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In proposing the ecosystem as an organizational frame-
work, Tansley (1935) explicitly included the effects of
humans. When asking, “Is man part of ‘nature’ or not?”,
Tansley concluded that “human activity finds its proper
place in ecology”. Nowhere is the human–nature inter-
face more unmistakable than in urban ecosystems, and
urban ecology has quickly become established within the
discipline of ecology (Grimm et al. 2013). This rapid
development was fueled by a need for multidisciplinary
approaches when studying the extraordinary rate of
urbanization and the complex connections among socio-
logical and environmental factors (Brennan 1999; Young
2009). Now, for the first time in recorded history, the
majority of people live in cities; moreover, with the char-
acteristics of cities foreshadowing those of the planet,
urban ecology is playing an increasingly important role
within ecology (Grimm et al. 2008a). Furthermore, urban
ecosystems will be on the “front lines” of environmental
and sociological change, providing urban ecologists with
new and exciting possibilities that combine research with
implementing urban design and planning, as well as
advocating policy.
Across the biological sciences, efforts to understand the
mechanisms that underlie patterns have found both inspi-
ration and a worthy challenge in the city (Grimm et al.
2008b). Human amplification of ecological processes in
urban areas tightens the links among these processes at
different biological, temporal, and spatial scales. For
example, disease spread through co-occurring populations
of wild animals and household pets depends on the behav-
ior, evolution, and immunological responses of both the
wild and the domesticated animals; local regulations; pet
owner behavior; and the habitat configuration, nutrient
availability, and climate of the broader ecosystem (Adler
and Tanner 2013). Ecologists have developed principles to
help understand such processes, but considering them
within an urban framework demands a more integrative
way of thinking (Grimm et al. 2000; Forman 2008a).
Pickett et al. (2008) characterized the challenges urban
ecologists face: “urban ecosystems are complex, dynamic
biological–physical–social entities, in which spatial het-
erogeneity and spatially localized feedbacks play a large
role”. Addressing such challenges requires interdiscipli-
nary thinking and, depending on the goals of a project,
CONCEPTS AND QUESTIONS
Urban ecology: advancing science and
society
Colby J Tanner1*, Frederick R Adler2, Nancy B Grimm3, Peter M Groffman4, Simon A Levin5,
Jason Munshi-South6, Diane E Pataki7, Mitchell Pavao-Zuckerman8, and William G Wilson9
Urban ecology has quickly become established as a central part of ecological thinking. As cities continue to
grow in size and number, two questions serve to unify this broad and multidisciplinary research landscape: (1)
how can urban ecology contribute to the science of ecology, and (2) how can urban ecology be applied to make
cities more livable and sustainable? In spite of the advances made thus far, there are many unexplored ways of
integrating the science and application of urban ecology. Although scientists assess and make predictions
regarding the connections between environmental and socioeconomic processes, practitioners involved in
real-world application deal with urban planning and with designing ecosystem services to improve living con-
ditions for all urban inhabitants and to make cities more sustainable. Research in urban ecosystems can be
developed from many different perspectives, and we suggest that each perspective has something to offer both
society and the science of ecology. We present several research perspectives and describe how these can inte-
grate conceptual and applied aspects to bridge the figurative gaps between trees, buildings, and people.
Front Ecol Environ 2014; 12(10): 574–581, doi:10.1890/140019
1School of Biological Sciences, University of Nebraska–Lincoln,
Lincoln, NE *(colbyjtanner@gmail.com); 2Departments of Biology
and Mathematics, University of Utah, Salt Lake City, UT; 3School of
Life Sciences, Arizona State University, Tempe, AZ; 4Cary Institute of
Ecosystem Studies, Millbrook, NY; 5Department of Ecology and
Evolutionary Biology, Princeton University, Princeton, NJ;
6Department of Biological Sciences, Fordham University, Bronx, NY;
7Department of Biology, University of Utah, Salt Lake City, UT;
8Biosphere 2 and Department of Ecology and Evolutionary Biology,
University of Arizona, Tucson, AZ; 9Department of Biology, Duke
University, Durham, NC
In a nutshell:
•Taking the emerging discipline of urban ecology “to the next
level” of scientific understanding and practical application
requires approaches that link the biophysical and social sci-
ences with planning, design, and management
•Integrating ideas and methods from various disciplines –
including infrastructure sciences; organismal and evolutionary
biology; critical geography; sociology; and ecosystem, behav-
ioral, and political ecology – has great potential in advancing
the field of urban ecology
CJ Tanner et al. Urban ecology: advancing science and society
575
© The Ecological Society of America www.frontiersinecology.org
perhaps also an interdisciplinary team
(Collins et al. 2000; Cid and Pouyat
2013; McHale et al. 2013). Indeed, an
urban ecology study might require a team
composed of ecologists, economists, soci-
ologists, meteorologists, hydrologists,
health-care professionals, landscape
designers, planners, and politicians (Bet-
tencourt and West 2010). Such broad,
multifaceted collaborations have made
remarkable progress in understanding
the holistic social–biophysical–ecologi-
cal processes of urbanization over the
past several decades (Pickett et al. 2008;
Collins et al. 2011; Grimm et al. 2013).
