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Contents lists available at ScienceDirect

Ecological Complexity
journal homepage: www.elsevier.com/locate/ecocom

Viewpoint

The dark side of the redundancy hypothesis and ecosystem

assessment
Federico Morellia,b,* , Piotr Tryjanowskic
a
Czech University of Life Sciences Prague, Faculty of Environmental Sciences, Department of Applied Geoinformatics and Spatial Planning, Kamck 129, CZ165 00 Prague 6, Czech Republic
b
Faculty of Biological Sciences, University of Zielona Gra, Prof. Szafrana St. 1, PL-65-516 Zielona Gra, Poland
c
 University of Life Sciences, Wojska Polskiego 71C, PL-60-625 Poznan
 , Poland
Institute of Zoology, Poznan

A R T I C L E I N F O

Article history:
Received 19 November 2015
Received in revised form 9 July 2016
Accepted 16 July 2016
Available online xxx
Keywords:
Complexity
Ecosystem stability
Environmental goods
Heterospecic interactions
Species redundancy
Species role

A B S T R A C T

Intricacy of biotic interactions (predator-prey relationship, strength of food web links and other type of
intra- and inter-specic relationships), as well as shifts in species functions in ecosystems, could affect
the accuracy of predictions derived from the theory of redundancy of species, when applied to
ecosystems assessment.
This opinion paper is based on three main considerations: 1) some fundamental differences between
ecological and engineering denition of redundancy, underlying the main concerns related to the use
frameworks derived from economical or engineering disciplines, as the ecosystems services paradigm; 2)
presence of empirical obstacles to establish whether two different species are fully or partially
redundant. When species redundancy in a particular community is estimated using a matrix with
species-specic functional traits, often forgetting potential biotic interactions not directly related to the
trophic chains or neglecting the variability in strengths of the links connecting these species; and 3)
recent evidence offered by studies that shed doubts on the validity of the ecological redundancy
hypothesis.
Finally, we claim that more attention must to be paid to intrinsic ecological aspects of ecosystem
components (per se values rather than derived values), and a precautionary principle is necessary for
decisions related to the assessment of ecosystems.
2016 Elsevier B.V. All rights reserved.

1. Ecological complexity: not only a question of numbers

In this note, we underline some issues related to the use (and
abuse) of a framework derived from disciplines such as economics
or engineering, in ecological studies. We highlight how the
complexity of biotic interactions could make the predictions
derived from the theory of redundancy less effective (Walker,
1992), when applied to ecosystem assessment. We claim that more
attention must to be paid to the intrinsic ecological aspects of
ecosystem components (per se values rather than derived values).
The study of ecosystems is an important applied discipline
suitable for understanding global change and human impact on
environments (Raudsepp-Hearne et al., 2010). For this reason,

* Corresponding author at: Czech University of Life Sciences Prague, Faculty of

Environmental Sciences, Department of Applied Geoinformatics and Spatial
Planning, Kamck 129, CZ-165 00 Prague 6, Czech Republic.
E-mail address: fmorellius@gmail.com (F. Morelli).

predominant directions in the study of ecosystems include a

growing focus on human-dominated landscapes and the development of the concept of ecosystem services for human resource
supply and well-being (Carpenter et al., 2009). Key ecosystem
processes include primary production, evapotranspiration, respiration, decomposition, secondary production, soil formation and
cation exchange, nutrient mineralization and immobilization, and
many others (Currie, 2011). Nowadays, a major challenge is to
ensure the functioning of ecosystem services to meet the needs of a
burgeoning world population of humans (Biggs et al., 2012;
Millennium Ecosystem Assessment, 2005). Ecosystem services are
by denition the benets that people obtain from ecosystems
(Millennium Ecosystem Assessment, 2005). However, even if
ecosystem services are a successful concept, they need much
deeper theoretical development (Currie, 2011; Morelli and Mller,
2015). One good example of the need for potentially better
understanding of the concept of ecosystem services is when this
concept is linked to the redundancy hypothesis, originally related

