it is mycorrhizal dependent.

In addition to these recommendations, metadata should include
temperature, soil/substrate type, pH, soil moisture content, soil
fertility, light intensity and experiment duration.

\par\null

Table 1: Key traits, their hypothesized function, and methods for trait
measurement\selectlanguage{english}
\begin{longtable}[]{@{}llllllll@{}}
\toprule
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Fungal and fungal mycorrhizal traits}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Hypothesized symbiotic effects}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
AM fungi\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Plant\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Soil*

~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Qualitative/quantitative (unit)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
References - methods on how to measure the traits\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
References**\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Spores}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Number\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
-fitness/competitive ability

-dispersal

-carbon storage\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

~

x

x

~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~

~

~

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Quantitative: number of spores/g soil

Spores/meter of mycelia\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Spores extracted by wet-sieving ((Gerdemann \& Nicolson, 1963)) and
sucrose gradient centrifugation (Ba, 1982) and counted under dissecting
microscope

~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Bever \emph{et al.}, 1996), (Chaudhary \emph{et al.}, 2020){}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
size - diameter\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
-fitness/competitive ability

-dispersal

-energy to support hyphal growth in absence of host

-carbon storage

-resistance to abiotic and biotic stress\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

~

x

x

~

~

x

x

~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~

~

~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~

~

~

~

~

~

x

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Quantitative: size measured in \selectlanguage{greek}µ\selectlanguage{english}m\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Spore diameter measured intact in water using a dissecting or optical
microscope

((Morton, 1995, 1996))\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Chaudhary \emph{et al.}, 2020; Deveautour \emph{et al.}, 2020){}

~

~\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
germination rate

\% of total\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
-fitness/competitive ability

-carbon storage\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~

~

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Quantitative: \% of germination\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Douds \& Schenck, 1991), Spores over filter paper in a soil-filled
Petri plate (Koske, 1981){}

~

~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Tommerup, 1983), (Maia \& Yano-Melo, 2001){}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
germination timing\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
-fitness/competitive ability

-resistance to abiotic and biotic stress

~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

~

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

~

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Quantitative - \% germination per unit of time

Qualitative - stratification needed\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Koske, 1981){}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Tommerup, 1984, 1985), (Koske \emph{et al.}, 1996) (Douds \& Schenck,
1991) (Juge \emph{et al.}, 2002){}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
color\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
- dispersal

- palatability

- UV protection

- germination duration

~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~

~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Qualitative: color based on CMYK color chart

~

Quantitative: RGB color channels extracted from digitized images (JPG,
TIF), calculation of luminance and saturation\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Spores observed under a dissecting microscope and compared with color
chart (Morton, 1996) or imaged and analyzed using computer software
(Deveautour et al. 2020)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Deveautour et al. (2020),(Zanne \emph{et al.}, 2020)\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
ornamentation\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
-dispersal

-resistance to abiotic and biotic stress\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Qualitative: type of ornamentation. Quantitative: size in \selectlanguage{greek}µ\selectlanguage{english}m\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Spores mounted on slides and observed under microscope

Koske and Walker, 1985\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Chaudhary et al. (2020)

~\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
wall thickness\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
- dispersal

-palatability

- carbon storage

- resistance to abiotic and biotic stress\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

x

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~

~

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Quantitative: thickness in \selectlanguage{greek}µ\selectlanguage{english}m\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Spores mounted on slides and thickness measured under microscope Morton
(1995, 1996)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Pawlowska \emph{et al.}, 1999){}

(Moore \emph{et al.}, 1985) {}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
wall number

~

~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
-dispersal

-resistance to abiotic and biotic stress\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Quantitative: number of walls\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Spores mounted on slides and observed under microscope Morton (1995,
1996)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Walker, 1983){}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
sporocarps\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
-dispersal\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Size and existence of fungal peridium on the sporocarp surface.\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Sporocarps are measured under a microscope for size (Redecker \emph{et
al.}, 2007) and existence of peridium (Sch\selectlanguage{ngerman}üßler \emph{et al.},
2011){}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Mangan \& Adler, 2002){}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
wall chemical receptors\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
- perception of host/soil environmental cues affecting germination\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Knowledge gap (develop a method to identify specific receptors)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Knowledge gap (are there specific receptors on the spore wall that
trigger germination?)\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Spore nuclear content\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
-spore viability and germination

- colonization ability after dispersal\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

~

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

~

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Number of nuclei per spore\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Confocal microscopy, Flow cytometry\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Kokkoris \emph{et al.}, 2020, 2021){}

