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

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

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

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

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

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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Functional structure of weeds 481

include the benets of weeds for other components of because they act dierently on species. Local conditions,
the local biodiversity, the regulation of crop production such as climate and soil conditions or history of land
and the aesthetic or conservation values of particular use, are independent state factors. In addition to driving
species. We examine below how this general framework weed assembly directly, they also modulate the eect of
applies to weed communities and suggest directions for crop abiotic factors on all local organisms (Fig. 2B,
future research considering actual issues of weed con- Chapin et al., 2002). For example, climatic condi-
trol, but also the contribution weed ecology could have tions at a site constrain the period of growth of all
to community ecology. For an easier reading, the organisms, whatever the kind of agricultural practices.
numbers in Fig. 1 correspond to sections of the text. As a consequence, interactions between weeds and
crop plants or other organisms respond to the crop
abiotic factors, whose impact is constrained by local
Environmental drivers acting on weeds
Two categories of environmental drivers act on weed These two groups of factors dier in their impact on
diversity (Fig. 1); some are related to local conditions, weed distribution. The larger inuence of soil conditions
whereas others depend on crop type and management. than of climate on weed distribution has been well
We suggest to name the second category of factors crop documented (references in Maillet, 1981; Lososova
abiotic factors instead of management factors to focus et al., 2004; Fried et al., 2008; Andreasen & Skovgaard,
on their eect on the environmental and ecological 2009; Hanzlik & Gerowitt, 2011). As an example, a
conditions (e.g. soil properties and resources, length of recent analysis of weed species of 700 arable elds in
growing season) and not on the dierent kinds of France showed that weed communities on basic clay
techniques that are used to manage weeds. Although soils were distinct from those on acidic sandy soils,
these two groups of factors have been traditionally whereas the inuence of climate and geographical region
merged into a single category forming the abiotic was identied mainly through relationships with precip-
environment (Fig. 2A, Booth et al., 2003), we propose itation and longitude (Fried et al., 2008). However,
to separate them into two hierarchical groups (Fig. 2B) major variations in weed composition between elds are
also associated with management factors: current or
preceding crop type, land drainage and occurrence of
A tillage (Andersson & Milberg, 1998; Fried et al., 2008;
Crop 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
Abiotic constrain the growth and phenology of weeds (Lososova
environment 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
B Local conditions:
climate, sol, history of crops in the rotation (Smith & Gross, 2007; Meiss
et al., 2010). More specically, a large body of literature
Crop Abiotic factors: is dedicated to the description of weed communities
Preceding crop type
Fertilisation, watering associated with each crop (for examples from recent
Tillage, herbicides
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
Weeds Animals the impact of management on the environment. How-
ever, these descriptions often relate an environmental
impact to a given management practice without dening
Fig. 2 Two representations of the relationships between crop and
weeds in an agro-ecosystem. (A) In the traditional representation the corresponding intensity and frequency of distur-
(Booth et al., 2003), the main interaction is between crop and weed bance and or constraint. This makes the comparison of
plants that are similarly impacted by all other biotic and abiotic the impacts of dierent techniques and combinations
components of the ecosystem. (B) In this study, we distinguish the thereof on weed trait distributions among dierent
local conditions that are state factors constraining all components
situations (e.g. among crops in a same area or among
of the system and crop abiotic factors, corresponding to the
impact of management practices on the local environment that varieties of a given crop plant) and their prediction
depend on the crop type and that are modulated by local under future conditions rather dicult. If we want
conditions. to understand the quantitative relationships between

