tor Oikos 122: 967976, 2013

di
doi: 10.1111/j.1600-0706.2012.00114.x

s
E 2012 The Authors. Oikos 2012 Nordic Society Oikos
OIKOS
Subject Editor: Richard Michalet. Accepted 20 September 2012
Ch
oice

Bottomup and topdown forces structuring consumer communities

in an experimental grassland

Michael Rzanny, Annely Kuu and Winfried Voigt

M. Rzanny (michael.rzanny@uni-jena.de) and W. Voigt, Inst. of Ecology, Friedrich Schiller Univ., DE-07743 Jena, Germany. MR also at:
Chair of Land Improvement and Nature Conservation, Technische Univ. Dresden, DE-01737 Tharandt, Germany. A. Kuu, Tartu College,
Tallinn Univ. of Technology, EE-51008 Tartu, Estonia.

The interplay between bottom-up and top-down effects is certainly a general manifestation of any changes
in both species abundances and diversity. Summary variables, such as species numbers, diversity indices or
lumped species abundances provide too limited information about highly complex ecosystems. In contrast,
Synthesis

species by species analyses of ecological communities comprising hundreds of species are inevitably only snap-
shot-like and lack generality in explaining processes within communities.
Our synthesis, based on species matrices of functional groups of all trophic levels, simplifies community
complexity to a manageable degree while retaining full species-specific information. Taking into account plant
species richness, plant biomass, soil properties and relevant spatial scales, we decompose variance of abundance in
consumer functional groups to determine the direction and the magnitude of community controlling processes.

After decades of intensive research, the relative importance of topdown and bottomup control for structuring eco-
logical communities is still a particularly disputed issue among ecologists. In our study, we determine the relative role
of bottomup and topdown forces in structuring the composition of 13 arthropod functional groups (FG) comprising
dierent trophic consumer levels. Based on species-specic plant biomass and arthropod abundance data from 50 plots
of a grassland biodiversity experiment, we quantied the proportions of bottomup and topdown forces on consumer
FG composition while taking into account direct and indirect eects of plant diversity, functional diversity, community
biomass, soil properties and spatial arrangement of these plots. Variance partitioning using partial redundancy
analysis explained 2144% of total variation in arthropod functional group composition. Plant-mediated bottomup
forces accounted for the major part of the explainable variation within the composition of all FGs. Predator-mediated
topdown forces, however, were much weaker, yet inuenced the majority of consumer FGs. Plant functional group
composition, notably legume composition, had the most important impact on virtually all consumer FGs. Compared to
plant species richness and plant functional group richness, plant community biomass explained a much higher proportion
of variation in consumer community composition.

Natural communities are typically structured in concert
by various biotic and abiotic forces (Hunter and Price
1992). Biotic forces are further subdivided into topdown
(predator controlled) and bottomup (resource controlled)
forces. Determining the relative importance of these forces
challenged ecologists for decades (Hairston et al. 1960,
Hunter and Price 1992). A number of studies have focused
on the relative roles of topdown and bottomup eects
in dierent ecological systems and from dierent perspec-
tives (Dyer and Letourneau 2003, Gruner 2004, Hoekman
2007, Gripenberg and Roslin 2007). Many experimental
studies evaluated their relative impact on the level of indi-
viduals (Kos et al. 2011), in very simple model ecosystems
(Hoekman 2010) or relied on experimental micro- or meso-
cosms (Aquilino et al. 2005). However, due to the enormous
complexity arising from multiple and parallel direct inter-
action chains, accompanied by a complex array of indirect
interactions, there is still remarkably little consensus on the
relative roles of bottomup and topdown forces on multi-
species community level (Boyer et al. 2003, Gruner 2004,
Halpern et al. 2006). Additionally, abiotic factors modify
interactions among species and so indirectly change the
strength of bottomup and topdown eects (Gripenberg
and Roslin 2007, Alberti et al. 2010, Hoekman 2010).
Biodiversityecosystem functioning research contributed
valuable insights into the question of how communities are
regulated and revealed the important role of plant diversity
on arthropod diversity in grassland ecosystems (Koricheva
et al. 2000, Haddad et al. 2001, 2009, Scherber et al. 2010,
Cardinale et al. 2011). However, these studies consider uni-
variate descriptors such as species diversity and functional
richness and neglect dierences in community composition.
Community composition refers to the number, identity
and abundance of species representing this community