http://dx.doi.org/10.1016/j.ecocom.2016.07.005
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to the concept of ecosystem stability, but currently extended also

to ecosystem assessment. The denition of ecological redundancy is based on the functional traits of species present in a given
community. But this approach has proven to provide an incomplete framework for ecosystem assessments due to the complexity
of ecological systems.
Even if a variety of denitions of complexity exists, the
ecological complexity is mainly associated with two aspects:
complicatedness (dened as the quantity of components), and
hierarchical complexity (the levels of arrangement in the
components) (Kolasa, 2005; Ricard, 2003; Van de Vijver et al.,
2003). This statement means that the ecological complexity is not a
simple count of components or relations among them, but also a
question about the strength of bonds among components (Kolasa,
2005). The main type of biotic interactions constrains the spatial
distribution of species/individuals across several mechanisms,
such as predation, competition, resource-consumer interactions,
host-parasite interactions, mutualism and facilitation (Bascompte,
2009; van Dam, 2009; Wisz et al., 2013b). We dene the
complexity of biotic interactions among species, populations or
individuals as the intricacy of biotic relationships at different
levels: from the complexity of the food web to the variability in
each single biotic interaction: i.e. the non-linearity of predatorprey dependencies and their temporal uctuation, vast array of
feeding interactions, interaction of horizontal and vertical diversity
(Duffy et al., 2007), antagonistic and mutualistic networks, as well
as other biotic interactions unrelated to food chains and cascades
of interactions (Allesina and Tang, 2012; Askeyev et al., 2010;
Bondavalli and Bodini, 2014; Eklf et al., 2013; Jochner et al., 2016).
For Bak et al. (1988) the species in an ecosystem support each
other in several ways, which cannot be understood by studying the
individual constituents in isolation: a point of view strongly linked
to complexity theory. Complexity theory emphasizes self-organization of the components of an ecosystem, and supports the idea
that there is no central control or master plan (Currie, 2011).
Organism assemblages fulll a critical set of ecological functions
for ecosystems (Barbet-Massin and Jetz, 2015). However, each
particular species plays a different role in the ecosystem; a role
that could be more or less unique (or redundant) (Naeem, 1998).
Increasingly, the role of biodiversity on ecosystems requires a clear
understanding of the roles of species richness and species
composition in communities (Downing and Leibold, 2002).It has
been demonstrated that diversity in traits among species and
derived functional diversity controls ecosystem functioning more

often than does species number (Mulder et al., 2012; SchererLorenzen, 2005).
2. Diversity and ecosystems stability: persistence, resistance
and resilience
The relationship between diversity and stability of ecosystems
has fascinated ecologists for many years (Borrelli et al., 2015;
McCann, 2000; Namba, 2015; Schleuning et al., 2015). The
diversity-stability-hypothesis has a long history from its initial
proposition (Elton, 1958; MacArthur, 1955; Odum, 1953), with a
temporary disappearance of consensus (May, 1973), to more recent
evidence suggesting that diversity begets stability (Kinzig et al.,
2002; McCann, 2000). Currently, more evidence supports the
hypothesis that the stability increases following the number of
links in a food web, while restricted diet reduces stability. If the
number of prey for each species remains constant, more species in
the community will increase stability (MacArthur, 1955). However,
nowadays the debate over the relationship between complexity of
ecosystems and ecological stability is far from settled. Some recent
papers, in fact, did indicate how the question is hard and how the
relationships between stability and complexity' could be positive,
neutral, or even inverse (Allesina and Tang, 2015, 2012; Altena
et al., 2016; Namba, 2015).
Following Scherer-Lorenzen (2005), ecosystem stability is
divided into three aspects: (i) persistence: the tendency to exist
in the same state through time; (ii) resistance: capability to remain
unchanged in the face of external pressures or disturbances; and
(iii) resilience: ability to return to its original or equilibrium state
after it has been displaced from it by external pressures. The term
resilience was introduced to the eld of ecology in the 1970s by
Holling (1973) as a measure of the persistence of systems and of
their ability to absorb change and disturbance and still maintain
the same relationships between populations or state variables.
Resilience is often dened as the ability of a system to absorb
disturbances and still retain its basic function and structure
(Walker and Salt, 2006) and the capacity to change in order to
maintain the same identity (Folke et al., 2010) (Fig. 1B). However,
stability can also be dened as the size of perturbation needed to
cause a change in identity of the system, while resilience can be
associated to the elasticity of the system: the rate of change of an
output divided by the rate of change of an input during
perturbation. Under these statements, ecosystems can be classied
into 4 groups: brittle (high resilience, low stability); exible (low

