DOI: 10.1111/j.1365-3180.2012.00941.

Trait-based approaches to unravelling the assembly

of weed communities and their impact on
agro-ecosystem functioning
M-L NAVAS
Montpellier SupAgro, Centre dEcologie Fonctionnelle et Evolutive (UMR 5175), Montpellier Cedex 5, France

Received 21 December 2011

Revised version accepted 28 June 2012
Subject Editor: Jonathan Storkey, Rothamsted Research, UK

and the role thereof in the assembly of weed commu-

Summary
The trait-based approach to plant functional ecology
has gained considerable attention over the last two
decades, allowing ecologists to address questions relat-
ing to species distribution, community assembly and
ecosystem functioning. We show here how this approach
can be used to address these issues for weed ecology in a
new way, allowing research to shift from purely weed
control issues to a more global understanding of the
impact of weed communities on the agro-ecosystem. We
review how weed species are sorted by environmental
factors and management according to the value of traits
nities. How weed trait values and their distribution
within communities aect agro-ecosystem processes is
discussed in relation to loss of crop production. We also
introduce the question of the impact of weed functional
structure on ecosystem services and suggest some
directions for research at species, community and
agro-ecosystem levels.
Keywords: agroecology, biodiversity, community struc-
ture, ecosystem processes, functional structure, plant
functional trait.

NAVAS M-L (2012). Trait-based approaches to unravelling the assembly of weed communities and their impact on
agro-ecosystem functioning. Weed Research 52, 479488.

species (Marshall et al., 2003; Petit et al., 2010). As a

Introduction
Weed research is facing new challenges because of
changes in agricultural management, climate and policy
(Fernandez-Quintanilla et al., 2008; Petit et al., 2010).
A priority in meeting these challenges is to take into
account changes in crop-weed oras, for example the
highly specialised ora in intensively managed crops
may shift to a highly diverse communities in places
where herbicide use is reduced (Aubertot et al., 2007;
Wezel et al., 2009). Moreover, the negative perception of
crop weeds that was universal until recently is being
challenged by the recognition of the positive impact
weeds can have on local biodiversity and ecosystem
functioning, and also of the conservation value of some
consequence, weed research needs to shift in emphasis
from purely weed control issues, based on the popula-
tion dynamics of a few major species, to a more global
understanding of the impact of weed communities on
the agro-ecosystem.
Shifting weed research from the population level to
communities and ecosystems requires a change in
scientic perspective, because the objectives and theo-
retical frameworks of population biology and commu-
nity ecology are clearly distinct. To characterise complex
communities, weed ecology must revisit issues of com-
munity ecology, such as the identication of assembly
rules of communities, that is, the processes related to the
co-existence of species, the recognition of groups of

Correspondence: Marie-Laure Navas, Montpellier SupAgro, Centre dEcologie Fonctionnelle et Evolutive (UMR 5175), 1919 route de Mende, 34293
Montpellier Cedex 5, France. Tel: (+33) 499 61 24 57; Fax: (+33) 499 61 24 26; E-mail: navas@supagro.inra.fr

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Weed Research  2012 European Weed Research Society Weed Research 52, 479488
480 M-L Navas

species that respond similarly to a set of environmental
conditions or management practices, or that similarly
aect the ecosystem (Booth & Swanton, 2002; Weiher
et al., 2011). Adapting these questions to weed ecology
is not straightforward because of the specic character-
istics of weed communities compared with other types of
vegetation (Cousens & Mortimer, 1995; Booth et al.,
2003). Weeds are highly adaptive organisms, because of
the large unpredictability of cultivated areas that
strongly interact with crop plants, and can recover from
the seed bank or through vigorous vegetative production
after destruction of plants by agricultural practices.
They form communities characterised by a fast turnover,
whose structure depends on current conditions and the
legacy of previous land use. These characteristics suggest
that both the ways in which the environment and the
crop conditions control weeds need to be taken into
account when analysing the structure of weed commu-
nities and their impact on the functioning of the agro-
ecosystem. The aim of this article is to show that trait-
based approaches give the opportunity to address these
new questions for weed ecology.
Trait-based approaches are built on the idea that a
trait is measured at the level of an individual organism
(see McGill et al., 2006; Lavorel et al., 2007), with the
following denition: any morphological, physiological
or phenological feature measurable at the individual
level, from the cell to the whole-organism level (Violle
et al., 2007). The cornerstone of the approach is the
responseeect framework proposed by Lavorel and
Garnier (2002) and further rened by Suding et al.
(2008): environmental drivers act as lters sorting
species according to the value of traits (so-called
response traits), which results in a functional diversity
of communities depending on the type and strength of
these lters. In turn, that functional diversity of com-
munities, assessed by their distribution of traits, has
various impacts (via so-called eect traits) on ecosystem
processes and services (Diaz et al., 2007b). The use of
this framework has led to universal, mechanistically
sound assessments of community structure and func-
tioning, explaining their increasing use for a large range
of situations and organisms (for a review of recent
papers see Garnier & Navas, 2012). A simplied scheme
of this framework incorporating services provided by
weeds is presented in Fig. 1. Environmental drivers
relate to local conditions and crop management that act
on the functional structure of weeds; agro-ecosystem
processes are related to crop production, trophic webs
and resource cycles, whereas agro-ecosystem services

Environmental drivers
1 Local conditions :
o Climate, soil and history
Crop abiotic factors:
o Crop type (sowing date,
herbicide, fertilization)
o Preceding crop type
o Tillage (date, depth)

2
Agro-ecosystem services
Shelter and food for non pests 5 Functional
(including pollinators)
Parasite trapping structure of weeds 3
Soil cover Distribution of traits
Biodiversity conservation
Cultural services

4
Agro-ecosystem processes
Biomass production
Trophic webs (pests, non pests,
disease agent of crops)
Biogeochemical cycles
Water cycle

Fig. 1 The functional structure of a community of weeds depends on the individual responses of species to environmental drivers
because of local conditions and crop abiotic factors (see text for precisions) according to their value of response traits. It also impacts
processes at the agro-ecosystem level and service supply through effect traits. Although this framework is valid for any kind of ecosystem, it
is illustrated here for weed communities, as part of the agro-ecosystem, with examples given in the text. Numbers refer to the different
sections of the article. After Diaz et al. (2007b).