967

Figure 1. Conceptual ecosystem diagram describing the hypo-
thesized and tested dependencies among the dierent ecosystem
components. Horizontal dependencies (within trophic level)
were considered for the carnivores to account for common trophic
relationships among the carnivorous functional groups. For func-
tional group abbreviations see Table 1. SR: Plant species richness,
FR: Number of plant functional groups.
(Legendre et al. 2005). One of the very few studies using
plant community composition to explain arthropod com-
position in multispecies communities showed that plant
composition is consistently the most important predictor
of community composition for several arthropod taxa
(Schaers et al. 2008). Another study aimed to disentangle
the eects of productivity, plant diversity and plant composi-
tion on arthropod abundance in mountain grasslands and
likewise found plant species composition to be the most
important predictor of arthropod abundance (Perner et al.
2005). Productivity aected arthropod abundance to a
lesser extent, while plant diversity eects were not detected.
Moreover, there is evidence from a recent experiment in
saltmarshes showing strong eects of increased plant bio-
mass production on arthropod community composition,
independent of plant species richness (Wimp et al. 2010).
Ecological communities may be perceived as systems of
functional groups (FG), interacting with each other and
with the abiotic environment (Fig. 1, Voigt et al. 2007,
Blaum et al. 2011, Rzanny and Voigt 2012). Changes in the
composition of plant functional groups, plant species
richness, functional richness, or community biomass are
expected to aect the composition of higher trophic levels,
i.e. bottomup forces. In the same way, microclimate,
depending on soil conditions, and spatial relationships
among the plots may aect arthropod composition.
Herbivorous and detritivorous species may act as food
sources for carnivorous species and, hence, will aect their
composition. The carnivores in turn will aect herbivore and
detritivore composition, in fact, exerting topdown forces
on these trophic levels.
We consider these multiple and intricate relationships
using an ecosystem scale approach covering a multitude of
factors potentially aecting arthropod composition at dier-
ent spatio-temporal scales (Fig. 1). We aim to resolve the
inconsistent relationships revealed by dierent studies
968
analyzing the role of plant species richness, functional rich-
ness, and plant biomass for the composition of arthropod
functional groups and compare the impact of these variables
with the impact of the composition of functional groups
from all trophic levels. To this end, we used data from a
controlled biodiversity experiment (The Jena experiment)
minimizing environmental gradients inherent to previous
observational approaches.
Specically we asked: 1) which abiotic or biotic variable
groups best explain the composition of dierent functional
groups in arthropod communities? 2) how is the composition
of specic arthropod functional groups aected by the com-
position of the functional groups of other trophic levels, i.e. to
what extent do bottomup or topdown forces inuence the
composition of specic consumer functional groups? 3) is the
composition of dierent arthropod functional groups aected
by plant species richness, plant functional group richness and
community biomass and to what extent are these eects inde-
pendent from the eects of plant community composition?
Material and methods
Experimental design the Jena experiment
All data were collected in 2005 on 50 grassland plots
(each 400 m) of the Jena experiment (detailed experimen-
tal design in Roscher et al. 2004). The experimental plots are
located in the oodplain of the river Saale at the northern
edge of Jena, Germany (5055N, 1135E; 135 m a.s.l.).
The plots were established from seeds on a former arable
eld in spring 2002. To maintain target plant species
composition, the plots were manually weeded (April and
July) and mown twice a year (June and September), and
remained unfertilized during the period of data collection.
The particular plant species mixtures were randomly selected
from a species pool comprising 60 plant species typically
occurring in semi-natural mesophillic grasslands in the
surrounding region. We used 16 monocultures, 16 four-
species mixtures, 14 sixteen-species mixtures and four
60-species mixtures for our analyses.
All plant species were a priori assigned to functional
groups: grasses (PGR, 16 species), legumes (PLG, 12 species),
tall herbs (PTH, 20 species) and small herbs (PSH, 12 spe-
cies), based on the results of a cluster analysis comprising
morphological, phenological and physiological traits (see
Roscher et al. 2004 for details). Species richness and the num-
ber of functional groups per plot (functional richness) were
varied as independently as possible (Roscher et al. 2004).
To take into account possible abiotic spatial patterns,
soil texture was determined using standard methods before
setting up the experiment (Roscher et al. 2004) and soil
moisture (gravimetric water content) was measured in
015 cm depth (Oelmann et al. 2011).
Plant sampling
Aboveground plant biomass was harvested between 30 May
and 8 June and again between 22 and 31 August (Weigelt
et al. 2010). This was done by clipping the vegetation 3 cm
above ground on four rectangles (each 0.20  0.50 cm)
positioned on predened locations. The coordinates of
these positions were assigned randomly prior to the harvest
in the inner area of each plot (at least three meters away
from the plot margins). Plant biomass was sorted to species
level and dried at 70C for 48 h. The dry weight in g m2
of each species was calculated as the mean of the four
subsamples per plot and was taken as a measure for plant
community biomass per plot (Weigelt et al. 2010).
Arthropod sampling
Arthropod abundance was estimated using two dierent
techniques. Two pitfall traps (4.5 cm diameter) were
placed near the centre of each plot to estimate activity
density of epigeic, ground-associated species. The traps were
emptied every three weeks during the vegetation period
(altogether six times). Only during the mowing periods, the
traps were closed for about two weeks. Population density of
vegetation-associated arthropods was measured using an
electric suction sampler in combination with a biocenometer
(bottomless, cube-shaped covering of gauze) covering an
area of 0.75  0.75 m. This method allows volume-specic
sampling and yields highly comparable arthropod abun-
dance data. Samples were taken on ve occasions from May
to October. The following taxa were sampled: Diptera,
Hymenoptera, Coleoptera, Araneae, Opiliones, Heteroptera,
Auchenorrhyncha and Orthoptera, representing a major
component of the entire aboveground fauna in these
grasslands. With the exception of some predominantly
parasitoid Hymenoptera, which were in most cases deter-
mined to the level of family or subfamily, all arthropods
were identied to species level. Only taxa with at least ten
sampled individuals were included in the analysis.
All arthropod species were assigned to functional
groups (Table 1) based on cluster analysis of feeding and life
history traits such as body mass, trophic level, feeding
strategy, degree of specialization or aerial mobility (Rzanny
and Voigt 2012). These functional groups reect the
basic trophic community organization in herbivorous, detri-
tivorous and carnivorous functional groups (Fig. 1). The
herbivorous functional groups comprise plant suckers
(HSU), seed predators (HSP), root feeders (HRF), special-
ized chewers (HSC) and endophytics (HEP). The detriti-
vores are subdivided into endophytic saprophages (DES)
and soil living saprophages (DSS). The carnivore trophic
level is formed by parasitoids (CPO), vegetation associated
suckers (CVS), ground living chewers (CGC), web spiders
(CWS), ground living suckers (CGS) and omnivores
(OMG) (Table 1). In combination with the plant func-
tional groups, which represent the producer trophic level,
we comprehend the whole community as a system of inter-
dependent functional groups (Fig. 1).
Spatial patterns
Arthropod composition may be aected by spatial patterns
across a variety of scales, ranging from broad geographical
gradients to ne scaled autocorrelation patterns. To account
for this, we used principal coordinates of neighbour matrices
(PCNM, Borcard and Legendre 2002, Borcard et al. 2004)
to describe the spatial relationships among the plots. This
method allows extracting spatial patterns across the whole
range of all perceivable scales of a given spatial structure. The
resulting orthogonal eigenvectors could be used as additive,
scale specic spatial descriptors in a canonical analysis.
PCNM analysis is based on the computation of princi-
pal coordinates of a truncated Euclidean distance matrix
describing the spatial relationships among the plots. The
geographical positions of each plot centre were used to calcu-
late this distance matrix. The resulting PCNM axes were
named following their order (PCNM 1-PCNM 30) and
used as independent spatial variables.
Data analysis
All data were divided into ve variable groups (Fig. 2).
Each functional group is represented as a species-by-plot
matrix containing the biomass (plants) or abundance
(arthropods) of each species (Fig. 2). The biomass-matrices
of the plant functional groups together with the experi-
mentally manipulated variables plant species richness (SR,
number of established plant species) and plant functional

