Decision-makers need to act now to halt biodiversity loss,
and ecologists must provide them with relevant species
interaction indicators to inform about community- and
ecosystem-level changes. Yet, the integration of ecological
networks into conservation is still virtually non-existent.
Here, we argue that existing data and methodologies are
sufficient to generate network information usable for
conservation, and to begin overcoming existing barriers to
the integration of network information and biodiversity
decision-making. Interaction network indicators must meet
criteria important to decision-makers and be tied to
specific conservation goals, which requires academics to
better engage with practitioners. We use network robustness
as an example of an already applicable indicator, and
showcase its potential with a reusable workflow to inform
decision-making.
Practitioners and scientists increasingly need
multi-species and whole-ecosystem indicators for
biodiversity conservation and management. Species
interaction networks hold a promising potential to fill
those needs.
Explicit and quantitative integration of network
indicators into conservation is still lacking due to
challenges with uncertainty and indicator accessibility to
practitioners. The resulting gap between network science and
management leads to decisions being made without considering
available scientific knowledge.
We need to start bridging network information into
biodiversity management, towards application. We can do this
now, building on existing metrics and available data as
starting points. We must accept data limitations and
uncertainty, and use what we have to establish an
operational framework and then focus on improving it with
better data and sampling programs.
Can
interaction network knowledge be quantitatively used for
biodiversity conservation and management?
The need to shift from single-species conservation
approaches to multi-species and whole ecosystem approaches
has long been recognized [1,2]. Network information can
provide a new perspective for whole ecosystem assessments in
biodiversity conservation and management. Preserving species
interactions can ensure long-term population persistence and
maintain ecosystem functions and services [3,4]. Focusing on
species interaction networks (see Glossary) as conservation targets
promotes the stability of populations and ecosystem
functions and minimises negative outcomes regarding species
extinctions [5–7]. Recent reviews list
specific interaction network metrics that
decision-makers can use [8]. Implicit network
information has already been integrated into conservation
planning, which should facilitate the uptake of
network-based biodiversity indicators in
decision-making [2,9,10]. For instance, network
information is implicitly integrated through the
consideration of keystone species that disproportionately
affect local communities (see Box 1).
Despite the potential benefits, conservation practices
rarely explicitly consider information derived from
ecological network metrics, and conservation policy and
practice still heavily focus on single species and habitats.
This is in part due to uncertainty, and in part due to the
choice of indicators. Uncertainty about network structure
and responses to human disturbances mirrors concerns in
macro-ecological and ecosystem models [11,12]. Additionally,
identifying which interaction network metrics are suitable
biodiversity indicators with clear interpretation for
conservation remains challenging.
Decision- and policy-makers must act now to bend the
curve of extinction and accelerate ecosystem recovery [13,14]. Ecologists need to
provide them with useful network and ecosystem-wide
information. For instance, protected area planning could
prioritise regions where mutualistic interaction partners or
prey and predators overlap [15], or where there is high
trophic diversity and redundancy, enhancing
robustness to extinctions [16]. Moreover, since
interaction network structure is linked to ecosystem
functioning and ecosystem service provision, focusing on
network metrics changes for conservation targets should
ensure ecosystem stability and service delivery [e.g.,
pollination, pest control, food production, 5,7,17]. Given the global goals
to maintain ecosystem services [Goal B of the Kunming-Montreal
Global Biodiversity Framework, 18], assessing changes in
network structure and stability should help managers and
decision-makers prioritise areas to maintain ecosystem
functioning and resilience [5,19].
Here, we identify the major challenges and opportunities
in incorporating information from species interaction
networks into biodiversity conservation and ecosystem
management. Despite these challenges, we need to start
integrating network concepts into management and
conservation in the face of global change, as we have
sufficient scientific evidence and tools to do so. Using
network robustness as an example, we show how simple
approaches and indicators can provide relevant information
for managers based on decision-making criteria, available
data, and reproducible workflows.