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Weed Research  2012 European Weed Research Society Weed Research 52, 479488

include the benets of weeds for other components of
the local biodiversity, the regulation of crop production
and the aesthetic or conservation values of particular
species. We examine below how this general framework
applies to weed communities and suggest directions for
future research considering actual issues of weed con-
trol, but also the contribution weed ecology could have
to community ecology. For an easier reading, the
numbers in Fig. 1 correspond to sections of the text.
Environmental drivers acting on weeds
Two categories of environmental drivers act on weed
diversity (Fig. 1); some are related to local conditions,
whereas others depend on crop type and management.
We suggest to name the second category of factors crop
abiotic factors instead of management factors to focus
on their eect on the environmental and ecological
conditions (e.g. soil properties and resources, length of
growing season) and not on the dierent kinds of
techniques that are used to manage weeds. Although
these two groups of factors have been traditionally
merged into a single category forming the abiotic
environment (Fig. 2A, Booth et al., 2003), we propose
to separate them into two hierarchical groups (Fig. 2B)
A tillage (Andersson & Milberg, 1998; Fried et al., 2008;
Crop
Animals
weeds
Abiotic
environment
B Local conditions:
climate, sol, history
Crop Abiotic factors:
Preceding crop type
Fertilisation, watering
Tillage, herbicides
Crop
Weeds Animals
Fig. 2 Two representations of the relationships between crop and
weeds in an agro-ecosystem. (A) In the traditional representation
(Booth et al., 2003), the main interaction is between crop and weed
plants that are similarly impacted by all other biotic and abiotic
components of the ecosystem. (B) In this study, we distinguish the
local conditions that are state factors constraining all components
of the system and crop abiotic factors, corresponding to the
impact of management practices on the local environment that
depend on the crop type and that are modulated by local
conditions.
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Functional structure of weeds 481
because they act dierently on species. Local conditions,
such as climate and soil conditions or history of land
use, are independent state factors. In addition to driving
weed assembly directly, they also modulate the eect of
crop abiotic factors on all local organisms (Fig. 2B,
Chapin et al., 2002). For example, climatic condi-
tions at a site constrain the period of growth of all
organisms, whatever the kind of agricultural practices.
As a consequence, interactions between weeds and
crop plants or other organisms respond to the crop
abiotic factors, whose impact is constrained by local
conditions.
These two groups of factors dier in their impact on
weed distribution. The larger inuence of soil conditions
than of climate on weed distribution has been well
documented (references in Maillet, 1981; Lososova
et al., 2004; Fried et al., 2008; Andreasen & Skovgaard,
2009; Hanzlik & Gerowitt, 2011). As an example, a
recent analysis of weed species of 700 arable elds in
France showed that weed communities on basic clay
soils were distinct from those on acidic sandy soils,
whereas the inuence of climate and geographical region
was identied mainly through relationships with precip-
itation and longitude (Fried et al., 2008). However,
major variations in weed composition between elds are
also associated with management factors: current or
preceding crop type, land drainage and occurrence of
Butler et al., 2009; Hawes et al., 2009; Cirujeda et al.,
2011). Current crop type includes the eects of sowing
season, herbicide families and fertilisation regimes that
constrain the growth and phenology of weeds (Lososova
et al., 2008; Gunton et al., 2011). Weed composition is
also impacted by crop rotation, but responds more to
the current identity of the crop than to the total number
of crops in the rotation (Smith & Gross, 2007; Meiss
et al., 2010). More specically, a large body of literature
is dedicated to the description of weed communities
associated with each crop (for examples from recent
work: soybean, De La Fuente et al., 2006; sugarcane,
Firehun & Tamado, 2006; temperate cereals, Potts et al.,
2010; Mediterranean cereals, Cirujeda et al., 2011).
Most of these studies include precise descriptions of
the impact of management on the environment. How-
ever, these descriptions often relate an environmental
impact to a given management practice without dening
the corresponding intensity and frequency of distur-
bance and or constraint. This makes the comparison of
the impacts of dierent techniques and combinations
thereof on weed trait distributions among dierent
situations (e.g. among crops in a same area or among
varieties of a given crop plant) and their prediction
under future conditions rather dicult. If we want
to understand the quantitative relationships between
482 M-L Navas
management practices and weed trait distribution, a
careful quantication of the impact of crop abiotic
factors on local environment is required, because the
interpretation of trait response depends on the local
value and range of the environmental factor under
scrutiny (McGill et al., 2006; Garnier et al., 2007). This
is one of the key reasons why the changes in environ-
ment because of all abiotic factors need to be quantied
with objective, comparable variables and methodologies.
Such an approach, leading to the proposal of standar-
dised procedures for characterising the dierences in
management, has been developed for grasslands and
pastures located throughout Europe (Garnier et al.,
2007). These dierences are mostly due to dierences in
defoliation regime, because of cutting or grazing by
dierent kinds of animals varying in number and period
of stay. Five components related to the intensity and
frequency of disturbance induced by defoliation were
dened, to compare the amount of produced and
removed biomass over the year, the dates and length
of periods of biomass removal. We recommend the use
of a similar procedure for standardising the impact of
major agricultural practices, such as dierent soil and
tillage practices, which result in dierent kinds and levels
of disturbance and or constraints.
Response traits of weeds
Traditionally, the rst step for identifying functional
groups is to detect emergent groups of species sharing
correlated traits that reect their adaptation to local
environment (Lavorel et al., 1997). The rst functional
classication of weeds was the ideal weed portrait
proposed by Baker (1974), Baker and Stebbins (1965).
In this proposal, weeds have a general purpose
genotype and a set of characteristics linked to weed-
iness: opportunistic germination of long-lived seeds,
high growth rate and short vegetative phase, self-
compatibility or non-specialised pollination systems,
production of a large number of seeds in a large range
of environments, dispersal over both long and short
distances, large competitive ability and vigorous vege-
tative reproduction for perennial species. Other classi-
cations based on ecological characteristics, such as
dispersal and reproductive patterns, demographical