Although our understanding of urban
ecosystems is expanding rapidly, how to
use this growing body of knowledge to
the benefit of society is much less well
established (Brennan 1999; Leach et al.
2010; Cote and Nightingale 2012).
As scientists continue to learn more
about the intended and unintended con-
sequences of engineering built environ-
ments, they are in a position to begin
asking what urban ecology can do for the
science of ecology (eg Collins et al.
2000) as well as for society (eg Young
and Wolf 2006; Pataki et al. 2011;
Douglas 2012). Specifically, how can urban ecologists pro-
mote ecological investigation and work with planners and
politicians to make cities more livable and sustainable?
Such questions can be asked from many different perspec-
tives (eg Grimm et al. 2008b; Beatley 2010; Cook and
Swyngedouw 2012; Pincetl 2012; Wheeler 2013; Childers
et al. 2014), so maintaining a diversity of approaches will
allow the field of urban ecology to continue to develop as
a science while simultaneously providing benefits to peo-
ple living in cities (Young and Wolf 2006).
However, a diverse combination of disciplines, goals,
and stakeholders can also make integrating and advanc-
ing science more difficult. Finding ways to integrate
research, planning, and citizen involvement is therefore
also vital for urban ecologists. We present five different
research perspectives on urban ecosystems (Figure 1),
show how each opens a potential new path forward for
the interested ecologist, and discuss how these perspec-
tives can improve our ecological understanding as well as
the livability of cities.
We argue that the success of urban ecology will be mea-
sured by the ability of urban ecologists to continue
advancing the science while simultaneously providing
tangible benefits to society. Future success therefore
depends on ecologists’ ability to include elements that are
unfamiliar to their discipline and often to science in gen-
eral. Ecologists – whether focused on behavior, communi-
ties, evolution, physiology, or ecosystems – are well posi-
tioned to address these challenges, given their training in
system approaches that can be directed to both the basic
and applied aspects of urban systems (Pickett and
Cadenasso 2006).
nFive perspectives of urban ecology research
The ecology and evolution of, and in, cities
The multitude of interactions between humans and
urban ecosystem function has been studied from many
angles. From an ecological perspective, the types and
degrees of disturbances associated with urban ecosystems
challenge scientists to develop principles to trace com-
plex feedbacks between human actions and their effects
on ecosystems and organisms. Two emerging ideas are
broadening the scope of this approach.
First, urban environments are not created de novo, but
are developed within the context of their region. As
global change accelerates, this context plays an impor-
tant role in the challenges associated with accelerated
urbanization (De Sherbinin et al. 2007), creating a situa-
tion in which urban areas could begin to face place-based
vulnerabilities to climate hazards that lie outside even
recent urban experience (Kunkel et al. 2010).
Furthermore, not all cities grow and develop in the same
way (McHale et al. 2013). As products of human plan-
ning within the context of broader regional scales, urban
Figure 1. An example of five different perspectives of urban ecosystems according to
ecological discipline. The combination of these perspectives will be vital in helping
urban ecologists to understand urbanization while also helping to make cities more
livable and sustainable.
Urban ecology: advancing science and society CJ Tanner et al.
576
www.frontiersinecology.org © The Ecological Society of America
areas are likely to change and respond to challenges in
different ways. These responses not only apply stress to
human infrastructure but also move ecosystems into new
and unfamiliar states. The science of urban ecology will
be well served by including these states as they are cre-
ated by human-induced changes at many scales.