Box 1 - Network information is already
implicitly considered in conservation and
decision-making
Explicitly considering interaction networks in
conservation and decision-making (i.e. by
monitoring and managing for network-derived
indicators) is not a drastic shift, as they are often
implicitly included in conservation decisions and recovery
plans. For example, the keystone species concept, frequently
mentioned in conservation literature [e.g., 2,20] and highlighted by
initiatives focused on rewilding and ecological restoration
[21,22], is linked to the
disproportionate effects some species have on their
(trophic) networks and ecosystem functioning [23, also see 24 for
the diverse roles of species identified as
keystones]. Similarly, several large carnivores have
been associated with trophic cascades, where effects of
predator declines propagated across food webs to herbivores,
mesopredators, and beyond [25]. This reflects network
consideration through species’ effects on others, even if
network-specific metrics are not explicitly quantified (see
network metrics in Glossary) and do not explicitly enter
planning or decision-making.
Importantly, keystone species are often tied to
quantified conservation targets, highlighting how the
concept is both accepted and used by practitioners. For
example the Recovery Strategy and Action Plan for the
Black-tailed Prairie Dog (Cynomys ludovicianus) in
Canada identifies it as a conservation priority due to its
keystone status – crucial for the recovery of the
Black-footed Ferret (Mustela nigripes) and a vital
food source for several other at-risk species [26]. Conservation targets
for Black-tailed Prairie Dogs in Canada include maintaining
a minimum area of occupancy of 1,400 ha across 20 colonies
and a minimum average population density of 7.5
individuals/ha by 2040, ensuring at least an 80% probability
of population persistence over 50 years [26; targets on which recovery of
the Black-footed ferret also depend].
The existing implicit consideration of network structure
in conservation targets can facilitate the uptake of new
network-based indicators by practitioners and
decision-makers. Other forms of network-thinking are also
part of management considerations, such as spatial
ecological networks planning [27] and ecosystem-based
management [11]. Explicitly considering
indicators of interaction networks will complement these
forms of network-thinking and enhance conservation
assessments to include ecosystem-wide components.
Challenges &
Opportunities
The explicit integration of network information into
management and conservation faces several challenges linked
to uncertainties and lack of interpretability and relevance
of network metrics for practitioners. These challenges will
hinder making effective decisions, for example on what
biodiversity and network-related metrics need to be measured
and monitored, what conservation targets and management
actions should be applied, how often to re-evaluate
decisions, etc. Hence, we can expect challenges at different
stages of management planning and decision-making [e.g. 28], such as the evaluation
of current conditions or upon decisions on possible actions
(e.g. responsive, preventative, etc.).
Uncertainty
Network Structure and
Composition
There is uncertainty in network structure, composition,
and variation across space and time, which affects
conservation assessments and actions [29,30]. Empirical studies on
networks are often spatially disjointed, biased
geographically and in the types of interactions, and rarely
replicated [31–33]. Sampling biases can
distort reported network patterns [34,35]. Terrestrial and
freshwater food webs are less studied than marine ones,
often with different research objectives [e.g.,
determining the effect of environmental factors, rather than
investigating management-related elements such as
sustainability, 31,36]. Such deficits of
information are problematic for decision-making, as it may
seem impossible to extract hard and transferable
(geographically or between ecosystems) guidelines for both
scientists and practitioners.
Despite these challenges, existing methodologies and data
can help integrate network information into conservation,
while empirical data continue to be gathered. Food webs can
be constructed from extensive, long-term monitoring datasets
to analyse network structure and stability [37,38]. As
binary interaction data are commonly
available, we can start ahead with these to establish
operational monitoring frameworks, while later integrating
uncertainties and flow-based data for a deeper and
error-informed understanding of ecological systems. Building
metawebs of all potential interactions in a
region or species pool, like the pan-European terrestrial
tetrapod metaweb [TETRA-EU, 39], can help inform
broad-scale assessments of network structure [40,41]. Metawebs have already
been used to derive spatially explicit network metrics and
generate conservation-relevant information [42–44]. For instance, Albouy
et al.[42] used a metaweb to
examine robustness to extinction scenarios for marine food
webs, showing higher robustness in coastal waters compared
to open waters and highlighting some potential to absorb
perturbations. Moreover, metaweb inference approaches allow
us to circumvent the lack of available local interaction
data [40] and, when used with
probabilistic networks, to integrate
uncertainty and variation in network structure across space
[45]. Network metrics and
their uncertainties can therefore be measured for
broad-scale assessments of variation in network structure,
and to derive network indicators that can be used to inform
decisions and planning (Boxes 2-3).
