DOI: 10.1111/1365-2664.
13039
EDITORIAL
Functional traits in agroecology: Advancing description and
prediction in agroecosystems
Adam R. Martin1|Marney E. Isaac1,2
1
Department of Physical and Environmental Sciences,and the Centre for Critical Development of Toronto Scarborough, Toronto, Ontario, Canada
2
Department of Geography,University of Toronto, Toronto, Ontario, Canada
Correspondence
Marney E. Isaac
Email: marney.isaac@utoronto.ca
1| THE IMPACTS AND PARADOXES OF
MO DERN AGRICULTURE
Agricultural expansion and intensification are contributing to the worlds
most pressing environmental issues including global climatic change,
widespread depletion and contamination of soil and water resources,
major disruptions in the Earths biogeochemical cycles and rates of spe-
cies extinctions that are conspicuous on geological time-scales (Burney,
Davis, & Lobell, 2010; Foley etal., 2005, 2011; Vitousek etal., 1997).
Additionally, dominant models of agricultural productionlargely envi-
sioned as intensively managed monocultureshave resulted in wide-
spread financial indebtedness of farmers, declining farmer autonomy in
management decision-making, and the loss of crop genetic resources
(Olson, Morris, & Mendez, 2012; Tomich etal., 2011). Paradoxically as
well, while intensive industrial agriculture has increased food produc-
tion rates to levels that are unprecedented in human history, they have
ultimately resulted in pervasive reductions in global food security and
sovereignty (Altieri & Toledo, 2011) to the extent that issues of food
production, distribution and nutrition occupy a central role in the United
Nations Sustainable Development Goals (SDGs; Prez-Escamilla, 2017).
2| FRAMING THE FIELD OF
AGROECOLOGY
Agroecology as a scientific discipline, an on-farm practice, and a social
or political movement is gaining momentum as a contemporary lens to
critique, evaluate and manage the myriad of global socio-economic and
environmental issues surrounding food production (Mendez, Bacon, &
Cohen, 2013). As a biophysical science, agroecology merges ecology,
global change science, plant biology and soil science in order to address
many of the most pressing issues of our time including food security,
biodiversity loss and climate change (Altieri, 1999; Altieri, Nicholls,
Henao, & Lana, 2015; Lin, 2011; Tomich etal., 2011; Tscharntke etal.,
J Appl Ecol. 2018;55:511. 2017 The Authors. Journal of Applied Ecology | 5
wileyonlinelibrary.com/journal/jpe
2012). But while agroecology is well positioned to address many of
these pressing global issues, the science of agroecology lacks a theo-
retical framework that allows for the development and testing of gen-
eralizable hypotheses; especially those that are broadly relevant for
farm-scale agricultural management, regional-level land-use planning
or international environmental policy. Agroecological research based
on a plant functional trait approach may provide such a generalizable
framework.
3|TRAIT-B ASED ECOLOGY FROM THEORY
TO APPLICATION
In the late 1970s through the 1990s, ecologists began to argue that
differences in ecological strategies among plants represent the basis
for explaining and predicting individual-, species-and ecosystem-level
responses to environmental change (Bazzaz, 1979; Bazzaz & Carlson,
1982; Givnish, 1988; Grime, 1977, 1979, 1988; Lambers & Poorter,
1992; Reich, Walters, & Ellsworth, 1992; Westoby, 1998; Westoby,
Falster, Moles, Vesk, & Wright, 2002). While such studies took a num-
ber of different approachese.g. the competition, stress, disturbance
(CSR) theory (Grime, 1977, 1988), or early-, mid-and late-successional
species classification schemes (e.g. Bazzaz, 1979)such studies were
broadly similar in that they focused on identifying a small number of
functional types that could be used to categorize the ecological dif-
ferences among a large number of plant species. In the 2000s, plant
ecologists began to refine these ideas, by evaluating the ecologically
meaningful characteristics of plants and plant partsor the functional
traitswhich both (1) quantitatively differentiate species from one an-
other along axes of ecological differences and/or functional biology
(Reich etal., 1999; Westoby, 1998; Westoby etal., 2002; Wright etal.,
2004, 2005), and (2) are most important in mechanistically predicting
plant responses to, and impacts on, surrounding environments (Lavorel
& Garnier, 2002).
2017 British Ecological Society
 | Journal of Applied Ecology
6 EDITORIAL
Thanks in part to the immense amount of species trait data that
emerged in the early 2000s (e.g. Wright etal., 2004), which has since
been consolidated into massive trait databases (e.g. Kattge etal.,
2011), studies differentiating trait-based ecological strategies have
now come to encapsulate thousands of plant species (Diaz etal.,
2016), and a global understanding of relationships between ecosys-
tem processes and plant functional biology is being defined and re-
fined (Diaz etal., 2004). On the basis of this rich history of theoretical
and empirical research, over the past 30 years, trait-based ecology has
emerged as one of the dominant paradigms in terrestrial ecology, and
the theory and principles of trait-based ecology are now widely used
to evaluate a remarkable range of questions and hypotheses in global
change biology (e.g. Garnier, Navas, & Grigulis, 2016).
