1878 NOTES Ecology, Vol. 94, No.

Ecology, 94(8), 2013, pp. 1878–1885

Ó 2013 by the Ecological Society of America

Functionally and phylogenetically diverse plant communities

key to soil biota
ALEXANDRU MILCU,1,2,18 ERIC ALLAN,3 CHRISTIANE ROSCHER,4,5 TANIA JENKINS,6 SEBASTIAN T. MEYER,7 DAN FLYNN,8
HOLGER BESSLER,9 FRANÇOIS BUSCOT,10,11 CHRISTOF ENGELS,9 MARLÉN GUBSCH,12 STEPHAN KÖNIG,10
ANNETT LIPOWSKY,8 JESSY LORANGER,13 CARSTEN RENKER,11 CHRISTOPH SCHERBER,14 BERNHARD SCHMID,8
ELISA THÉBAULT,15 TESFAYE WUBET,10 WOLFGANG W. WEISSER,7 STEFAN SCHEU,16 AND NICO EISENHAUER17
1
CNRS, Ecotron–UPS 3248, Campus Baillarguet, 34980, Montferrier-sur-Lez, France
2
Imperial College London, Division of Biology, Ascot SL5 7PY United Kingdom
3
Institute of Plant Sciences, University of Bern, Altenbergrain 21, 3013 Bern, Switzerland
4
UFZ, Helmholtz Centre for Environmental Research, Department of Community Ecology, Halle, Germany
5
Max Planck Institute for Biogeochemistry, Hans-Knöll Strasse 10, 07745 Jena, Germany
6
Department of Ecology and Evolution, University of Lausanne, Biophore-Quartier Sorge CH-1015 Lausanne, Switzerland
7
Technische Universität München, Department of Ecology and Ecosystem Management, Hans-Carl-von-Carlowitz-Platz 2,
85350 Freising, Germany
8
Institute of Evolutionary Biology and Environmental Studies, University of Zurich, Winterthurerstrasse 190,
CH-8057 Zurich, Switzerland
9
Humboldt-Universität zu Berlin, Institute of Plant Nutrition, Albrecht-Thaer-Weg 4, 14195 Berlin, Germany
10
UFZ, Helmholtz Centre for Environmental Research, Department of Soil Ecology, Theodor-Lieser-Straße 4, 06102 Halle, Germany
11
University of Leipzig, Institute of Biology, Johannis-Allee 21-23, 04103 Leipzig, Germany
12
Institute of Plant Sciences, ETH Zurich, Universitätsstrasse 2, 8092 Zurich, Switzerland
13
CNRS CEFE, UMR 5175, 1919 Route de Mende, 34293, Montpellier, France
14
Georg-August-University Göttingen, Agroecology, Department of Crop Sciences, Grisebachstrasse 6, 37077 Göttingen, Germany
15
CNRS, UMR 7618, Laboratoire ‘‘Bioge´ochimie et e´cologie des milieux continentaux,’’ 46 Rue d’Ulm, 75005 Paris, France
16
Georg-August-University, J.F. Blumenbach Institute of Zoology and Anthropology, Berliner Strasse 28, 37073 Göttingen, Germany
17
Friedrich-Schiller-University, Institute of Ecology, Dornburger Strasse 159, 07743 Jena, Germany

Abstract. Recent studies assessing the role of biological diversity for ecosystem
functioning indicate that the diversity of functional traits and the evolutionary history of
species in a community, not the number of taxonomic units, ultimately drives the biodiversity–
ecosystem-function relationship. Here, we simultaneously assessed the importance of plant
functional trait and phylogenetic diversity as predictors of major trophic groups of soil biota
(abundance and diversity), six years from the onset of a grassland biodiversity experiment.
Plant functional and phylogenetic diversity were generally better predictors of soil biota than
the traditionally used species or functional group richness. Functional diversity was a reliable
predictor for most biota, with the exception of soil microorganisms, which were better
predicted by phylogenetic diversity. These results provide empirical support for the idea that
the diversity of plant functional traits and the diversity of evolutionary lineages in a
community are important for maintaining higher abundances and diversity of soil
communities.
Key words: above–belowground interactions; biodiversity; functional diversity; functional traits; Jena
Experiment; phylogenetic diversity; plant species richness; soil fauna.