Fig. 1. The concept of engineering (A) and ecological (B) resilience illustrated by the simplication with the ball-and cup heuristic. The cup represents the region in the state
space, in which the system tends to remain, and includes all possible values of system variables of interest. The basin represents the different regimes that the system can
assume without loss of identity (source: Liao, 2012).

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resilience, high stability); robust (high resilience, high stability);

and fragile (low in both).
3. Ecological redundancy: is there an insurance for ecosystem
services?
In ecosystem assessments, ecological redundancy is dened as
functional compensation due to different drivers (species or
organism) performing similar functions in an ecosystem (Walker,
1992). In this way, particular species can cause aggregate stability
in an ecosystem, enhancing ecosystem resilience. In a given
assemblage, the loss of a few ecologically unique species is
expected to have a larger ecological impact than the loss of species
sharing very similar functional traits (Barbet-Massin and Jetz,
2015) (Fig. 2). For instance, if species playing equivalent roles (i.e.
are functionally redundant) go locally extinct, this would not cause
substantial loss in ecosystem function (Fonseca and Ganade, 2001).
Thus larger values of species richness offer functional redundancy
of assemblages and may thus buffer them against loss or
immigration of species (Lawton and Brown, 1993; MacArthur,
1955). Yet, the supposition made by Walker (1992) that greater
conservation efforts must be exerted in sites where redundancy of
biological composition is lower, based on the proposition that
species with high redundancy are a lower priority for conservation,
can be risky. This makes a dangerous link between ecological
restoration planning and ecosystem assessment, often used in the
paradigm of ecosystem services. Community ecological theory
plays an important role in the development of the science of
restoration ecology, offering the basis for natural resource
managers to restore damaged ecosystems (Palmer et al., 1997).
But the criteria concerning ecological restoration and conservation
are fundamentally based on conceptual or theoretical models of
nature, commonly adopted in assessment studies (Naveh, 1994;

Suding and Hobbs, 2009). We can highlight this fact using an

example: The term ecological functions (dened as pure
ecological phenomena), is precedent to the term ecosystem
services (De Groot et al., 2002), and it originally referred to the
role of processes and structures in the overall dynamics of
ecosystems. However, later it was used to describe the roles of
ecological structures and processes useful for human society, e.g. in
economic or social activities (De Groot, 1992). The shift of
ecological terminology to the social and economic elds has
accompanied the development of the ecosystem services approach
in recent decades (Prez-Soba et al., 2012). Furthermore, the
concept of biological or functional redundancy is increasingly
applied in studies on assessments of ecosystem services (Casatti
et al., 2015). Response diversity and functional redundancy work in
combination to enhance the resilience of ecosystem services
(Walker et al., 1999; Nystrm, 2006). In this optic, redundancy is
opposite to disparity and provides insurance for ecosystem
service provisioning by allowing some system elements to
compensate for the loss or failure of others (Biggs et al., 2012).
However, in our opinion, the use of the redundancy concept
applied in order to assess ecosystem services, remains unclear and
in need of stronger ecological support. The main issues remain the
lack of consensus about functional indices used for redundancy
calculation and the use of too technical denition, which can result
in misunderstandings. The denition of redundancy used by Biggs
et al. (2012): redundancy is a system property that describes the
replication of particular elements or pathways in a system, sounds
somewhat alien when speaking about species and ecosystems.
Moreover, identication of keystone entities for the assessment of
an ecosystem service based on redundancy values could be a
dangerous strategy. It is virtually impossible to extract a reliable
tool, supported by two concepts that are so vague: ecological
redundancy and benets provided by ecosystems to humans
(ecosystem services). As underlined above, the concept of
redundancy in ecosystem assessments needs more insights, in
order to prevent too simplistic solutions adopted in ecological
planning: Can the redundancy of species justify the replacement of
some of them in a given ecosystem or maybe mobilize large
conservation efforts?
4. Are species replaceable like gears?