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

management practices and weed trait distribution, a size, relatively large plant height and opportunistic
careful quantication of the impact of crop abiotic phenology (Storkey, 2006; Ryan et al., 2010; Gunton
factors on local environment is required, because the et al., 2011).
interpretation of trait response depends on the local A second set of studies characterised the ability of
value and range of the environmental factor under weeds to respond to fast changes in manmade selective
scrutiny (McGill et al., 2006; Garnier et al., 2007). This pressures (Navas, 1991). For example, species that
is one of the key reasons why the changes in environ- increased in occurrence in sunower crops during the
ment because of all abiotic factors need to be quantied last forty years belong to a sunower mimicking
with objective, comparable variables and methodologies. functional group: they are nitrophilous and heliophil-
Such an approach, leading to the proposal of standar- ous, tolerant to sunower herbicides and share a rapid
dised procedures for characterising the dierences in summer life cycle, independently of phylogeny (Fried
management, has been developed for grasslands and et al., 2009). By contrast, weeds occurring in less
pastures located throughout Europe (Garnier et al., intensive systems are generally short plants, producing
2007). These dierences are mostly due to dierences in big seeds and owering later than species found in more
defoliation regime, because of cutting or grazing by intensive systems (Lososova et al., 2006). These species
dierent kinds of animals varying in number and period are excluded from intensive systems because short plants
of stay. Five components related to the intensity and cannot be competitive enough to survive under highly
frequency of disturbance induced by defoliation were productive conditions and the few seedlings produced by
dened, to compare the amount of produced and big seeds have a low probability to avoid herbicide
removed biomass over the year, the dates and length spraying. Unsurprisingly, the same set of traits was
of periods of biomass removal. We recommend the use found to characterise weed species selected against over
of a similar procedure for standardising the impact of the past 60 years in cereal crops in the UK (Storkey
major agricultural practices, such as dierent soil and et al., 2010).
tillage practices, which result in dierent kinds and levels A third set of comparative studies identied weed
of disturbance and or constraints. traits responding to specic components of manage-
ment. Most of experiments have been performed with a
limited number of major species, and generalisation of
Response traits of weeds
results is not possible. However, the response of weed
Traditionally, the rst step for identifying functional communities to sowing time or tillage has been described
groups is to detect emergent groups of species sharing using comparative studies on germination syndrome.
correlated traits that reect their adaptation to local For example, changes in seed bank across time were
environment (Lavorel et al., 1997). The rst functional related to a limited number of seed traits: seed mortality
classication of weeds was the ideal weed portrait in the soil decreases exponentially with seed coat
proposed by Baker (1974), Baker and Stebbins (1965). thickness (Gardarin et al., 2010b), germination varies
In this proposal, weeds have a general purpose with mass, area and lipid content of seeds (Gardarin
genotype and a set of characteristics linked to weed- et al., 2011), whereas seedling emergence depends on
iness: opportunistic germination of long-lived seeds, seed mass and hypocotyl or epicotyl diameter and diers
high growth rate and short vegetative phase, self- among monocotyledon and dicotyledon species (Fayolle
compatibility or non-specialised pollination systems, et al., 2009; Gardarin et al., 2010a). The response to
production of a large number of seeds in a large range nutrient availability was characterised using a similar
of environments, dispersal over both long and short comparative approach: seed mass decreased as nutrient
distances, large competitive ability and vigorous vege- availability increased; plant height was unchanged in all
tative reproduction for perennial species. Other classi- fertilised treatments, with shorter plants being found in
cations based on ecological characteristics, such as unfertilised areas (Storkey et al., 2010). Weed response
dispersal and reproductive patterns, demographical to competition by crop plants was related to specic leaf
traits or adaptations to management practices, have area (the ratio of leaf area and biomass), a trait that
also been proposed as a basis for further identication of varies with light availability in crop weed mixture
potential weeds (Patterson, 1985; Noble, 1989). For (Violle et al., 2009; Fila & Sartorato, 2011). By contrast,
example, Newsome and Noble (1986) analysed 86 herbicide tolerance is most often described by genetic or
noxious weeds in Victoria (Australia) and recognised physiological characteristics of a few species, yet cannot
10 groups of species on the basis of 17 characteristics, be generalised. Comparative studies on the impact of
including longevity, size, origin, growth form, photo- leaf anatomy and architecture, cuticle thickness, plant
synthetic pathway and germination. More recently, trait growth and phenology are lacking (e.g. Hilgenfeld et al.,
analyses related weediness to short life cycle, low seed 2004; Baucom & Mauricio, 2008; Vila-Aiub et al., 2009),