INTRODUCTION 2009). Particularly, understanding the links between

Linking changes in community composition and
diversity between trophic levels presents a major
challenge for community and ecosystem ecology (Van
der Putten et al. 2001, Wardle et al. 2004, Haddad et al.
Manuscript received 5 November 2012; revised 30 January
2013; accepted 25 February 2013. Corresponding Editor: B. A.
Wardle.
18 E-mail: a.milcu@imperial.ac.uk
above- and belowground communities has emerged as
an important challenge given that soil biota are not just
a ‘‘black box’’ of highly redundant species and that they
drive a range of ecosystem functions (Scheu and Setälä
2002, Wardle et al. 2004). We are increasingly learning
that soil biota are closely interlinked with aboveground
communities and that there is a greater degree of
specificity between plants and soil organisms than was
previously assumed (Scheu 2001, Wardle et al. 2004,
 August 2013 NOTES
Bardgett and Wardle 2010). There is compelling
evidence that soil biota are responsive to the quality
and quantity of organic matter inputs as well as to
changes in micro-environmental conditions associated
with changes in plant diversity (Wardle et al. 2004,
Bardgett and Wardle 2010). Despite this, a large body of
literature suggests that soil biota may be less sensitive to
changes in plant diversity than aboveground biota
(Gastine et al. 2003, Scherber et al. 2010). However,
this conclusion was mainly based on short-term studies
that investigated only two facets of plant diversity, i.e.,
species and functional group richness (Eisenhauer et al.
2012).
Recent studies indicate that the diversity of functional
traits or the evolutionary history of a community, not
the number of taxonomic units, ultimately drives
biodiversity–ecosystem-functioning relationships (Ca-
dotte et al. 2009, Flynn et al. 2011). A trait is any
morphological, biochemical, behavioral, and phenolog-
ical characteristic of an individual that potentially
affects its performance and fitness (Petchey and Gaston
2002). Identifying the most relevant functional traits
underpinning the biodiversity–ecosystem-functioning
relationship can be challenging, with the results sensitive
to the number and choice of traits included in the
analyses (Petchey and Gaston 2006). Given these
potential limitations of trait-based approaches, phylo-
genetic diversity, the sum of the shared evolutionary
history in a community, has been proposed as a useful
proxy to describe the true functional diversity of a
community (Cadotte et al. 2009). Phylogenetic diversity
should affect ecosystem functioning if ecological dissim-
ilarity is correlated with evolutionary divergence,
meaning that the more phylogenetically divergent
species are present, the more likely it is that they have
dissimilar functional traits and occupy different niches,
thereby differentially impacting ecosystem functioning
(Felsenstein 1985, Maherali and Klironomos 2007).
Additionally, plant phylogenetic diversity may be
particularly important for higher trophic levels, and
more than just a proxy for functional diversity if plant
phylogeny reflects coevolutionary interactions between
plants and other organism groups (Dinnage et al. 2012).
Lately, functional and phylogenetic diversity have
been shown to be better predictors of primary produc-
tivity (Cadotte et al. 2009, Clark et al. 2012) and
arthropod diversity and abundances than were plant
species or functional group richness (Dinnage et al.
2012). However, we have remarkably little empirical
evidence on whether these indices are superior predictors
of soil biota than species and functional group richness.
In addition, information on the relevance of different
plant diversity indices may contribute to a better
understanding of how plant diversity effects cascade
into belowground food webs. Here we assess the
performance of several plant functional and phyloge-
1879
netic diversity metrics, alongside more conventional
metrics of plant community composition such as
realized plant species richness (Rdiv), functional group
richness (FG) and functional group biomass as deter-
minants of soil biota using data from one of the most
comprehensive biodiversity experiments so far, the Jena
Experiment (Roscher et al. 2004). We focused on the
abundances and diversity of soil biota collected from 82
grassland plots with experimentally manipulated plant
species (1, 2, 4, 8, 16, and 60) and functional group
richness (1, 2, 3, and 4), measured on the sixth year from
the onset of the diversity treatments.
METHODS
Study site and experimental design.—The experimental
site (50855 0 N, 11835 0 E, 130 m above sea level; mean
annual temperature 9.38C, mean annual precipitation
587 mm) was a former arable field located on the
floodplain of the Saale River, Jena, Germany. The
number of plant species, plant functional groups and
plant identity is controlled, in a randomized four block
design comprising 82 plots of 20 3 20 m. Plots were
established in May 2002 with 1, 2, 4, 8, 16, or 60
perennial grassland plant species typical for local
Arrhenatherum grasslands, with 16, 16, 16, 16, 14, and
4 replicates, respectively (see Roscher et al. [2004] for
details on experimental design). Plant compositions in
the plots were randomly chosen from a pool of 60
species and maintained by a combination of biannual
mowing, weeding, and herbicide applications.
Functional diversity estimated from traits.—We select-
ed 12 plant functional traits, based on literature-
informed knowledge, that affect soil biota and processes
through the quality and consistency of plant-derived
organic inputs as well as through changes of microhab-
itat environmental conditions. For each of the 60 plant
species, the traits were derived from in situ measure-
ments (shoot biomass dry mass [mg], biomass-to-N ratio
[mg N/g], shoot lignin and hemicelluloses content [%],
shoot C-to-N ratio, seed mass [mg], leaf area ratio [mm2/
mg dry mass], ability to fix atmospheric N2 [binary]) and
literature surveys (seasonality of foliage [ordinal; 1,
summer green; 2, partly evergreen; 3, evergreen], number
of known secondary compounds, rooting type [ordinal;
1, long-living primary root system; 2, secondary fibrous
roots in addition to the primary root system; 3, short-
living primary root system, extensive secondary root
system] and rooting depth [cm] as used by Roscher et al.
[2004]; see Appendix A for details on plant traits). The
traits were scaled to have a mean of zero and variance of
one. The resulting trait matrix was converted into a
Euclidean distance matrix and used to calculate the
distance based functional diversity metrics; for calculat-
ing functional diversity (FD) the distance matrix was
converted into a functional dendrogram by a UPGM
clustering analysis (Petchey and Gaston 2002; see also
 1880 NOTES
Appendix B). For each plant community, FD was
calculated as the branch length connecting the member
species of the respective community. In addition, Rao’s
quadratic diversity (Qr) (Rao 1982) was estimated as
done by Botta-Dukát (2005) using species percentage
cover (averaged for May and August 2008) to weight
branch lengths between species to generate an abun-
dance-weighted functional index that incorporates
information about functional richness as well as
functional evenness of a community.
Phylogenetic diversity.—A phylogeny of all 60 species
in the Jena Experiment species pool was constructed
based on four genes, using Bayesian methods (for
details, see Allan et al. [2013] and Appendix C). Two
measures of phylogenetic diversity were calculated from
this phylogeny: mean pairwise distance (MPD) and
mean nearest neighbor distance (MNND). MPD mea-
sures the mean phylogenetic distance between all pairs of
species (close and distant relatives) and is affected by the
number of deeper splits in the phylogeny. MNND
measures the mean distance between each species and its
closest relative and therefore measures diversity only at
the tips of the phylogeny. However, several communities
had high MPD but low MNND, providing the
justification for including both measures in our analyses.
Soil organism and plant sampling.—Five soil cores (5
cm diameter, 5 cm depth) were taken from each plot for
determining microbial biomass (Cmic). The pooled five
samples were homogenized, sieved (2 mm) to remove
larger roots, animals, and Cmic was measured using an
O2-microcompensation apparatus (Scheu 1992). Glu-
cose was added to saturate the catabolic enzymes of the
microorganisms (4 mg/g dry mass added as solution to
increase the water content to the water holding capacity
of the soil). The mean of the lowest three hourly
readings within the first 10 h was taken as maximum
initial respiratory response (MIRR; lL O2h1g soil dry
mass1) and microbial biomass (lg C/g soil dry mass)
was calculated as 38 3 MIRR (Beck et al. 1997).
Belowground macro- and mesofauna were Tullgren
extracted from one large (20 cm diameter, 20 cm depth)
and one small (5 cm diameter, 10 cm depth) soil cores
per plot. Diversity of arbuscular mycorrhizal fungi
(AMF) was analyzed using a molecular TaqMan qRT-
PCR assay designed after initial ITS PCR based
inventory of the AMF species list present at the site
(Konig et al. 2010). Plant species cover was estimated
from a 9-m2 subplot in each plot, whereas the
aboveground plant community biomass was the average
of the late May and late August (year 2008) harvests by
clipping the vegetation at 3 cm above ground in four
rectangles of 0.2 3 0.5 m per plot (see Weigelt [2010] for
methodological details on measurements of plant
biomass and cover estimates). Data on soil biota have
been used previously to explore the effects of plant
species and functional groups (Eisenhauer et al. 2010,
Ecology, Vol. 94, No. 8
Scherber et al. 2010), but have not been used to test
functional or phylogenetic diversity indices.
Statistical analyses.—We used path analysis, a partic-
ular case of structural equation modeling involving only
measured variables, to test the support for multiple
potential drivers while accounting for the unavoidable
colinearities among the explanatory variables (Grace
2006; see also Appendix D). As the calculation of the
abundance weighted functional and phylogenetic diversi-
ty indices was based on the realized species richness
(Rdiv) from the same year as the soil biota samplings
(year 2008), we therefore preferred to use the year 2008
Rdiv in the analyses (note that Rdiv is highly correlated
with sown species richness [Pearson’s r (realized, sown) ¼
0.99]). For each response variable (see Appendix E) a full
model of causal relationships was created including
simultaneously several hypothetical pathways through
which log-transformed (to reduce leverage and linearize
relationships) Rdiv could affect soil biota. The full model
(see Appendix F) included a direct pathway between Rdiv
and soil biota abundance/diversity and several indirect
pathways via functional diversity indices (FD and Qr),
phylogenetic diversity indices (MPD and MNND),
functional group richness (FG) and functional group
biomass (i.e., the biomass of legumes, grasses, small
herbs, and tall herbs). We included the path between Rdiv
and the other diversity measures because the Jena
Experiment manipulated species richness and therefore
variation in functional/phylogenetic diversity is caused by
variation in species richness and composition between
plots. Functional and phylogenetic diversity were in turn
hypothesized to affect soil biota directly or indirectly via
several measures of plant biomass (shoot, root, and total)
as plant productivity has been shown to be an important
driver of belowground communities (Spehn et al. 2000).
A continuous variable (percent clay content in the upper
10 cm soil layer ranging from 13.7% to 25.6% was
preferred to the block variable to account for the
variability in soil texture. The full models were simplified
by step-wise exclusion of variables with nonsignificant
regression weights and nonsignificant covariances as
estimated by AIC (Akaike information criterion) scores
until a minimal adequate model was achieved. Minimal
adequate models were indicated by non-significant
differences when comparing the predicted and observed
covariance matrices (v2 tests with P . 0.05), by lower
AIC, lower root mean squared error approximation
(RMSEA ,0.05) and higher comparative fit index (CFI
. 0.90) (Grace 2006, Arbuckle 2009). Path analysis was
performed using the SPPS Amos 18 statistical package
(Arbuckle 2009). Scatter plots with univariate linear
regression line were produced in R 2.15.0 (R Develop-
ment Core Team 2012) for visualizing the direction of the
relationship between the soil biota and the different
diversity indices (Appendix G). Data on abundance and
diversity of the soil biota was square-root transformed to
 August 2013 NOTES 1881

