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RESEARCH IN CONTEXT
Evidence for phylogenetic correlation of plant–AMF assemblages?
A. Montesinos-Navarro
1,2,
*, J. G. Segarra-Moragues
1
, A. Valiente-Banuet
2,3
and M. Verdu´
1
1
Centro de Investigaciones sobre Desertificacio´ n (CIDE, CSIC-UV-GV), Carretera de Moncada-Na´ quera Km 4.5 46113
Moncada, Valencia Spain,
2
Departamento de Ecologı´a de la Biodiversidad, Instituto de Ecologı´a, Universidad Nacional
Auto´ noma de Me´xico, AP 70-275, CP 04510, Me´xico, DF, Me´xico and
3
Centro de Ciencias de la Complejidad, Ciudad
Universitaria, Universidad Nacional Auto´ noma de Me´xico, 04510, DF, Me´xico
* For correspondence. E-mail ali.montesinos@gmail.com
Received: 26 February 2014 Returned for revision: 10 September 2014 Accepted: 9 October 2014
Background and Aims Specificity in biotic interactions is mediated’ by functional traits inducing shifts in the
community species composition. Functional traits are often evolutionarily conserved, resulting in closely related
species tending to interact with similar species. This tendency may initially shape the phylogenetic composition of
coexisting guilds, but other intraguild ecological processes may either blur or promote the mirroring of the phyloge-
netic compositions between guilds. The roles of intra- and interguild interactions in shaping the phylogenetic
community composition are largely unknown, beyond the mere selectivity in the interguild interactions. Plant facili-
tation is a phylogenetically structured species-specific process involving interactions not only between the same
guild of plants, but also between plants and other guilds such as arbuscular mycorrhizal fungi (AMF). In this study
it is hypothesized that reciprocal plant–AMF interactions will leave an interdependent phylogenetic signal in the
community composition of both plants and AMF.
Methods A correlation was used to test for a relationship between the phylogenetic composition of plant and
AMF assemblages in a patchy xeric shrubland environment shaped by plant facilitation. In addition, a null model
was used to test whether this correlation can be solely explained by selectivity in plant–AMF interactions.
Key Results A significant correlation was observed between the phylogenetic composition of plant and AMF as-
semblages. Plant phylogenetic composition in a patch was related to the predominance of plant species with high
nursery quality that can influence the community assembly. AMF phylogenetic composition was related to the
AMF phylogenetic diversity in each patch.
Conclusions This study shows that shifts in the phylogenetic composition of plants and AMF assemblages do not
occur independently. It is suggested that besides selectivity in plant–AMF interactions, inter-related succession dy-
namics of plants and AMF within patches could be an ecological mechanism driving community assembly. Future
lines of research might explore whether interlinked above- and below-ground dynamics could be occurring across
multiple guilds simultaneously.
Key words: Arbuscular mycorrhizal fungi, AMF, biotic interactions, community assemblages, facilitation, phylo-
genetic composition, plant guilds, vegetation patches, xeric shrubland.
INTRODUCTION
Biotic interactions between guilds are determinant forces as-
sembling communities. It is well known that specificity in these
interactions, when they occur across guilds (e.g. predators and
preys, host and parasites, plants and soil micro-organisms), can
induce reciprocal shifts in each assemblage, ultimately affecting
the community species composition (Janzen, 1970;Connell,
1971;Packer and Clay, 2000;Dickie et al.,2002;Hart et al.,
2003;Wehner et al., 2011).
Biotic interactions are mediated by functional traits, and
most functional traits are evolutionarily conserved. Therefore,
when ecologically relevant functional traits are not available, or
when there is an interest in all traits, beyond single trait rela-
tionships, we can use the shared evolutionary history of species
(phylogenetic relatedness) as a proxy for functional similarity.
As a consequence of trait conservatism, closely related species
tend to interact with similar species (i.e. phylogenetic signal in
the interaction; Go´mez et al.,2010), and this phylogenetic sig-
nal can shape the phylogenetic composition of assemblages of
coexisting guilds (Sargent et al., 2011;Waterman et al.,2011).