From an applied perspective, trait-
based ecology has more
recently been embraced as a critical means by which scientists can
test hypotheses on, and recommend management of, managed ter-
restrial ecosystems (e.g. Cadotte, 2011 and references therein). The
transition of trait-based ecology into resource management fields was
facilitated by clear empirical evidence that plant functional traits, and
functional trait diversity are strong mechanistic predictors of ecosys-
tem functioning (Cadotte, Cavender-Bares, Tilman, & Oakley, 2009;
reviewed by Garnier etal., 2016). Scientists are now adopting trait-
based approaches to research in many applied fields including resto-
ration ecology (e.g. Funk, Cleland, Suding, & Zavaleta, 2008; Laughlin,
2014), urban ecology (Duncan etal., 2011) and more recently, agro-
ecology (Garnier & Navas, 2012; Martin & Isaac, 2015; Milla, Osborne,
Turcotte, & Violle, 2015; Wood etal., 2015).
4| TRAIT-B ASED AGROECOLOGY
Opinions that trait-based ecology might be efficacious in addressing
major questions in agricultural systems first emerged over 5years ago
in a review by Garnier and Navas (2012) that summarized the con-
cepts, tools and applications of functional ecology within agroecology.
Then in 2015, three review papers published near-simultaneously
(Martin & Isaac, 2015; Milla etal., 2015; Wood etal., 2015) summa-
rized in greater detail the types of broad agroecological questions,
hypotheses and management decisions that might be informed by
trait-based research. Specifically, these papers reviewed how trait-
based ecology could address questions and hypotheses surrounding
the ecological consequences of crop breeding (Milla, Morente-Lopez,
Alonso-Rodrigo, Martin-Robles, & Chapin, 2014; Milla etal., 2015);
how functional traits underpin crop yield and yield models (Gagliardi,
Martin, Virginio, Rapidel, & Isaac, 2015); how functional traits influ-
ence agricultural nutrient cycling (Garcia-Palacios etal., 2013); and
how functional diversity inform an understanding of agricultural con-
tributions to global net primary productivity (Monfreda, Ramankutty,
& Foley, 2008). But while these and other examples were drawn from
the literature, these review papers (Martin & Isaac, 2015; Milla etal.,
2015; Wood etal., 2015) also highlighted the lack of research employ-
ing trait-based approaches to test hypotheses on linkages between
on-farm diversity and agroecosystem functioning.
This lack of research requires some qualification. Plant breeders, crop
biologists, agronomists, ecophysiologists and farmers have certainly al-
located considerable attention to understanding and selecting for crops
that express certain reproductive, leaf, root, phenological and chemical
traits (e.g. Meister, Rajani, Ruzicka, & Schachtman, 2014; Meyer, DuVal,
& Jensen, 2012). However, this vast amount of scientific and informal
on-farm research tends to focus on understanding and managing link-
ages between only certain functional traitsthose forming part of a crop
domestication syndrome (Meyer etal., 2012)and yield.
In comparison, trait-based agroecology as envisioned by applied
ecologists (Garnier & Navas, 2012; Martin & Isaac, 2015; Milla etal.,
2015; Wood etal., 2015), and expanded upon by the research in this
Special Feature, seeks to understand how knowledge of the variation
in the functional traits and trait diversity of both crop and non-crop
species can be used to predict, manage and enhance multiple critical
ecosystems functions other than yield alone. Discussing the value and
potential application of trait-based agroecology must deal with issues
of yield to some degree, as it is the key ecosystem service upon which
many livelihoods are predicated. But this Special Feature has a broader
goal, in seeking to significantly advance a trait-based understanding of
how a wider range of above-and below-ground agroecosystem func-
tions are linked to functional trait diversity among and within both crop
and non-crop species.
This Special Feature includes seven studies that employ functional
trait-
based approaches, to frame, test and interpret management-
relevant hypotheses in agroecosystems. In doing so, cumulatively
these studies provide arguably the single largest empirical and con-
ceptual contribution to a trait-based understanding of agroecological
structure and function. Specifically, these studies provide novel in-
sights and evaluations into a wide range of agroecological questions,
including how functional trait diversity is driving agricultural nutrient
cycles and below-
ground processes (Blesh, 2017; Pommier etal.,
2017); how knowledge of functional traits can inform the restoration
of degraded agricultural land (Lohbeck, Winowiecki, Aynekulu, Okia, &
Vgen, 2017); how functional traits can be used to infer the ecological
consequences of crop domestication (Roucou etal., 2017); how func-
tional diversity-based approaches can be used to assess nutritional
diversity in agricultural systems (Wood, 2017); and lastly, how farm-
ers value, and make management decisions based on, functional traits
of crop and non-crop species (Damour, Navas, & Garnier, 2017; Isaac
etal., 2017).