FIG. 1. Minimal adequate models for the effects of multiple plant community predictors on the abundance and diversity of
various groups of belowground organisms (see Appendix F for the maximal model). Solid arrows show significant relationships
(pathways) between variables, dotted arrows indicate a nonsignificant relationship, and numbers next to arrows show standardized
parameter estimates (i.e., standardized regression weights). Circles (e1–e6) indicate error terms, and double-headed arrows indicate
significant correlations between the error terms. Squared multiple correlations (R 2) for the predicted/dependent group of soil biota
is given on the box of the dependent variable. Abbreviations are: Rdiv, realized plant species richness; Cmic, microbial biomass;
AMFdiv, arbuscular mycorrhizal fungal diversity; CollembolaAB, abundance of Collembola; OrbatidaAB, abundance of
Oribatida; TotalmesofaunaAB, total mesofauna invertebrate abundance; MacrosaprotrophAB, macrofauna decomposer
invertebrate abundance; MacrosaprotrophDIV, macrofauna decomposer invertebrate species richness; MacroherbivoreAB,
macrofauna herbivore invertebrate; MacroherbivoreDIV, macrofauna herbivore invertebrate species richness; Totalmacrofau-
naAB, total macrofauna abundance; TotalmacrofaunaDIV, total macrofauna species richness; FD, functional diversity; Qr, Rao’s
quadratic diversity; MPD, mean pairwise phylogenetic distance; MNND, mean nearest neighbor phylogenetic distance; LegBM,
biomass of legumes; RootBM, root biomass; ShootBM, shoot biomass. No adequate models with significant regression weights
could be obtained for abundance of Gamasida, abundance of Symphyla, macrofauna predator abundance, and macrofauna
predator species richness (see Appendices H and I for full results of the path analysis).