Table 1. Functional group codes including species richness and total abundance of arthropods considered for the analysis.

Trophic level Code Functional group Important taxa No. of taxa Total abundance
Plants PSH small herbs misc. 12
PGR grasses Poaceae 16
PTH tall herbs misc. 19
PLG legumes Fabaceae 12
Herbivores HSU plant suckers Auchenorrhyncha, Heteroptera 44 9550
HSP seed predators Carabidae 25 4072
HRF root feeders Coleoptera 16 15 690
HSC specialized chewers Coleoptera 20 5076
HEP endophytics Diptera 26 2930
Carnivores CPO parasitic species Hymenoptera, Diptera 35 8196
CVS vegetation associated suckers Diptera 11 1580
CGC ground living chewers Carabidae, Staphylinidae 25 2313
CWS web spiders Araneae 33 4860
CGS ground living suckers Araneae 29 11 225
OMG omnivores misc. 28 12 514
Detritivores DES endophytic saprophages Diptera 5 382
DSS soil living saprophages Diptera, Coleoptera 25 3270

969
Figure 2. Overview of the analysis steps and derivation of the ve variable groups. Species-by-plot matrices of the plant functional
groups contain species-specic plant biomass, the species by plot matrices contain species abundance data. The variable groups plants,
carnivores, spatial patterns and soil were used as predictor variables for all consumer functional groups, while herbivore functional group
composition was a used as predictor variable for only the carnivore functional groups.

