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com

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Global ecological regionalization: from biogeography to

ecosystem services
Yanxu Liu1, Bojie Fu1,2, Shuai Wang1 and Wenwu Zhao1

Ecological regionalization is not only an abstraction of ecological regionalization has been considered as a meth-
understanding nature but also a form of management guidance odology of grouping site-level ecosystems on the macro-
toward achieving environmental sustainability. The existing scale [3]. The result is usually expressed as an ecoregion,
global ecoregion scheme successfully contributes to the and some other naming conventions, such as ecozone and
conservation of global and regional biodiversity. As highlighted ecological zone, also exist for the early stage [4,5]. As a
by the conceptual framework of the Intergovernmental Platform spatial generalization of ecological hierarchy, an ecore-
on Biodiversity and Ecosystem Services (IPBES), an integrated gion is not only a theoretical summary of the laws of
understanding of biodiversity and ecosystem services is nature but also a spatial planning tool to guide sustainable
required. In this new age, we propose a concept of global natural resource management [6].
ecological functional regionalization, which should be
separated from the former idea of biogeographical An ecoregion is defined based on the purpose of regionali-
regionalization. With the consideration of key geographical zation. In the famous first scheme of global ecological
characteristics and ecosystem services, the views of global regionalization, Bailey defined ecoregions as ‘large por-
environmental change and human activities are included in this tions of the Earth’s surface over which the ecosystems have
new framework. However, key questions still exist about the characteristics in common’ [2]. This definition identifies
principles, indicators, methods and applications of a global the original purpose of ecological regionalization as an
ecological functional regionalization. In the next studies, global explanation of the natural rule of the distribution of
ecological regionalization should focus more on the ecosystems. With the particular objective of biodiversity
uncertainties of spatial datasets and mathematical models. conservation, Olson et al. defined ecoregions as ‘relatively
Addresses large units of land containing a distinct assemblage of
1
State Key Laboratory of Earth Surface Processes and Resource natural communities and species, with boundaries that
Ecology, Faculty of Geographical Science, Beijing Normal University,
100875 Beijing, China
approximate the original extent of natural communities
2
State Key Laboratory of Urban and Regional Ecology, Research Center prior to major land-use change’ [1]. This terrestrial
for Eco-Environmental Sciences, Chinese Academy of Sciences, ecological regionalization is supported by the World Wild-
100085 Beijing, China life Fund (WWF) and has made a profound influence;
additionally, this framework is largely followed by more
Corresponding author: Fu, Bojie ([email protected])
recent ecological regionalization schemes and several
institutions, such as The Nature Conservancy (TNC).
Current Opinion in Environmental Sustainability 2018, 33[1_TD$IF]:1–[2_TD$IF]8 Compared with Bailey’s ecoregion scheme, the WWF
This review comes from a themed issue on System dynamics and ecoregions provide more specific boundaries for global
sustainability biodiversity hotspots and identify conservation priorities
Edited by Bojie Fu and Yongping Wei [7,8]. Along with the WWF ecoregion framework, the
For a complete overview see the Issue and the Editorial
global ecological regionalization focuses on freshwater
[9], coastal and shelf areas [10], and pelagic oceans [11].
Available online 16th May 2018
In addition to the focus on integrated ecosystems, the
Received: 30 October 2017; Accepted: 12 February 2018 detailed global biogeographical regionalizations, such as
global zoogeographical regionalization, have revealed
https://doi.org/10.1016/j.cosust.2018.02.002 additional information on macro-ecological investigations
and animal biodiversity conservation [12–14].
1877-3435/ã 2018 Elsevier B.V. All rights reserved.