reduce heteroscedasticity of error variances, whereas
microbial biomass was log-transformed to improve
normality. Although variables such as total mesofauna
and macrofauna abundances and diversity are not
independent from the comprising subordinate groups,
we consider them informative because independent data
sets with abundances and diversity of multiple trophic
levels of soil biota as well as in-situ collected plant
functional traits are rarely available. In addition, it is
unknown whether diversity and abundance of these
groups respond similarly to functional and phylogenetic
plant diversity.
RESULTS
Plant functional or phylogenetic diversity indices were
retained as significant predictors of soil biota diversity
and abundance in 10 out of the 11 minimal adequate
path analysis models, while direct effects of realized
species richness (Rdiv) were retained in just two models
(Fig. 1, Appendixes H and I). Rdiv is correlated with
 1882 NOTES
most of the other diversity measures (Appendix D),
therefore the path between Rdiv and the other diversity
measures was retained in almost all models; however the
lack of direct paths between Rdiv and soil biota diversity
and abundance indicates that Rdiv is a less important
predictor for soil biota than functional/phylogenetic
diversity. No minimal adequate models could be
achieved for the abundances of gamasid mites and
Symphyla or for the diversity and abundance of
macropredators. Rao’s quadratic diversity (Qr) was
retained as the sole predictor in the models predicting
the abundance and diversity of herbivores (Fig. 1) and
the diversity of macrofauna. Functional diversity (FD)
was retained in six of the eleven adequate models and
affected soil biota directly but also indirectly (via plant
shoot or root biomass). Note that direct paths indicate
effect pathways unrelated to plant biomass. Plant shoot
biomass was retained as a significant predictor in the
models for microbial biomass and the total abundance
of macrofauna (Fig. 1), which was in turn affected by
the amount of legume biomass and FD. Root biomass
was retained in the models for predicting the abundance
of Collembola, mesofauna and macrosaprotrophs and
the diversity of macrosaprotrophs. Phylogenetic diver-
sity was retained in the models predicting microbial
biomass (retained MPD) and the diversity of arbuscular
mycorrhizae (retained MNND). In all minimal adequate
models, the standardized and unstandardized regression
weights (Fig. 1, Appendixes H and I) indicate that
higher functional or phylogenetic diversity led to higher
abundances and diversity of the analyzed taxonomic
groups (Appendix G) and increased plant root and
shoot biomass.
DISCUSSION
Functional and phylogenetic diversity indices have
been proposed as a pragmatic and more accurate way of
capturing potential niche complementarity in a commu-
nity (Cadotte et al. 2009, Clark et al. 2012). While it
often has been shown that plant functional and
phylogenetic diversity drives aboveground communities
and processes (Cadotte et al. 2009, 2012, Flynn et al.
2011), little empirical evidence is available showing that
it is also a key determinant of soil communities. The
findings of this study provide strong evidence that
belowground communities increase in complexity (abun-
dances and species richness) in response to increased
plant functional diversity. The results also show that
functional and phylogenetic diversity metrics outweigh
the traditionally used species and functional group
richness as predictors of soil taxa abundance and
diversity. Measuring multiple functional traits should
provide a higher resolution picture of potential niche
complementarity in a community beyond what species
or functional group richness can explain since not every
species increases functional diversity by an identical
Ecology, Vol. 94, No. 8
amount (Petchey and Gaston 2002). Furthermore,
additional mechanisms emerging from niche comple-
mentarity such as increased microhabitat heterogeneity
(Eisenhauer et al. 2011), substrate diversity (Spehn et al.
2000), and asynchronous population fluctuations
(Roscher et al. 2011), all of which have been shown to
contribute to increased ecosystem stability (Naeem and
Li 1997, Milcu et al. 2010), can be better captured by
functional diversity (Flynn et al. 2011, Cadotte et al.
2012). This is in contrast to previous studies showing
that soil organisms mainly respond to the presence of
certain plant functional groups (e.g., N2 fixers [Spehn et
al. 2000, Milcu et al. 2008]) and underlines the
importance of considering multiple plant traits in
functional metrics aiming to predict belowground
communities.
Recent studies have emphasized the importance of the
diversity of plant evolutionary lineages for ecosystem
functioning and diversity of arthropods (Cadotte et al.
2012, Dinnage et al. 2012), and here we show that the
evolutionary history of the plant community (measured
as phylogenetic diversity) also drives mycorrhizal diver-
sity and soil microbial biomass. This supports the
existence of strong coevolutionary links between soil
microorganisms and plants and the existence of specific
associations between plants and microorganisms (Rey-
nolds et al. 2003, Eisenhauer et al. 2010). Arbuscular
mycorrhizae are obligate biotroph symbionts that form
tight associations with their host plants, and recent
studies using network theory to link the diversity of host
plants and of arbuscular mycorrhizae found a nested
relationship pattern suggesting strong specialization
(Montesinos-Navarro et al. 2012). As these groups are
important for plant nutrient uptake and decomposition,
and a high mycorrhizal diversity has been shown to be
associated with higher plant biomass production (Mahe-
rali and Klironomos 2007), their greater abundance in
phylogenetically diverse communities might also partially
explain the increase in plant biomass with plant
phylogenetic diversity (Cadotte et al. 2009).
In our study, FD and Qr were superior predictors of
soil biota for meso- and macrofauna, which suggests that
these groups instead respond to the diversity in traits
affecting plant resource quality and micro-environmental
changes. This suggests that different soil organisms are
specialized on different types of plant resources and plant
communities with higher diversity of resources support
more abundant and diverse meso- and macrofauna.
Plant traits are not stable plant characteristics, but vary
with growing seasons, environmental (Ackerly and
Cornwell 2007) and diversity gradients (Gubsch et al.
2011), or even at a given time point within populations
(Albert et al. 2010, Clark 2010). In spite of efforts to
incorporate intraspecific trait variation in measures of
functional diversity (Albert et al. 2010) and the relative
amount of variation in different traits (Clark et al. 2012), it
 August 2013 NOTES
is unclear how this variation could best be included in
modeling relationships between functional trait diversity
and processes related to other trophic levels, which also
are not constant temporally. It is usually assumed in trait-
based approaches that within-species trait variation is
smaller than between-species trait variation (McGill et al.
2006). Nevertheless, we cannot exclude the possibility that
missing or weak relationships between plant functional
diversity and soil biota were due to not incorporating
intraspecific trait variation. However, the many significant
effects of functional trait diversity that we did find suggest
that the majority of belowground organism groups
respond to interspecific variation in functional traits.
Predator trophic levels (gamasid mites and macro-
predators), did not respond to any of our measures of
plant diversity. This is in line with previous studies
indicating that effects of species diversity of one trophic
level become weaker with the trophic distance (De Deyn
et al. 2004, Scherber et al. 2010). Moreover, it suggests
that the failure to detect bottom-up effects of plant
species richness on predatory trophic levels is not simply
due to overlooking indirect effects via functional and
phylogenetic diversity. Instead, top-down (Haddad et al.
2009) or environmental drivers probably shaped the
abundance and diversity of predatory communities.
The multiple pathways through which species richness
can affect the diversity and abundance of soil fauna
emphasizes the multidimensionality of factors involved in
plant–soil interactions. While many of the plant func-
tional and phylogenetic effects where direct, in some cases
they were mediated by plant productivity, e.g., soil
microbial biomass increased with shoot biomass (which
in turn was affected by the presence of legumes and
functional diversity), but also with phylogenetic diversity.
Shoot biomass has previously been suggested to affect
microbial biomass through the amount of root exudates
entering the soil (Eisenhauer et al. 2010). Furthermore,
root biomass was also retained in several models as an
indirect pathway through which functional diversity
affected abundances and diversity of saprotrophic fauna,
in line with previous studies underlining the role of the
quantity of belowground inputs for decomposer food
webs (Pollierer et al. 2007, Bardgett and Wardle 2010).
Overall, the relationship between functional diversity
and belowground communities may still seem somewhat
weaker when compared with aboveground food webs
(Scherber et al. 2010). However, it is highly unlikely that
there will ever be a ‘‘perfect’’ index of functional diversity
able to equally predict the response of all trophic levels
(Petchey and Gaston 2006). Moreover, the heterogeneous
nature of the soil environment at different spatial and
temporal scales poses a major challenge for ecologists
trying to quantify the importance of biotic determinants
of soil communities and very likely explains the low effect