(Bianciotto \emph{et al.}, 1995; Marleau \emph{et al.}, 2011){}

~\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Elemental composition\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
-Energy support for hyphal growth\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Elemental composition\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Proton-induced X-ray emission\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Hammer \emph{et al.}, 2011){}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Extraradical Hyphae}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
length\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
-- nutrient/water acquisition

-carbon storage

-soil aggregation

-plant productivity

-resistance to abiotic and biotic stress\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

~

x

x

~

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

~

x

x

x

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

~

x

x

x

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Quantitative: hypha length in m/g dry soil\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Miller \emph{et al.}, 1995){}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Wilson \emph{et al.}, 2009){}

(Johnson \emph{et al.}, 2015){}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
architecture (branching rate, anastomoses rate,

absorptive/runner hypha)

~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
-- nutrient/water acquisition

-carbon storage

-soil aggregation

-plant productivity

-resistance to abiotic and biotic stress\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

~

x

x

x

x

~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

~

x

x

x

x

~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

~

x

x

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Qualitative

~

~

~

Quantitative: Using image analysis in \emph{in vitro} systems\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Friese \& Allen, 1991) or (Bago \emph{et al.}, 1998b) in monoxenic
conditions

~

(Hammer \emph{et al.}, 2023){}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Knowledge gap (e.g., is hyphal architecture akin to root architecture
for nutrient acquisition? how does hyphal architecture influence soil
aggregate stability?)\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
inter-host connection\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
- transfer of nutrient/water/signals among hosts

~-resistance to abiotic and biotic stress

~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

~

~

x

~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

~

~

x

~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Quantitative: amount of nutrient/signal transferred

Number of hosts connected by the same fungus\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Weremijewicz \& Janos, 2019){}

(Frey \& Sch\selectlanguage{ngerman}üepp, 1993){}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Knowledge gap (e.g., are there fungi that interconnect more hosts than
others? can common mycorrhizal networks provide additional pathogen
protection?)\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
hyphal diameter\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
--carbon storage

- resistance to abiotic and biotic stress

- palatability\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

x

~

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~

x

~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

~

~

~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Quantitative: in \selectlanguage{greek}µ\selectlanguage{english}m\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Friese \& Allen, 1991){}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Klironomos \& Kendrick, 1996){}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
growth rate\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
-- nutrient/water acquisition

- carbon storage

- resistance to abiotic and biotic stress\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

~

x

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

~

x

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

~

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Quantitative: hyphal growth in mm/day\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Sch\selectlanguage{ngerman}ütz \emph{et al.}, 2022){}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Jakobsen \emph{et al.}, 1992b){}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
hyphal lifespan/turnover\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
- carbon storage

- nutrient/water acquisition\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Qualitative\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Pepe \emph{et al.}, 2018){}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Pepe \emph{et al.}, 2018){}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Genetic organization/hyphal fusion (homokaryon/dikaryon)

(note: also applies to spores)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
-fitness

-- nutrient/water acquisition

-carbon storage

-soil aggregation

-plant productivity

-resistance to abiotic and biotic stress

- hyphal network interconnectedness\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

x

~

x

x

x

x

~

~

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~

x

~

x

x

x

x

~

~

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~

~

~

x

x

~

~

~

~

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Quantitative ddPCR: number of nuclei?\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Cornell \emph{et al.}, 2022){}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Cornell \emph{et al.}, 2022){}

(Serghi \emph{et al.}, 2021){}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Exudation rate/leakiness\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
-carbon storage

-influence soil pH and fertility

-soil aggregation

-resistance to abiotic and biotic stress\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

x

~

x

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

x

~

x

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

x

~

x

~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Quantitative: measure release of a molecule in \selectlanguage{greek}µ\selectlanguage{english}M\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Tawaraya \emph{et al.}, 2006){}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Tawaraya \emph{et al.}, 2006){}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
absorptive capacity\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
-- nutrient/water acquisition\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Quantitative: mol m\textsuperscript{-1} s\textsuperscript{-1} or \% of
nutrient taken up\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Frey \& Sch\selectlanguage{ngerman}üepp, 1993), (Jakobsen \emph{et al.}, 1992a){}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Frey \& Sch\selectlanguage{ngerman}üepp, 1993){}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
color\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
-resistance to abiotic and biotic stress\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Qualitative: color described by CMYK model\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(de la Providencia \emph{et al.}, 2005) using transformed roots and
(Koske, 1981) using spores over filter paper on soil-filled Petri
plate\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Knowledge gap (are darker hyphae, more melanized, more resistant to
fungivores?)\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
wall/membrane chemical composition{}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
-resistance to abiotic and biotic stress

-nutrient/water acquisition

-fungal recognition (anastomosis)

-palatability

~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

~

x

~

x

~

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~

~

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

~

x

~

~

~

~

~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Quantitative: in \selectlanguage{greek}µ\selectlanguage{english}g\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Bethlenfalvay \emph{et al.}, 1981) for chitin.