Therefore, in addition to developing principles that help
to generalize urban ecosystems, it is equally important for
urban ecologists to investigate and predict the ways in
which cities differ globally.
Second, as they become larger, older, and more inter-
connected, cities have the potential to act as hotspots of
microevolution; examples include rapid evolution in
response to antagonistic selective pressures (eg antimi-
crobials, pesticides, and hunting), pollution, and frag-
mentation (Vandergast et al. 2007; Cheptou et al. 2008;
Whitehead et al. 2010). From the perspective of ecologi-
cal communities, the complex effects of fragmentation
can reshape intraspecific competition and interspecific
interactions, which in turn cause changes in dispersal,
competitive behavior, and social behavior. These
hotspots could provide ideal opportunities to observe
and understand evolution in the “new wild”, including
the eco-evolutionary feedbacks between urban organ-
isms and their environments. In addition, as with pre-
dicting how cities will differentially respond to global
change, predicting evolutionary responses on a local
scale will help planners and designers more effectively
manage future climate hazards and less-desirable urban
organisms.
The ecology of urban infrastructure
The study of ecology in cities has often focused on non-
human organisms and remnant habitats, and how they
respond to human-induced changes around them.
However, the built environment itself is an
important part of the urban ecosystem.
Understanding this “gray infrastructure” (ie
man-made components of urban ecosystems)
on its own terms is an emerging frontier for
ecologists, and one that is tied to questions of
design and engineering.
Under a shifting climate regime, infrastruc-
ture will be increasingly stressed not only by
altered nutrient, material, and water flows,
but also by the effects of invasive species.
Understanding how urban ecosystems
respond to stress requires the inclusion of all
infrastructure, including the built environ-
ment. Aging and degradation of urban infra-
structure comes with ecological and eco-
nomic costs, as well as with design and
planning opportunities (Figure 2; Kaushal
and Belt 2012). Although creating infra-
structure has traditionally been the purview
of engineers and designers, evaluating how
biophysical and socioeconomic environments interact
with design must become part of the broader science of
urban ecology (Grimm et al. 2008b). For instance, urban
ecologists will be called on to evaluate how different
combinations of gray and green (ie biologically derived
components of urban ecosystems) infrastructure affect
stormwater runoff to control flooding and erosion, main-
tain nutrient retention and cycling, and provide other
services such as recreation (Collins et al. 2011; Pincetl
2012). Evaluation of the functioning of the built environ-
ment can capitalize on “designed experiments”, wherein
scientists work with landscape designers to give urban
ecologists avenues to simultaneously create and evaluate
designs in a controlled manner (Felson et al. 2013; Ahern
et al. 2014). These types of experiments will further an
understanding of complex ecological concepts and pro-
vide applied solutions.
In addition to shifting biogeophysical contexts,
processes within the urban ecosystem can change with
city size. From this perspective, research on the size of
habitat patches, which has played a key role in ecological
thinking, can be applied to cities. Larger cities may pro-
vide a greater range of public goods (eg services, parks,
roads, and airports) with lower per capita infrastructure
needs, but also bring public ills (eg congestion, pollution,
disease, and crime; Bettencourt 2013). The expansion of
cities is generally associated with an increase in social
quantities (eg wages and innovation), as well as eco-
nomic inequality and segregation among urban inhabi-
tants, emphasizing the importance of understanding how
heterogeneous characteristics of a city scale with size.
Including ecological and sociological theory and applied
ecology within urban ecosystems will lead to system com-
ponents, from the buildings and pipes in the urban core
to the exurban fringe, being treated as part of the whole
(Forman 2008b).
Figure 2. As aging urban infrastructure degrades, many ecological, social, and
economic problems, as well as opportunities, may arise. Urban ecologists will be
called on to help develop science and policy to maximize the opportunities for
urban inhabitants equitably and in sustainable ways.
CJ Tanner
CJ Tanner et al. Urban ecology: advancing science and society
577
© The Ecological Society of America www.frontiersinecology.org
The science of nature within cities
Just as urban ecosystem science brings the built environ-
ment into the purview of ecological thinking, it recog-
nizes human well-being as a focal point of urban planning
and design, and acknowledges that issues such as health,
happiness, comfort, safety, and security are inexorably
tied to humans’ connections with the natural world. As
cities continue to grow and transition from the sanitary
ideal of the past century to a sustainable mix of various
colored infrastructures (eg gray man-made components,
green vegetation, brown soils, and blue water), a new
understanding of the role that nature plays in urban soci-
ety will need to be developed (Pincetl 2012; Grimm et al.