The availability of the global ecoregion schemes is largely

dependent on the original goal of ecological regionaliza-
tion. Beyond the conservation of biodiversity, there are
questions on what ‘nature’s contributions to people’ is,
Introduction and this content is usually called ecosystem services [15].
The natural resources on our planet are not isolated, and For a more concise definition, ecosystem services are the
recognizing the hierarchal spatial relationship of the direct and indirect contributions of ecosystems to human
distribution of natural resources should be a means of well-being [16]. With this perspective, scholars in natural
guidance for conservation planning at global and regional and social sciences should focus not only on the status of
scales [1,2]. Focusing on the hierarchy of ecosystems, biodiversity elements but also on the benefits received

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2 System dynamics and sustainability

from biodiversity, namely, ecosystem services [17]. The scheme [34]. In addition to the thought patterns of
Intergovernmental Platform on Biodiversity and Ecosys- Dokuchaev [35], Merriam [36], Herbertson [37], Köppen
tem Services (IPBES) has been accordingly established [38], Tansley [39], Holdridge [40,41], Isachenko [42] and
and aims to discover how changes in biodiversity impact other pioneers [43–45], the production of a map of global
ecosystem services and human well-being [18]. The goal biogeographic provinces from Dasmann [46] and Udvardy
of the IPBES platform is to contain both ‘conservation [34] has provided substantial guidance for the subse-
and sustainable use of biodiversity’ and ‘long-term human quent schemes of global biogeographical regionalization.
well-being and sustainable development’, as well as to However, because they are limited by spatial resolution,
provide support in bridging the gap between knowledge these achievements can hardly be applied to the conser-
and policy [19,20]. vation of regional biodiversity. Nevertheless, the theo-
retical foundations of these milestones have largely con-
Under these backgrounds, several questions emerge in tributed to the formation of a global ecoregion map.
global ecological regionalization: firstly, are the current
ecoregion schemes able to provide conservation guidance The WWF ecoregions do not refer to Bailey’s ecoregion
for both biodiversity and ecosystem services, and if not, map because the organization has a more specific focus on
why [21]? Secondly, which type of ecological regionali- conservation planning. This scheme sub-divides the global
zation is required under the interaction between the territorial land into 8 realms, 14 biomes, and finally, 867 ecor-
biophysical processes and social processes of global envi- egions that are partly dependent on Dasmann’s and Udvar-
ronmental change [22]? Finally, from a methodological dy’s biotic provinces [1]. This scheme has been cited more
perspective, how could ecological regionalization be than 2000 times in 16 years, which makes it the leading
effective for the sustainability of the coupled human biogeographical regionalization of the 21st century. Accord-
and natural systems [23]? The arguments for these ques- ing to the effectiveness and the significant contribution of
tions form the purpose of this article. Specifically, three WWF ecoregions (loosely combined within Bailey’s ecor-
objectives are attained in this article: to briefly explain the egion in the United States in TNC) on biodiversity conser-
history of the development of biogeographical regionali- vation [47–50], the map has been updated several times. In
zation; to specify the conception of the consideration of the recent version of Ecoregions 2017, 846 terrestrial ecor-
ecosystem services into regionalization; and to pose key egions are identified as units for the goal of protecting half of
questions and a case framework for the future of ecologi- the terrestrial realm by 2050 [51].
cal functional regionalization.
Different from the biodiversity-oriented biogeographical
Contributions of biogeographical regionalization including freshwater ecoregions [9],
regionalization marine ecoregions [10], pelagic provinces [11], and zoo-
From the perspective of biodiversity conservation, ecologi- geographic regions [12], the term global urban ecoregion
cal regionalization is specifically called biogeographical has appeared as a result of the consideration of human
regionalization in this article, and it provides both theoretical disturbance, in the form of urban construction, on the
and practical contributions to scientists and policy makers. environment [52]. In terms of the boundary of WWF
On one side, ecoregions provide a holistic framework to ecoregions, the global human pressures on land and
comparatively analyze environmental problems based on within protected areas have been evaluated [53]. From
the integration of disparate knowledge of both geography the perspective of global change, an indicator framework
and ecology [24]. On the other side, the ecoregions within that introduced socioeconomic conditions into biodiver-
specific objectives are results of applied science and provide sity conservation planning was established [54]. Accord-
direct benefits for environmental management [25,26]. ingly, it is agreed upon that research on the quantification
From a philosophical perspective, ecological regionalization of human activity should be strengthened for global
is sometimes an objective and a form of deductive science biodiversity conservation. Nevertheless, the consider-
with a cognitive perspective; at other times, it is an inductive ation of the impacts of human activity on biodiversity
and subjective art of planning with a management perspec- conservation is far from complete. In addition, from the
tive — which perspective to take is dependent on the opposite perspective, the benefits biodiversity conserva-
application of the ecoregion [27]. According to the various tion impose on human activity also requires further
perspectives in ecological regionalization, the contributions quantification. Thus, the following question emerges:
of existing ecoregion schemes are inconsistent. can the existing biogeographical regionalization schemes
effectively combine the inclusion of anthropogenic influ-
The first global ecoregion map from Bailey [28,29] had a ences and requirements?
profound influence in the 20th century [30–32]. Because
systematic criteria for sub-dividing a landscape into eco- Consideration of ecosystem services in
systems was provided in the scheme [33], much more regionalization
spatially explicit boundaries were received in Bailey’s In fact, the abovementioned question was answered more
ecoregion compared with the former province-level than ten years ago with the quantitative comparison on