traits or adaptations to management practices, have
also been proposed as a basis for further identication of
potential weeds (Patterson, 1985; Noble, 1989). For
example, Newsome and Noble (1986) analysed 86
noxious weeds in Victoria (Australia) and recognised
10 groups of species on the basis of 17 characteristics,
including longevity, size, origin, growth form, photo-
synthetic pathway and germination. More recently, trait
analyses related weediness to short life cycle, low seed
Weed Research  2012 European Weed Research Society Weed Research 52, 479488
size, relatively large plant height and opportunistic
phenology (Storkey, 2006; Ryan et al., 2010; Gunton
et al., 2011).
A second set of studies characterised the ability of
weeds to respond to fast changes in manmade selective
pressures (Navas, 1991). For example, species that
increased in occurrence in sunower crops during the
last forty years belong to a sunower mimicking
functional group: they are nitrophilous and heliophil-
ous, tolerant to sunower herbicides and share a rapid
summer life cycle, independently of phylogeny (Fried
et al., 2009). By contrast, weeds occurring in less
intensive systems are generally short plants, producing
big seeds and owering later than species found in more
intensive systems (Lososova et al., 2006). These species
are excluded from intensive systems because short plants
cannot be competitive enough to survive under highly
productive conditions and the few seedlings produced by
big seeds have a low probability to avoid herbicide
spraying. Unsurprisingly, the same set of traits was
found to characterise weed species selected against over
the past 60 years in cereal crops in the UK (Storkey
et al., 2010).
A third set of comparative studies identied weed
traits responding to specic components of manage-
ment. Most of experiments have been performed with a
limited number of major species, and generalisation of
results is not possible. However, the response of weed
communities to sowing time or tillage has been described
using comparative studies on germination syndrome.
For example, changes in seed bank across time were
related to a limited number of seed traits: seed mortality
in the soil decreases exponentially with seed coat
thickness (Gardarin et al., 2010b), germination varies
with mass, area and lipid content of seeds (Gardarin
et al., 2011), whereas seedling emergence depends on
seed mass and hypocotyl or epicotyl diameter and diers
among monocotyledon and dicotyledon species (Fayolle
et al., 2009; Gardarin et al., 2010a). The response to
nutrient availability was characterised using a similar
comparative approach: seed mass decreased as nutrient
availability increased; plant height was unchanged in all
fertilised treatments, with shorter plants being found in
unfertilised areas (Storkey et al., 2010). Weed response
to competition by crop plants was related to specic leaf
area (the ratio of leaf area and biomass), a trait that
varies with light availability in crop weed mixture
(Violle et al., 2009; Fila & Sartorato, 2011). By contrast,
herbicide tolerance is most often described by genetic or
physiological characteristics of a few species, yet cannot
be generalised. Comparative studies on the impact of
leaf anatomy and architecture, cuticle thickness, plant
growth and phenology are lacking (e.g. Hilgenfeld et al.,
2004; Baucom & Mauricio, 2008; Vila-Aiub et al., 2009),
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despite the fact that they could help to rank species
according to herbicide tolerance in less intensively
managed situations.
Functional structure of weeds
The functional diversity of a community can be
described through the distribution of traits. A rst
descriptor of this structure estimates the average value
of traits in the community as:
X n
CWM pi  traiti 1 documented for other crop abiotic factors, once they are
i1
where CWM represents the community-weighted mean,
pi and traiti are, respectively, the proportion and the
trait value for species i, and n is the total number of species
in the community (Garnier et al., 2004). This calculation is
based on the mass ratio hypothesis (Grime, 1998), which
proposes that the controls on function by species are
proportional to their abundance. A second descriptor
assesses the functional divergence of traits of species
present in a community. Many different indices are used
(reviewed in Garnier & Navas, 2012); most are based on the
sum of or average functional distances between species
pairs, distances between species along a hierarchical clas-
sification or on the distribution of abundances along
functional trait axes. Recent developments in community
ecology emphasise the relevance of these descriptors for
assessing the assembly of species (Cornwell & Ackerly,
2009). The deviation of observed mean, range and diver-
gence of traits compared with values generated with a null
model can reveal the filtering effect of environmental factors
on traits and the nature of the associated processes (Weiher
& Keddy, 1995; Grime, 2006). Descriptors of functional
structure are also used to assess ecosystem services (see
below). So far, few calculations of CWM of traits of crop
weeds or functional divergence have been published (Stor-
key et al., 2010), reflecting the poor understanding we have
of the functional structure of weed communities.
When analysing the functional structure of weeds
found in a eld, there is a question of where to include
crop plants in the analysis as a driver or component of
the ora. We argue that crop plants that may display a
range of phenotypes, despite genetic homogeneity, in
response to local environmental heterogeneity, represent
a local biotic lter acting on weed species via their
impact on local resource level and environmental
conditions (Fig. 2B). This ltering eect constrains the
values of traits of weeds. For example, weeds mimicking
crop plants with slight dierences in trait values (e.g.
earlier owering phenology or with brittle owering
stems, Navas, 1991; Linghwa & Morishima, 1997; Fried
et al., 2009) are an example of responses to such a
ltering eect, revealing niche dierentiation between
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Functional structure of weeds 483
crop and weed plants. We suggest here that constraints
on weed traits by crop functional structure occur also in
more complex situations, as suggested by a recent
comparative study of weed communities occurring in
dierent crops (Gunton et al., 2011). In this study,
weeds occurring in crops with late sowing date were
taller, established and owered later and had a shorter
period of owering than other weed species. Further-
more, the mean and range of date of emergence of
weeds, respectively, increases and decreases with crop
sowing date. We suspect that similar control could be
evaluated with standardised methods, as suggested
earlier. For example, we suggest that CWM of crop
height at the time of canopy closure is of major
importance to restrict weed diversity, as this trait
describes the use of light and water by the community
throughout the growing season (Violle et al., 2009).
Effect traits and agro-ecosystem processes
The most documented eect of weeds on the agro-
ecosystem is the depression of crop production. Because