richness (FR, number of established plant functional groups)
and complemented by total aboveground plant biomass
(biomass) formed the predictor variable group plants. The
herbivore and detritivore functional group matrices together
formed the predictor variable group herbivores. They were
combined in this group because both are placed as an inter-
mediate level between producers and carnivores (Fig. 1).
Both act as food source for the carnivores and at the same
time consume dead or living plant biomass. Carnivore func-
tional group abundance matrices were considered as carni-
vores. All PCNMs with positive eigenvalues entered the
analysis as spatial patterns. The soil variables form the last
variable group soil. All variable groups entered the analysis
separately and all groups were independently subjected to
variable selection in order to determine the overlapping parts
of variation accounted for by these dierent groups.
The consumer communities are allowed to colonize the
plots freely from the surrounding area and are subject
to environmental lters and particular dispersal abilities of
the component species, which may result in positive or neg-
ative spatial autocorrelation patterns. In contrast, the plant
composition of each plot is determined by the experimental
design and any species, which is not part of this planned
composition, is weeded out from the respective plot. There-
fore, we explicitly did not consider any eects
on the plant communities, as our experimental design is
not suitable to detect such eects (Fig. 1). As a rst step, all
variable groups had to be processed in dierent ways to
make them suitable for variance partitioning (Fig. 2). The
dierent data sets were log-transformed (all functional
groups matrices) or standardized by adjustment to stan-
dard deviation (experimental variables, soil variables). The
functional group matrices were subsequently Hellinger-
transformed (Legendre and Gallagher 2001) to render
the species data suitable for redundancy analysis. All func-
tional group matrices were condensed to linear combina-
tions using principal component analysis (PCA). However,
we considered dierent numbers of ordination axes in
the plant and consumer functional groups. Each plant
functional group was reduced to six PCA-axes to ensure
that they comprise all potentially important information.
When all variables forming these groups were analyzed
simultaneously in a global redundancy analysis, extracting
more than six axes substantially increased multicollinear-
ity among the predictor plant functional groups (e.g.
highest variance ination factor [VIF] when 6 axes per
functional group were used: VIF  2.9). The same proce-
dure was applied to consumer functional groups. However,
the rst PCA-axes of the consumer functional groups
showed particularly strong correlations (r  0.7) resulting in
increased variance ination (for three axes per functional
group maximum VIF  8.9). Data exploration showed that