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Weed Research  2012 European Weed Research Society Weed Research 52, 479488
Functional structure of weeds 483

despite the fact that they could help to rank species crop and weed plants. We suggest here that constraints
according to herbicide tolerance in less intensively on weed traits by crop functional structure occur also in
managed situations. more complex situations, as suggested by a recent
comparative study of weed communities occurring in
dierent crops (Gunton et al., 2011). In this study,
Functional structure of weeds
weeds occurring in crops with late sowing date were
The functional diversity of a community can be taller, established and owered later and had a shorter
described through the distribution of traits. A rst period of owering than other weed species. Further-
descriptor of this structure estimates the average value more, the mean and range of date of emergence of
of traits in the community as: weeds, respectively, increases and decreases with crop
X n sowing date. We suspect that similar control could be
CWM pi  traiti 1 documented for other crop abiotic factors, once they are
evaluated with standardised methods, as suggested
where CWM represents the community-weighted mean, earlier. For example, we suggest that CWM of crop
pi and traiti are, respectively, the proportion and the height at the time of canopy closure is of major
trait value for species i, and n is the total number of species importance to restrict weed diversity, as this trait
in the community (Garnier et al., 2004). This calculation is describes the use of light and water by the community
based on the mass ratio hypothesis (Grime, 1998), which throughout the growing season (Violle et al., 2009).
proposes that the controls on function by species are
proportional to their abundance. A second descriptor
Effect traits and agro-ecosystem processes
assesses the functional divergence of traits of species
present in a community. Many different indices are used The most documented eect of weeds on the agro-
(reviewed in Garnier & Navas, 2012); most are based on the ecosystem is the depression of crop production. Because
sum of or average functional distances between species the prediction of lost crop production is a major
pairs, distances between species along a hierarchical clas- economic issue, it has given rise to a signicant
sification or on the distribution of abundances along modelling eort since the early 70s (Doyle, 1997),
functional trait axes. Recent developments in community especially for elds invaded by major noxious species.
ecology emphasise the relevance of these descriptors for In this case, predictions have been based on the
assessing the assembly of species (Cornwell & Ackerly, assessment of changes in demography of the weed in
2009). The deviation of observed mean, range and diver- response to dierent management systems (Petit et al.,
gence of traits compared with values generated with a null 2010). Most of these models are deterministic and
model can reveal the filtering effect of environmental factors require a large experimental eort to assess a large
on traits and the nature of the associated processes (Weiher number of parameters, explaining why they have been
& Keddy, 1995; Grime, 2006). Descriptors of functional developed for only a few species. Recent demo-
structure are also used to assess ecosystem services (see graphical models group species into functional types
below). So far, few calculations of CWM of traits of crop (Storkey & Cussans, 2007; Gardarin et al., 2011), but
weeds or functional divergence have been published (Stor- the complexity of the models impedes further extension
key et al., 2010), reflecting the poor understanding we have to diverse weed communities.
of the functional structure of weed communities. An alternative approach is to characterise the more
When analysing the functional structure of weeds general competitive eect of weeds on crops. The
found in a eld, there is a question of where to include signicant relationship linking competitive eect to
crop plants in the analysis as a driver or component of size-related traits (Gaudet & Keddy, 1988; Keddy &
the ora. We argue that crop plants that may display a Shipley, 1989) has been conrmed for weed species:
range of phenotypes, despite genetic homogeneity, in comparative studies related seed size or plant height to
response to local environmental heterogeneity, represent competitive eect (Goldberg & Fleetwood, 1987; Gold-
a local biotic lter acting on weed species via their berg & Landa, 1991; Torner et al., 2000; Storkey et al.,
impact on local resource level and environmental 2010). Yet, these traits have not been used in models
conditions (Fig. 2B). This ltering eect constrains the predicting competition outcomes. These models are
values of traits of weeds. For example, weeds mimicking based on the fact that weed size relative to that of the
crop plants with slight dierences in trait values (e.g. crop, measured at an early stage depending on weed and
earlier owering phenology or with brittle owering crop phenology, is the best predictor of the nal crop
stems, Navas, 1991; Linghwa & Morishima, 1997; Fried yield (e.g. Krop & Spitters, 1992; Booth & Swanton,
et al., 2009) are an example of responses to such a 2002). For example, beet yield was accurately predicted
ltering eect, revealing niche dierentiation between by the relative leaf area of Chenopodium album measured