the prediction of lost crop production is a major
economic issue, it has given rise to a signicant
modelling eort since the early 70s (Doyle, 1997),
especially for elds invaded by major noxious species.
In this case, predictions have been based on the
assessment of changes in demography of the weed in
response to dierent management systems (Petit et al.,
2010). Most of these models are deterministic and
require a large experimental eort to assess a large
number of parameters, explaining why they have been
developed for only a few species. Recent demo-
graphical models group species into functional types
(Storkey & Cussans, 2007; Gardarin et al., 2011), but
the complexity of the models impedes further extension
to diverse weed communities.
An alternative approach is to characterise the more
general competitive eect of weeds on crops. The
signicant relationship linking competitive eect to
size-related traits (Gaudet & Keddy, 1988; Keddy &
Shipley, 1989) has been conrmed for weed species:
comparative studies related seed size or plant height to
competitive eect (Goldberg & Fleetwood, 1987; Gold-
berg & Landa, 1991; Torner et al., 2000; Storkey et al.,
2010). Yet, these traits have not been used in models
predicting competition outcomes. These models are
based on the fact that weed size relative to that of the
crop, measured at an early stage depending on weed and
crop phenology, is the best predictor of the nal crop
yield (e.g. Krop & Spitters, 1992; Booth & Swanton,
2002). For example, beet yield was accurately predicted
by the relative leaf area of Chenopodium album measured
484 M-L Navas
30 days after weed emergence (Krop & Spitters, 1991).
Recently, the dierence in plant height among weed and
crop plants assessed soon after weed emergence was
related to the intensity of competition and crop yield
and also to the reproductive performance of weeds
(McDonald et al., 2010). These results can be explained
in the light of recent theoretical ecological studies:
(i) traits of interacting plants are tightly connected, and
the competitive response of a plant varies with the
competitive eect of its neighbours (Wang et al., 2010);
(ii) the dierences in size-related traits among interacting
plants explain the co-existence of species in communities
regenerated from seeds (Turnbull et al., 2004) and
(iii) the distribution of trait values for the whole
community formed by weed and crop plants reects
the competition at the site (McGill et al., 2006).
Although simple to parameterise and ecologically
sound, these models lack accuracy, because demograph-
ical changes of weeds are not taken into account (but see
McDonald et al., 2010). Therefore, there is an urgent
need to identify traits related to demography, because
only a few studies addressed this question. Some traits
have been proposed for tropical trees (Garnier & Navas,
2012), and a recent modelling study characterised which
life-history traits of weeds are more sensitive to man-
agement (Colbach et al., 2010).
The predictions by these two kinds of models are,
however, of low value when the functional diversity of
weeds is high under extensive management systems.
Some authors have recently suggested that in such
conditions, the ability of weeds to draw on resource
pools complementarily to that of crop plants, which can
be assessed by dierences in root depth and architecture
(Freckleton & Watkinson, 2001; Deen et al., 2003; Smith
et al., 2010), might be related to lower weed-crop
competition (Smith et al., 2010). This means that the
functional divergence of the crop-weed community
dened by traits related to resource use could be of
major importance for predicting crop yield. A high
divergence inducing complementarity in resource use by
weeds and crop across time or space, in relation to niche
dierentiation, should result in a reduced impact of
weeds on crops. This suggests that the limiting similarity
eect, one hypothesis explaining co-existence of species
in spontaneous communities [co-existing species slightly
dier in functioning then have separated niche dimen-
sions (Wilson, 2011)], should also prevail in managed
places.
There are probably other eect traits of weeds that
could relate to major ecosystem processes. For example,
(i) occurrence of herbivores is generally related to tissue
chemistry, leaf morphology and seed production,
(ii) pollinator provision is explained by oral traits such
as accessibility, attractiveness and nectar production and
Weed Research  2012 European Weed Research Society Weed Research 52, 479488
(iii) soil protection depends on canopy size or architec-
ture or growth form (De Bello et al., 2010). However,
quantitative links between these services and weed traits
have yet to be fully explored.
Weed traits and services
The recent emergence of the concept of ecosystem
services has led to the recognition of the positive impact
weeds can have on agro-ecosystem. To that aim, a
framework recognising three categories of weeds,
involved in the whole range of ecosystem services, has
been recently proposed (Moonen & Ba`rberi, 2008):
(i) species with high intrinsic conservation value or
because of sheltered biodiversity, (ii) species contribut-
ing to the production of food for animals, or to services
of regulation such as pollination or limitation of soil
erosion and (iii) species with no intrinsic value but that
are used as indicators of high-quality practices. The
next step is to recognise traits related to properties and
processes that have substantial contribution to services,
following the pioneering framework proposed by De
Bello et al. (2010). Although the positive impact of
weeds on local biodiversity is well known (Ba`rberi et al.,
2010), no clear set of traits has been related to these
services. A recent study performed in oilseed rape, beet
and maize crops in the UK showed that functional types
of weeds, identied by categorical traits, can be associ-
ated with those of non-pest invertebrate species (Hawes
et al., 2009). A similar result was found in Mediterra-
nean winter wheat elds when classifying weeds into
three functional groups (Caballero-Lopez et al., 2010).
On the other hand, the ability of weeds to host other
organisms, which can be positive when weeds act as trap
crops attracting pests (Ba`rberi et al., 2010), or negative
when they host disease agents (Franke et al., 2009), has
not been clearly depicted by traits. An exception is the
identication of shifted phenology of weeds that host
fungi or virus comparatively to crops, allowing the
disease agent to persist between two periods of crop
production (Navas et al., 1998).
Once eect traits are identied, which is something
still to be done for weeds, it is necessary to evaluate the
shortest set of traits that provision a given service at a
site. To that aim, Diaz et al. (2007a) proposed a
hierarchical procedure that should be used in agro-
ecosystems. Once the eect of the environment on
services is identied, CWM and functional divergence of
eect traits are assessed, then the impact of some species
on services, for example the dominant when the biomass
ratio hypothesis is valid (Grime, 1998), as it is the case
when assessing services related to resource use (Garnier
et al., 2004), is quantied. This procedure was used rst
for grasslands and allowed to recognise the best com-
 2012 The Author