2013). Just as urban ecosystems result from the interac-
tion between human design and a larger regional context,
so cities themselves can be thought of as “reconstructed
nature” (Pincetl 2012), with both shared and unique
characteristics (Figure 3). In fact, cities offer urban inhab-
itants multiple ways of being engaged with nature accord-
ing to their preferences and social, cultural, and environ-
mental conditions (Standish et al. 2013).
Although interactions between humans and their sur-
rounding flora have been assumed to be important, the
actual geographical relationships between humans and
plants are still surprisingly vague (Head and Atchison
2009). There is increasing evidence of physiologically
and psychologically important interactions between
humans, their socioeconomic status, and their surround-
ing environment (Heynen et al. 2006; Wolch et al. 2014).
Excessive heat and poor air quality have negative conse-
quences for human health, and a lack of trees in the envi-
ronment, often associated with low socioeconomic status,
translates into sociological, physiological, and psycholog-
ical costs (Figure 4; reviewed in Wolch et al. 2014).
Furthermore, there are potential human physical and psy-
chological benefits to having access to green space with
high biodiversity (Fuller et al. 2007), although the bene-
fits of actual diversity versus perceived diversity are still
unclear (Dallimer et al. 2012). In addition to the direct
health benefits associated with green space, stewardship
of urban green space resulting from an interest in health
and recovery can be associated with social, natural, and
economic capital (Burls 2007). In this way, urban green
space simultaneously provides both individual and com-
mon goods. However, not all urban human–flora interac-
tions are positive. For instance, urban parks may be asso-
ciated with personal safety issues (reviewed in
McCormack et al. 2010), and pollen from urban plants
can cause respiratory problems (Lyytimäki et al. 2008). By
gaining a better understanding of the relationship
between nature and city residents, urban ecologists can
help design cities that optimize both infrastructure and
ecosystem services.
The importance of an intersection between the city
and the natural world for urban inhabitants (eg gardens
and parks) is a central theme of “middle nature”, which
conceptually describes turning nature into culture and
providing access to all by merging the natural world and
the built environment (Cosgrove 1993). Middle nature
therefore is a way of considering the many roles of culti-
vated nature in the city. Some of these roles involve the
benefits that nature provides to humans within the built
environment (ie ecosystem services). But how these ser-
vices are measured is open to interpretation (Reyers et al.
2013) and further depends on how such services are
defined as well as on humans’ experience with the natural
world (Krasny et al. 2014). The way in which people
inhabit this world in turn shapes the broader phenomena
of urban culture, economics, and governance, each of
which varies dramatically at global and local scales. Yet
scientists know surprisingly little about how these feed-
backs filter from the urban ecosystem back into human
experience, let alone society. Surveys have shown that
individuals are happier in the short term when they have
higher financial incomes; but at larger temporal and spa-
Figure 3. Urban ecosystems are an amalgamation of blue, gray, and green infrastructures, with tight spatial and cultural links.
Understanding the ecological effects of these links, and how these effects feed back to the well-being of urban inhabitants, depends on
many historical and place-based contexts. For example, the cities of Vancouver, Canada (a) and Cape Town, South Africa (b) are
both composed of infrastructural and ecological components similar to those of all cities, yet each city has unique characteristics that
have emerged from its geography and history.
(a) (b)
CJ Tanner
CJ Tanner
Urban ecology: advancing science and society CJ Tanner et al.
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www.frontiersinecology.org © The Ecological Society of America
tial scales, human life satisfaction is more strongly corre-
lated with social capital (ie social cohesion and personal
investment in the community) and access to quality out-
door recreation (Vemuri et al. 2011; Hager et al. 2013).
How satisfaction with the environment feeds back to
modify society remains to be seen.
The behavior and evolutionary ecology of urban
humans
The social, political, and economic structure of urban
societies largely determines the way in which humans
create the urban environment. But what do ecologists
have to offer in terms of understanding phenomena that
have long been the subjects of history, economics, and
political science? Below, we spell out two of the reasons
that ecological thinking does indeed have a place in ana-
lyzing the dynamics of urban humans.