As new empirical data becomes available, predictions can be
evaluated, refined, and become more informative [46]. We discuss the
challenges surrounding their validation in our Concluding Remarks.
Network
Responses to Environmental Change
Uncertainty exists in how networks will respond to
environmental changes and disturbances, particularly for
interaction rewiring and changes in
interaction strength. Questions remain on the extent of
rewiring due to species turnover versus prey switching and
behavioural adaptation, and how these changes will propagate
across trophic levels.
While data gaps exist, modelling and inference can
explore the limits of network rewiring under current or
future conditions (Box 3).
Rewiring potential is likely captured in existing and
inferred metawebs [47], which can be combined
with simulations to anticipate network changes. For
instance, Dansereau et al.’s [45] approach can be extended
to explore climate change impacts on network structure,
given the dual uncertainty in species interactions and
future species ranges. Importantly, network models (and
information) do not need well-constrained or low uncertainty
predictions before they can inform management decisions on
interventions like species eradication, especially if they
tend to correctly identify whether effects on other species
will be positive or negative [48]. Model uncertainty can
also be high despite high quality data [48]. Regardless of its
generality, this suggests that the performance of a model
should be monitored whenever new data are added. Similar
trends of model change in performance with additional data
have been reported in the study of species distributions
[49].
Approaches to include specific types of network response
uncertainty in conservation and management have also been
proposed. Van Kleunen et al.[50] suggested a multi-step
framework for decision-making under uncertainty for species
introduction into ecological networks, based on conservation
decision theory. This framework includes: the identification
of management objectives, the evaluation of outcomes for
management (including multiple outcomes, evaluation of
trade-offs, and assessment of uncertainty), and the
improvement of future predictions through an adaptive
management framework. Van Kleunen et al.’s [50] decision-making approach
can be applied now, despite uncertainties, to guide
management of species introductions.
Compounding
Uncertainty in Change Types
There is compounding uncertainty in the type and strength
of change applied to a network. Climate uncertainty, for
instance, results from uncertainty in future greenhouse
gases emissions (i.e. emission scenario uncertainty), in
climate processes (general circulation model uncertainty)
and their stochasticity (model run uncertainty). For
networks, we add uncertainty in changes resulting from
disturbance regimes (e.g. fire, drought, pests) and in
species distribution predictions [which can
result from direct impacts of abiotic change, of disturbance
regimes and of biotic changes that may be linked to network
structure itself, 51,52]. If accounted for
simultaneously, these uncertainties will inevitably lead to
high variance in predicted network responses.
We can estimate some uncertainty through
hindcasting: past environmental changes are
used to predict changes in network metrics that are
cross-validated against observed past or current networks.
Fisheries data, for instance, allow reconstructing
well-resolved networks over time, which can be related to
known environmental changes [53–55] and be used to calibrate
predictive network models, like Bayesian networks [56]. Hindcasting models,
used as ex-ante scenarios of change, have been successfully
used to simulate and assess the effectiveness of
conservation actions on ecosystem services [57].
Simulating scenarios of change can also help delimit the
possible changes in network structure [Box 3, 58]. When combined with
metrics of network change and sensitivity to disturbance,
these projections can be used to identify
target areas that show fragility to an array of scenarios
and are of special concern, or that show less fragility and
could be considered refugia. They can also highlight
problematic or incomplete sampling. Projections will also
serve to perform validation and assess indicator behaviour
in an empirical setting, whether through existing data or
hindcasting exercises, which could lead to network-specific
monitoring programs.