bination of environmental factors and functional diver-
sity components acting on ecosystem processes and
services.
The identication of services provided by weeds
introduces complexity in the denition of control
strategies, as the balance between the negative and
positive impacts weeds have on the dierent components
of the agro-ecosystem needs to be assessed. Although
there are some exceptions, weed species cannot be
grouped into two contrasted groups, the rst including
highly competitive species without conservation value
and a second one formed of species with opposite
characteristics. The reality is in between, as documented
for common weed species of UK, for which no clear
correlation exists between harmfulness and positive
eects on biodiversity (Storkey, 2006). Therefore, pro-
posing to manage elds to keep only species with
positive eects appears to be unrealistic (Storkey &
Westbury, 2007). Furthermore, dening which weeds are
acceptable because they have more positive than nega-
tive eects remains a challenge, because the former
eects remain poorly understood and largely unquanti-
ed. Potentially, functional traits could be used as a
common metric for trading o the positive and negative
impacts of contrasting weed communities.
Conclusion: directions for weed research
Applying trait-based approaches to weed ecology raises
a number of challenges that are somewhat dierent to
those proposed for other kinds of vegetation (see
Garnier & Navas, 2012, for a general discussion). The
most important dierences can be summarised as
follows:
1 Concerning plant functioning, whether any easily
measured traits can be related to the various aspects
of the regeneration niche and to demographical
parameters, such as population birth and death rate,
appears as a central issue for better prediction of weed
demography under contrasted management scenarios.
We suspect that such research could be a major
contribution to community ecology, because weed
species are characterised by rapidly adapting life-
history traits in response to varying environmental
conditions.
2 At the community level, understanding the mecha-
nisms through which species traits determine func-
tional structure will require theoretical, experimental
and modelling approaches. There is a need to
better identify and quantify the crop abiotic factors
acting on weeds, and weed response traits, by
performing comparative experiments. Working with
the crop-weed system gives two opportunities for
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Functional structure of weeds 485
community ecology. First, this will improve the
understanding of the impact of disturbances on
community assembly. If the impact of resource
change on traits is rather well documented, there is
still no general understanding of the impact of
disturbances on functional structure of plants (see
Garnier & Navas, 2012, for a general discussion). The
rather specic disturbances occurring in an agro-
ecosystem (e.g. soil tillage and herbicide application)
compared with other systems (e.g. re, ooding and
herbivory) should allow the general quantication of
processes linked to disturbance. Second, the crop-
weed system is an ideal one to test the hypothesis that
co-existence among species is governed by their values
of traits, especially those related to competition,
because of the asymmetrical relationships between
the two kinds of plants.
3 At the agro-ecosystem level, the understanding of
how and which traits (in terms of values and ranges)
affect processes, properties and services remains very
preliminary. The originality of weeds compared with
other vegetation of agricultural interest, such as
grasslands, is that the balance between their impact
on ecosystem can vary from purely negative (if only
loss of crops is taken into account) to purely positive
(if only services on biodiversity are taken into
account), allowing the characterisation of the whole
range of trade-offs.
In conclusion, trait-based approaches should give new
tools for predicting weed community assembly and
impact on agro-ecosystem services, especially in com-
plex communities where a detailed mechanistic and
modelling approach based on in-depth knowledge
of all organisms involved will probably not be
tractable.
Acknowledgements
The idea of this study came after the publication of a
review on trait-based approach for agro-ecology
(Garnier & Navas, 2012) including a very short section
on crop weeds. I also thank S Gaba, B Chauvel, G Fried
and E Kazakou for very interesting discussions on some
of the ideas developed here. Many thanks also to
referees and editor who carefully read a former version
and provide very useful recommendations for improving
the ideas.
References
ANDERSSON TN & MILBERG P (1998) Weed ora and the
relative importance of site, crop, crop rotation, and nitrogen.
Weed Science 46, 3038.
486 M-L Navas
ANDREASEN C & SKOVGAARD IBM (2009) Crop and soil factors
of importance for the distribution of plant species on arable
elds in Denmark. Agriculture Ecosystems & Environment
133, 6167.
AUBERTOT J-N, BARBIER J-M, CARPENTIER A et al. (2007)
Pesticides, Agriculture et environnement. Reduire lutilisation
des pesticides et en limiter les impacts environnementaux.
Expertise scientique collective Inra-Cemagref (decembre
2005). Quae, Paris.
BAKER HG (1974) The evolution of weeds. Annual Review of
Ecology and Systematics 5, 124.