As a result of simultaneous processes of coevolution, niche dif-
ferentiation and niche conservatism among closely related taxa,
phylogenetic distance among host organisms may affect the
communities of their mutualistic associates (Cavender-Bares
et al.,2009). However, although most research has focused on
interguild interaction selectivity, other ecological processes,
such as enhanced coexistence through competitive intransitivity
(Laird and Schamp, 2006), or intraguild competition among rel-
atives within a host (Aure´lien et al., 2013), may blur or promote
the mirroring of the phylogenetic compositions between the in-
teracting guilds. Nevertheless, less attention has been paid to
these other local-scale processes that also determine the phylo-
genetic community structure.
The species composition of a community is the product of a
plethora of ecological processes occurring in that particular
community (e.g. Thonar et al.,2014). For a given guild, the
phylogenetic species composition encapsulates the phyloge-
netic neighbourhood for each species within an assemblage
V
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(Pillar and Duarte, 2010). The characterization of the phyloge-
netic composition at small scales allows the comparison of mul-
tiple communities where both intra- and interguild interactions
can shape the final phylogenetic composition of species beyond
the mere selectivity in the interguild interactions. To do so, the
final phylogenetic composition of each guild can be considered
across a set of natural communities (or neighbourhoods) (Pillar
and Duarte, 2010) where multiple ecological processes occur
simultaneously.
Patchy environments provide an appropriate framework to
test for related phylogenetic composition between guild assem-
blages. A well-known patchy environment resulting from
ecological interactions is that generated by plant–plant facilita-
tion interactions. Facilitation is a key process for seedling estab-
lishment, which leads to an aggregated spatial distribution of
species, hereafter vegetation patches (McAuliffe, 1988;Eccles
et al., 1999;Valiente-Banuet and Verdu´, 2007;Castillo et al.,
2010). Plant facilitation is a phylogenetically structured spe-
cies-specific process involving both above- (plant–plant) and
below-ground [plant–arbuscular mycorrhizal fungi (AMF)] in-
teractions (Valiente-Banuet and Verdu´, 2008,2013;
Montesinos-Navarro et al.,2012a). The initial coexistence
pattern of plant and AMF determined by selectivity in the inter-
guild interactions (Montesinos-Navarro et al., 2012b) is later
modified by different ecological processes and indirect interac-
tions emerging in multispecific vegetation patches (Castillo
et al.,2010;Beltra´n et al.,2012;Oviedo et al., 2013).
Accordingly, although a phylogenetic signal in plant–AMF
interaction (Montesinos-Navarro et al., 2012a) suggests a mir-
roring between the plant and AMF phylogenetic composition,
this pattern might be blurred or promoted by other ecological
processes. The combination of multiple ecological processes
occurring in vegetation patches can result in a local species
(plant and AMF) pool (i.e. the taxa in a given patch) which
might be a non-random sample of the regional species pool
(i.e. the sum of taxa observed across patches). A local species
pool might not be solely predicted based on the specificity of
the interactions between guilds (e.g. plant–AMF) as intraguild
ecological processes such as competitive exclusion or competi-
tion alleviation (Aure´lien et al.,2013;Thonar et al.,2014)may
be operating within a patch. These ecological processes are
influenced by species traits. Regarding AMF, functional com-
plementarity among taxa can promote plant coexistence
through pathogen protection and increase of nutrient absorption
rates (Van der Heijden et al., 1998;Hart et al.,2003;Powell
et al., 2009;Wagg et al.,2011). Regarding plant assemblages,
plant coexistence can be mediated by facilitation, where nurse
plants enhance the microhabitat for the establishment of facili-
tated plants. Plants with high nursery quality have traits typical
of early successional species such as drought tolerance,
high growth rates, lower shoot:root ratio and nitrogen
fixation (Valiente-Banuet et al., 2006;Butterfield and
Briggs, 2011;Cadotte and Strauss, 2011). Interestingly, AMF
are often present from the beginning of primary plant succes-
sion and show different relationships with pioneer and late-
successional plant species (Kikvidze et al.,2010). Altogether,
these findings suggest that reciprocal plant–AMF interactions
are involved in the ecological mechanisms determining the
community succession and therefore community phylogenetic
composition.