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484 M-L Navas

30 days after weed emergence (Krop & Spitters, 1991). (iii) soil protection depends on canopy size or architec-
Recently, the dierence in plant height among weed and ture or growth form (De Bello et al., 2010). However,
crop plants assessed soon after weed emergence was quantitative links between these services and weed traits
related to the intensity of competition and crop yield have yet to be fully explored.
and also to the reproductive performance of weeds
(McDonald et al., 2010). These results can be explained
Weed traits and services
in the light of recent theoretical ecological studies:
(i) traits of interacting plants are tightly connected, and The recent emergence of the concept of ecosystem
the competitive response of a plant varies with the services has led to the recognition of the positive impact
competitive eect of its neighbours (Wang et al., 2010); weeds can have on agro-ecosystem. To that aim, a
(ii) the dierences in size-related traits among interacting framework recognising three categories of weeds,
plants explain the co-existence of species in communities involved in the whole range of ecosystem services, has
regenerated from seeds (Turnbull et al., 2004) and been recently proposed (Moonen & Ba`rberi, 2008):
(iii) the distribution of trait values for the whole (i) species with high intrinsic conservation value or
community formed by weed and crop plants reects because of sheltered biodiversity, (ii) species contribut-
the competition at the site (McGill et al., 2006). ing to the production of food for animals, or to services
Although simple to parameterise and ecologically of regulation such as pollination or limitation of soil
sound, these models lack accuracy, because demograph- erosion and (iii) species with no intrinsic value but that
ical changes of weeds are not taken into account (but see are used as indicators of high-quality practices. The
McDonald et al., 2010). Therefore, there is an urgent next step is to recognise traits related to properties and
need to identify traits related to demography, because processes that have substantial contribution to services,
only a few studies addressed this question. Some traits following the pioneering framework proposed by De
have been proposed for tropical trees (Garnier & Navas, Bello et al. (2010). Although the positive impact of
2012), and a recent modelling study characterised which weeds on local biodiversity is well known (Ba`rberi et al.,
life-history traits of weeds are more sensitive to man- 2010), no clear set of traits has been related to these
agement (Colbach et al., 2010). services. A recent study performed in oilseed rape, beet
The predictions by these two kinds of models are, and maize crops in the UK showed that functional types
however, of low value when the functional diversity of of weeds, identied by categorical traits, can be associ-
weeds is high under extensive management systems. ated with those of non-pest invertebrate species (Hawes
Some authors have recently suggested that in such et al., 2009). A similar result was found in Mediterra-
conditions, the ability of weeds to draw on resource nean winter wheat elds when classifying weeds into
pools complementarily to that of crop plants, which can three functional groups (Caballero-Lopez et al., 2010).
be assessed by dierences in root depth and architecture On the other hand, the ability of weeds to host other
(Freckleton & Watkinson, 2001; Deen et al., 2003; Smith organisms, which can be positive when weeds act as trap
et al., 2010), might be related to lower weed-crop crops attracting pests (Ba`rberi et al., 2010), or negative
competition (Smith et al., 2010). This means that the when they host disease agents (Franke et al., 2009), has
functional divergence of the crop-weed community not been clearly depicted by traits. An exception is the
dened by traits related to resource use could be of identication of shifted phenology of weeds that host
major importance for predicting crop yield. A high fungi or virus comparatively to crops, allowing the
divergence inducing complementarity in resource use by disease agent to persist between two periods of crop
weeds and crop across time or space, in relation to niche production (Navas et al., 1998).
dierentiation, should result in a reduced impact of Once eect traits are identied, which is something
weeds on crops. This suggests that the limiting similarity still to be done for weeds, it is necessary to evaluate the
eect, one hypothesis explaining co-existence of species shortest set of traits that provision a given service at a
in spontaneous communities [co-existing species slightly site. To that aim, Diaz et al. (2007a) proposed a
dier in functioning then have separated niche dimen- hierarchical procedure that should be used in agro-
sions (Wilson, 2011)], should also prevail in managed ecosystems. Once the eect of the environment on
places. services is identied, CWM and functional divergence of
There are probably other eect traits of weeds that eect traits are assessed, then the impact of some species
could relate to major ecosystem processes. For example, on services, for example the dominant when the biomass
(i) occurrence of herbivores is generally related to tissue ratio hypothesis is valid (Grime, 1998), as it is the case
chemistry, leaf morphology and seed production, when assessing services related to resource use (Garnier
(ii) pollinator provision is explained by oral traits such et al., 2004), is quantied. This procedure was used rst
as accessibility, attractiveness and nectar production and for grasslands and allowed to recognise the best com-