BAKER HG & STEBBINS GL (1965) The Genetics of Colonizing
Species. Academic Press, New York & London.
BA`RBERI P, BURGIO G, DINELLI G et al. (2010) Functional
biodiversity in the agricultural landscape: relationships
between weeds and arthropod fauna. Weed Research 50,
388401.
BAUCOM RS & MAURICIO R (2008) Constraints on the
evolution of tolerance to herbicide in the common morning
glory: resistance and tolerance are mutually exclusive.
Evolution 62, 28422854.
BOOTH BD & SWANTON CJ (2002) Assembly theory applied to
weed communities. Weed Science 50, 213.
BOOTH BD, MURPHY SD & SWANTON CJ (2003) Weed Ecology
in Natural and Agricultural Systems. CABI Publishing,
Oxon, UK.
BUTLER SJ, BROOKS D, FEBER RE, STORKEY J, VICKERY JA &
NORRIS K (2009) A cross-taxonomic index for quantifying
the health of farmland biodiversity. Journal of Applied
Ecology 46, 11541162.
CABALLERO-LOPEZ B, BLANCO-MORENO JM, PEREZ N et al.
(2010) A functional approach to assessing plant-arthropod
interaction in winter wheat. Agriculture Ecosystems &
Environment 137, 288293.
CHAPIN FS III, MATSON PA & MOONEY HA (2002) Principles of
Terrestrial Ecosystem Ecology. Springer-Verlag, New-York.
CIRUJEDA A, AIBAR J & ZARAGOZA C (2011) Remarkable
changes of weed species in Spanish cereal elds from 1976 to
2007. Agronomy for Sustainable Development 31, 675688.
COLBACH N, DARMENCY H & TRICAULT Y (2010) Identifying
key life-traits for the dynamics and gene ow in a weedy crop
relative: sensitivity analysis of the GENESYS simulation
model for weed beet (Beta vulgaris ssp. vulgaris). Ecological
Modelling 221, 225237.
CORNWELL WK & ACKERLY D (2009) Community assembly
and shifts in plant trait distributions across an environ-
mental gradient in coastal California. Ecological
Monographs 79, 109126.
COUSENS R & MORTIMER M (1995) Dynamics of Weed
Populations, 1st edn. Cambridge University Press,
Cambridge, UK.
DE BELLO F, LAVOREL S, DIAZ S et al. (2010) Towards an
assessment of multiple ecosystem processes and services via
functional traits. Biodiversity and Conservation 19, 28732893.
DE LA FUENTE EB, SUAREZ SA & GHERSA CM (2006) Soybean
weed community composition and richness between 1995
and 2003 in the Rolling Pampas (Argentina). Agriculture
Ecosystems & Environment 115, 229236.
DEEN W, COUSENS R, WARRINGA J et al. (2003) An evaluation
of four crop: weed competition models using a common data
set. Weed Research 43, 116129.
Weed Research  2012 European Weed Research Society Weed Research 52, 479488
DIAZ S, LAVOREL S, BELLO FD, QUETIER F, GRIGULIS K &
ROBSON TM (2007a) Incorporating plant functional diversity
eects in ecosystem service assessments. Proceedings of the
National Academy of Sciences of the United States of America
104, 2068420689.
DIAZ S, LAVOREL S, CHAPIN FS, TECCO PA, GURVICH DE &
GRIGULIS K (2007b) Functional diversity at the crossroads
between ecosystem functioning and environmental lters.
In: Terrestrial Ecosystems in a Changing World (eds JG
CANADELL, D PATAKI & L PITELKA), 8191. Springer-Verlag,
Berlin, Heidelberg.
DOYLE CJ (1997) A review of the use of models of weed control
in integrated crop protection. Agriculture Ecosystems &
Environment 64, 165172.
FAYOLLE A, VIOLLE C & NAVAS ML (2009) Dierential impacts
of plant interactions on herbaceous species recruitment:
disentangling factors controlling emergence, survival and
growth of seedlings. Oecologia 159, 817825.
FERNANDEZ-QUINTANILLA C, QUADRANTI M, KUDSK P &
BA`RBERI P (2008) Which future for weed science? Weed
Research 48, 297301.
FILA G & SARTORATO I (2011) Using leaf mass per area as
predictor of light interception and absorption in crop weed
monoculture or mixed stands. Agricultural and Forest
Meteorology 151, 575584.
FIREHUN Y & TAMADO T (2006) Weed ora in the Rift
Valley sugarcane plantations of Ethiopia as inuenced by
soil types and agronomic practises. Weed Biology and
Management 6, 139150.
FRANKE AC, LOTZ LAP, van der Burg WJ & OVERBEEK LV
(2009) The role of arable weed seeds for agroecosystem
functioning. Weed Research 49, 131141.
FRECKLETON RP & WATKINSON AR (2001) Asymmetric
competition between plant species. Functional Ecology 15,
615623.
FRIED G, NORTON LR & REBOUD X (2008) Environmental and
management factors determining weed species composition
and diversity in France. Agriculture Ecosystems & Environ-
ment 128, 6876.
FRIED G, CHAUVEL B & REBOUD X (2009) A functional analysis
of large-scale temporal shifts from 1970 to 2000 in weed
assemblages of sunower crops in France. Journal of
Vegetation Science 20, 4958.
GARDARIN A, DURR C & COLBACH N (2010a) Eects
of seed depth and soil aggregates on the emer-
gence of weeds with contrasting seed traits. Weed Research
50, 91101.
GARDARIN A, DURR C, MANNINO MR, BUSSET H & COLBACH N
(2010b) Seed mortality in the soil is related to seed coat
thickness. Seed Science Research 20, 243256.
GARDARIN A, DAURR C & COLBACH N (2011) Prediction of
germination rates of weed species: relationships between
germination speed parameters and species traits. Ecological
Modelling 222, 626636.
GARNIER E & NAVAS M-L (2012) A trait-based approach to
comparative functional plant ecology: concepts, methods
and applications for agroecology. A review. Agronomy for
Sustainable Development 32, 365399.
GARNIER E, CORTEZ J, BILLES G et al. (2004) Plant functional
markers capture ecosystem properties during secondary
succession. Ecology 85, 26302637.
 2012 The Author