Fig. 2. Schematic example of potential effects caused by removal of one redundant

or non-redundant species on the stability of an ecosystem. The cases A and B are the
expected effects as predicted by engineering interpretation of redundancy theory in
ecosystem assessment (stability or instability). The case C proposes also the effects
of positive heterospecic interactions among species (or components), driving a
potential instability of the ecosystem when removing one redundant component.
The classication of a species as redundant based only on species traits (the most
often used approach) is insufcient to guarantee the real effect of the removal of
that species from the ecosystem, due to possible neglected biotic interactions
(Heikkinen et al., 2007; Morelli and Tryjanowski, 2014), temporal changes in the
species interactions (Saavedra et al., 2015), changes in the interaction strength
(Bondavalli and Bodini, 2014) or cascade effects (Altieri et al., 2007).

Some researchers recently focused on the relationships

between functional redundancy and functional diversity, developing several indices useful for studying communities (de Bello et al.,
2007; Petchey et al., 2007). The functional diversity within each
species assemblage is assessed based on the functional traits of the
species present and the combination of functional traits values in
the community (Maire et al., 2015; Petchey and Gaston, 2006). The
same species traits are the elements used to assess the functional
redundancy in a given species assemblage. However, some studies
showed controversial ndings about the right value of the
redundancy hypothesis when tested in natural communities or
real study case: no correspondence between high redundancy and
higher diversity in some marine ecosystems (Worm et al., 2006),
less redundancy than expected in some plant species assemblages
in a restoration faunal habitat study (Moir et al., 2010), and also
some evidence that species with similar functional traits in
ecosystems have biotic interactions more characterized by
complementary, rather than redundancy (Thibault et al., 2010).
We consider that the redundancy hypothesis neglects some
important notions of biotic interactions. For instance, when species
redundancy in a particular community is estimated using a matrix
with species-specic functional traits, the potential biotic interactions unrelated to trophic chains are often forgotten (Farnsworth
et al., 2012). During recent years, ecologists started to emphasize

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the importance of heterospecic biotic interactions on species