5|WHAT DO WE KNOW ABOUT
FUNCTIONAL TRAITS IN CROPS AND
AGROECOSYSTEMS?
In this Special Feature, Damour etal. (2017) make a major concep-
tual contribution to a broader understanding of the potential applica-
tions of trait-based agroecology, though a revised response/effect
framework that aids researchers in hypothesizing linkages and feed-
backs between plant functional diversity and ecosystem processes
(Lavorel & Garnier, 2002). In their revised trait-
based response
EDITORIAL
and effect framework, Damour etal. (2017) outline hypotheses on
how, and at which stages of management, knowledge of plant func-
tional traits enters into agroecological management prescriptions.
Damour etal. (2017) outline a key differentiation that functional
diversity in agroecosystems (and in turn rates of agroecosystem
service provisioning) is comprised of both planned and spontane-
ous plant diversity, drawn from a pool of available crop cultivars,
and non-crop species (such as weeds) drawn from regional species
pool respectively. The processes that result in these different func-
tional diversity components are governed by a set of longer term
strategic management decisions, shorter term on-farm tactical deci-
sions, as well as multiple interacting socio-economic and biophysi-
cal constraints operating in different farms, regions or landscapes.
Damour etal. (2017) then discuss how all of these factors lead to
prospective differences in the functional profiles of the targeted
agroecological plant communities (i.e. plant assemblages envisioned
by farmers when making management decisions) vs. the realized
plant communities (i.e. plant communities actually emerging follow-
ing interactions between management decisions and the biophysical
environment). In hypothesizing a sequence under which these dif-
ferent process occur, as well as the factors governing each stage,
Damour etal. (2017) contribute a theoretical framework that can be
employed to evaluate multiple linkages between different stages of
management, on-farm environmental change, functional trait diver-
sity, and the structure and functioning of agroecosystems.
6|FUNCTIONAL TRAIT ECOLOGY AND
CROP DOMESTICATION
Prior reviews and research in the field of trait-based agroecology
have highlighted how intense artificial selection has resulted in major
shifts in functional trait syndromes of crops (Martin etal., 2017;
Milla etal., 2014, 2015), and how this variation then governs rates
of agroecosystem functioning (e.g. Garcia-Palacios etal., 2013). Of all
themes addressed in the trait-based agroecology literature to date,
this area has arguably the strongest empirical support from stud-
ies comparing functional trait syndromes in crops vs. wild ancestors
(e.g. Milla etal., 2014). In this Special Feature, Roucou etal. (2017)
make a major contribution to the theme of understanding the traits
other than those strictly related to yield that have also been selected
for during the process of crop domestication. Through a compara-
tive evaluation of above-and below-ground functional trait variation
among 40 different accessions of wheat (Triticum turgidum), Roucou
etal. (2017) provide strong empirical support for the hypothesis that
due to evolutionary constraints in functional trait trade-offs, crop
breeding has (likely inadvertently) also selected for suites of traits
that favour high rates of resource acquisition. Their contribution is
essential in defining a more comprehensive understanding of the eco-
logical consequences of an expanded crop domestication syndrome
(cf. Meyer etal., 2012), and presents lines of evidence that support
the concept of functional trait-based screening for future crops (Milla
etal., 2015). More practically, their research presents novel ideas
Journal of Applied Ecology |
7
surrounding how complementary functional trait syndromes can
contribute to the ecologically based selection of genotypes for inter-
cropped agroecosystems.
7|FUNCTIONAL TRAIT ECOLOGY FOR
MULTI-F UNCTIONALITY IN
AGROECOSYSTEMS
In the applied ecology literature, studies that evaluate the complex
linkages between functional traits, functional diversity and ecosys-
tem functioning have recently begun to explore concepts surround-
ing multi-functionality: the idea that multiple ecosystem functions
should be considered as a management goal, vs. one or a narrow set
of functions (Gamfeldt, Hillebrand, & Jonsson, 2008). The concept of
multi-functionality has clear implications for trait-based agroecology,
where managers may manipulate on-farm diversity to enhance certain
ecosystem services beyond yield alone, or alternatively, minimize the
net-negative trade-offs that might occur among ecosystem functions
(Finney & Kaye, 2017; Schipanski etal., 2014).
Research by Blesh (2017) addresses this theme by evaluating hy-
potheses on how functional diversity in temperate North American
organic polyculture systems influences multiple agroecosystem func-
tions. Blesh (2017) contributes novel empirical evidence on how the
on-
farm selection of crops with complementary functional traits,
across a gradient of management-relevant environmental conditions,
enhances the multi-functionality of agroecosystems including biologi-
cal nitrogen (N) fixation (BNF), soil N retention and weed suppression.