Interpretability and
relevance
Network metrics are often not intuitive or deemed
relevant for practitioners and decision-makers; many are
complex and may not show clear relationships with ecosystem-
and species-level responses, particularly in applied
contexts. For instance, omnivory and network motifs are tied
to food web robustness and extinction risks [59,60], highlighting their
ecological relevance. On the other hand, while network
nestedness indicates a buffer against extinctions and
fluctuations in mutualistic networks, this is less clear in
antagonistic networks [7]. Connectance has also
been tied in contrasting ways to network stability, with
higher connectance leading to increases or decreases of
invasion success rates given invader trophic levels [61], or linked to higher
robustness to extinction, but larger extinction cascades
[62].
Moreover, not all network metrics are suitable as
conservation indicators, nor do they need to be. Several
have been reviewed for their relevance and limitations in
achieving conservation goals [63,
see Table 1 therein]. For example, prioritising
trophic networks with stabilising motifs when selecting
protected areas can help achieve ecological resilience goals
[63]. This information can
already be used towards conservation planning but it needs
to be both accepted by and available to decision-makers and
managers.
First, metrics must meet decision-makers’ criteria. The
ROARS (Relevant, Objective, Available, Realistic, Specific)
and SMART (Specific, Measurable, Achievable, Replicable,
Time-bound) criteria [8, see Table 3 therein]
focus on the decision-makers’ receptiveness to suggested
indicators during the selection, paving a way to communicate
network information with stakeholders and embed network
indicators in ecological monitoring and ecosystem health
assessments. Network indicators will therefore need to be
evaluated in terms of usefulness to achieve conservation
goals [63]and
decision-maker receptiveness [as in 8], as we move towards
developing ecosystem management and monitoring frameworks
that quantitatively and explicitly embed network indicators
(see example in Box 2).
Second, network ecologists have the opportunity to expand
their focus from the development of mathematical tools,
theory and theoretical validation, to involving
decision-makers and meeting their needs [64]. Consensus for
conservation goals can be achieved through mixed
methodology, such as iterative and anonymous Delphi panels
[see
65 for applications in
ecology]. Engaging stakeholders in this way would
ultimately provide valuable guidance to prioritise new
fundamental research questions and methodological
development. Although they do not ultimately make the
decisions, network ecologists must be proactive in this
process, especially given the limited time and staffing
resources across many institutions where decisions are made.
This process takes time and co-production effort, and needs
to be promoted by academics who can guide and support
practitioners in designing management strategies and making
conservation decisions using network information. Academics
place a strong focus on the development of tools and
knowledge, but ensuring their adoption (particularly for
non-academics) will require delivering them in a form that
can instantly be used with minimal additional work [66].
Finally, network ecologists can take concrete steps to
ensure that network-based measures are perceived as relevant
by decision-makers. Workshops and stakeholder involvement
are essential to bridge the gap between science and practice
[66] and can facilitate
choosing appropriate metrics [8]. Involving a wide-range
of ecosystem-management players, and creating new
opportunities to actively involve stakeholders in deciding
how network information can be applied, will be key to
ensure receptiveness and a speedy uptake of indicators for
management planning and actions.
Forecasting changes in network structure
under environmental and management scenarios (Box 3)
and linking network indicators to ecosystem services [17] can enhance
receptiveness, especially if we clearly demonstrate that
forecasts work well. This will provide essential information
on risks, on boundaries of change given environmental
conditions, and on the effectiveness of certain management
actions in achieving conservation targets [67].
Box 2 - Assessing the relevance of a
potential network indicator for decision-making
Network metrics should be evaluated using criteria
important to decision-makers to ensure their relevance as
indicators and encourage adoption. In addition to the ROARS
and SMART criteria, Fath et al.[8] suggest that effective
indicators should also “describ[e] directional change
[of ecosystems], [be] easily communicable to managers and
policy makers, [be] integrative and indicative to a known
response to a disturbance” [as per 68], and provide insight to
ecosystem functioning and services.