sizes detected in some studies so far (Gastine et al. 2003,
Viketoft et al. 2009). Despite some limitation of the used
1883
metrics (e.g., unknown effects of plant functional trait
variation), the results provide strong evidence that soil
biota are responsive to facets of plant diversity such as
functional and phylogenetic diversity and add to the
mounting evidence that plant diversity is a key driver of
belowground communities and ecosystem functioning
(Zak et al. 2003, Milcu et al. 2008, Eisenhauer et al. 2012).
ACKNOWLEDGMENTS
Deutsche Forschungsgemeinschaft (DFG) is acknowledged
for funding the Jena Experiment (FOR 456) and Nico
Eisenhauer (Ei 862/1 and Ei 862/2). A. Milcu is grateful to
Georgina Mace and Jacques Roy for their support.
LITERATURE CITED
Ackerly, D. D., and W. Cornwell. 2007. A trait-based approach
to community assembly: partitioning of species trait values
into within-and among-community components. Ecology
Letters 10:135–145.
Albert, C. H., W. Thuiller, N. G. Yoccoz, A. Soudant, F.
Boucher, P. Saccone, and S. Lavorel. 2010. Intraspecific
functional variability: extent, structure and sources of
variation. Journal of Ecology 98:604–613.
Allan, E., T. Jenkins, A. J. F. Fergus, C. Roscher, M. Fischer,
J. Petermann, W. W. Weisser, and B. Schmid. 2013.
Experimental plant communities develop phylogenetically
overdispersed abundance distributions during assembly.
Ecology 94:465–477.
Arbuckle, J. 2009. Amos 18 user’s guide. SPSS Incorporated,
Armonk, New York, USA.
Bardgett, R. D., and D. A. Wardle. 2010. Aboveground-
belowground linkages: biotic interactions, ecosystem process-
es, and global change. Oxford University Press, Oxford, UK.
Beck, T., R. Joergensen, E. Kandeler, F. Makeschin, E. Nuss,
H. Oberholzer, and S. Scheu. 1997. An inter-laboratory
comparison of ten different ways of measuring soil microbial
biomass C. Soil Biology and Biochemistry 29:1023–1032.
Botta Dukát, Z. 2005. Rao’s quadratic entropy as a measure of
functional diversity based on multiple traits. Journal of
Vegetation Science 16:533–540.
Cadotte, M., J. Cavender-Bares, D. Tilman, and T. Oakley.
2009. Using phylogenetic, functional and trait diversity to
understand patterns of plant community productivity. PLoS
ONE 4:843–845.
Cadotte, M., R. Dinnage, and G. D. Tilman. 2012. Phyloge-
netic diversity promotes ecosystem stability. Ecology 93:223–
233.
Clark, C. M., D. F. B. Flynn, B. J. Butterfield, and P. B. Reich.
2012. Testing the link between functional diversity and
ecosystem functioning in a Minnesota grassland experiment.
PLoS ONE 7:e52821.
Clark, J. S. 2010. Individuals and the variation needed for high
species diversity in forest trees. Science 327:1129–1132.
De Deyn, G. B., C. E. Raaijmakers, J. Van Ruijven, F.
Berendse, and W. H. Van Der Putten. 2004. Plant species
identity and diversity effects on different trophic levels of
nematodes in the soil food web. Oikos 106:576–586.
Dinnage, R., M. W. Cadotte, N. M. Haddad, G. M. Crutsinger,
and D. Tilman. 2012. Diversity of plant evolutionary lineages
promotes arthropod diversity. Ecology Letters 15:1308–1317.
Eisenhauer, N., H. Beßler, C. Engels, G. Gleixner, M.
Habekost, A. Milcu, S. Partsch, A. Sabais, C. Scherber,
and S. Steinbeiss. 2010. Plant diversity effects on soil
microorganisms support the singular hypothesis. Ecology
91:485–496.
Eisenhauer, N., A. Milcu, E. Allan, N. Nitschke, C. Scherber,
V. Temperton, A. Weigelt, W. W. Weisser, and S. Scheu.
 1884 NOTES
2011. Impact of above and below ground invertebrates on
temporal and spatial stability of grassland of different
diversity. Journal of Ecology 99:572–582.
Eisenhauer, N., P. B. Reich, and S. Scheu. 2012. Increasing
plant diversity effects on productivity with time due to
delayed soil biota effects on plants. Basic and Applied
Ecology 13:571–578.
Felsenstein, J. 1985. Phylogenies and the comparative method.
American Naturalist 125:1–15.
Flynn, D., N. Mirotchnick, M. Jain, M. Palmer, and S. Naeem.
2011. Functional and phylogenetic diversity as predictors of
biodiversity-ecosystem function relationships. Ecology 92:
1573–1581.
Gastine, A., M. Scherer-Lorenzen, and P. Leadley. 2003. No
consistent effects of plant diversity on root biomass, soil
biota and soil abiotic conditions in temperate grassland
communities. Applied Soil Ecology 24:101–111.
Grace, J. B. 2006. Structural equation modeling and natural
systems. Cambridge University Press, Cambridge, UK.
Gubsch, M., N. Buchmann, B. Schmid, E.-D. Schulze, A.
Lipowsky, and C. Roscher. 2011. Differential effects of plant
diversity on functional trait variation of grass species. Annals
of Botany 107:157–169.
Haddad, N. M., G. M. Crutsinger, K. Gross, J. Haarstad,
J. M. H. Knops, and D. Tilman. 2009. Plant species loss
decreases arthropod diversity and shifts trophic structure.
Ecology Letters 12:1029–1039.
Konig, S., T. Wubet, C. F. Dormann, S. Hempel, C. Renker,
and F. Buscot. 2010. TaqMan real-time PCR assays to assess
arbuscular mycorrhizal responses to field manipulation of
grassland biodiversity: effects of soil characteristics, plant
species richness, and functional traits. Applied and Environ-
mental Microbiology 76:3765.
Maherali, H., and J. N. Klironomos. 2007. Influence of
phylogeny on fungal community assembly and ecosystem
functioning. Science 316:1746.
McGill, B. J., B. J. Enquist, E. Weiher, and M. Westoby. 2006.
Rebuilding community ecology from functional traits.
Trends in Ecology and Evolution 21:178–185.
Milcu, A., S. Partsch, C. Scherber, W. W. Weisser, and S.
Scheu. 2008. Earthworms and legumes control litter decom-
position in a plant diversity gradient. Ecology 89:1872–1882.
Milcu, A., E. Thebault, S. Scheu, and N. Eisenhauer. 2010.
Plant diversity enhances the reliability of belowground
processes. Soil Biology and Biochemistry 42:2102–2110.
Montesinos-Navarro, A., J. G. Segarra-Moragues, A. Valiente-
Banuet, and M. Verdú. 2012. The network structure of plant–
arbuscular mycorrhizal fungi. New Phytologist 194:536–547.
Naeem, S., and S. Li. 1997. Biodiversity enhances ecosystem
reliability. Nature 390:507–509.
Petchey, O. L., and K. J. Gaston. 2002. Functional diversity
(FD), species richness and community composition. Ecology
Letters 5:402–411.
Petchey, O. L., and K. J. Gaston. 2006. Functional diversity:
back to basics and looking forward. Ecology Letters 9:741–
758.
Pollierer, M., R. Langel, C. Körner, M. Maraun, and S. Scheu.
2007. The underestimated importance of belowground
carbon input for forest soil animal food webs. Ecology
Letters 10:729–736.
SUPPLEMENTAL MATERIAL
Appendix A
A table with the means and standard deviations of the 12 functional traits derived from the 60 species present in the Jena
Experiment (Ecological Archives E094-170-A1).
Ecology, Vol. 94, No. 8
R Development Core Team. 2012. R: a language and
environment for statistical computing. R Foundation for
Statistical Computing, Vienna, Austria. www.r-project.org
Rao, C. R. 1982. Diversity and dissimilarity coefficients—a
unified approach. Theoretical Population Biology 21:24–43.
Reynolds, H. L., A. Packer, J. D. Bever, and K. Clay. 2003.
Grassroots ecology: plant–microbe–soil interactions as driv-
ers of plant community structure and dynamics. Ecology 84:
2281–2291.
Roscher, C., J. Schumacher, J. Baade, W. Wilcke, G. Gleixner,
W. W. Weisser, B. Schmid, and E. D. Schulze. 2004. The role
of biodiversity for element cycling and trophic interactions:
an experimental approach in a grassland community. Basic
and Applied Ecology 5:107–121.
Roscher, C., A. Weigelt, R. Proulx, E. Marquard, J.
Schumacher, W. W. Weisser, and B. Schmid. 2011. Identi-
fying population- and community-level mechanisms of
diversity–stability relationships in experimental grasslands.
Journal of Ecology 99:1460–1469.
Scherber, C., N. Eisenhauer, W. W. Weisser, B. Schmid, W.
Voigt, M. Fischer, E. D. Schulze, C. Roscher, A. Weigelt,
and E. Allan. 2010. Bottom-up effects of plant diversity on
multitrophic interactions in a biodiversity experiment.
Nature 440:553–556.
Scheu, S. 1992. Automated measurement of the respiratory
response of soil microcompartments: active microbial bio-
mass in earthworm feaces. Soil Biology and Biochemistry 24:
1113–1118.
Scheu, S. 2001. Plants and generalist predators as mediators
between the below-ground and the above-ground system.
Basic and Applied Ecology 2:3–13.
Scheu, S., and H. Setälä. 2002. Multitrophic interactions in
decomposer food webs. Pages 223–264 in B. A. Hawkins and
T. Tscharntke, editors. Multitrophic level interactions. Cam-
bridge University Press, Cambridge, UK.
Spehn, E. M., J. Joshi, B. Schmid, J. Alphei, and C. Körner.
2000. Plant diversity effects on soil heterotrophic activity in
experimental grassland ecosystems. Plant and Soil 224:217–
230.
Van der Putten, W. H., L. E. M. Vet, J. A. Harvey, and F. L.
Wackers. 2001. Linking above-and belowground multitro-
phic interactions of plants, herbivores, pathogens, and their
antagonists. Trends in Ecology and Evolution 16:547–554.
Viketoft, M., J. Bengtsson, B. Sohlenius, M. P. Berg, O.
Petchey, C. Palmborg, and K. Huss-Danell. 2009. Long-term
effects of plant diversity and composition on soil nematode
communities in model grasslands. Ecology 90:90–99.
Wardle, D. A., R. D. Bardgett, J. N. Klironomos, H. Setälä,
W. H. Van Der Putten, and D. H. Wall. 2004. Ecological
linkages between aboveground and belowground biota.
Science 304:1629.
Weigelt, A., E. Marquard, V. M. Temperton, C. Roscher, C.
Scherber, P. N. Mwangi, S. Felten, N. Buchmann, B. Schmid,
and E. D. Schulze. 2010. The Jena Experiment: six years of
data from a grassland biodiversity experiment. Ecology 91:
930–931.
Zak, D. R., W. E. Holmes, D. C. White, A. D. Peacock, and D.
Tilman. 2003. Plant diversity, soil microbial communities,
and ecosystem function: are there any links? Ecology 84:
2042–2050.
 August 2013 NOTES 1885