Blesh (2017) also contributes an important conceptual advancement
to applied trait-based ecology, through a more detailed discussion
on BNF in functional ecology. On the one hand, trait-based studies
commonly view the ability to fix atmospheric N as a binary functional
trait, most widely observed in plants in the Fabaceae family (Perez-
Harguindeguy etal., 2013); however, Blesh (2017) points to the fact
that in agroecosystems, quantitative rates of BNF should be viewed
as a key ecosystem function. The work of Blesh (2017) therefore in-
dicates that certain conventions in trait-based ecology surrounding
how plant functional traits are defined need to be revisited when
applied in agroecological contexts. From a management perspective,
Blesh (2017) makes an important contribution to understanding how
inter-and intraspecific trait variation across environmental gradients
and management systems plays a key role in mediating relationships
between functional diversity on agroecosystem functioning.
Of all the potential applications of trait-
based agroecology,
understanding how traits mediate multiple ecological interactions
that impact different aspects of nutrient cycles is arguably of the
highest importance to farmers and land managers. At the same time,
a key theme at the leading edge of trait-based ecology is better un-
derstanding how to characterize the diversity and role of functional
traits in microbial species (Green, Bohannan, & Whitaker, 2008).
Pommier etal. (2017) address both of these themes, by offering a
robust example of how a functional trait-based approach provides
new insights into our understanding and prediction of linkages
 | Journal of Applied Ecology
8 EDITORIAL
between microbial diversity and nutrient cycles. Pommier etal.
(2017) distil trait-mediated interactions among plant and microbial
species that affect nitrogen (N) dynamics in European-
managed
grasslands agroecosystems, showing that variation in nitrate and
ammonium leaching and soil organic matter content is largely ex-
plained by microbial traits including the maximum rate of nitrifica-
tion and fungi: bacteria ratios. To date, this contribution by Pommier
etal. (2017) is among very few studies that incorporate diversity
of microbial traits with soil and plant metrics, representing an im-
portant advancement in the understanding of microbial-controlled
ecosystem services in agroecosystems.
8| FUNCTIONAL TRAIT ECOLOGY FOR
LAND MANAGEMENT AFTER THE CROPS
HAVE GONE
Agricultural abandonment has also emerged as a major global socio-
economic and land-
use trend, with major implications for human
well-being, global economic systems and the Earths biophysical en-
vironment. In turn, restoration of agricultural lands is now emerging
as both a challenge and opportunity for conservation biology (e.g.
Queiroz, Beilin, Folke, & Lindborg, 2014), climate change mitigation
(e.g. Pan etal., 2011) and the revitalization of other ecological pro-
cesses such as nutrient cycling (e.g. Knops & Tilman, 2000). Trait-
based approaches to the planning and managing of biodiversity and
ecosystem functioning in abandoned agricultural lands represent a
major emerging vanguard for policymakers and practitioners.
In this Special Feature, Lohbeck etal. (2017) provide empirical ev-
idence on how plant functional traits and trait diversity is linked with
the restoration of ecosystem functions in degraded agricultural lands.
Lohbeck etal. (2017) show that enhancement of functional diversity
in degraded East African agroecosystems is positively associated with
key parameters of soil remediation including increased organic carbon
stocks and decreased soil erosion, while the abundance of invasive
species is negatively associated with these parameters. Lohbeck etal.
(2017) then suggest that strategies for restoring degraded agricultural
land should be informed by principles of functional ecology, with con-
crete policy-and practice-relevant suggestions that include farmer-
lead tree selection during restoration processes.
9| FUNCTIONAL TRAIT ECOLOGY FOR
AGRICULTURAL LIVELIHOODS AND POLICY
While the conceptual (Damour etal., 2017) and empirical (Blesh, 2017;
Lohbeck etal., 2017; Pommier etal., 2017; Roucou etal., 2017) studies
in this Special Feature address major gaps in the scientific literature of
agroecology, there remains an major elephant in the trait-based agro-
ecology room: do farmers and policymakers base their decisions on
the same traits that ecologists consider vital to ecosystem functioning?
Earlier papers have summarized the scant evidence that farmers
make decisions based on functional ecology prioritized traits (Martin &
Isaac, 2015). But here, Isaac etal. (2017) tackle this challenging theme
directly by analysing farmer perceptions on how variation in the leaf
economics traits of coffee (Coffea arabica) influence the management
and understanding of agroecosystem functions. Through in-depth in-
terviews and surveys, coupled with a visual elicitation tool dubbed the
leaf book, Isaac etal. (2017) show that farmers diagnose major man-
agement concerns based on, and adjust their practices at least in part
according to, intraspecific variation in leaf economics traits. This research
represents a major contribution to the literature on how farmers em-
ploy an understanding of plant functional traits other than yield alone
to manage agroecological systems. From a wider ecological perspective,
this research stands as the first to extend a major theme in the functional
ecology literaturethe concept of trait trade-offs and spectra (Wright
etal., 2004)into the sphere of local ecological knowledge. In theoriz-
ing the presence of a Farmer Economics Spectrum, Isaac etal. (2017)
contribute a conceptual advancement through their hypothesis that land
managers acknowledge and make decisions based on the trait trade-offs
that underpin inter-and intraspecific variation in plant functional biology.