First, many ecologists work comfortably with complex
feedbacks among processes that occur at different organi-
zational levels, from physiology to climate. For example,
the urban heat island effect involves processes ranging
from the physics of surfaces, sunlight, and water, to the
physiology of plants and the behavior of humans – and
each of these processes can be decoupled temporally and
spatially. Understanding the consequences of urban heat
islands for plants and animals requires an understanding
of their behavioral, physiological, and possibly evolution-
ary responses, including a positive feedback from reduced
evapotranspiration and increased air conditioning, back
to increased temperatures. Placing humans within a
socioecological context requires an understanding of
these types of feedbacks. The physiological stresses of
urban heat islands are accentuated in lower-income areas,
which typically have reduced tree cover (reviewed in
Wilson 2011 and Wolch et al. 2014). Behavioral
responses are more limited in these areas because poverty
reduces opportunities for active or passive cooling, for
example through the use of swimming pools (Harlan et al.
2007). Urban heat islands are thus embedded in a succes-
sion of feedbacks involving human society, with poorer
areas becoming trapped in a local warming cycle. These
feedbacks, with their ties to nature within cities, exem-
plify the difficulties that ecologists face when including
political and socioeconomic issues in their analyses
(Wolch et al. 2014). In a given city, how do ecologists
effectively persuade elected municipal officials that more
trees need to be planted in response to voters demanding
a higher standard of living? Furthermore, what types of
trees should be planted, how many, and where? Or rather
how should urban areas be designed to optimize the
human–environment relationship (Pataki et al. 2011)?
Only by making known the explicit connections between
the natural environment and quality of life will ecologists
be able to help in the design of urban areas most affected
by this relationship.
Second, ecology, like all of biology, is unified through
evolutionary thinking. From an evolutionary perspective,
early humans generally used group living strategies to deal
with the selective pressures of finding resources, gaining
knowledge, and obtaining protection; humans have
Figure 4. Many aspects of human physical and mental health
can be linked to something as basic as whether or not an urban
resident’s view of the urban ecosystem (a) includes (b) or
excludes (c) a view with trees. Perhaps even more important for
urban ecologists is answering questions regarding what types of
trees should be planted where to optimize the human–
environment interaction for all members of the urban ecosystem,
and how related services can be provided where they are most
needed.
(a) (b)
(c)
CJ Tanner
CJ TannerKSV Photography/www.iStockphoto.com
CJ Tanner et al. Urban ecology: advancing science and society
579
© The Ecological Society of America www.frontiersinecology.org
therefore evolved a wide range of psychological mecha-
nisms to collectively solve these challenges (Van Vugt
and Kameda 2013). As the ancestors of modern humans
developed new technologies, leading to intensification
(eg agriculture), the effects of niche construction and
social structure on group cohesion, information sharing,
and workforce specialization were key components of
their success (Johnson and Earle 2000). In principle,
modern humans evolved from ancestors that used social
cohesion to solve problems relating to uncertainty in
their environment (Larson and Christensen 1993; Van
Vugt and Kameda 2013). As social cohesion in urban
areas disintegrates, largely due to a combination of
socioeconomic and environmental pressures, cities are
becoming increasingly prone to crime (Kuo and Sullivan
2001, reviewed in Brennan 1999), again linking nature
and health with human behavior. In many areas, govern-
mental intervention plans overlook these underlying
determinants (Jütersonke et al. 2009), thereby complicat-
ing the integration of science and policy. Before dis-
cussing this issue, however, we must first address how
groups of individuals living in proximity agree on a set of
common goals (Levin 1999). Urban ecologists are posi-
tioned to use research and principles from voting theory
and collective decision making, as well as from behav-
ioral and evolutionary ecology, to understand governance
in cities where humans and non-human animals share
common environmental and economic resources
(Ehrlich and Levin 2005).
Citizen and stakeholder science
Many successful ecologists think about their study sys-
tems from the perspectives of the organisms or ecosystems
that they study (eg “thinking like a mountain”; Leopold
1949). As humans become more integrated into ecologi-
cal studies as both agents and subjects, ecologists find
themselves in the unusual position of needing to think
about the urban environment as would non-scientists.