Table I illustrates the process of detailing how a
potential network indicator meets the criteria mentioned
previously, using trophic network robustness to species
extinctions (hereafter robustness) as an example. Evaluating
network metrics in this way is crucial for making them more
relevant and acceptable to decision-makers, as it
demonstrates why and how the indicator can be used
effectively. We emphasize that such evaluation should be
done with other network metrics to facilitate uptake by
practitioners and decision-makers.
We chose robustness as it can be a useful indicator of
ecosystem integrity and stability to environmental change
given data we already have. The structural stability of
trophic networks is closely linked to the stability of
ecosystem functioning [see review by 69], with trophic
interactions considered as ecosystem functions and services
(e.g., top-down pest control by predators). Here we show a
formulation of robustness derived from earlier work [70–72] that reflects the
capacity of a network (or the ecosystem it represents) to
withstand cascading extinctions:
where ‘no. secondary extinctions’ are
extinctions due to the loss of prey species and
‘secondary consumers’ are species that
consume other species in the network (calculated as network
species richness minus the number of basal
species). ‘Initial’ refers to before extinctions
took place.
Robustness is easy to interpret (see Specific in
Tbl. 1) and to calculate using binary trophic networks,
which are more commonly available and can be constructed
from existing trophic metawebs – this allows us to derive
initial (even if coarse) estimates of robustness at large,
regional and local scales (see references in Tbl. 1). It
also relates to ecological issues that have a firm place in
ecosystem management and conservation, and resonate with
decision-makers – numerous directives, policies and
management frameworks focus on avoiding species extinctions
(see examples in Tbl. 1). By showing here how robustness
meets decision-making criteria, we highlight a process
transferable to other network metrics to identify the most
applicable ones for biodiversity conservation and
management.
Table 1: Relevance of a network indicator
for decision-making. Dale & Beyler’s [68], ROARS (Relevant,
Objective, Available, Realistic, Specific) and SMART
(Specific, Measurable, Achievable, Replicable, Time-bound)
criteria for good ecological network indicators, as
described by Fath et al.[8], and how they apply to
robustness of trophic (binary) networks.
Robustness measures loss of species with respect to a
given (pre-disturbance) species composition.
Easily communicable to managers and policy makers
The relationship between robustness and species
extinctions is intuitive and easy to understand.
See also entry for “Relevant” below.
Integrative and indicative to a known response to a
disturbance
Trophic networks summarise the energy flows in an
ecosystem; their structural stability is linked to stability
of ecosystem functioning [69].
Robustness measures trophic network responses to
disturbances that lead to cascading species
extinctions.
ROARS
Relevant
It relates to an important part of an objective or
output
Preventing species extinctions is at the heart of
numerous conservation policies, directives and frameworks
[e.g.,
73,74–76].
Objective
Based on facts, rather than feelings or impressions and
thus measurable
Robustness is based on assessments of species
composition pre- and post- disturbance.
Available
Data should be readily available or reasonably
measurable
At the regional scale, available metawebs [e.g., 39,53] can be combined with
species range data (e.g., IUCNii and GBIFiii) and scenarios of change to
assess robustness (see Box 3).
Sub-regional/local scale assessments are possible in
locations with monitoring data [e.g., 37,38].
Realistic
It should not be too difficult or too expensive to
collect the information
Marine and freshwater network data are already being
collected as part of monitoring programs and fisheries
activities;
Terrestrial metawebs exist [39] or can be inferred [77]
Methodology to calculate robustness is not overly complex
and can be pipelined (see Box 3).
Specific
The measured changes should be expressed in precise
terms
Robustness is calculated as 1 minus the ratio of
secondary extinctions to the initial number of secondary
consumers. It is scaled from 0-1, with 1 indicating maximum
robustness (no secondary extinctions) and 0 indicating no
robustness (all secondary consumers went secondarily extinct
due to loss of feeding resources).
SMART
Specific
Measured changes should be expressed in precise terms
and suggest the direction of actions
See entry for “Specific” above.
Maps of robustness can indicate hotspots and priority
areas for conservation.