Wood (2017) also addresses the policy relevance of functional
trait-based agroecology, through a novel empirical analysis that ex-
tends methods in functional ecologynamely functional diversity met-
ricsinto the realm of human nutrition. Broadly, Wood (2017) presents
arguments that trait-based agroecology must focus beyond the traits
that mediate biophysical interactions, and instead also consider traits
that are empirically linked with human well-being or other social as-
pects of agroecosystems. Wood (2017) then illustrates the value of
this concept, by employing principles of functional trait diversity to de-
rive a new nutritional diversity metric: potential nutritional adequacy
(PNA). This metric represents a major advance in how plant functional
trait diversity can be empirically linked with the nutritional outcomes
associated with enhancing on-farm diversity. In his application of this
new metric using data from Senegal, Wood (2017) shows that nei-
ther intensification nor diversification represent a singular approach
to increasing PNA in small-holder agroecosystems; this represents a
more measured and policy-realistic approach to addressing questions
of food security, as compared to arguments that fall on one side of the
intensification vs. extensification debate (Foley etal., 2011). Wood
(2017) then proposes that PNA could be paired with other indicators
of environmental outcomes as a means to assess the joint benefits
of, or prospective trade-offs among, the provisioning of food security,
food sovereignty and other ecosystem services in agroecosystems.
10|FUNCTIONAL TRAIT ECOLOGY
FOR THE FUTURE OF SUSTAINABLE
AGRICULTURE
The research in this Special Feature demonstrates how a functional
trait-based approach represents meaningful advances in integrating
leading edge ecological sciences into an understanding of the link-
ages between agricultural diversity and agroecosystem functioning.
The papers here address themes in agriculture that span from genetic
to regional scales of investigation. Taken as a whole, we hope that
EDITORIAL
they represent at least a small step in having trait-based agroecology
contribute to the sustainable management of agricultural systems, the
restoration of degraded agricultural lands, the understanding of the
crop nutrition-food security nexus and the overall enhancement of
agroecosystem functioning. Based on the empirical and conceptual
advances put forth in this Special Feature, clearly there is a role for a
trait-based approach to agroecology in guiding the composition and
function of sustainable agroecosystems.
Contemporary sustainable agriculture is more relevant than ever,
particularly under the United Nations SDGs launched in 2015: 17 global
goals with 169 total targets, designed to frame a global sustainable de-
velopment agenda through the year 2030. Generally, the SDGs fall into
the two primary realms: socio-economic development such as human
health, education, social justice and inequality, and environmental sus-
tainability including clean water provisioning, climate change mitiga-
tion and adaptation, and terrestrial and aquatic resource conservation.
Many of the SDGs straddle this broad divide, perhaps no more than
SDG 2 that calls for End hunger, achieve food security and improved
nutrition, and promote sustainable agriculture. This SDG, which entails
targets surrounding agricultural production and sustainability, provides
a clear opportunity for trait-based agroecology to inform a better un-
derstanding of the types of farming systems that can move towards
meeting these targets. For instance, SDG 2 Target 2.4 calls for the im-
plementation of resilient agricultural practices that increase produc-
tivity and production, that help maintain ecosystems, that strengthen
capacity for adaptation to climate change, extreme weather, drought,
flooding and other disasters, and that progressively improve land and
soil quality. Sustainable Development Goal 2 Target 2.5 calls for the
maintenance of a genetic diversity of seeds, cultivated plants, farmed
and domesticated animals and their related wild species, including
through soundly managed and diversified seed and plant banks.
If not explicitly, the promise of trait-based agroecology is well-
encapsulated by language within the SDGs that indicates a strong role
for agroecology (and applied ecology more broadly) in helping to address
some of the worlds most pressing socio-economic and environmental
concerns. In agriculture broadly, the realized and potential impacts of
climate change, depletion of soil resources, as well as the loss of crop
and non-crop diversity, require new approaches to frame, test and pre-
scribe management for resilient agroecosystems. Trait-based ecology
has advanced our understanding of the causes and consequences of
shifts in ecological functioning of natural ecosystems, and based on the
research in this Special Feature, represents a novel means of advancing
our understanding of agroecosystem structure, function and perfor-
mance. Most importantly, as this Special Feature demonstrates, trait-
based approaches can play a key role in diagnosing cropenvironment
interactions, predicting crop and non-crop interactions, understanding
future impacts of crop domestication and, ultimately, prescribing eco-
logically sound and sustainable management practices.