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 Global ecological functional regionalization Liu et al. 3

global biodiversity conservation and ecosystem services. Figure 1

Depending on the estimated value of terrestrial ecosys-
tem services, Turner et al. found that global biodiversity
conservation priorities were not spatially concordant with Nature
ecosystem services in a large area [55]. Then, Naidoo et al.
Biodiversity and
similarly concluded that the global biodiversity regions ecosystems Nature’s contribution to people
provide no more ecosystem services than do random
regions [21]. The regional studies also concluded that Ecosystems
services
Good quality of life

regulating services were not efficiently addressed in pro- Human

Biogeographical wellbeing
tected areas [56,57]. From another perspective that con- regionalization
fronted the large influence on global biodiversity caused Ecological
by humans, anthropogenic biomes were proposed, and functional Ecological
regionalization policy
they considered the importance of human population instruments
density and land use, which are spatially different within
the biomes of existing biogeographical regionalization
schemes [58]. Considering the human demands for eco-
Current Opinion in Environmental Sustainability
system services, as well as the human influences on
ecosystem pattern and process, spatially understanding
them is required in ecosystem management. However, The conceptual position of ecological functional regionalization, as
indicated by the Intergovernmental Platform on Biodiversity and
the spatial allocation of these demands and influences is Ecosystem Services (IPBES) framework.
still required and has not been provided by biogeograph-
ical regionalization.
neglected in ecological functional regionalization. A
As a link between ecosystems and human beings, the recommended thought is to separate ecological func-
importance of ecosystem services is undisputed [59]. tional regionalization into two steps. The first is to
Global ecosystem services have been estimated and finish biogeographical regionalization, and the second
mapped in several ways during the last 20 years [60]. is to regionalize ecosystem services based on the
However, with the exception of quantifying ecosystem boundaries of biogeographical regionalization; this
services, the methods used to integrate different types of prevents the boundaries from being disordered in
ecosystem services are still lacking. The trade-offs and the intersection of the two types of regionalization.
dominance among ecosystem services have not yet been Therefore, the priority of either conserving biodiver-
spatially clear. Similar to the objective of biodiversity sity or maintaining ecosystem services can be spa-
conservation in biogeographical regionalization, the global tially identified. Moreover, according to ecosystem
spatial allocation of typical ecosystem services is necessary function is a concept linking ecosystem process and
to support the sustainable management of ecosystem ecosystem service, the term ‘function’ is applied
services and to maintain and enhance the ecological ben- instead of ‘service’ because of ‘ecosystem service
efits to human beings. Because it is separated from the regionalization’ may narrow the research scope.
former ecological regionalization that focused on biogeog- (2) Ecological functional regionalization includes not
raphy, and because it is identified by the IPBES frame- only the ecological requirements of humans but also
work, we call this spatial allocation of ecosystem services the anthropic influences on ecosystems. Although the
ecological functional regionalization (Figure 1). The bio- potential biogeographic characteristics provide guid-
geographical regionalization, ecological functional region- ance to the objective of biodiversity conservation, the
alization, and ecological policy instruments are all envi- present anthropic influences on ecosystems should
ronmental management tools and in cascade relationship. not be neglected in regionalization. Within the ulti-
The importance of proposing these concepts can be mate goal of long-term human well-being and sus-
described by the following three aspects. tainable development, the issue is not the necessity of
(1) Ecological functional regionalization is a spatial inte- anthropic influences; rather, the issue is how to spa-
gration of global biodiversity conservation and eco- tially delineate anthropic influences. To avoid con-
system services. From the guidance of the IPBES, founding anthropic influences with the conservation
biodiversity and ecosystem services should not be of biodiversity and ecosystem services, sub-ecore-
considered separately. Therefore, compared with the gions are recommended as to represent the division
objective of biodiversity conservation in biogeograph- of different levels of influence caused by human
ical regionalization, ecological functional regionaliza- activities. As a consequence, the conservation meth-
tion should not limit its objective to maintaining ods among regions that are dominated by urbanized
ecosystem services. Although capital ecosystem ser- landscapes, cultivated landscapes and natural land-
vices are highlighted in ecological functional region- scapes will be separated, contributing to the goal of
alization, the biogeographic characteristics are not targeted sustainable environmental management.