Appendix B
Dendrogram representing the relationships between the 60 species present in the Jena Experiment based on functional traits
(Ecological Archives E094-170-A2).

Appendix C
Maximum clade-credibility phylogeny of the 60 species in the Jena Experiment (Ecological Archives E094-170-A3).

Appendix D
Correlation matrixes of predictors (Ecological Archives E094-170-A4).

Appendix E
A table presenting the different groups of soil biota sampled in year 2008 (Ecological Archives E094-170-A5).

Appendix F
A schematic of the maximal model used in structural equation modeling (Ecological Archives E094-170-A6).

Appendix G
Scatter plots with linear regression line for visualizing the direction of the relationship between the soil biota and the different
diversity indices (Ecological Archives E094-170-A7).

Appendix H
Model fit estimates for the minimal adequate models (Ecological Archives E094-170-A8).

Appendix I
Standardized and unstandardized maximum-likelihood estimates for the minimal adequate structural equation models
(Ecological Archives E094-170-A9).
Milcu et al. 2013

Appendices

Appendix A
Table with the means and standard deviatoins (SD) of the 12 functional traits derived from
the 60 species present in the Jena Experiment.

Appendix B
Dendrogram representing the relationships between the 60 species present in the Jena
Experiment based on functional traits.

Appendix C
Maximum clade-credibility phylogeny of the 60 species in the Jena Experiment.

Appendix D
Correlation matrixes of predictors.

Appendix E
Table with the different groups of soil biota sampled in year 2008.

Appendix F
Schematic of the maximal model used in structural equation modelling.

Appendix G
Scatter plots with linear regression line for visualising the direction of the relationship
between the soil biota and the different diversity indices

Appendix H
Model fit estimates for the minimal adequate models.

Appendix I
Standardised and unstandardized maximum-likelihood estimates for the minimal adequate
structural equation models.

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Appendix A
Means and standard deviations (SD) of the 12 functional traits derived from the 60 species
present in the Jena Experiment. Trait data were collected in monocultures and averaged for
the two sampling periods (May and August 2006, corresponding to the time of biomass
harvest) with the exception of six species, which were sampled in 2008. For details on the in-
situ collected plant trait data see Roscher et al. (2012). Literature sourced data was presented
in Roscher et al. (2004) and the data on secondary compounds in Loranger et al. (2013).

Functional trait Type (Unit) Mean SD Source

shoot biomass continous (mg dry weight) 600.75 805.49 in-situ
biomass-to-N ratio Continous (mg N g-1) 48.03 21.60 in-situ
2 -1
leaf area ratio continous (mm mg dry weight) 12.69 6.15 in-situ
seed mass continous (mg) 2.30 3.61 in-situ
hemicellulose content percentage 16.36 5.14 in-situ
lignin content percentage 8.16 4.73 in-situ
C-to-N ratio ratio 26.52 11.19 in-situ
sesonality of foliage ordinal 2.22 0.87 literature
rooting type ordinal 2.33 0.75 literature
ability to fix nitrogen binary (presence / absence) 0.20 0.40 literature
known secondary comounds counts 6.72 8.57 literature
rooting depth continous (cm) 0.70 0.27 literature

Additional literature cited

Loranger, J., S. Meyer, B. Shipley, J. Kattge, H. Kern, C. Roscher, and W. Weisser. 2013.

Predicting invertebrate herbivory from plant traits: evidence from 51 grassland species in

experimental monocultures. Ecology 93, 2674–2682.

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Appendix B
Dendrogram representing the relationships between the 60 species present in the Jena
Experiment based on functional traits. Different functional groups are colored: graminoids in
green, legumes in yellow, small herbs in red and tall herbs in blue.

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Appendix C
Maximum clade-credibility phylogeny of the 60 species in the Jena Experiment as published
in Allan et al. (2012). Different functional groups are colored: graminoids in green, legumes
in yellow, small herbs in red and tall herbs in blue. 95% confidence intervals for node ages
are shown. Congeners were used for Onobrychis viciifolia (O. montana) and for Pimpinella
major (P. saxifraga). Node support was high: 66% of nodes gave a posterior probability of 1
and a further 23% a posterior probability >0.97. Only 7 nodes were less well-supported, 4 in
the Poaceae, plus the placement of Bellis perennis (0.64), Rumex acetosa (0.68) and the node
between Cardamine pratensis and Geranium pretense (0.82).

Ranunculus acris
Ranunculus repens
Cardamine pratensis
Geranium pratense
Onobrychis viciifolia
Trifolium dubium
Trifolium campestre
Trifolium pratense
Trifolium repens
Trifolium hybridum
Trifolium fragiferum
Medicago lupulina
Medicago varia
Vicia cracca
Lathyrus pratensis
Lotus corniculatus
Sanguisorba officinalis
Rumex acetosa
Primula veris
Galium mollugo
Veronica chamaedrys
Plantago media
Plantago lanceolata
Ajuga reptans
Prunella vulgaris
Glechoma hederacea
Knautia arvensis
Campanula patula
Leucanthemum vulgare
Achillea millefolium
Bellis perennis
Crepis biennis
Tragopogon pratensis
Centaurea jacea
Cirsium oleraceum
Leontodon autumnalis
Taraxacum officinale
Leontodon hispidus
Pimpinella major
Heracleum sphondylium
Pastinaca sativa
Carum carvi
Anthriscus sylvestris
Daucus carota
Bromus hordeaceus
Bromus erectus
Anthoxanthum odoratum
Trisetum flavescens
Arrhenatherum elatius
Avenula pubescens
Phleum pratense
Alopecurus pratensis
Poa pratensis
Poa trivialis
Cynosurus cristatus
Dactylis glomerata
Festuca pratensis
Festuca rubra
Holcus lanatus
Luzula campestris

120 100 80 60 40 20 0
Branch lengths in millions of years

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Appendix D

Correlation matrix of the predictors. Acronyms: realized plant species richness (Rdiv),

biomass of legumes (legBM), biomass of grasses (grBM), biomass of small herbs (shBM),

biomass of tall herbs (thBM), functional diversity (FD), Rao’s quadratic diversity (Qr), Mean

Pairwise phylogenetic Distance (MPD) and Mean Nearest Neighbour phylogenetic Distance

(MNND).

Rdiv grBM shBM thBM legBM FD Qr MPD MNND

Rdiv 1.00 0.35 0.41 0.25 0.17 0.90 0.78 0.55 0.05
grBM 0.35 1.00 -0.11 -0.15 -0.12 0.39 0.22 -0.05 -0.26
shBM 0.41 -0.11 1.00 -0.18 0.17 0.37 0.37 0.35 0.12
thBM 0.25 -0.15 -0.18 1.00 -0.02 0.25 0.17 0.27 0.20
legBM 0.17 -0.12 0.17 -0.02 1.00 0.21 0.17 0.18 0.13
FD 0.90 0.39 0.37 0.25 0.21 1.00 0.64 0.41 -0.05
Qr 0.78 0.22 0.37 0.17 0.17 0.64 1.00 0.45 0.05
MPD 0.55 -0.05 0.35 0.27 0.18 0.41 0.45 1.00 0.82
MNND 0.05 -0.26 0.12 0.20 0.13 -0.05 0.05 0.82 1.00
Correlations of predictors marked red are significant at p < 0.05

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Appendix E
List of acronyms for the different groups of soil biota sampled in year 2008. AB and DIV refer to abundances and species richness, respectively.
Data on species richness was not available for the mesofauna groups.
Acronym Description
Cmic Microbial biomass
AMFDIV Arbuscular mycorrhizal fungal species richness
CollembolaAB Abundance of Collembola
OribatidaAB Abundance of Oribatida
GamasidaAB Abundance of Gamasida
SymphylaAB Abundance of Symphyla
TotalmesofaunaAB Total mesofauna invertebrate abundance including Collembola, Oribatida, Gamasida and Symphyla
MacrosaprotrophAB Macrofauna decomposer invertebrate abundance, including Oligocheta, Isopoda, Diplopoda, Diplura and Protura
MacrosaprotrophDIV Macrofauna decomposer invertebrate species richness
MacroherbivoreAB Macrofauna herbivore invertebrate abundance including Stylomatophora, Thysanoptera, Coleoptera, Lepidoptera,
Heteroptera, Sternorrhyncha and Auchenorrhyncha
MacroherbivoreDIV Macrofauna herbivore invertebrate species richness
MacropredatorsAB Macrofauna predator abundance including Linyphiidae, Hahniidae, Lycosidae, Tetragnathidae, Thomisidae,
Theridiidae, Dictynidae, Gnaphosidae, Chilopoda and Colleoptera
MacropredatorsDIV Macrofauna predator species richness
TotalmacrofaunaAB Total macrofauna abundance including Macrosaprotroph, Macroherbivores and Macropredators
TotalmacrofaunaDIV Total macrofauna species richness

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Appendix F

Schematic of the maximal model used in the path analysis. The maximal model included

simultaneously all measures of functional (FD and Qr) and phylogenetic (MPD and MNND)

diversity as well as any additional explanatory indices (i.e. legBM, grBM, shBM, thBM and

FG). Measurements of plant net primarily productivity (shoot, root and total plant biomass)

and soil texture have also been introduced in the model and retained in the minimal adequate

models if significant.