ACKNOWLE DG E MEN TS
The authors thank Marc W. Cadotte, Jos Barlow, Alice Plane and
Kirsty Lucas for guidance, advice and technical assistance in
Journal of Applied Ecology
9|
organizing this Special Feature. The authors also thank all authors
for their contributions (and patience), and also extend their grati-
tude to the participants of the special session on Functional traits in
tropical agroecology at the 2016 Annual Meeting of the Association
of Tropical Biology and Conservation, where earlier discussions on
trait-based agroecology were held. M.E.I. is supported by a Natural
Sciences and Engineering Research Council of Canada (NSERC)
Discovery Grant, as well the Canada Research Chairs program.
DATA ACC ES S I B I L I T Y
Data have not been archived because this article does not contain
data.
REFERENCES
Altieri, M. A. (1999). The ecological role of biodiversity in agroecosystems.
Agriculture Ecosystems & Environment, 74, 1931.
Altieri, M. A., Nicholls, C. I., Henao, A., & Lana, M. A. (2015). Agroecology
and the design of climate change-resilient farming systems. Agronomy
for Sustainable Development, 35, 869890.
Altieri, M. A., & Toledo, V. M. (2011). The agroecological revolution in Latin
America: Rescuing nature, ensuring food sovereignty and empowering
peasants. Journal of Peasant Studies, 38, 587612.
Bazzaz, F. A. (1979). Physiological ecology of plant succession. Annual
Review of Ecology and Systematics, 10, 351371.
Bazzaz, F. A., & Carlson, R. W. (1982). Photosynthetic acclimation to vari-
ability in the light environment of early and late successional plants.
Oecologia, 54, 313316.
Blesh, J. (2017). Functional traits in cover crop mixtures: Biological nitrogen
fixation and multifunctionality. Journal of Applied Ecology, 55, 3848.
Burney, J. A., Davis, S. J., & Lobell, D. B. (2010). Greenhouse gas mitigation
by agricultural intensification. Proceedings of the National Academy of
Sciences of the USA, 107, 1205212057.
Cadotte, M. W. (2011). The new diversity: Management gains through in-
sights into the functional diversity of communities. Journal of Applied
Ecology, 48, 10671069.
Cadotte, M. W., Cavender-Bares, J., Tilman, D., & Oakley, T. H. (2009).
Using phylogenetic, functional and trait diversity to understand pat-
terns of plant community productivity. PLoS ONE, 4, e5695.
Damour, G., Navas, M.-L., & Garnier, E. (2017). A revised trait-based frame-
work for agroecosystems including decision rules. Journal of Applied
Ecology, 55, 12-24.
Diaz, S., Hodgson, J. G., Thompson, K., Cabido, M., Cornelissen, J. H. C., Jalili,
A., Zak, M. R. (2004). The plant traits that drive ecosystems: Evidence
from three continents. Journal of Vegetation Science, 15, 295304.
Diaz, S., Kattge, J., Cornelissen, J. H. C., Wright, I. J., Lavorel, S., Dray, S.,
Gorne, L. D. (2016). The global spectrum of plant form and function.
Nature, 529, U167U173.
Duncan, R. P., Clemants, S. E., Corlett, R. T., Hahs, A. K., McCarthy, M. A.,
McDonnell, M. J., Williams, N. S. G. (2011). Plant traits and extinc-
tion in urban areas: A meta-analysis of 11 cities. Global Ecology and
Biogeography, 20, 509519.
Finney, D. M., & Kaye, J. P. (2017). Functional diversity in cover crop poly-
cultures increases multifunctionality of an agricultural system. Journal
of Applied Ecology, 54, 509517.
Foley, J. A., DeFries, R., Asner, G. P., Barford, C., Bonan, G., Carpenter, S.
R., Snyder, P. K. (2005). Global consequences of land use. Science,
309, 570574.
Foley, J. A., Ramankutty, N., Brauman, K. A., Cassidy, E. S., Gerber, J. S.,
Johnston, M., Zaks, D. P. M. (2011). Solutions for a cultivated planet.
Nature, 478, 337342.
 |Journal of Applied Ecology
10
Funk, J. L., Cleland, E. E., Suding, K. N., & Zavaleta, E. S. (2008). Restoration
through reassembly: Plant traits and invasion resistance. Trends in
Ecology & Evolution, 23, 695703.
Gagliardi, S., Martin, A. R., Virginio, E. D., Rapidel, B., & Isaac, M. E. (2015).
Intraspecific leaf economic trait variation partially explains coffee
performance across agroforestry management regimes. Agriculture
Ecosystems & Environment, 200, 151160.
Gamfeldt, L., Hillebrand, H., & Jonsson, P. R. (2008). Multiple functions in-
crease the importance of biodiversity for overall ecosystem function-
ing. Ecology, 89, 12231231.