In this study we use previous data on plant community facili-
tation interactions (Valiente-Banuet and Verdu´, 2007,2008)
and AMF sequences (Montesinos-Navarro et al., 2012b)toas-
sess the relevance of interguild interactions in the assembly of
natural communities by using vegetation patches resulting from
plant–plant facilitation. We specifically test if there is a rela-
tionship between the phylogenetic composition of plant and
AMF which cannot be solely explained by the selectivity in
plant–AMF interactions. We discuss possible assembly mecha-
nisms linked to plant and AMF traits driving successional tra-
jectories of vegetation patches.
MATERIALS AND METHODS
Study system
The study area is a xeric shrubland in the Valley of Zapotitla´n
(1820N, 9728W), central Mexico, in which the AMF regional
species pool (i.e. the total sum of taxa linked to plants across
patches) was obtained (Montesinos-Navarro et al., 2012b). This
system is dominated by the columnar cactus Neobuxbaumia
tetetzo (J.M. Coult.) Backeb., and other species of
Asparagaceae, Fabaceae and Asteraceae. Plants are spatially
aggregated in discrete vegetation patches with areas ranging
from 1 to 5 m
2
. Patches are surrounded by open space, but plant
species can easily disperse from one patch to another. Most
(97 %) of the plant species in this system require facilitation to
establish (Valiente-Banuet and Verdu´, 2007), leading to an ab-
sence of plants in between vegetation patches as a generalized
pattern in the system. Vegetation patches are initiated by a
nurse species that facilitates seedlings of other species; how-
ever, some patches may harbour only adult plants of facilitated
species which are interpreted as late successional stages after
the death of the nurse starting plant. From the data published in
Montesinos-Navarro et al. (2012b), we selected only those veg-
etation patches in which complete information for all the plant
species within each patch was available. A total of 17 patches
were selected, sampled along two plots of 500 m
2
, each captur-
ing a representative sample of the plant species in the commu-
nity and reflecting their relative abundances (Supplementary
Data Table S1). In each patch, the species composition was re-
corded (a total of 39 plant individuals and 23 plant species),
and the AMF community harboured in each plant within the
patch was obtained from Montesinos-Navarro et al.(2012b)
(see more details about the root sampling in Montesinos-
Navarro et al.,2012b).
Phylogenetic community composition
Phylogenetic trees of plants and AMF were obtained from
those published in Montesinos-Navarro et al.(2012b;GenBank
accession numbers are given in Supplementary Data Table S1).
The phylogenetic community composition of plants and AMF
was independently characterized using analytical tools recently
developed in a meta-community framework (Pillar and Duarte,
2010). Following Pillar and Duarte (2010), AMF sequences
were used to calculate, after fuzzy weighting, species phyloge-
netic composition through an index (Pmatrix) that character-
izes the phylogenetic neighbourhood for each species within
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each patch. Two Pmatrices were independently calculated for
plant and AMF. The elements of the Pmatrix (patch taxa)
provide a characterization of the phylogenetic composition of
the assemblage. Pmatrices were calculated using the SYNCSA
package implemented in R (Debastiani and Pillar, 2012). As
suggested by Pillar and Duarte (2010), Pmatrices can be used
to explore phylogenetic patterns at the community level by us-
ing ordination techniques. The numbers of axes that capture at
least 90% of the variation in the phylogenetic community com-
position of plants and AMF were calculated, and a non-para-
metric multivariate analysis of variance (ANOVA; Anderson,
2001) was used to test if the phylogenetic composition of plants
could explain the phylogenetic composition of AMF (‘adonis’
routine of the Vegan package of R; Oksanen et al., 2013).
Afterwards, we focused on the first principal component (PC1)
of plants and AMF in order to be able to provide an ecological
explanation for the main variation axis. Principal co-ordinate
analyses (PCoAs) were conducted with Euclidean, Maximum,
Manhattan and Minkowski distances. Because all four distance
indices were consistent, hereafter we refer to Euclidean distance
results only, equivalent to using principal component analyses
(PCAs). PCAs were performed on the Pmatrix of plant and
AMF independently. The magnitude of each species loading in-
dicates the relative contribution of that species to differentiate
patches along the PC1 axis (patch scores). Patch scores were
calculated considering plants (plant PC1 scores) and AMF
(AMF PC1 scores) independently. Accordingly, a patch score
along the plant PC1 will compile information about the phylo-
genetic composition of the plants in that patch, and the patch
score of AMF PC1 will compile information about the phyloge-
netic composition of the AMF assembly in the same patch.