(Frey \emph{et al.}, 1992, 1994) for chitin and ergosterol (Butler \&
Lachance, 1986) for melanin; (Harrison \& Vanbuuren, 1995) for P
transporters\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Deveautour \emph{et al.}, 2020){}

~\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
pattern of anastomosis\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
- fungal recognition

- fitness\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Quantitative: number of anastomosis per hyphal length (cm) or percentage
of anastomosis (\%)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(de la Providencia \emph{et al.}, 2005){}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(de la Providencia \emph{et al.}, 2005){}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Intraradical Hyphae}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
-hyphal thickness\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
-Resource flux/exchange

-resistance to abiotic and biotic stress\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

~

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

~

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Quantitative: in \selectlanguage{greek}µ\selectlanguage{english}m\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Abbott, 1982)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Knowledge gap (e.g., are thicker hyphae more resistant to pathogens? Is
there a tradeoff between nutrient transfer and biotic resistance in
terms of hyphal thickness?)\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
-pattern of

colonization (localized / widespread)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
-Resource flux/exchange

-resistance to abiotic and biotic stress\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

~

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

~

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Qualitative\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Dickson, 2004){}

(McGONIGLE \emph{et al.}, 1990) {}

(Abbott, 1982) {}

~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Knowledge gap (e.g., is resource exchange more/less efficient when
colonization is localized or widespread?)\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
-rate of root colonization\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
-Resource flux/exchange

-resistance to abiotic and biotic stress\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Quantitative: \% of root colonization over time\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Dickson, 2004){}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Campo \emph{et al.}, 2020)\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Arbuscules}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
- architecture (Paris/Arum type)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
- resource flux/exchange

-resistance to abiotic and biotic stress

~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

~

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

~

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Qualitative\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Dickson, 2004){}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(van Aarle \emph{et al.}, 2005){}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
- turnover rate\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
-resource flux/ exchange\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Quantitative: number of days\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Alexander \emph{et al.}, 1989)), (Toth \& Miller, 1984)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Knowledge gap (e.g., are some arbuscules more short lived than others?
How does arbuscule turnover affect resource exchange?)\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
-number\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
-resource flux/ exchange\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Quantitative: number of arbuscules\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Quantification of Arbuscules Using Morphometric Cytology - (Toth, 1992)

~

Magnified~ intersections~ method - (McGONIGLE \emph{et al.}, 1990)

~

Image analysis - (Smith \& Dickson, 1991)

~

Direct count - (Menge \emph{et al.}, 1978){}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Koch \emph{et al.}, 2017){}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Vesicles}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
-size and form (globose/lobbed)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
- carbon storage

- resistance to abiotic and biotic stress\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Quantitative: in \selectlanguage{greek}µ\selectlanguage{english}m (for size). Qualitative (for form)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Abbott, 1982)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Knowledge gap (e.g., are fungi with larger vesicles more resistant to
stress?)\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
-number\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
- carbon storage\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Quantitative: number per root length\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Abbott, 1982), (Menge \emph{et al.}, 1978){}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Kobae \emph{et al.}, 2016)\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
-chemical composition (C/lipid storage)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
- carbon storage\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Quantitative: \% total lipids/ fatty acids~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Jabaji-Hare \emph{et al.}, 1984){}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Jabaji-Hare \emph{et al.}, 1984){}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Turnover rate

~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
- carbon storage\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Quantitative: number of days\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Knowledge gap (adapt the method used for arbuscules)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Knowledge gap (e.g., are some vesicles more short lived than others? How
does vesicle turnover affect C storage?)\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Other genetic traits}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Genome size\textbf{}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
-reproductive rate

- survival\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Whole genome sequencing

Flow cytometry\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Sperschneider et al., 2023

~

~(Hosny \emph{et al.}, 1998){}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
GC content of the genome\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
- mycorrhizal host response

-host preference\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x

~

x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
x\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
~\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Whole genome sequencing

Knowledge gap (missing genomes across phylogeny)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
(Malar C \emph{et al.}, 2022){}\strut
\end{minipage}\tabularnewline
\bottomrule
\end{longtable}

*abiotic and/or biotic

**Examples/papers showing the relationship between measuring the trait
its function. References might not necessarily represent best practice.

~



\par\null\par\null

\textbf{References}

\textbf{van Aarle IM} , \textbf{Cavagnaro TR} , \textbf{Smith SE}
,\textbf{Smith FA} , \textbf{Dickson S} . \textbf{2005} . Metabolic
activity of Glomus intraradices in Arum- and Paris-type arbuscular
mycorrhizal colonization. \emph{The New phytologist} \textbf{166} :
611--618.