Networks with high robustness will indicate ecosystems
whose structure is more stable and that could be managed as
“safety nets” and/or with more liberal use. Those with low
robustness should be further assessed for their uniqueness
(e.g,. uniqueness of species composition and interactions,
of habitat type, etc.) to plan conservation
actions.
Measurable
Indicators should be related to things that can be
measured in an unambiguous way
In an empirical setting, there may be ambiguity in
determining whether an extinction was secondary (due to loss
of other species in the network) or primary (due to, e.g.,
loss of climate suitability).
In a modelling setting secondary and primary
extinctions can be determined. Null models can be
used to test whether projected extinctions significantly
deviate from random.
Uncertainty in both network species composition and
structure will need to be recognised and accounted for
explicitly whenever possible [e.g., 45]
Achievable
Indicators should be reasonable and possible to reach,
and therefore sensitive to changes
See entry for “Available” above.
Hindcasting and historical observational data can be used
to gauge the sensitivity of robustness to past environmental
change.
Forecasting data can be used to assess robustness
boundaries to expected changes and complemented with
monitoring data to verify how networks are responding to
change.
Replicable
Measurements should be the same when made by different
people using the same method
Transparent and freely accessible pipelines can be
developed and automated to ensure repeatability.
Time-bound
There should be a time limit within which changes are
expected and measured
This likely depends on the species and type of
environmental changes considered, given different life cycle
histories and species’ sensitivities to change.
Box 3 - Illustration of an accessible and
reproducible workflow to inform decision-making using
network robustness
Effective decision-making requires indicators based on
accessible and reproducible workflows that evaluate a range
of scenarios. Keeping trophic network robustness as our
example, we demonstrate how such a workflow can be built
using different network disturbance scenarios and
open-access data (Fig. 1). By using extreme scenarios, we
can explore the boundaries of robustness to projected
environmental change. The framework can be applied spatially
to identify target areas for management and conservation
action (Fig. 2), or to single well-resolved networks
(e.g. local scale).
Workflow steps:
Build local ‘baseline networks’ by combining a regional
metaweb of interactions with ‘baseline’ local species
presence/absence information (‘baseline’ on Fig. 1 referring
to any reference period) – species that interact in the
metaweb and are locally present will appear and interact in
the local network;
For each baseline network, calculate the number of
secondary consumers and other relevant network metrics
(e.g., species and average trophic level, connectance,
etc.);
Build local ‘disturbed networks’ by combining the
regional metaweb with species ranges projected under
different scenarios;
Calculate and map robustness and other network metrics
(Fig. 2).
Figure 1: Workflow to
calculate robustness. Simple network metrics like
robustness can be incorporated into workflows to assess
potential ecosystem fragility to scenarios of disturbance
and inform management and decision-making at large
scales.
Here we illustrate this workflow using a worked example
with pan-European tetrapod trophic networks. We explore the
boundaries of network robustness by using two extreme
scenarios: worst-case climate change (CMIP5 RCP 8.5,
equivalent to CMIP6 SSP5-8.5), and failure to protect
endangered species (loss of all species with IUCN status of
Critically Endangered, Endangered, and Vulnerable, across
their entire range). The scenarios caused changes in species
composition due to climate-driven range shifts (‘climate
change’ in Fig. 2) or to targeted species removals (‘IUCN
extinctions’). Two extinction outcomes were possible:
species became primarily extinct when predicted to be absent
from a pixel due to future climatic conditions or due to
targeted removal, or secondarily extinct when the pixel was
climatically suitable but had too few prey items. Following
the workflow above, we used a metaweb adapted from TETRA-EU
[78] build baseline and
disturbed local networks [using projected species
distributions based on habitat preferences and
presence-absence data from 79], calculate the number of
secondary consumers (from baseline networks) and secondary
extinctions (from disturbed networks), then calculate and
map robustness (see Supplemental Information online for full
workflow details).