assemblages in ecosystems (Mod et al., 2015; Wisz et al., 2013a).
One way to measure the heterospecic biotic interactions is across
spatial correlations among the distribution of species. The
correlation among the occurrence of several species in the same
ecosystem might result from co-evolutionary relationships, shared
habitat preferences (e.g. habitat characteristics, abundance of
particular food resources, etc.), and/or due to heterospecic
attraction (Thomson et al., 2003; Mnkknen et al., 1999). Even
a cascade of biotic interactions could help to clarify the pattern of
several species (Duffy et al., 2007; Mnkknen et al., 1999). In fact,
many studies highlighted the importance of positive heterospecic
interactions in species distribution, as complement to environmental characteristics (Bogliani et al., 1999; Heikkinen et al., 2007;
Hromada et al., 2008; Morelli and Tryjanowski, 2015; Tryjanowski,
2001; Wiklund, 1979). And these biotic interactions have both
ecological and evolutionary effects, because the actions of one
component can affect the interactions among other components of
the community (Mller, 2008).
Currently the notion of ecological redundancy does not
consider the multivariate concept in order to handle the
complexity of the ecosystems, and overlooks the fact that different
species can carry out different functions at any one time (Gamfeldt
et al., 2008; Maire et al., 2015). From this point of view, the
redundancy theory applied to assess ecosystem functions or
services looks like a too simplistic perception. The possibility to
replace a single ecosystem component (or species) by another,
considered redundant, could be a demonstration of nearsightedness of ecological vision. A too simplistic approach, as proposed by
redundancy theory, can cause a series of unexpected effects and
several wrong predictions in applied ecology. Different causes can
contribute to the multidimensional effects of each species in an
ecosystem, independent of its redundancy level. Many measures of
biological redundancy, based on biological traits of species (diet,
dispersal ability, breeding habitat selection) are insufcient to
dene the complex role of species in the community. More
information is necessary in order to dene this role: Abundance of
individuals, connectivity with other populations, and even
heterospecic biotic interactions not only related to the food
web. In fact, it was recently found that rare species are also rarer
than expected by chance, and for this reason any theoretical
supposition about ranking of species on the basis of their
functional redundancy has to be strongly linked to the local/
regional abundance of species (Mouillot et al., 2013). Yet,
functionally redundant species have similar functional traits
(Carmona et al., 2012), but the perceived redundancy can be a
mirage: strongly affected by the traits used to measure it (Petchey
et al., 2007), and changing in time if species shift their ecological
functions (Gamfeldt et al., 2008).
Yodzis (1981) already highlighted how models structured from
compiled food-web relationships, with plausible interaction
strengths, are generally more stable than randomly constructed
food webs, underlying the importance of interaction strength on
the overall stability of ecosystems. Other recent studies emphasized the importance of considering not only the direction of links,
but even combinations between the type of connections (functional or redundant) and the strength of these interactions, that
articulate the food web, constituting crucial principles in order to
understand the stability of these networks (Bondavalli and Bodini,
2014; Tang et al., 2014). In a similar way as proposed by Yodzis
(1981) for food-webs, the principles behind the insurance
hypothesis proposes that biodiversity buffers ecosystem processes against environmental changes such as global warming, because
different species respond differently to such changes. This will
result in functional compensation among species and hence more
stable community properties (Yachi and Loreau, 1999).