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4 System dynamics and sustainability

(3) Ecological functional regionalization tends to be rela- a critical technical process in regionalization is deter-
tively robust to global environmental change. The val- mining which indicators are necessary and which
uation of ecosystem services will never be a constant datasets are effective. For example, not all ecosystem
value. The significant changes in climate, such as change services can be mapped precisely. As guidance for the
in precipitation, would triggers change in terrestrial spatial conservation of ecosystem services, an uncer-
vegetation, and thus, change in ecosystem services. tain map of ecosystem services would largely reduce
Because an annual update of the ecological regionaliza- the effectiveness of ecological functional regionaliza-
tion scheme is unpractical, global environmental change tion in environmental policy making.
should be embodied in the ecological functional region- (3) How can a relatively accurate spatial boundary that is
alization. Similar to the attribution of anthropic influ- based on mathematical models be produced in global
ences, the simultaneous consideration of environmental ecological functional regionalization? For the various
change combining with both biodiversity conservation indicators, a classification based on several mathemat-
and ecosystem services would confound the naming of ical models would result in a more accurate spatial
ecoregions. Therefore, sub-ecoregions are recom- boundary than would manually merging the patches.
mended to separate the patterns of significant improve- However, which model is most suitable for regionali-
ment and degradation within dominant ecosystem ser- zation is still unknown; additionally, how to dissolve
vices, according to both the ecosystem services the classification pixels into ecoregions is also
estimation for the past decades, and the ecosystem unknown. Regionalization is an abstraction of known
services forecast based on the climate change scenarios. spatial elements, rather than an estimation of
This information should be an important supplementa- unknown spatial elements. Thus, one of the obstacles
tion for non-specialists who will apply the ecoregion of regionalization is the different mathematical vali-
schemes to sustainable environmental management. dation from landscape classification. The manage-
ment-oriented ecoregion scheme is not completely
Key questions in ecological functional derived from mathematical principles, and from the
regionalization perspective of future boundary planning, a boundary
As a new idea, the conceptual system of ecological func- is hard to validate at the present moment.
tional regionalization must be continued. Several key (4) How can we ensure the ecoregions in global ecologi-
questions must be answered before this conception can cal functional regionalization will be suitable for
be applied in practice. Focusing on the ultimate goal of practical environmental management? Ecological
integrating long-term human well-being and sustainable functional regionalization is proposed from a stand-
development with biogeographic characteristics and eco- point of differentiated management. Targeted objec-
system services, implementing global ecological func- tives should be determined based on the dominant
tional regionalization should be more complex than the functionality of each ecoregion. The sustainable
implementation of the biogeographical regionalization in thresholds for the different types of ecoregions obvi-
the last decade. In this article, we propose four questions ously vary. Therefore, to ensure the practicality of
to start the scientific discussion on the implementation of different thresholds, the bottom line of each ecore-
global ecological functional regionalization. gion’s functionality should be identified first. Then,
the next question should involve how to achieve the
(1) What are the indispensable and dispensable principles in targeted objectives. The enlightenment of planetary
global ecological functional regionalization? In the cen- boundaries can provide the guidance for the determi-
tennial inheritance of various types of regionalization [35– nation of sustainable thresholds [61]. We believe the
42], the principles are inconsistent. There is no doubt that ecoregions of ecological functional regionalization
all regionalization schemes are comprehensive, and the could provide a spatially consistent regional context
regions are named based on the dominant factor. Never- for the determination of targeted objectives.
theless, what is the next important principle, and what
principles are not important? For regionalization princi- We have tried to provide a case response for the first three
ples, there is not a ‘the more, the better’ theorem. Rather, questions as one of the understandings of ecological
ecoregions should be concise and practical, and the functional regionalization (Figure 2). The existing region-
unnecessary principles should be pruned after a deep alization schemes, such as Ecoregions 2017 and Bailey’s
discussion on the potential fields of application relevant to ecoregions, can provide basal support to the determina-
ecological functional regionalization. tion of principles and indicators in global ecological
(2) What are the key spatial indicators in global ecological functional regionalization. Based on these guidelines,
functional regionalization? Although there are more the ecological functional regionalization is decomposed
indicators, a more comprehensive framework with a into 4 steps, as follows.
redundant indicator system would be difficult to
effectively compose. The uncertainty would increase (1) In addition to the fundamental principles of compre-
when low-quality datasets were involved. Therefore, hensiveness and relative uniformity of dominant