In this example, most networks were very robust to
extinctions driven by a) climate change or b) the removal of
endangered species listed in IUCN, but several networks in
Northern Europe, Crete and in the Canary Islands show lower
robustness to targeted IUCN extinctions (Fig. 2 b). For the
purpose of this illustration, we show median robustness
values per ecoregion [defined by 80], which represent
geographically meaningful boundaries for species and
interaction composition [81] and simultaneously
highlight a regional-level at which robustness can be used
to inform policy-making (see Supplemental Information,
Figure S2 for pixel values). We note that this is a
conceptual illustration to present robustness as an example
of a readily applicable indicator given the data we already
have. Yet, further analyses could be focused on
investigating which species are projected to be lost, their
roles in the networks and best strategies to protect these
networks from a multispecies perspective.
Antunes et al.[17] proposed a similar
workflow to calculate network-provided Nature’s
contributions to people, but our framework involves
methodological approaches that are less sophisticated and
data-hungry. We emphasize that presenting a fully worked
example for potential network indicators, as we do here with
an accessible automated pipeline [82], is a transparent and
practical way to not only encourage the development and
sharing of reusable analyses, but also to facilitate and
accelerate uptake by practitioners, managers and
decision-makers.
Figure 2: Robustness
of European vertebrate networks to disturbance scenarios per
ecoregion. Extreme scenarios of climate change and
of species extinctions can be used to explore (lower)
boundaries of network robustness and identify areas where we
may expect a high number of cascading (secondary)
extinctions and, consequently, larger disruptions to
ecosystem functioning and services. Lower limit was set to
0.80 for illustration purposes.
Concluding remarks
Ecological networks already can and should be used as
indicators in biodiversity conservation and ecosystem
management. Sufficient data is available for initial
assessments of network structure and responses to change.
Additionally, we have relevant network indicators for
ecosystem management and conservation that can be weaved
into management frameworks and monitoring programs. Starting
now ensures that future data will be useful to detect
network changes and to address current knowledge gaps.
We recognize that the lack of empirical support for
theory and scenarios of network responses (including
robustness) to environmental change can refrain academics
from providing guidance to practitioners. Robustness and
extinction studies usually rely on simulations to
investigate effects of species loss (rather than
observations or experimental removals) and predictions
remain mostly untested in the field [83,
see Table 1 therein for some empirical validation
examples]. Overcoming this barrier will require
setting up programs that go beyond documenting networks and
towards empirical measurements of network responses to
realistic disturbances. Moreover, empirical and monitoring
programs will need to collect and integrate network
information across multiple scales, as management actions
and policy-making differ between regional and local levels.
Yet, despite this and other limitations (i.e., data,
uncertainty, and interpretability challenges highlighted
above), we believe the field is sufficiently mature to make
recommendations for ecosystem management and conservation as
these programs are implemented.
We envision five important aspects for future directions
(see also Outstanding
Questions). First, developments addressing evaluation,
propagation, and communication of uncertainty in network
structure and metrics are needed. These will be key to a)
integrate uncertainty into management frameworks and move
towards more transparent and informed decisions, but also to
b) use existing tools and data to compare known network and
ecosystem changes with predictions (e.g. hindcasting),
estimate boundaries of future network changes (e.g.
forecasting), and assess the usefulness of network metrics
as indicators of future change. Second, network
considerations will need to be explicit in future sampling
and monitoring designs, and in ecosystem conservation
regulations and decisions. Third, current data, network
models and indicators need to be more widely assessed for
their usefulness for ecosystem management, which should
actively involve stakeholders. Fourth, empirical programs
focused on testing and measuring network (metrics’)
responses to change, and across scales, will need to be set
up. Finally, incorporating network information explicitly
into conservation will require developing network-based
targets—specific, quantified metric values to aim for or
avoid (thresholds) based on whole network
characteristics.
Outstanding questions
How variable is network structure across space and time
and does it influence the usefulness of network metrics as
indicators of ecosystem functioning and stability?
What network metrics are ubiquitous, reliable and
applicable indicators of ecosystem functioning and
stability?
How much can we expect networks to change given
uncertainty in future environmental conditions?
How can current and future monitoring programs be
improved to sample network information relevant for
management?
How can we put in place a strong empirical program to
validate network indicators, which for now heavily rely on
simulations?