Patches harbouring species with opposite loading signs will
tend to be located in opposite extremes of the PC1 axis. The re-
lationship between plant and AMF phylogenetic composition
was tested by correlating plant PC1 scores and AMF PC1
scores. A potential spatial autocorrelation was tested using a
Mantel test with 9999 permutations to correlate plant PC1 and
AMF PC1 with the spatial distance between patches. PCAs and
correlation analyses were performed using the software R.
In order to provide an ecological interpretation of the plant
PC1 axis, plant species loadings were correlated with their nurs-
ery quality, which is a proxy of multiple functional traits poten-
tially influencing biotic interactions and community
assemblages. The nursery quality of plant species
(Supplementary Data Table S2) was characterized from the in-
formation available from previous studies in this system
(Valiente-Banuet and Verdu´ , 2007,2008). Nursery quality was
estimated as the number of seedlings growing underneath a
nurse which survive until the adult stage. In order to be consid-
ered as facilitation, the number of seedlings underneath a nurse
needs to differ from the number expected in a random distribu-
tion, considering the proportions of the area occupied by each
plant cover vs. open space (see more details on the estimation
of this parameter in Valiente-Banuet and Verdu´, 2007,2008).
Plant species loadings were correlated with their level of nurs-
ery quality using a non-parametric Spearman correlation, as
nursery quality was not normally distributed.
Regarding AMF, in which most DNA sequences cannot be
confidently assigned to known species, functional traits cannot
be used. Consequently, the ecological interpretation of the
AMF PC1 axis was based on its relationship with phylogenetic
diversity, assuming trait conservatism (Van der Heijden et al.,
1998;Powell et al.,2009). The mean phylogenetic diversity
was characterized by the mean pairwise phylogenetic distance
(MPD) among every pair of AMF sequences coexisting in a
patch. The MPD of each patch was obtained as a sub-set of the
matrix of phylogenetic distances between all AMF sequences
in the community. The relationship between phylogenetic com-
position and phylogenetic diversity was assessed using a non-
parametric Spearman correlation between AMF PC1 scores and
MPD of AMF sequences across patches.
Finally, the relative contribution of plant MPD, nursery qual-
ity of the patch (estimated as the community-weighted mean:
Lavorel et al., 2008), plant and AMF phylogenetic composition
(plant PC1 and AMF PC1) and the interaction between plant
phylogenetic diversity and nursery quality on AMF phyloge-
netic diversity (AMF MPD) was estimated by means of a
non-parametric ANOVA based on permutation tests using the
‘adonis’ function (Oksanen et al., 2013). The community-
weighted mean was determined using the ‘functcomp’ function
of the PD package of R (Laliberte´ et al., 2014).
Null model to discard patterns due to plant–AMF specificity
A significant relationship between plant and AMF phyloge-
netic compositions may just reflect the specificity in
plant–AMF interactions described at the species level
(Montesinos-Navarro et al.,2012b). In order to discard this pos-
sibility, a matrix of 576 AMF sequences by 23 plant species
present in the community was generated, with ‘1’ for the al-
lowed plant–AMF interactions representing the specificity in
the interactions and ‘0’ for the forbidden interactions. From the
576 allowed interactions, the same number of interactions as
that observed in each patch was randomly selected, to generate
17 ‘theoretical patches’. In this way, the plant–AMF specificity
pattern and the number of interactions per patch were main-
tained, but the phylogenetic composition in each patch was ran-
domized. A correspondence between plant and AMF
phylogenetic composition was tested for the set of 17 ‘theoreti-
cal patches’ as explained above for the real phylogenetic com-
positions. This process was repeated 1000 times.