970
the rst axes were always strongly correlated to the commu-
nity biomass of the plots. Multicollinear predictor variables
within a variable group would reduce the interpretability
of the variable selection results. The selection of a correlated
variable may obscure the causal eect variable within a
group of variables (Graham 2003) and therefore, impede
the potential to reveal mechanistic explanations. Therefore,
we statistically controlled for the eect of community
biomass on the PCA scores of all consumer functional
groups through the calculation of partial principal compo-
nent analysis (pPCA), i.e. community biomass was
included as a covariable when calculating the PCA scores.
In doing so, we eectively partial out the eect of biomass
on the composition of the consumer functional groups, which
we intend to use as explanatory variables. On the other hand,
the eect of community biomass on functional group com-
position enters the analysis separately via the variable group
plants. These adjusted pPCA-axes generally show lower cor-
relations among each other (maximum VIF  5.4) and nota-
bly reduces correlations among the rst axes. However,
including more than three pPCA axes per functional group
would again increase multicollinearity to critical levels.
To reduce the possibility that important information is
missed by only considering three (consumers) or six (plants)
composition axes, all species within one trophic level
(i.e. without distinction in functional groups) were used to
calculate as many pPCA (or PCA in the case of plants) axes
as necessary to explain at least 80% of the total variation
within this trophic level. This set of axes was always analyzed
in the same way as the FG-resolved dataset to ensure that
no important information was lost through the inadequate
representation of variation in the ordination axes.
In the second step, all variable groups were subjected to a
forward selection procedure (Blanchet et al. 2008) except
the experimentally manipulated variables SR and FR as
well as biomass, because we wanted to determine and
compare their direct impact, relative to the other variables for
each functional group. This forward selection procedure
involved two stopping criteria, which has been shown to
greatly reduce type I error and reports reliable portions of
explained variance, two severe problems inherent to the classi-
cal approach (Blanchet et al. 2008). Prior to the forward selec-
tion, a global test (redundancy analysis) of the entire variable
group on the response functional group was performed. Only
when this test was signicant, we proceeded with forward
selection. Selection of variables was stopped when either the
contribution of the included variable was not signicant or the
explained variation, through the selected subset, exceeded the
variation accounted for by the entire subset.
As a nal step, the subsets of the selected variables were
used for the partitioning of variation using redundancy
analysis (Fig. 2, Legendre et al. 2005, Peres-Neto et al.
2006). We then calculated the unique and shared eects of
the dierent subsets on each functional group response
matrix and tested the unique fraction for statistical signi-
cance using a permutation test with 9999 permutations
(function anova.cca in package vegan, Oksanen et al. 2010).
Here, we tested the unique eect of the particular variable
group, independent of the eect of any other variables
considered in the variation partitioning. Finally, the inde-
pendent and shared eects of all selected functional
Table 2. Structure of the different pRDA-models explaining the inde-
pendent fraction of variation in the composition of (a) herbivorous/
detritivorous functional groups (FG) and (b) carnivorous functional
groups. All respective variables were only included in the pRDA
if they had a signicant effect on the particular functional group
composition. Abbreviations: SR: plant species richness; FR: plant
functional richness.
Predictor variables Control; effects partialled out in pRDA
(a) Herbivorous/detritivorous functional groups
Plant FG carnivore composition, soil structure, spatial
composition patterns
Biomass, SR, FR plant composition, carnivore composition,
soil structure, spatial patterns
Carnivore FG plant composition, soil structure, spatial
composition patterns
Spatial patterns plant composition, carnivore composition,
soil structure
Soil structure plant composition, carnivore composition,
spatial patterns
(b) Carnivorous functional groups
Plant FG carnivore composition, soil, spatial patterns
composition
Biomass, SR, FR plant composition, herbivore composition,
carnivore composition, soil structure,
spatial patterns
Carnivore FG plant composition, carnivore composition,
composition herbivore composition, soil structure,
spatial patterns
Herbivore FG plant composition, carnivore composition,
composition soil structure, spatial patterns
Spatial patterns plant composition, carnivore composition,
soil structure
Soil structure plant composition, carnivore composition,
spatial patterns
group compositions and the selected abiotic variable groups
were determined separately through incorporating all
signicant variables in turn as covariables in the pRDA
(Table 2). Independent and shared eects of plant com-
position species richness, plant functional richness and
community biomass were separately determined for each
consumer functional group. The composition of carnivore
functional groups was used as a potential predictor for
herbivore functional group composition and all con-
sumer functional groups (excluding the respective response
functional group from the predictor variable set) were used
as predictors for the carnivore functional groups (Table 2).
The variable groups plants, spatial patterns and soil
were exclusively used in an explanatory role, while the com-
munity composition of the arthropod functional groups were
used both as predictor and explanatory variables (Table 2).
The amount of explained variance depends, to a certain
degree, on the number of species within this group. In
order to compare the relative importance of the dierent
fractions, we always show and compare these fractions rela-
tive to the total amount of variation explained by all
variables together. We report unbiased adjusted R values
; Peres-Neto et al. 2006), which takes the number of
(R adj
included variables and the sample size into account and
prevents an overestimation of the fraction of explained
may take negative values if a variable explains
variance. Radj
less variation than a random variable would explain. In
this case, the fraction of explained variance was set to zero
(Peres-Neto et al. 2006).
971
 Our approach is essentially correlative. The identied vari-
ance partitions, together with their associated p-values, repre-
sent estimates as observational evidence that a certain source
of variation is important in structuring the composition of a
certain arthropod functional group. The variance partitioning
approach used here does not explicitly address trophic cascades
between the functional groups of adjacent trophic levels.
Instead, it quanties the strength and direction of dependency
between two functional groups. Species are simultaneously
subjected to topdown and bottomup forces (Hunter and
Price 1992). This idea is taken into account by using species
composition matrices as explanatory variables (Legendre and
Gallagher 2001, Halpern et al. 2006) and considering them
reciprocally as either dependent or independent variables. In
contrast to lumped species abundance commonly used in uni-
variate approaches (Koricheva et al. 2000, Haddad et al. 2001,
Scherber et al. 2010), the composition of certain functional
groups at dierent trophic levels represents a much more
detailed measure of heterogeneity between the plots. We gen-
erally did not account for multiple testing because of the
exploratory character of this study and following the mathe-
matical and logical arguments of Moran (2003).
All statistical analyses have been performed using the
statistical software R 2.12.0 (R Development Core Team)
using the packages vegan (Oksanen et al. 2010), and packfor
(Dray et al. 2009).
Results
In total, 81 658 individuals belonging to 322 arthropod taxa
were considered for the analysis. Root feeders, with 15 690
sampled individuals, formed the most abundant functional
group, while detritivorous endophytics with 382 sampled
Figure 3. Relative fraction of variance explained (R adj) by the dierent variable groups for all functional groups. The small black bars at
the bottom represent the absolute total explainable variance for each functional group (axis title and scale at the bottom right).
972
individuals were the functional group with the lowest total