Garcia-Palacios, P., Milla, R., Delgado-Baquerizo, M., Martin-Robles, N.,
Alvaro-Sanchez, M., & Wall, D. H. (2013). Side-effects of plant domes-
tication: Ecosystem impacts of changes in litter quality. New Phytologist,
198, 504513.
Garnier, E., & Navas, M. L. (2012). A trait-based approach to comparative
functional plant ecology: Concepts, methods and applications for agro-
ecology. A review. Agronomy for Sustainable Development, 32, 365399.
Garnier, E., Navas, M. L., & Grigulis, K. (2016). Plant functional diversity:
Organism traits, community structure, and ecosystem properties. Oxford,
UK: Oxford University Press.
Givnish, T. J. (1988). Adaptation to sun and shade A whole-plant perspec-
tive. Australian Journal of Plant Physiology, 15, 6392.
Green, J. L., Bohannan, B. J. M., & Whitaker, R. J. (2008). Microbial biogeog-
raphy: From taxonomy to traits. Science, 320, 10391043.
Grime, J. P. (1977). Evidence for the existence of three primary strate-
gies in plants and its relevance to ecological and evolutionary theory.
American Naturalist, 111, 11691194.
Grime, J. P. (1979). Plant strategies and vegetation processes. New York, NY:
John Wiley and Sons.
Grime, J. P. (1988). The CSR model of primary plant strategiesorigins, im-
plications and tests. In L. Gottlieb & S. Jain (Eds.), Plant evolutionary biol-
ogy (pp. 371393). Dordrecht, the Netherlands: Springer Netherlands.
Isaac, M. E., Cerda, R., Rapidel, B., Martin, A. R., Dickinson, A. K., & Sibelet,
N. (2017). Farmer perception and utilization of leaf functional traits
inmanaging agroecosystems. Journal of Applied Ecology, 55, 69-80.
Kattge, J., Diaz, S., Lavorel, S., Prentice, C., Leadley, P., Bonisch, G., Wirth,
C. (2011). TRY A global database of plant traits. Global Change Biology,
17, 29052935.
Knops, J. M. H., & Tilman, D. (2000). Dynamics of soil nitrogen and carbon
accumulation for 61 years after agricultural abandonment. Ecology, 81,
8898.
Lambers, H., & Poorter, H. (1992). Inherent variation in growth-rate be-
tween higher-plants A search for physiological causes and ecological
consequences. Advances in Ecological Research, 23, 187261.
Laughlin, D. C. (2014). Applying trait-based models to achieve functional
targets for theory-driven ecological restoration. Ecology Letters, 17,
771784.
Lavorel, S., & Garnier, E. (2002). Predicting changes in community compo-
sition and ecosystem functioning from plant traits: Revisiting the Holy
Grail. Functional Ecology, 16, 545556.
Lin, B. B. (2011). Resilience in agriculture through crop diversification: Adaptive
management for environmental change. BioScience, 61, 183193.
Lohbeck, M., Winowiecki, L., Aynekulu, E., Okia, C., & Vgen, T. (2017).
Trait-based approaches for guiding the restoration of degraded agri-
cultural landscapes in East Africa. Journal of Applied Ecology, 55, 59-68.
Martin, A. R., & Isaac, M. E. (2015). Plant functional traits in agroeco-
systems: A blueprint for research. Journal of Applied Ecology, 52,
14251435.
Martin, A. R., Rapidel, B., Roupsard, O., van den Meersche, K., de M. Virginio Filho,
K. Isaac, M. E. (2017). Intraspecific trait variation across multiple scales: The
leaf economics spectrum in coffee. Functional Ecology, 31, 604612.
Meister, R., Rajani, M. S., Ruzicka, D., & Schachtman, D. P. (2014). Challenges
of modifying root traits in crops for agriculture. Trends in Plant Science,
19, 779788.
EDITORIAL
Mendez, V. E., Bacon, C. M., & Cohen, R. (2013). Agroecology as a transdis-
ciplinary, participatory, and action-oriented approach. Agroecology and
Sustainable Food Systems, 37, 318.
Meyer, R. S., DuVal, A. E., & Jensen, H. R. (2012). Patterns and processes
in crop domestication: An historical review and quantitative analysis of
203 global food crops. New Phytologist, 196, 2948.
Milla, R., Morente-Lopez, J., Alonso-Rodrigo, J. M., Martin-Robles, N., &
Chapin, F. S. (2014). Shifts and disruptions in resource-use trait syn-
dromes during the evolution of herbaceous crops. Proceedings of the
Royal Society B, 281, pii: 20141429.
Milla, R., Osborne, C. P., Turcotte, M. M., & Violle, C. (2015). Plant domes-
tication through an ecological lens. Trends in Ecology & Evolution, 30,
463469.