RESULTS
The phylogenetic composition of plants and AMF assemblages
in each patch was characterized by means of plant PCA and
AMF PCA, respectively. Four and two PC axes captured at
least 90% of the variation of the phylogenetic composition of
plants and AMF, respectively. Plant PC1 explained 52 % and
AMF PC1 88% of the total variance of phylogenetic composi-
tion among patches (note that these two PCs come from inde-
pendent PCA analyses). There was no significant spatial
autocorrelation in either plant PC1 (r¼010; P-value ¼098)
or AMF PC1 (r¼006; P-value ¼015). The phylogenetic
composition of plants was not significantly correlated with the
number of plant species in the vegetation patch (r¼037;
P-value ¼017). Similarly, the AMF phylogenetic composition
was not significantly correlated with the number of AMF se-
quences found in the patch (r¼024; P-value ¼035).
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The AMF phylogenetic composition (AMF PC1) showed a
linear positive relationship with AMF phylogenetic diversity
(i.e. MPD) across patches (rho ¼089; P<0001; Fig. 1).
There was a negative correlation between plant species loading
in plant PC1 and the nursery quality of plant species
(rho ¼065; P<001, Supplementary Data Table S2).
Finally, there was a significant relationship between the phylo-
genetic composition of plants (plant PCA) and AMF (AMF
PCA) assemblages across patches. This relationship was signifi-
cant when considering all PC axes accumulating at least 90 %
of the total variation of plants (plant PC1–PC4) and AMF
(AMF PC1–PC2) (R
2
¼049; P-value ¼0038), and also when
considering the correlation between just the first principal axes
for plant (plant PC1) and AMF (AMF PC1) (R
2
¼024;
P-value ¼004; Fig. 2). This significant relationship between
plant and AMF phylogenetic compositions was not just a reflec-
tion of the specificity in plant–AMF interactions because the
significant correlation between plant PC1 and AMF PC1 disap-
pears in 903 of the 1000 randomly assembled communities in
which specificity in plant–AMF interactions was preserved.
The model constructed to quantify the effect of the relation-
ship of several plant and AMF variables on the mean phyloge-
netic diversity of AMF (AMF MPD) showed significant
contributions of the phylogenetic composition of AMF (39 %)
and plants (28%), and of the interaction between plant phyloge-
netic diversity and patch nursery quality (10 %) (Table 1). In
addition, there was still 18 % of non-explained variation. Plant
phylogenetic diversity and nursery quality independently did
not explain AMF phylogenetic diversity. A high plant phyloge-
netic diversity could result from distantly related species with
either low or high nursery quality, and these differences can in-
fluence AMF phylogenetic diversity. Similarly, a patch with a
high nursery quality community-weighted mean can harbour
different plant phylogenetic diversity, which will also condition
–100 1020304050
0
0⋅1
0⋅2
0⋅3
AMF phylo
g
enetic composition (AMF PC1 scores)
AMF phylogenetic diversity (MPD)
FIG. 1. Linear regression between phylogenetic diversity and phylogenetic com-
position of arbuscular mycorrhizal fungi (AMF) assemblages (i.e. patches).
Phylogenetic diversity is measured as mean phylogenetic distance between each
pairwise AMF DNA sequence present in each patch. Phylogenetic composition
in each patch is characterized as the first principal component of the Pmatrix
described by Pillar and Duarte (2010) which represents the phylogenetically
weighted taxa composition of each vegetation patch.