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Appendix G. Scatterplots with linear regression line for visualising the direction of the relationship between the soil biota and the different
diversity indices. A localy-eighted regression procedure (loess) has been also plotted for estimating a regression surface (the grey surface
alongside the regressin line) by a multivariage smooting procedure fitting a linear function of the independent variables in a moving fashion that
is analogous to how a moving average is computed for a time series. For brevity we chose to present the log species richness (Rdiv) and
functional group richness (FG) alongside the best functional diversity (FD or Qr) and phylogenetic diversity (MPD or MNND) predictors.
Several groups of soi biota where no diversity predictors are significant are not presented. * P , 0.05; ** P , 0.01; *** P , 0.001; ns, not
significant. The graphs and associated statistics were performed in R 2.15 ( R Development Core Team, 2012).

**, r2= 0.10 ***, r2= 0.17 *, r2= 0.07 ***, r2= 0.15

ns, r2= 0.00 ns, r2= 0.03 ns, r2= 0.00 *, r2= 0.05

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**, r2= 0.11 ns, r2= 0.02 *, r2= 0.07 ns, r2= 0.05

*, r2= 0.08 ns, r2= 0.00 *, r2= 0.06 ns, r2= 0.01

ns, r2= 0.03 ns, r2= 0.02 *, r2= 0.05 ns, r2= 0.03

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*, r2= 0.07

ns, r2= 0.03 ns, r2= 0.03 *, r2= 0.06

*, r2= 0.07 *, r2= 0.06 ***, r2= 0.16 **, r2= 0.09

ns, r2= 0.02 ns, r2= 0.02 *, r2= 0.06 ns, r2= 0.03

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*, r2= 0.06 ns, r2= 0.00 +, r2= 0.04 ns, r2= 0.01

+, r2= 0.04 +, r2= 0.04 ns, r2= 0.03

ns, r2= 0.02

ns, r2= 0.02 ns, r2= 0.01 ns, r2= 0.02 ns, r2= 0.02

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Appendix H

Results of structural equation modelling outputs for the most parsimonious adequate models. See Fig. 1 for the schematics of the adequate

models and Appendix F in Supporting Information for the schematic of the maximal model; AIC: Akaike information criterion; RMSEA: root

mean squared error approximation; CFI: comparative fit index; NA: not applicable. No adequate models with significant regression weights

could be obtained for GamasidAB, SymphylaAB, MacropredatorAB and MacropredatorDIV (see Appendix E for the list of acronyms). Note

that here P-values >0.05 imply that the departure of the data from the model is not significant hence, the models are considered adequate. For

OribatidaAB probability levels could not be computed as the model was saturated.

Model χ2 df P AIC* RMSEA CFI

Cmic 6.47 9 0.840 54.42 0 1
AMFDIV 0.40 3 0.939 22.40 0 1
CollembolaAB 2.52 2 0.284 26.51 0.04 1
OribatidaeAB NA 0 NA 10.00 0.10 1
TotalmesofaunaAB 0.85 1 0.357 28.00 0 1
MacrosaprotrophAB 1.74 1 0.187 27.74 0.06 0.99
MacrosaprotrophDIV 0.01 1 0.935 26.01 0 1
MacroherbivoreAB 0.24 1 0.620 16.24 0 1
MacroherbivoreDIV 0.40 1 0.525 16.40 0 1
TotalmacrofaunaAB 5.51 9 0.788 41.51 0 1
TotalmacrofaunaDIV 0.36 1 0.548 16.36 0 1
* The AIC of the maximal models varied between 250.64 and 247.49 depending on the dependent variable.

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Appendix I. Table outputs with maximum likelihood estimates of the minimum adequate

structural equation models.

Microbial biomass (Cmic)

Regression Weights:

Estimate S.E. C.R. P

FD <--- Rdiv 28.454 1.559 18.256 ***
leg <--- Rdiv .377 .129 2.924 .003
MPD <--- Rdiv 116.201 20.385 5.700 ***
shootBM <--- leg 39.867 12.304 3.240 .001
shootBM <--- FD 1.949 .466 4.185 ***
Cmic <--- MPD .000 .000 3.786 ***
Cmic <--- clay .016 .002 7.004 ***
Cmic <--- shootBM .000 .000 2.208 .027

Standardized Regression Weights:

Estimate
FD <--- Rdiv .901
leg <--- Rdiv .315
MPD <--- Rdiv .543
shootBM <--- leg .313
shootBM <--- FD .405
Cmic <--- MPD .320
Cmic <--- clay .567
Cmic <--- shootBM .186

Intercepts:

Estimate S.E. C.R. P

FD -5.301 1.169 -4.534 ***
leg .291 .097 3.009 .003
MPD 77.888 15.291 5.094 ***
shootBM 43.918 9.422 4.661 ***
clay 21.581 .397 54.391 ***
Cmic 2.527 .051 49.590 ***

Variances:

Estimate S.E. C.R. P

e1 <--> e3 10.679 4.053 2.635 .008
e1 <--> e2 -122.552 48.586 -2.522 .012

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Arbuscular mycorrhizal fungal diversity (AMFDIV)

Regression Weights:

Estimate S.E. C.R. P

MNND <--- Rdiv -4.045 21.328 -.190 .850
AMFdiv <--- clay .027 .011 2.359 .018
AMFdiv <--- MNND .001 .001 1.948 .051

Standardized Regression Weights:

Estimate
MNND <--- Rdiv -.022
AMFDIV <--- clay .258
AMFDIV <--- MNND .213

Intercepts:

Estimate S.E. C.R. P

MNND 100.757 16.296 6.183 ***
clay 21.455 .406 52.868 ***
AMFDIV 1.924 .254 7.566 ***

Variances:

Estimate S.E. C.R. P

Rdiv .171 .028 6.105 ***
e2 12.275 2.011 6.105 ***
e1 5791.766 948.761 6.105 ***
e3 .120 .020 6.105 ***

Abundance of Collembola (CollembolaAB)

Regression Weights:

Estimate S.E. C.R. P

FD <--- Rdiv 28.629 1.578 18.138 ***
rootBM <--- Rdiv 324.351 98.496 3.293 ***
rootBM <--- FD -7.174 3.103 -2.312 .021
CollembolaAB <--- rootBM .063 .024 2.671 .008

Standardized Regression Weights:

Estimate
FD <--- Rdiv .902

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Estimate
rootBM <--- Rdiv .809
rootBM <--- FD -.568
CollembolaAB <--- rootBM .294

Intercepts:

Estimate S.E. C.R. P

FD -5.308 1.179 -4.501 ***
rootBM 300.159 35.811 8.382 ***
CollembolaAB 53.573 10.515 5.095 ***

Variances:

Estimate S.E. C.R. P

Rdiv .177 .029 6.145 ***
e1 33.247 5.410 6.145 ***
e2 24172.051 3933.459 6.145 ***
e3 1197.611 194.884 6.145 ***