Paradoxically, thinking like a typical urban human might
be more difficult for an urban ecologist than thinking like
a deer in a forest. Every urban resident brings a unique
objective, perspective, history, and set of needs – deter-
mining the identity and location of the stakeholders most
in need of science and policy integration is a daunting but
essential part of urban ecological studies.
Involving these stakeholders as partners in research,
particularly with regard to sustainability and access to
ecosystem services, generates useful questions and inter-
actions (Groffman et al. 2010). The model of “civic ecol-
ogy” – where scientists work with urban residents to
develop questions and methods, collect and interpret
data, and ultimately translate these data into policy rec-
ommendations – can be far more effective than the more
traditional one-way dissemination of knowledge from sci-
entists to the general public (Krasny and Tidball 2012).
This comparison between information sharing and infor-
mation flow is critical to integrating science and policy in
urban ecosystems. If obtaining funding, performing
research, and publishing results are not having an effect
“on the ground”, then collaborating with stakeholders
could provide a mechanism to do so.
Even for general science questions that do not directly
involve people, urban ecology is an ideal setting for new
collaborations and teaching opportunities. For instance,
exclosure experiments, residential development, and the
evolution of urban organisms all provide opportunities for
involving stakeholders, including students and citizen
scientists (Dickinson et al. 2012). As working with stake-
holders becomes more “normal” for urban ecology, prob-
lem solving by integrating science and policy will be one
of the primary advances.
nConclusions
The term “ecology” was coined by Ernst Haeckel to mean
“the study of the house or habitation”, whereas “econom-
ics” has the related but more directed meaning of “the art
of managing a household”. For most people, the city is
now their “house”. Just as architects cannot design build-
ings without considering the people who inhabit them,
policy makers cannot manage cities without a holistic
understanding of how urbanization affects both human
and non-human residents. Furthermore, as urban popula-
tions grow and infrastructure ages, studies that can best
determine how to provide cost-effective services to the
urban inhabitants most in need will be in high demand.
Ecology encompasses a broad spectrum of disciplines; with
their diverse set of skills and interests, ecologists will be
able to provide links between the natural and built envi-
ronments. To ensure this outcome, urban ecologists must
find ways of continuing to promote the science of ecology
while also becoming more involved with planners and
policy makers. As demonstrated here, although the
research perspectives of urban ecology can be extremely
varied, they share a common, inclusive theme of promot-
ing science and benefiting society. The difficulties of inte-
grating science and policy can be amplified when, as is
often the case, stakeholders have differing agendas. For
example, how do we balance the need for managing the
“global commons” with the need for addressing the
increasingly imperative “brown issues” of pollution and
land degradation (Brennan 1999)? And how can we pro-
mote the protection of endangered species in an area
while ignoring the health and environmental problems
faced by the people living there (Hardoy and
Satterthwaite 1991)? Such questions are further compli-
cated when the associated costs and benefits vary with
temporal and spatial scales. Environmentalists are occa-
sionally accused of caring more about trees than people;
perhaps when we fail to include people as part of our stud-
ies, this may appear equally true of ecologists. By
approaching urban ecosystems with a more inclusive per-
spective, which seeks to integrate both conceptual and
Urban ecology: advancing science and society CJ Tanner et al.
applied aspects, urban ecologists will be better prepared to
help bridge the gaps between trees, buildings, and people.
The German writer Goethe (1749–1832) supposedly
said, “A poem is just as much a part of nature as a tree”.
The built environment, nature in its many forms, and
humans all share a complex network of interdependent
connections that provide both academic and applied
challenges. To advance the science of ecology, as well as
promote benefits to society, urban ecologists must
approach their discipline in a similarly inclusive manner,
ready to incorporate unfamiliar areas ranging from eco-
nomics to politics. But such roles can be challenging. For
example, how will advocating policy affect scientists’
ability (real or perceived) to remain objective? For now
we can only say, “let your conscience be your guide”, and
suggest this as another area where collaborative research
would be useful. Although making these links is challeng-
ing, Goethe’s observation highlights a key element of
modern urban ecology and suggests that interfacing with
unfamiliar disciplines will help us to understand and more
effectively manage the urban ecosystems that are coming
to dominate our planet.
nAcknowledgements
This manuscript is the culmination of a series of Ignite
sessions given at the Ecological Society of America’s 2013
Annual Meeting.
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