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Functional structure of weeds 485

bination of environmental factors and functional diver- community ecology. First, this will improve the
sity components acting on ecosystem processes and understanding of the impact of disturbances on
services. community assembly. If the impact of resource
The identication of services provided by weeds change on traits is rather well documented, there is
introduces complexity in the denition of control still no general understanding of the impact of
strategies, as the balance between the negative and disturbances on functional structure of plants (see
positive impacts weeds have on the dierent components Garnier & Navas, 2012, for a general discussion). The
of the agro-ecosystem needs to be assessed. Although rather specic disturbances occurring in an agro-
there are some exceptions, weed species cannot be ecosystem (e.g. soil tillage and herbicide application)
grouped into two contrasted groups, the rst including compared with other systems (e.g. re, ooding and
highly competitive species without conservation value herbivory) should allow the general quantication of
and a second one formed of species with opposite processes linked to disturbance. Second, the crop-
characteristics. The reality is in between, as documented weed system is an ideal one to test the hypothesis that
for common weed species of UK, for which no clear co-existence among species is governed by their values
correlation exists between harmfulness and positive of traits, especially those related to competition,
eects on biodiversity (Storkey, 2006). Therefore, pro- because of the asymmetrical relationships between
posing to manage elds to keep only species with the two kinds of plants.
positive eects appears to be unrealistic (Storkey & 3 At the agro-ecosystem level, the understanding of
Westbury, 2007). Furthermore, dening which weeds are how and which traits (in terms of values and ranges)
acceptable because they have more positive than nega- affect processes, properties and services remains very
tive eects remains a challenge, because the former preliminary. The originality of weeds compared with
eects remain poorly understood and largely unquanti- other vegetation of agricultural interest, such as
ed. Potentially, functional traits could be used as a grasslands, is that the balance between their impact
common metric for trading o the positive and negative on ecosystem can vary from purely negative (if only
impacts of contrasting weed communities. 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
Conclusion: directions for weed research
range of trade-offs.
Applying trait-based approaches to weed ecology raises
In conclusion, trait-based approaches should give new
a number of challenges that are somewhat dierent to
tools for predicting weed community assembly and
those proposed for other kinds of vegetation (see
impact on agro-ecosystem services, especially in com-
Garnier & Navas, 2012, for a general discussion). The
plex communities where a detailed mechanistic and
most important dierences can be summarised as
modelling approach based on in-depth knowledge
of all organisms involved will probably not be
1 Concerning plant functioning, whether any easily tractable.
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 The idea of this study came after the publication of a
demography under contrasted management scenarios. review on trait-based approach for agro-ecology
We suspect that such research could be a major (Garnier & Navas, 2012) including a very short section
contribution to community ecology, because weed on crop weeds. I also thank S Gaba, B Chauvel, G Fried
species are characterised by rapidly adapting life- and E Kazakou for very interesting discussions on some
history traits in response to varying environmental of the ideas developed here. Many thanks also to
conditions. referees and editor who carefully read a former version
2 At the community level, understanding the mecha- and provide very useful recommendations for improving
nisms through which species traits determine func- the ideas.
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 ANDERSSON TN & MILBERG P (1998) Weed ora and the
performing comparative experiments. Working with relative importance of site, crop, crop rotation, and nitrogen.
the crop-weed system gives two opportunities for Weed Science 46, 3038.

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486 M-L Navas

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