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 Global ecological functional regionalization Liu et al. 5

Figure 2

Methods
Principles
Unsupervised classification: such
Foundational: comprehensiveness, as K-Means clustering, iterative
relative uniformity of dominant factors self organizing data analysis
techniques algorithm
Essential: coordination of functionality,
Supervised classification: such as
stability of temporal tendency
support vector machine, random
forest, Bayes classifier
Dispensable: unanimous occurrence,
spatial continuity
Elimination: area threshold of
discrete patches
Existing regionalization schemes, such
as Ecoregions2017, Bailey’s ecoregions

Indicators Calibrations
Response tendencies: Dominant factor from
Geographical elements: improvement / degradation each method
terrain, climate, soil, of geographical elements
vegetation, etc. and ecosystem services
Region Subregion The quantified degree
Ecosystem services: Human activities: of the dominance
carbon fixation, soil population, gross domestic
retention, water product, construction land,
conservation, etc. nighttime light, etc. The preliminary scheme

Global ecological functional

regionalization scheme

Current Opinion in Environmental Sustainability

An example of simplified technical routes of ecological functional regionalization.

factors, at least two principles are essential in (2) Four types of indicators have been introduced into
global ecological functional regionalization. The first global ecological functional regionalization. We
principle is the coordination of functionality. From named them geographical elements, ecosystem ser-
the multifunctional landscape perspective, the func- vices, response tendencies and human activities. The
tionalities zoned in the same ecoregion could confuse common geographical elements such as terrain, cli-
the stakeholders, as the enhancement of one ecosys- mate, soil, vegetation are, without a doubt, the basic
tem service could trigger the loss of another ecosys- indicators. The improved global spatial datasets such
tem service. The other principle is the stability of as SRTM (terrain), MSWEP (precipitation), Soil
temporal tendencies. In other words, the regions with Grids/HWSD (soil), and MODIS NDVI/LAI/NPP
unstable ecosystem services should be identified. (vegetation) can elaborately express the indicators.
Confusion between improving and degrading ecosys- The ecosystem services at relatively high resolutions
tem services would hinder the judgment of policy are the key indicators for ecological functional region-
makers. In addition, some principles may be dispens- alization. Some regulating services such as carbon
able. The unanimous occurrence or absence in a fixation, soil retention and water conservation are
geological period may not effectively support the recommend. The above two types of indicators form
separation of today’s ecosystem services. To maintain the indicator system for the division of ecoregions.
the spatial continuity of each ecoregion would largely Although cultural services are certainly important,
increase the number of units, which is unsuitable for spatially separating cultural services is more uncertain
environmental management. than spatially estimating ecological functions. The