Abundace of Oribatid mites (OribatidaAB)

Regression Weights:

Estimate S.E. C.R. P

OribatidaAB <--- Rdiv 34.033 11.409 2.983 .003

Standardized Regression Weights:

Estimate
OribatidaAB <--- Rdiv .325

Intercepts:

Estimate S.E. C.R. P

OribatidaAB 53.720 8.524 6.302 ***

Variances:

Estimate S.E. C.R. P Label

Rdiv .177 .029 6.145 ***

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Total mesofauna invertebrate abundance (TotalmesofaunaAB)

Regression Weights:

Estimate S.E. C.R. P

FD <--- Rdiv 28.453 1.567 18.153 ***
rootBM <--- Rdiv 303.236 98.056 3.092 .002
rootBM <--- FD -6.680 3.105 -2.152 .031
TotalmesofaunaAB <--- rootBM .065 .031 2.071 .038
TotalmesofaunaAB <--- FD .744 .398 1.870 .062

Standardized Regression Weights:

Estimate
FD <--- Rdiv .901
rootBM <--- Rdiv .758
rootBM <--- FD -.528
TotalmesofaunaAB <--- rootBM .226
TotalmesofaunaAB <--- FD .204

Intercepts:

Estimate S.E. C.R. P

FD -5.272 1.178 -4.475 ***
rootBM 304.207 35.944 8.463 ***
TotalmesofaunaAB 77.756 14.065 5.528 ***

Variances:

Estimate S.E. C.R. P

Rdiv .177 .029 6.186 ***
e1 33.225 5.371 6.186 ***
e2 24506.447 3961.818 6.186 ***
e3 2082.479 336.663 6.186 ***

Total saprotroph mesofauna abundance (MacrosaprotrophAB)

Regression Weights:

Estimate S.E. C.R. P

FD <--- Rdiv 28.637 1.583 18.086 ***
rootBM <--- Rdiv 295.812 98.880 2.992 .003
rootBM <--- FD -6.535 3.108 -2.102 .036
MacrosaprotrophAB <--- Rdiv 4.794 2.075 2.311 .021
MacrosaprotrophAB <--- rootBM -.010 .005 -1.874 .061

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Standardized Regression Weights:

Estimate
FD <--- Rdiv .900
rootBM <--- Rdiv .735
rootBM <--- FD -.517
MacrosaprotrophAB <--- Rdiv .263
MacrosaprotrophAB <--- rootBM -.213

Intercepts:

Estimate S.E. C.R. P

FD -5.470 1.195 -4.576 ***
rootBM 307.712 36.684 8.388 ***
MacrosaprotrophAB 10.127 2.326 4.353 ***

Variances:

Estimate S.E. C.R. P

Rdiv .172 .028 6.186 ***
e1 33.055 5.344 6.186 ***
e2 24439.434 3950.985 6.186 ***
e3 52.609 8.505 6.186 ***

Macrosaprprotroph diversity (MacrosaprotrophDIV)

Regression Weights:

Estimate S.E. C.R. P

FD <--- Rdiv 28.637 1.583 18.086 ***
rootBM <--- Rdiv 295.812 98.880 2.992 .003
rootBM <--- FD -6.535 3.108 -2.102 .036
MacrosaprotrophDIV <--- FD .025 .008 3.088 .002
MacrosaprotrophDIV <--- rootBM -.001 .001 -2.194 .028

Standardized Regression Weights:

Estimate
FD <--- Rdiv .900
rootBM <--- Rdiv .735
rootBM <--- FD -.517
MacrosaprotrophDIV <--- FD .330
MacrosaprotrophDIV <--- rootBM -.235

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Intercepts:

Estimate S.E. C.R. P

FD -5.470 1.195 -4.576 ***
rootBM 307.712 36.684 8.388 ***
MacrosaprotrophDIV 2.236 .287 7.791 ***

Variances:

Estimate S.E. C.R. P

Rdiv .172 .028 6.186 ***
e1 33.055 5.344 6.186 ***
e2 24439.434 3950.985 6.186 ***
e3 .846 .137 6.186 ***

Macroherbivore abundance (MacroherbivoreAB)

Regression Weights:

Estimate S.E. C.R. P

Qr <--- Rdiv 8.477 .800 10.595 ***
MacroherbivoreAB <--- Qr .055 .022 2.537 .011

Standardized Regression Weights:

Estimate
Qr <--- Rdiv .771
MacroherbivoreAB <--- Qr .279

Intercepts:

Estimate S.E. C.R. P

Qr .255 .604 .423 .673
MacroherbivoreAB 1.710 .157 10.915 ***

Variances:

Estimate S.E. C.R. P

Rdiv .172 .028 6.186 ***
e1 8.438 1.364 6.186 ***
e2 .749 .121 6.186 ***

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Macroherbivore diversity (MacroherbivoreDIV)

Regression Weights:

Estimate S.E. C.R. P

Qr <--- Rdiv 8.477 .800 10.595 ***
MacroherbivoreDIV <--- Qr .047 .012 3.827 ***

Standardized Regression Weights:

Estimate
Qr <--- Rdiv .771
MacroherbivoreDIV <--- Qr .401

Intercepts:

Estimate S.E. C.R. P

Qr .255 .604 .423 .673
MacroherbivoreDIV 1.262 .089 14.112 ***

Variances:

Estimate S.E. C.R. P

Rdiv .172 .028 6.186 ***
e1 8.438 1.364 6.186 ***
e2 .244 .039 6.186 ***

Total macrofauna abundance (TotalmacrofaunaAB)

Regression Weights:

Estimate S.E. C.R. P

leg <--- Rdiv .408 .129 3.148 .002
FD <--- Rdiv 28.637 1.583 18.086 ***
shootBM <--- FD 1.945 .475 4.098 ***
shootBM <--- leg 39.982 12.542 3.188 .001
TotalmacrofaunaAB <--- shootBM .004 .003 1.651 .099
TotalmacrofaunaAB <--- clay -.127 .050 -2.537 .011

Standardized Regression Weights:

Estimate
leg <--- Rdiv .339
FD <--- Rdiv .900
shootBM <--- FD .401

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Estimate
shootBM <--- leg .312
TotalmacrofaunaAB <--- shootBM .178
TotalmacrofaunaAB <--- clay -.274

Intercepts:

Estimate S.E. C.R. P

FD -5.470 1.195 -4.576 ***
leg .262 .098 2.685 .007
shootBM 43.949 9.488 4.632 ***
clay 21.528 .398 54.029 ***
TotalmacrofaunaAB 7.223 1.115 6.475 ***

Variances:

Estimate S.E. C.R. P

Rdiv .172 .028 6.186 ***
e1 33.055 5.344 6.186 ***
e2 .221 .036 6.186 ***
e5 12.150 1.964 6.186 ***
e3 2725.403 440.600 6.186 ***
e4 2.317 .375 6.186 ***

Total macrofauna diversity (TotalmacrofaunaDIV)

Regression Weights:

Estimate S.E. C.R. P

Qr <--- Rdiv 8.477 .800 10.595 ***
TotalmacrofaunaDIV <--- Qr .033 .015 2.172 .030

Standardized Regression Weights:

Estimate
Qr <--- Rdiv .771
TotalmacrofaunaDIV <--- Qr .241

Intercepts:

Estimate S.E. C.R. P

Qr .255 .604 .423 .673
TotalmacrofaunaDIV 2.638 .111 23.772 ***

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Variances:

Estimate S.E. C.R. P

Rdiv .172 .028 6.186 ***
e1 8.438 1.364 6.186 ***
e2 .376 .061 6.186 ***

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