abundance (Table 1). The pPCA axes used as explanatory vari-
ables accounted for between 72% and 88% (six axes; plants)
and between 39% and 84% (three axes; consumers) of the
original variation of each functional group. Using variance par-
titioning, between 20.7% (CWS) and 43.6% (DES) of the
total variation in FG composition could be explained (Fig. 3).
Effects of the variable groups on the composition of
functional groups
The variable group plants was generally the most impor-
tant variable group for the species composition of all con-
sumer functional groups (Fig. 3). Plants explain a highly
signicant fraction of variance in each functional group.
However, strong dierences in variance attributed to these
relative fractions existed between the dierent functional
groups. The average relative proportion of variation in her-
bivore functional group composition due to plants was
0.58  0.059 (arithmetic mean  SE), for the detritivore
functional groups 0.62  0.109 and 0.49  0.042 for the
carnivore functional groups.
The composition of herbivore/detritivore functional
groups had signicant, unique eects on parasitoids and
vegetation associated suckers (Fig. 3) and some rather
small and insignicant eects on web spiders, omnivores and
ground associated chewers.
The variable group carnivores explained signicant pro-
portions of variation in the composition of most herbivore
and detritivore functional groups and to a lesser extend of
some carnivore functional groups (Fig. 3). The average rela-
tive proportion of variation explained by carnivore functional
groups in the composition of the herbivore and detritivore
functional groups was 0.16  0.046 and 0.07  0.031 in the
carnivore functional groups. Again, there are strong dier-
ences among the dierent functional groups.
The PCNM scores, signicantly aected the composition of
ground associated suckers, web spiders and root feeders, while
these eects were shared with other sources of variation in seed
predators and omnivores. PCNM 7 and PCNM 1 were selected
as signicant predictors in three functional groups (OMG,
HRF, CGS and HSP, HRR, CGS), followed by PCNM 2,
which was selected two times (CWS, CGS). PCNM1 and
PCNM2 relate to broad-scale spatial gradients. PCNM7 repre-
sents a circular spatial pattern with similar plots within the outer
margins and the interior plots of the experimental area, but
great dierences between the outer and interior plots of the
experimental eld site (Supplementary material Appendix 1).
Soil explained independently signicant fractions of
variation in the detritivorous functional groups (DSS:
0.19; DES: 0.10) and also aected some functional groups
of the other trophic levels (Fig. 3).
Compositional effects among functional groups
Legumes represent the most important functional group,
explaining the largest, highly signicant independent frac-
tion of total explainable variance in composition through-
out all consumer functional groups (Fig. 4, Supplementary
material Appendix 2). Here, the relative fractions of explained
variance range from 0.16 (HSU) to 0.45 (OMG). Legume
composition explains an average fraction of 0.22  0.030 of
the total explainable herbivore functional group variation
and a markedly higher relative fraction in the detritivore
functional groups (0.37  0.006) and the carnivore func-
tional groups (0.34  0.037). The composition of tall herbs
predict high and signicant fractions of variance in all her-
bivorous functional groups (average: 0.16  0.012) but less
so in only some carnivorous functional groups (0.027  0.014)
and in one of the two detritivorous functional groups (0.041)
(Fig. 4). Grass composition explained signicant indepen-
dent fractions of variance for most functional groups, with
the largest eect on plant suckers. A similar pattern appears
for the small herbs, which show highly signicant impacts
on some functional groups, but none on others (Fig. 4).
The herbivore functional groups explain much less
variance of carnivore composition compared to the plant
functional groups. They show strong uctuations regarding
their predictive power for the carnivores (Fig. 4). While
the endophytics explain signicant variations in the compo-
sition of parasitoids and vegetation associated suckers,
the specialized chewers were not selected in any of the for-
ward selection procedures. The compositions of plant suck-
ers and soil living saprophages account for a signicant
proportion of parasitoid composition.
The carnivore functional groups, again, show very dier-
ent and specic contributions to the explained variance in
the response functional groups (Fig. 4). However, there are
only two cases where the composition of carnivore functional
groups explains a signicant fraction of variation in another
carnivore functional group. In contrast, all carnivore func-
tional groups explain a signicant proportion of variation in
the composition of all but one herbivore functional group.
Soil living saprophages is the only detritivorous functional
group which is aected by a carnivore functional group (CPO).
Effects of plant species richness, functional richness
and community biomass
Community biomass, plant species richness and plant func-
tional richness together explained signicant proportions
of variance in the composition of all arthropod functional
groups (except FR in DES; Fig. 4). Community biomass
accounted for a higher relative fraction of variance in commu-
nity composition of all consumer functional groups when
compared to plant species richness and functional richness.
The relative variation, attributed to community biomass, shares
a large fraction of explained variation with other variables and
has a signicant, independent eect on the composition of two
carnivorous functional groups: web spiders and vegetation
associated suckers. Considering the sum of the independent
and the shared fraction, results in high proportions of variance
explained in detritivorous (0.34  0.003) and carnivorous
functional groups (0.33  0.073) and only about the half in
herbivorous functional groups (0.16  0.011).
Plant species richness showed a signicant, independent
eect on only root feeders. Functional richness on the other
hand, showed a signicant, independent eect on the vegeta-
tion associated suckers (Fig. 4). Nonetheless, both variables
share a large part of variation with other predictors. Jointly with
the shared fraction, species richness explained an average pro-
portion of 0.12  0.021 in the herbivore functional group
composition, 0.18  0.027 in detritivore functional group
composition and 0.16  0.025 in carnivore functional group
composition. Plant functional richness explains 0.11  0.017 in
herbivore functional groups, 0.10  0.065 in detritivore func-
tional groups and 0.18  0.068 carnivore functional groups
when the shared fractions of variance are taken into account.
Discussion
Our results provide the rst general overview of the
relative importance of dierent biotic and abiotic forces in
determining multitrophic functional group composition in
experimental grassland communities. We could experimen-
tally support the view that plants (i.e. bottomup forces)
predominantly determine consumer community composi-
tion of various trophic levels (White 1978, Halaj and Wise
2001, Kagata and Ohgushi 2006, Randlkofer et al. 2010).
Contributions of different variable groups to the
composition of consumer functional groups
Our approach allows the disentanglement of the indepen-
dent and the shared eects of dierent variable groups. We
considered and quantied abiotic and spatial heterogeneity
and showed that they play a minor role in structuring most
consumer functional groups on the scale of our designed
experiment. The results show that although bottomup
eects account for a major proportion of variation in the
composition of consumers from all trophic levels, likewise,
topdown forces exerted by carnivores have a signicant
impact on most herbivorous functional groups (Hunter and
Price 1992, Gruner 2004, Hoekman 2007).
Some recent studies found no support for the presence
of topdown forces on consumer groups in experimental
973
Figure 4. Relative fraction of variance (R adj) in consumer functional group composition explained by the dierent predictor variables
(plant functional group composition, plant biomass (biomass), plant species richness (SR), number of plant functional groups (FR ),
herbivore functional group composition (only for carnivore functional groups) and carnivore functional groups). Dark colored fractions
of the bars represent independent eects, light colored fractions represent shared eects. Consumer functional groups, independently
and signicantly (p  0.05) aected by the respective variable are marked by an asterisk.