Cathestecum brevifolium 0⋅02
Bouteloua gracilis 0⋅03
Agavemacroacantha 0⋅07
Ruelliahirsuto–glandulosa 0⋅14
Justiciamexicana 0⋅16
Solanumtrydinamum 0⋅14
Ipomoea sp. 0⋅14
Mammillariacarnea 0⋅24
Mammillaria collina 0⋅24
Coryphantha pallida 0⋅23
Neobuxbaumiatetezto 0⋅23
Allionia incarnata 0⋅26
Mascagnia seleriana –0⋅18
Ditaxisguatemalensis –0⋅22
Senna wislizenii –0⋅28
Caesalpinia melanadenia –0⋅28
Calliandra eryophylla –0⋅28
Acaciaconstricta –0⋅28
Mimosaluisana –0⋅28
Dalea sp. –0⋅28
Eysenhardtia polystachya –0⋅27
Cardiospermum halicacabun –0⋅08
Thompsonella minutiflora –0⋅01
Plant species PC1 loadings
Plant nursery quality
AMF ph
y
lo
g
enetic diversit
y
–100 1020304050
–4
–2
0
2
4
AMF phylogenetic composition (AMF PC1 scores)
Plant phylogenetic composition (Plant PC1 scores)
Nurse Facilitated
HighLow
FIG. 2. Relationship between the phylogenetically weighted taxa composition of
plant and arbuscular mycorrhizal fungi (AMF) across vegetation patches. The
phylogenetic composition of plants and AMF in each patch is characterized as
the first principal component of the Pmatrix described by Pillar and Duarte
(2010). Plant and AMF PC1 represent the phylogenetic composition of the as-
semblages of the plant and AMF, respectively. Extreme values of the plant PC1
correspond to patches in which either nurse (high negative loading values) or fa-
cilitated (high positive loading values) species predominate. Extreme values of
the AMF PC1 correspond to patches with low and high AMF phylogenetic
diversity.
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AMF phylogenetic diversity. Accordingly, the interaction be-
tween plant phylogenetic diversity and patch nursery quality
better explained the phylogenetic composition of AMF, while
both effects independently were non-significant. The positive
significant interaction between these two terms indicated that
AMF MPD was promoted by the combination of high nurse
quality and phylogenetic diversity.
DISCUSSION
This study shows a correlation between the phylogenetic com-
positions of two interacting guild assemblages across a patchy
environment (Fig. 2). This correlation cannot be solely ex-
plained by plant–AMF specificity, and we suggest other ecolog-
ical processes potentially shaping the phylogenetic composition
of the vegetation patches.
Plant phylogenetic composition across patches is influenced
by the predominance of plant species with the lowest nursery
quality (e.g. Cactaceae, Solanaceae or Convolvulaceae), which
contribute to the highest values of the plant PC1, and the pre-
dominance of plant species with the highest nursery quality
(e.g. Fabaceae), which contribute to the lowest values of the
plant PC1. In addition, low values of the plant PC1 correspond
to high values of the AMF PC1 (Fig. 2), which in turn are
patches with a high phylogenetic diversity of AMF (Fig. 1).
This result, together with the significant interaction showing
that the combination of plant high nursery quality and phyloge-
netic diversity promotes AMF MPD, suggests that patches con-
taining predominantly diverse plant species with high nursery
quality (e.g. Fabaceae) tend to have a high AMF phylogenetic
diversity. In contrast, patches in which species with low nursery
quality predominate such as the Cactaceae, Mammillaria col-
lina,M. carnea,Neobuxbaumia tetetzo and Coryphanta pallida;
or the Acanthaceae, Ruellia hirsuto-glandulosa, tend to have a
low phylogenetic diversity of AMF.
Taxa composition can be crucial in plant–AMF interactions
as species traits, such as nursery quality, a functional character-
istic which encapsulates different traits (Valiente-Banuet et al.,
2006), may be influencing patterns of co-occurrence of AMF in
the patch. Fabaceae have been shown to be important nurses in
other systems (Barnes and Archer, 1996;Flores and Jurado,
2003;Liphadzi and Reinhardt, 2006), especially in semi-arid
environments (Flores and Jurado, 2003;Bashan et al.,2009;
Muro-Pe´rez et al.,2012). Their association with nitrogen-fixing
bacteria has been suggested as a potential mechanism underly-
ing colonization behaviour (Cadotte and Strauss, 2011), and a
tendency to act as a nurse for other species in the community.
In addition, the presence and the identity of AMF can alter the
competitive interactions between plant species (Van der
Heijden et al., 1998;Scheublin et al., 2007), and the Fabaceae
tend to obtain a greater benefit from AMF than other plants
(Scheublin et al., 2007). Our results suggest that the capability
to interact with phylogenetically distant AMF might be a poten-
tially relevant trait in defining the nursery quality of a plant spe-
cies in semi-arid plant communities, but experimental
approaches will be necessary to test this hypothesis.