5. Engineering redundancy vs. ecological redundancy: so close,

so far away
Some recent studies showed how living organisms and life in
general can be conceived as an integrated information processing
system (Farnsworth et al., 2013). Traditionally, biotic systems are
considered more complex than abiotic counterpart, with more
predictable interactions among components. However, also
engineered or abiotic systems could become complex. Some
researchers argue that the critical difference between abiotic
complex systems and biotic ones is the phenomenon of downward
causation (for more details, see Auletta et al., 2008; Davies, 2012;
Ellis, 2012). Furthermore, complex natural systems show different
behavior than engineering systems: for example, enhancing
resistance to one disturbance in complex adaptive systems often
creates vulnerabilities to others (Holling and Meffe, 1996). This
notion, of course, could be achieved in some cases, even for
engineering systems.
In technology design, mainly in informatics sciences, scientists
have focused in recent years on failure tolerance (redundancy
theory applied to informatics) (Li et al., 2010; Owre et al., 1995).
However, the current tendency is to focus on a completely different
kind of paradigm: failure prediction or failure acceptance
(Ravisankar et al., 2010; Wang et al., 2014). In informatics,
increasing redundancy causes the code (with the information or
message) to become larger and more fault-tolerant, and it takes
less effort to understand the message. But there exists a penalty to
pay for redundancy: the increase in bandwidth required and the
additional effort. In nature, all systems tend to minimize the cost/
benet ratio, and there does not exist a general message to be
transmitted, as in informatics. In information theory, each
component (or symbol) serves to transmit a message, that many
times is the sum of all components (e.g. the meaning of the
discourse), while in ecology, each component (or species) could be
considered a nal message in itself. For this reason, the hypothesis
of accumulation of redundant pieces (species) in an ecosystem
looks more like a human artifact than an ecological condition. In
the same way, although short-term or single measures in several
analyses may suggest redundancy in ecosystem components, some
studies support the longstanding dictum that niches of coexisting
species are often similar but not identical, reducing the potential
redundancy values among species (Thibault et al., 2010). Still, the
addition of similar species in a particular ecosystem can provide
contrasting effects, demonstrating for example how the interference of redundant species (predators) can decrease or increase the
predation rates, depending on the characteristics of predators and
their potential for intra-guild predation (Griffen and Byers, 2006).
Moreover, Polak et al. (2015) showed how ecological planning only
using an ecosystem-based approach
based on environmental
surrogates at various scales, and trying to represent biodiversity as
a whole, including species and processes (Margules et al., 2002)
does not provide the most efcient outcomes due to lack of
insurance for all species requirements.
Partially, we can guess the causes of this mismatch between
engineering and ecological systems: we know that the functional
diversity approach is essential in order to comprehend the patterns
of species co-occurrence, the community assembly rules and the
role of different traits on ecosystem functioning (Laureto et al.,
2015). The denition of species redundancy is based on the same
species traits or function traits, that are the bricks used in the study
of functional diversity. But recent studies, exploring and quantifying the importance of multiple functional diversity components as
drivers of species coexistence, showed how it is important to
consider a multi-trait approach, and also focus on measures of
within-species, between-species and total functional diversity
variance (in other words, try to partition the functional diversity

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in each community) (de Bello et al., 2010; Le Bagousse-Pinguet

et al., 2014). In effect, recognizing the conceptual complexity of
functional diversity, some researchers proposed to adopt a unied
framework that implement the Hutchinsonian concept of the niche
as a probabilistic hyper-volume for the calculation of functional
diversity (Colwell and Rangel, 2009; Hutchinson, 1957); a method
that authors called the trait probability density (TPD) approach
(Carmona et al., 2016).
6. Final considerations
In summary, a limited approach, as suggested by the
assessment of species redundancy focusing only on species traits
or function traits (currently the most used approach), neglecting
or minimizing the importance of interaction strengths in food
webs, multiple-interactions (Mller, 2008) or even other types of
biotic interactions, can constitute further evidence of a too
anthropocentric approach (science is made by humans, after all),
but capable of distorting the vision or paradigm on which science
tries to focus. For this reason it is important to highlight the
fundamental differences between engineering resilience and
ecological resilience (Holling, 1996; Liao, 2012), trying to avoid
the use of two interpretations that can cause an unreliable
evaluation of the role of species and their importance for the
stability of the ecosystem, leading to a wrong assessment and
suggesting contrasting measures for management. With this
aim, we provide in Table 1 a synthesis of the main characteristics,
as well as the expected responses of the systems facing changes,
in order to clarify how engineering and ecological resilience
differ.
The concept of redundancy applied in ecology is also backed by
another misleading concept: effectiveness or functioning of the
entire system (ecosystem). But functioning is only apparent or
provisional in ecosystems, because it is continuously evolving.
Evolution teaches us that all balances are only temporary. In this
comment we propose to pay more attention to the potential and