grasslands (Scherber et al. 2010), whereas, earlier experi-
mental and observational studies corroborated the presence
of topdown forces structuring herbivore communities in
grasslands (Koricheva et al. 2000, Gruner 2004, Perner
et al. 2005, Borer et al. 2006). Using path analysis, Koricheva
et al. (2000) found that herbivore abundance was more
strongly aected by parasitoid abundance than by plant
diversity, indicating topdown forces acting on the herbi-
vores. A eld study, conducted in montane European
grasslands, found that the highest proportion of variation
in herbivores was explained by the (summed) abundances
of carnivorous functional groups (Perner et al. 2005), like-
wise indicating the presence of topdown forces. Unlike
the study of Koricheva et al. (2000) and Scherber et al.
(2010), we used the exact species composition of trait-based
functional groups as response variable in a multivariate
framework. This oers a higher resolution of the relation-
ships among functionally dierent species groups. Indeed,
there are marked dierences in the response of functional
groups (Fig. 2, 3). Some functional groups showed no
signicant response to the composition of carnivore func-
tional groups (HRF, DES, CVS, OMG) indicating limited
topdown forces exerted on them. In contrast, carnivore
composition explained about 31% of the total explained
variation of seed predators and about 25% of the total
explained variation of plant suckers (Fig. 3) and hence imply
the presence of topdown forces. As indirect plant biomass
eects on carnivores are accounted for by pPCA-axes and
other potential common sources of variation are fully con-
sidered using the variance partitioning approach, these
eects are direct and not mediated by correlations with
other variables.