Interestingly, AMF can be present from the beginning of
plant succession and relate differently to pioneer and late-suc-
cessional plant species (Kikvidze et al.,2010). This suggests
that reciprocal plant–AMF interactions may be involved in eco-
logical mechanisms of succession within a patch. The variation
in phylogenetic composition of plant and AMF across patches
could represent the inter-related replacement dynamics of
plants and AMF within a vegetation patch over time (i.e. suc-
cession). During the successional history of a patch, it might
shift from harbouring mainly species with high nursery quality,
which are the only pioneer species in this system (Valiente-
Banuet and Verdu´, 2008), to harbouring mainly facilitated spe-
cies upon the death of the species with high nursery quality.
Our results are consistent with the hypothesis that this shift
from the predominance of nurse to facilitated species within a
patch might be coupled with a reduction in below-ground phy-
logenetic diversity. However, further experiments are necessary
to test this hypothesis. We show that shifts in the phylogenetic
composition of the two guild assemblages are not independent
but there remains variation not explained by this relationship,
suggesting that other indirect effects such as interactions with
other guilds, not considered here, can also be influencing this
dynamic. In addition, we have focused on the traits related to
the PC axes that explain most of variation (PC1 of plants and
AMF), but other traits related to less important sources of varia-
tion can be driving other ecological processes represented in
the remaining axes. Furthermore, our results are based on the
most abundant group of AMF, Glomus, but the consideration of
other Glomeromycota could reveal new ecological processes
and functions that would provide a more complete understand-
ing of plant–AMF interactions.
In summary, we show that shifts in the phylogenetic compo-
sition of plants and AMF assemblages do not occur indepen-
dently, providing indirect evidence for the potential relevance
of interguild interactions in community assemblage processes.
We suggest that besides selectivity in plant–AMF interactions,
inter-related succession dynamics of plants and AMF within
patches could be an ecological mechanism leading to the ob-
served pattern. The consideration of other coexisting guilds in
these patches and the design of experimental approaches to test
for the indirect evidence provided here are necessary for a bet-
ter understanding of the role of interguild biotic interactions in
community assemblage. Considering the increasing feasibility
of exploring the community structure of soil micro-organisms
using next-generation sequencing tools, promising future lines
of research might explore whether there is evidence of
TABLE 1. Non-parametric multiple regression to test for multiple
effects on AMF phylogenetic diversity (AMF MPD)
Source Estimates d.f. FR
2
P-value
Plant MPD 3910
6
122003 061
Patch nursery quality 2910
2
100000 098
Plant PC1 1210
2
1165028 0005
AMF PC1 4010
3
1230039 0002
Plant MPD patch nursery quality 102 10
4
160010 0031
Residuals 11 018
Total 16
The model tests for the relative contribution of plant mean phylogenetic
distance (plant MPD), community-weighted mean of nursery quality (patch
nursery quality), plant and AMF phylogenetic composition (plant PC1 and
AMF PC1) and the interaction of plant phylogenetic diversity and nursery
quality.
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interlinked dynamics above- and below-ground, taking into ac-
count multiple guilds simultaneously.
SUPPLEMENTARY DATA
Supplementary data are available online at www.aob.oxford-
journals.org and consist of the following. Table S1:AMF
GenBank accession number, plant host species and PC1 load-
ing. Table S2: nursery quality score and PC1 loading for plant
species.
ACKNOWLEDGEMENTS
We thank J. P. Castillo, M. Morales, C. Silva and L. Sortibra´n
for help with field samplings, and S. Donat and M. Morales
for laboratory assistance. Three anonymous referees provided
valuable input that considerably improved the manuscript.
This work was funded by AECID (Projects A017475/08,
A023461/09), DGAPA-UNAM (Project IN-202811-3; IN-
213414-3), CYTED (Accio´n 409AC0369) and MICINN
(CGL2011-29585-C02-01). A.M.N. was supported by a
DGAPA-UNAM post-doctoral fellowship and an Early Career
Project Grant from the BES (3975-4849), and J.G.S.-M. was
supported by a ‘Ramo´ n y Cajal’ post-doctoral contract from
MICINN.
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