Monfreda, C., Ramankutty, N., & Foley, J. A. (2008). Farming the planet: 2.
Geographic distribution of crop areas, yields, physiological types, and
net primary production in the year 2000. Global Biogeochemical Cycles,
22, GB1022. https://doi.org/10.1029/2007GB002947
Olson, M. B., Morris, K. S., & Mendez, V. E. (2012). Cultivation of maize
landraces by small-scale shade coffee farmers in western El Salvador.
Agricultural Systems, 111, 6374.
Pan, Y. D., Birdsey, R. A., Fang, J. Y., Houghton, R., Kauppi, P. E., Kurz, W. A.,
Hayes, D. (2011). A large and persistent carbon sink in the worlds
forests. Science, 333, 988993.
Prez-Escamilla, R. (2017). Food security and the 20152030 sustain-
able development goals: From human to planetary health. Current
Developments in Nutrition, 1, e000513.
Perez-Harguindeguy, N., Diaz, S., Garnier, E., Lavorel, S., Poorter, H.,
Jaureguiberry, P., Cornelissen, J. H. C. (2013). New handbook for
standardised measurement of plant functional traits worldwide.
Australian Journal of Botany, 61, 167234.
Pommier, T., Cantarel, A. A. M., Grigulis, K., Lavorel, S., Legay, N.,
Baxendale, C., Clment, J.-C. (2017). The added value of including
key microbial traits to determine nitrogen-related ecosystem ser-
vices in managed grasslands. Journal of Applied Ecology, 55, 4958.
Queiroz, C., Beilin, R., Folke, C., & Lindborg, R. (2014). Farmland abandon-
ment: Threat or opportunity for biodiversity conservation? A global re-
view. Frontiers in Ecology and the Environment, 12, 288296.
Reich, P. B., Ellsworth, D. S., Walters, M. B., Vose, J. M., Gresham, C., Volin,
J. C., & Bowman, W. D. (1999). Generality of leaf trait relationships: A
test across six biomes. Ecology, 80, 19551969.
Reich, P. B., Walters, M. B., & Ellsworth, D. S. (1992). Leaf life-span in rela-
tion to leaf, plant, and stand characteristics among diverse ecosystems.
Ecological Monographs, 62, 365392.
Roucou, A., Violle, C., Fort, F., Roumet, P., Ecarnot, M., & Vile, D. (2017).
Shifts in plant functional strategies over the course of wheat domesti-
cation. Journal of Applied Ecology, 55, 2537.
Schipanski, M. E., Barbercheck, M., Douglas, M. R., Finney, D. M., Haider,
K., Kaye, J. P., White, C. (2014). A framework for evaluating ecosys-
tem services provided by cover crops in agroecosystems. Agricultural
Systems, 125, 1222.
Tomich, T. P., Brodt, S., Ferris, H., Galt, R., Horwath, W. R., Kebreab,
E., Yang, L. (2011). Agroecology: A review from a global-c hange
perspective. Annual Review of Environment and Resources, 36,
193222.
Tscharntke, T., Clough, Y., Wanger, T. C., Jackson, L., Motzke, I., Perfecto, I.,
Whitbread, A. (2012). Global food security, biodiversity conservation
and the future of agricultural intensification. Biological Conservation,
151, 5359.
Vitousek, P. M., Aber, J. D., Howarth, R. W., Likens, G. E., Matson, P. A.,
Schindler, D. W., Tilman, D. (1997). Human alteration of the global
nitrogen cycle: Sources and consequences. Ecological Applications, 7,
737750.
Westoby, M. (1998). A leaf- height- seed (LHS) plant ecology strategy
scheme. Plant and Soil, 199, 213227.
EDITORIAL
Westoby, M., Falster, D. S., Moles, A. T., Vesk, P. A., & Wright, I. J. (2002).
Plant ecological strategies: Some leading dimensions of variation be-
tween species. Annual Review of Ecology and Systematics, 33, 125159.
Wood, S. A. (2017). Nutritional functional trait diversity of crops in south-
eastern Senegal. Journal of Applied Ecology, 55, 81-91.
Wood, S. A., Karp, D. S., DeClerck, F., Kremen, C., Naeem, S., & Palm, C. A.
(2015). Functional traits in agriculture: Agrobiodiversity and ecosystem
services. Trends in Ecology & Evolution, 30, 531539.
Journal of Applied Ecology |
11
Wright, I. J., Reich, P. B., Cornelissen, J. H. C., Falster, D. S., Garnier, E.,
Hikosaka, K., Westoby, M. (2005). Assessing the generality of global
leaf trait relationships. New Phytologist, 166, 485496.
Wright, I. J., Reich, P. B., Westoby, M., Ackerly, D. D., Baruch, Z., Bongers,
F., Villar, R. (2004). The worldwide leaf economics spectrum. Nature,
428, 821827.