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6 System dynamics and sustainability

spatialized anthropic information has been intro- findings of the complicate relationships among the spatial
duced in the human activities indicator. The datasets elements are still hidden in the centurial topic of regional-
of DMSP-OLS (nighttime lights) and SSP database ization. Thus, this article is intended to begin the discus-
(population and economy) are widely used. Accord- sion, rather than to provide a definite solution to sustain-
ingly, the indicators of response tendencies and able environmental management.
human activities form the indicator system for sub-
ecoregion divisions. The improvement or degradation Conclusions
of geographical elements and ecosystem services are Ecological regionalization is an abstraction of site-level
separated. The construction land and nighttime light ecosystems on the macroscale, which not only contributes
are considered in combination with population and to the understanding of nature but also provides differ-
gross domestic product. entiated guidance to sustainable environmental manage-
(3) The mathematical methods for regionalization are ment. Today’s global ecoregion scheme offers explicit
still lacking, whereas the spatial classification models guidelines for biodiversity conservation, but it still faces
are abundant. Both unsupervised and supervised obstacles in the inclusion of the improvement of human
classifications can be introduced into regionalization. well-being. After affirming the great achievements in
Although supervised classification needs training biogeographical regionalization, we have proposed the
examples, these points can be manually extracted concept of ecological functional regionalization. With
from a presupposed naming system. For example, the key consideration of ecosystem services, the views
the extreme combinational relationship among eco- of global environmental change and human activities
system services is known and can be identified by the should be followed in ecological functional regionaliza-
dominant ecosystem service. Then, the unknown tion. Four key questions are provided in the consideration
combinations can be classified from the identified of the principles, indicators, methods and applications of
training examples. Because the result of classification global ecological functional regionalization. We provide
is on the pixel level, the definition of area threshold possible answers to some of these questions to set a
used to eliminate discrete patches is a critical step. If starting point for further discussion.
thresholds are determined without comparing the
various results, the output is unreliable, since neither In the centurial evolution of the numerous types of region-
a fragmentized pattern nor a homogeneous large unit alization, the new age of mass data provides new oppor-
would be suitable in regionalization. tunities to global regionalization. In some sense, global
(4) Calibration of the boundary should not be neglected in ecological functional regionalization is an easy task, as
regionalization. However, after elimination, the exact abundant spatial datasets can provide various types of
spatial boundary of an ecoregion can be hard to find in information. However, in another sense, global ecological
the abstraction of the characteristics. A relatively pre- functional regionalization is more difficult than the histori-
cise scheme that can highlight the dominant factors cal tasks. The uncertainty of these datasets and the meth-
should be selected. The dominant factor from each ods of regionalization may result in huge controversies over
method cannot be completely consistent because the the precision of the result. In our opinion, compared with a
foundational principle of ecological functional region- loose zoning that is based on several common indicators, a
alization is assumed to be the relative uniformity of scheme that stands up to argument is what we need in this
dominant factors. The higher the degree of dominance, new age. In other words, a misguided scheme is easy to
the higher the applicability of the method. Thus, the extract but is no better than nothing, and sometimes, it can
preliminary scheme can be identified from the various cause negative effects. In conclusion, thinking compre-
intermediate results derived from different methods. hensively and working cautiously are what we need in the
next global ecological regionalization.
In fact, we should acknowledge that there may be no
perfect scheme to resolve all the questions proposed above Acknowledgement
or to cope with all the environmental concerns in practice. This research was financially supported by the National Key Research and
However, we know that regionalization research will pro- Development Program of China (No. 2017YFA0604701) and China
Postdoctoral Science Foundation.
ceed forward in two directions: more specific and more
comprehensive directions. The specific ecological region-
alization may target the conservation of critical types of References and recommended reading
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