variable relationships among ecosystem components (species),

when using tools developed to assess biotic communities based on
redundancy approach: More attention must be paid to considering
multidimensional species niches that can constrain species
interactions when assessing the extent to which one species can
substitute for another (Yachi and Loreau, 2007), because the
functional complementarity could be a more appropriate descriptor of species assemblages than functional redundancy (Thibault
et al., 2010). On the other hand, the effects of the replacement of a
redundant species by another can vary under some conditions. The
relative functional importance of different species (rare or
common species) on ecosystem multifunctionality changes even
depending on the function considered (Pendleton et al., 2014;
Soliveres et al., 2015). However, even if it is possible to establish
and quantify the relationship of redundancies among parts of the
ecosystem, this redundancy can change over time, because many
species interactions embedded in ecological communities are not
permanent, but characterized by temporal changes that follow
abiotic and biotic variation (see some examples in Gamfeldt et al.
(2008) and Saavedra et al. (2015)). Thus, any calculation about
redundancy can only be relative to the process at hand, at a given
time, and it can be biased by the multidimensional role of the
species in an ecosystem and an incomplete denition of the
multiple species-trait dimensions (Boersma et al., 2016).
In practical terms, we suggest that redundancy assessment of
species needs to incorporate a more complete list of heterospecic
biotic interactions and potential cascade effects of interactions, as
well as the explicit integration of the strengths among links
connecting ecosystem components, in order to achieve a most
complete vision of the role of each species in the mosaic of an
ecosystem. More attention must be paid to the per se values of each
ecosystem component, in contrast to the derived values, as the
values relative to their contribution to provisioning of ecosystem
services. Simultaneously, we emphasize how a precautionary
principle to decisions affecting the natural world is necessary
(Gascon et al., 2015), mainly when ecosystems are assessed on the

Table 1
Highlights of the main differences related to the redundancy hypothesis when applied to engineering systems and ecological systems (ecosystems): characteristics, expected
reactions to a change, interaction among components, and effects of the elimination of a component.
Characteristic

Engineering redundancy

Ecological redundancy

Type of stability

stable equilibrium (Holling, 1996)

System probabilities of failures

Time of recovery to normal levels of
functionality after a failure
System predictability after change
System probabilities of return to the initial
state after the disturbance
Expected behavior of the system after the
disturbance
Expected redundancy
Stability key
Fluctuation of the system (effects)
Number of regimes that the system can
assume
Role between redundant components
Redundant component function in the
system
Interpretation of redundancy of system

low (Wang and Blackmore, 2009)

quick (Liao, 2012)

instable equilibrium, multiple-equilibrium state (Scheffer

et al., 2001)
high (Liao, 2012)
slow (Holling, 1973)

high
high

low (Liao, 2012)

null or very low (Folke et al., 2010; Holling, 1973)

recovery: back to the original condition (Wang and

Blackmore, 2009)
remain unchanged (recovery capacity)
consistency (Liao, 2012)
deviant effect
mono-regime (Fig. 1A)

reorganization: transformation into a new (equivalent)

condition (Gunderson and Holling, 2002)
survive after change (reorganization capacity)
persistency (Liao, 2012)
potential evolutionary effect
multiple-regime (Fig. 1B)

identical (Holling, 1996)

duplicate (Liao, 2012)

similar, compatible (Holling, 1996)

diversity and replicate (Adger et al., 2005)

the extent to which system components are substitutable

(Liao, 2012)
Interactions between components
null or clearly dened
Removal of a redundant component (effects) replace or elimination of one element = reduction on the
overall redundancy of the system
Addition of a redundant component (effects) addition of one element = increasing the overall
redundancy of the system

ability to absorb disturbances and reorganize to persist

(Holling, 1996)
undened, potential biotic interactions
extinction of one species (loss of evolutionary distinctiveness)
(McCann, 2000)
invasion of one species (cascade effect on ecosystem)
(McCann, 2000)

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basis of different paradigms. In fact, socio-economic tools

(ecosystem services approach) plus tools derived by
engineering (redundancy theory) are insufcient for assessing
the complexity of ecosystem relationships: a more ecological
vision is needed.
Acknowledgements
We thank A. P. Mller, M. Moretti, Y. Benedetti and L. Brotons for
their valuable contributions during the writing of this manuscript.
We are also grateful to J. Kolasa and the anonymous reviewers for
their important suggestions, that were very useful in order to
clarify difcult concepts. Finally, we are very grateful to the prof.
Emma Nelson, University of Liverpool, for the nal English proof
reading service.
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