974
 In contrast to a previous study by Crist et al. (2006),
spatial variation played a comparably minor role in structur-
ing the consumer communities (Fig. 3). These dierences
were likely caused by methodological dierences and the
larger plot size of 20  20 m in our experiment. In our
analysis, the global test of all PCNMs excludes the spatial
predictor variable set in many cases. The selected eigen-
vectors PCNM 1 and PCNM2 represent large-scale spatial
gradients (Supplementary material Appendix 2). PCNM7
may reect an invasion pattern, showing the colonization
process from the surrounding meadows is, at least for three
functional groups (CGS, OMG, HRF), still in progress.
However, the nearly complete absence of signicant ne-
scale patterns indicates that the large plot size of 400 m
ensures the development of separate communities on each
plot with autocorrelation playing a minor role in structuring
the consumer communities (Bommarco and Banks 2003).
The impact of soil variables is most pronounced for
both detritivorous functional groups. This was expected and
is in line with Perner et al. (2005) who identied soil and
management variables together with plant composition as
the most important predictor for the composition of detri-
tivorous functional groups.
The importance of legumes
The importance of legumes in structuring consumer species
composition and interactions has been stressed earlier
(Koricheva et al. 2000, Spehn et al. 2002, Scherber et al.
2006, Chen et al. 2010). Koricheva et al. (2000) found
the presence of legumes to be the major determinant of
abundance for various arthropod groups in experimental
grasslands and stated that the eects of plant diversity on
consumer species were mostly indirect and mediated
via plant biomass. The presence of legumes and plant
diversity are inevitably linked to plant community biomass
(Spehn et al. 2002, Cardinale et al. 2011), but the relation-
ship between these factors and the mechanisms causing com-
munity response to changes in plant composition are dicult
to disentangle. In our analysis, this relationship is reected in
a strong correlation between the rst ordination axis, repre-
senting the strongest gradient in legume composition and
plant community biomass (Pearsons correlation r  0.78).
This strong relationship mirrors the interdependence of these
two variables and renders impossible a statistical separation
of their relative contributions to changes in consumer func-
tional group composition. However, it has been shown
experimentally that increased primary production alone may
change the structure and composition of terrestrial arthro-
pod communities in monocultures (Wimp et al. 2010).
The role of plant species richness, plant functional
richness and plant biomass in relation to functional
group composition
Plant species richness and plant functional richness are
weaker predictors of consumer functional group composi-
tion in relation to plant composition. Legume composition
alone explains more variation in the composition of each
functional group than species richness or functional richness
(Fig. 4). Also taking into account the amount of explained
variance of the remaining plant functional groups, it indicates
that arthropods from all trophic levels are much more inu-
enced by species composition than by species diversity. This is
in line with other studies (Perner et al. 2005, Beals 2006,
Schaers et al. 2008, Vehvilinen et al. 2008, Zhang and
Adams 2011) and emphasizes that interactions between plants
and consumers are determined primarily by the identity and
abundance of particular species (Randlkofer et al. 2010).
Plant biomass has stronger eects for most functional
groups rather than plant species richness and plant functional
richness (Fig. 4). Interestingly, Scherber et al. (2010) found no
signicant eect of plant biomass on any considered consumer
group, while plant species richness directly aected plant bio-
mass and herbivore abundance. These authors considered
summed abundances of three trophically distinct groups,
while we used thirteen species-resolved functional group
abundance matrices as response variables. This raises the ques-
tion of the appropriate scale to tackle such questions and indi-
cates that strong simplications may obscure eects that are
present on a ner scale. However, using a very similar approach
to the study of Scherber et al. (2010), Koricheva et al. (2000)
found plant diversity eects on arthropod abundance being
mostly indirectly mediated by changes in plant biomass.
Conclusions
Plant composition is by far the most important predictor
for the composition of all consumer functional groups irre-
spective of their trophic level. Plant species composition
integrates several abiotic and biotic environmental dier-
ences among ecological communities (Schaers et al. 2008)
and hence is a very sensitive ecosystem descriptor. We
strongly encourage ecologists to consider community com-
position when attempting to interpret the relative role of the
dierent forces structuring ecological communities. Our
results imply that invertebrate communities are mainly
structured by bottomup forces, while additionally, top
down forces aect nearly all herbivorous functional groups.
We quantied and directly compared the impact of plant spe-
cies richness, functional richness and community biomass on
the composition of functional rather than taxonomic groups.
Our results reveal that community biomass is consistently
more decisive for the species composition of all consumer
functional groups than it is plant species richness per se.
Acknowledgements We thank all colleagues and people who
conceived and maintained the Jena Experiment and provided the
basic plant and soil texture data, in particular C. Roscher, A. Weigelt
and J. Baade. We are grateful to R. Mller who has a share in
the eld data acquisition and kindly contributed data for some
arthropod taxa, and Y. Oelmann and W. Wilcke who provided
data on soil moisture. The Jena Experiment has been funded by
the Deutsche Forschungsgemeinschaft (DFG, Wi 1601/4-1,2,
FOR 456). MR acknowledges funding by the German Federal
Foundation for Environment (DBU). We are very grateful to
suggestions from J. Engel, M. F. Gizzie-Voigt and C. Mulder
which substantially improved this manuscript.
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