GARNIER E, LAVOREL S, ANSQUER P et al. (2007) Assessing the
eects of land-use change on plant traits, communities and
ecosystem functioning in grasslands: a standardized
methodology and lessons from an application to 11
European sites. Annals of Botany 99, 967985.
GAUDET CL & KEDDY PA (1988) A comparative approach
to predicting competitive ability from plant traits. Nature
334, 242243.
GOLDBERG DE & FLEETWOOD L (1987) Competitive eect
and response in four annual plants. Journal of Ecology 75,
11311143.
GOLDBERG DE & LANDA K (1991) Competitive eect and
response hierarchies and correlated traits in the early stages
of competition. Journal of Ecology 79, 10131030.
GRIME JP (1998) Benets of plant diversity to ecosystems:
immediate, lter and founder eects. Journal of Ecology 86,
902910.
GRIME JP (2006) Trait convergence and trait divergence in
herbaceous plant communities: mechanisms and conse-
quences. Journal of Vegetation Science 17, 255260.
GUNTON RM, PETIT S & GABA S (2011) Functional traits
relating arable weed communities to crop characteristics.
Journal of Vegetation Science 22, 541550.
HANZLIK K & GEROWITT B (2011) The importance of climate, site
and management on weed vegetation in oilseed rape in Ger-
many. Agriculture Ecosystems & Environment 141, 323331.
HAWES C, HAUGHTON AJ, BOHAN DA & SQUIRE GR (2009)
Functional approaches for assessing plant and invertebrate
abundance patterns in arable systems. Basic and Applied
Ecology 10, 3442.
HILGENFELD KL, MARTIN AR, MORTENSEN DA & MASON SC
(2004) Weed management in a glyphosate resistant
soybean system: weed species shifts. Weed Technology
18, 284291.
KEDDY PA & SHIPLEY B (1989) Competitive hierarchies in
herbaceous plant communities. Oikos 54, 234241.
KROPFF MJ & SPITTERS CJT (1991) A simple model of crop
loss by weed competition from early observations on relative
leaf area of the weeds. Weed Research 31, 97105.
KROPFF MJ & SPITTERS CJT (1992) An ecophysiological model
for interspecic competition, applied to the inuence of
Chenopodium album L. on sugar-beet. 1. Model description
and parameterization. Weed Research 32, 437450.
LAVOREL S & GARNIER E (2002) Predicting changes in
community composition and ecosystem functioning from
plant traits: revisiting the Holy Grail. Functional Ecology 16,
545556.
LAVOREL S, MCINTYRE S, LANDSBERG J & FORBES TDA (1997)
Plant functional classications: from general groups to
specic groups based on response to disturbance. Trends in
Ecology & Evolution 12, 474478.
LAVOREL S, DIAZ S, CORNELISSEN H et al. (2007) Plant
Functional Types: Are We Getting Any Closer to the Holy
Grail? In: Terrestrial Ecosystems in a Changing World (eds
JG CANADELL, D PATAKI & L PITELKA), 150164. Springer
Verlag, Berlin, Heidelberg, Germany.
LINGHWA T & MORISHIMA H (1997) Genetic characterization of
weedy rices and the inference on their origins. Breeding
Science 47, 153160.
LOSOSOVA Z, CHYTRY M, CIMALOVA S et al. (2004) Weed
vegetation of arable land in Central Europe: gradients of
 2012 The Author
Weed Research  2012 European Weed Research Society Weed Research 52, 479488
Functional structure of weeds 487
diversity and species composition. Journal of Vegetation
Science 15, 415422.
LOSOSOVA Z, CHYTRY M, KUHN I et al. (2006) Patterns of plant
traits in annual vegetation of man-made habitats in central
Europe. Perspectives in Plant Ecology, Evolution and
Systematics 8, 6981.
LOSOSOVA Z, CHYTRY M & KUEHN I (2008) Plant
attributes determining the regional abundance of weeds
on central European arable land. Journal of Biogeography
35, 177187.
MAILLET J (1981) Evolution de la ore adventice dans le
Montpellierais sous la pression des techniques culturales.
Docteur Ingenieur, Universite des Sciences et Techniques du
Languedoc, Montpellier.
MARSHALL EJP, BROWN VK, BOATMAN ND et al. (2003) The
role of weeds in supporting biological diversity within crop
elds. Weed Research 43, 7789.
MCDONALD AJ, RIHA SJ & DITOMMASO A (2010) Early season
height dierences as robust predictors of weed growth
potential in maize: new avenues for adaptive management?
Weed Research 50, 110119.
MCGILL BJ, ENQUIST BJ, WEIHER E & WESTOBY M (2006)
Rebuilding community ecology from functional traits.
Trends in Ecology & Evolution 21, 178185.
MEISS H, MEDIENE S, WALDHARDT R et al. (2010) Perennial
lucerne aects weed community trajectories in grain crop
rotations. Weed Research 50, 331340.
MOONEN AC & BA`RBERI P (2008) Functional biodiversity:
an agroecosystem approach. Agriculture Ecosystems &
Environment 127, 721.
NAVAS ML (1991) Using plant-population biology in weed
research a strategy to improve weed management. Weed
Research 31, 171179.
NAVAS ML, FRIESS N & MAILLET J (1998) Inuence of
cucumber mosaic virus infection on the growth response
of Portulaca oleracea (purslane) and Stellaria media (chick-
weed) to nitrogen availability. New Phytologist 139, 301309.
NEWSOME AE & NOBLE IR (1986) Ecological and physiological
characters of invading species. In: Ecology of Biological
Invasions (eds RH GROVES & JJ BURDON), 120. Cambridge
University Press, New York, USA.
NOBLE I (1989) Attributes of invaders and the invading process:
terrestrial and vascular plants. In: Biological Invasions: A
Global Perspective (eds JA DRAKE, HA MOONEY, FD CASTRI
& et al.), 301314. John Wiley & Sons, Chichester, UK.
PATTERSON DT (1985) Comparative Ecophysiology of Weeds
and Crops. CRC Press, Boca Raton, FL, USA.
PETIT S, BOURSAULT A, GUILLOUX ML, MUNIER-JOLAIN N &
REBOUD X (2010) Weeds in agricultural landscapes. A
review. Agronomy for Sustainable Development 31, 309317.
POTTS GR, EWALD JA & AEBISCHER NJ (2010) Long-term
changes in the ora of the cereal ecosystem on the Sussex
Downs, England, focusing on the years 19682005. Journal
of Applied Ecology 47, 215226.
RYAN MR, SMITH RG, MIRSKY SB, MORTENSEN DA & SEIDEL
R (2010) Management lters and species traits: weed
community assembly in long-term organic and conventional
systems. Weed Science 58, 265277.
SMITH RG & GROSS KL (2007) Assembly of weed communities
along a crop diversity gradient. Journal of Applied Ecology
44, 10461056.
488 M-L Navas
SMITH RG, MORTENSEN DA & RYAN MR (2010) A new
hypothesis for the functional role of diversity in mediating
resource pools and weed-crop competition in agroecosys-
tems. Weed Research 50, 3748.
STORKEY J (2006) A functional group approach to the
management of UK arable weeds to support biological
diversity. Weed Research 46, 513522.
STORKEY J & CUSSANS JW (2007) Reconciling the conservation
of in-eld biodiversity with crop production using a simu-
lation model of weed growth and competition. Agriculture
Ecosystems & Environment 122, 173182.
STORKEY J & WESTBURY DB (2007) Managing arable weeds for
biodiversity. Pest Management Science 63, 517523.
STORKEY J, MOSS SR & CUSSANS JW (2010) Using assembly
theory to explain changes in a weed ora in response to
agricultural intensication. Weed Science 58, 3946.
SUDING KN, LAVOREL S, CHAPIN FS et al. (2008) Scaling
environmental change through the community-level: a trait-
based response-and-eect framework for plants. Global
Change Biology 14, 11251140.
TORNER C, DEL ARCO MJS, SATORRE E & FERNANDEZ
QUINTANILLA C (2000) A comparison of the growth patterns
and the competitive ability of four annual weeds. Agronomie
20, 147156.
TURNBULL LA, COOMES D, HECTOR A & REES M (2004) Seed
mass and the competition colonization trade-o:
competitive interactions and spatial patterns in a guild of
annual plants. Journal of Ecology 92, 97109.
Weed Research  2012 European Weed Research Society Weed Research 52, 479488
VILA-AIUB MM, NEVE P & POWLES SB (2009) Evidence for an
ecological cost of enhanced herbicide metabolism in Lolium
rigidum. Journal of Ecology 97, 772780.
VIOLLE C, NAVAS M-L, VILE D et al. (2007) Let the concept of
trait be functional!. Oikos 116, 882892.
VIOLLE C, GARNIER E, LECOEUR J et al. (2009) Competition,
traits and resource depletion in plant communities.
Oecologia 160, 747755.
WANG P, STIEGLITZ T, ZHOU DW & CAHILL JF (2010)
Are competitive eect and response two sides of the
same coin, or fundamentally dierent? Functional Ecology
24, 196207.
WEIHER E & KEDDY PA (1995) Assembly rules, null models,
and trait dispersion: new questions front old patterns. Oikos
74, 159164.
WEIHER E, FREUND D, BUNTON T, STEFANSKI A, LEE T &
BENTIVENGA S (2011) Advances, challenges and a developing
synthesis of ecological community assembly theory.
Philosophical Transactions of the Royal Society of London.
Series B, Biological Sciences 366, 24032413.
WEZEL A, BELLON S, DORE T, FRANCIS C, VALLOD D & DAVID
C (2009) Agroecology as a science, a movement and a
practice. A review. Agronomy for Sustainable Development
29, 203515.
WILSON JB (2011) The twelve theories of co-existence
in plant communities: the doubtful, the impor-
tant and the unexplored. Journal of Vegetation Science 22,
184195.
 2012 The Author