\textbf{Abbott LK} . \textbf{1982} . Comparative Anatomy of
Vesicular-Arbuscular Mycorrhizas Formed on Subterranean
Clover.\emph{Australian journal of botany} \textbf{30} : 485--499.

\textbf{Aguilar-Trigueros CA} , \textbf{Hempel S} , \textbf{Powell JR}
,\textbf{Cornwell WK} , \textbf{Rillig MC} . \textbf{2019} . Bridging
reproductive and microbial ecology: a case study in arbuscular
mycorrhizal fungi. \emph{The ISME journal} \textbf{13} : 873--884.

\textbf{Aguilar-Trigueros CA} , \textbf{Krah F-S} , \textbf{Cornwell WK}
,\textbf{Zanne AE} , \textbf{Abrego N} , \textbf{Anderson IC}
,\textbf{Andrew CJ} , \textbf{Baldrian P} , \selectlanguage{ngerman}\textbf{Bässler C}
,\textbf{Bissett A} , \emph{et al.} \textbf{2023} . Symbiotic status
alters fungal eco-evolutionary offspring trajectories. \emph{Ecology
letters}\textbf{26} : 1523--1534.

\textbf{Alexander T} , \textbf{Toth R} , \textbf{Meier R} ,
\textbf{Weber HC} . \textbf{1989} . Dynamics of arbuscule development
and degeneration in onion, bean, and tomato with reference to
vesicular--arbuscular mycorrhizae in grasses. \emph{Canadian journal of
botany. Journal canadien de botanique} \textbf{67} : 2505--2513.

\textbf{Ames RN} , \textbf{Mihara KL} , \textbf{Bethlenfalvay GJ}
.\textbf{1987} . The Establishment of Microorganisms in
Vesicular-Arbuscular Mycorrhizal and Control Treatments. \emph{Biology
and fertility of soils} \textbf{3} : 217--223.

\textbf{Antunes PM} , \textbf{Koyama A} . \textbf{2016} . Mycorrhizas as
Nutrient and Energy Pumps of Soil Food Webs: Multitrophic Interactions
and Feedbacks. In: Mycorrhizal Mediation of Soil. Elsevier, 149--173.

\textbf{Avio L} , \textbf{Pellegrino E} , \textbf{Bonari E}
,\textbf{Giovannetti M} . \textbf{2006} . Functional diversity of
arbuscular mycorrhizal fungal isolates in relation to extraradical
mycelial networks. \emph{The New phytologist} \textbf{172} : 347--357.

\textbf{Ba D} . \textbf{1982} . Method for the recovery and quantitative
estimation of propagules from soil. \emph{Methods and principles of
mycorrhizal research} : 29--36.

\textbf{Babikova Z} , \textbf{Gilbert L} , \textbf{Bruce TJA}
,\textbf{Birkett M} , \textbf{Caulfield JC} , \textbf{Woodcock C}
,\textbf{Pickett JA} , \textbf{Johnson D} . \textbf{2013} . Underground
signals carried through common mycelial networks warn neighbouring
plants of aphid attack. \emph{Ecology letters} \textbf{16} : 835--843.

\textbf{Bago B} , \textbf{Azcón-Aguilar C} , \textbf{Goulet A}
,\textbf{Piché Y} . \textbf{1998a} . Branched absorbing structures
(BAS): a feature of the extraradical mycelium of symbiotic arbuscular
mycorrhizal fungi. \emph{The New phytologist} \textbf{139} : 375--388.

\textbf{Bago B} , \textbf{Azcón-Aguilar C} , \textbf{Piché Y}
.\textbf{1998b} . Architecture and developmental dynamics of the
external mycelium of the arbuscular mycorrhizal fungus Glomus
intraradices grown under monoxenic conditions. \emph{Mycologia}
\textbf{90} : 52--62.

\textbf{Barto EK} , \textbf{Hilker M} , \textbf{Müller F} ,
\textbf{Mohney BK} , \textbf{Weidenhamer JD} , \textbf{Rillig MC} .
\textbf{2011} . The fungal fast lane: common mycorrhizal networks extend
bioactive zones of allelochemicals in soils. \emph{PloS one} \textbf{6}
: e27195.

\textbf{Behm JE} , \textbf{Kiers ET} . \textbf{2014} . A phenotypic
plasticity framework for assessing intraspecific variation in arbuscular
mycorrhizal fungal traits. \emph{The Journal of ecology} \textbf{102} :
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