How should we implement coordinated monitoring of
network indicators across multiple scales? Can the same
indicators be used to inform at broad, regional and local
scales?
Glossary
Basal species: species that do not feed
on other species in a trophic network; e.g. primary
producers.
Binary and probabilistic networks:
networks where links represent either the presence or
absence of an interaction between species, or its
probability.
Forecasting: using current (known)
conditions to predict future conditions of a system or
events.
Hindcasting: using current (known)
conditions to predict past conditions of a system or
events.
Metaweb: all potential interactions in a
region or species pool. Metawebs can be either binary or
probabilistic, and are mostly common for trophic,
mutualistic and parasitism networks. Due to their potential
nature, they provide an “upper ceiling” of species
interactions.
Monitoring programs: established
long-term programs to track biodiversity status and changes.
Data collected in situ through sampling or using
remote sensing can be used in the calculation of
biodiversity indicators and support decision-making.
Network indicators: network metrics with
a clear interpretation and potential use for biodiversity
conservation and management. This includes meeting criteria
important for decision-making (e.g. ROARS, SMART). Here, we
use trophic network robustness as an example of a useful
indicator.
Network metrics: measurements made on
networks, their nodes and links, regarding their
composition, structure or properties pertaining to node or
link importance. Common examples include number of links
(interactions) and nodes (species), connectance, nestedness,
trophic level, centrality, omnivory and network motifs.
Primary extinctions: extinctions
directly due to disturbances. In our scenarios disturbances
were changes in species climate suitability or the failure
to protect endangered species.
Projection: a model prediction based on
novel data (data beyond the fitting dataset) or scenarios,
not necessarily tied to future or past conditions. For
instance, a network prediction in a new location or with a
different set of species.
Rewiring: changes in the interaction
structure of a network. For instance, disturbances,
environmental change, and addition or loss of species can
lead to gain, loss, and reorganization of interactions.
Robustness: capacity of a network (or
the ecosystem it represents) to withstand species
extinctions following a disturbance. Robustness can be
measured in multiple ways. Here we measure robustness as 1
minus the ratio of secondary extinctions to the initial
number of secondary consumers, following concepts of
robustness by Dunne et al.[70].
Secondary consumers: species that
consume other species in the network (calculated as network
species richness minus the number of basal species).
Secondary extinctions: extinctions due
to the loss of prey species.
Species interaction networks: networks
assessing the ecological links and relationships between
species, highlighting how they are interconnected and
influence each other. Links can be trophic (representing
feeding links), flow-based (representing transfers of
energy, matter, or resources), and mutualistic
(e.g. pollination), among others.
Acknowledgements
G.D. is funded by the NSERC Postgraduate Scholarship –
Doctoral (grant ES D – 558643), the FRQNT doctoral
scholarship (grant no. 301750), and the NSERC CREATE BIOS2
program. T.P. is funded by the Wellcome Trust
(223764/Z/21/Z), NSERC through the Discovery Grant and
Discovery Accelerator Supplements programs, and the Courtois
Foundation. WT, LOC and LM acknowledge support from the
European Union’s Horizon Europe under grant agreement number
101060429 (project NaturaConnect). JMM acknowledges the
support of Horizon Europe project BIOcean5D (award number
101059915) and the French Agence Nationale de la Recherche
through LabEx TULIP (ANR-10-LABX-41). GFF and WT acknowledge
the support of Biodiversa+, the European Biodiversity
Partnership, co-funded by the European Commission (grant
agreement no. 101052342 ‘PrioritIce’).
Executive Secretary of the
Convention on Biological Diversity (2022) Expert input
to the Post-2020 Global Biodiversity Framework:
Transformative actions on all drivers of biodiversity loss
are urgently required to achieve the global goals by
2050
CBD (2022) Decision adopted by
the conference of the parties to the convention on
biological diversity 15/4. Kunming-montreal global
biodiversity framework
O’Connor, L. (2022) Accounting for
food webs to conserve biodiversity and nature’s
contributions to people in the context of global changes.
PhD thesis, Université Grenoble Alpes