Guidelines Connect Conserv Plann EU
Guidelines Connect Conserv Plann EU
DOI: https://doi.org/10.3897/arphapreprints.e129021
00.00.2023
Author-formatted document posted on 06/06/2024. DOI: https://doi.org/10.3897/arphapreprints.e129021
D6.1 Guidelines for connectivity conservation and planning in Europe with supporting web-based inventory and
databases
29.03.2024
NaturaConnect receives funding under the European Union’s Horizon Europe research and
innovation programme under grant agreement number 101060429.
naturaconnect.eu
Scientific coordinator: Piero Visconti, PhD, IIASA
Type: HORIZON Innovation Actions
Call: HORIZON-CL6-2021-BIODIV-01
The contents of this material are the sole responsibility of the NaturaConnect consortium
and do not necessarily reflect the opinion of the European Union. This report reflects the
version finalised and submitted to the European Commission on 29.03.2024. Further
changes to the report may be integrated following review from the European Commission.
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D6.1 Guidelines for connectivity conservation and planning in Europe with supporting web-based inventory and
databases
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Author-formatted document posted on 06/06/2024. DOI: https://doi.org/10.3897/arphapreprints.e129021
D6.1 Guidelines for connectivity conservation and planning in Europe with supporting web-based inventory and
databases
29.03.2024
Deliverable description This guidelines will be the product of tasks 6.1 and 6.2
In 6.1 we will co-design a framework for multi-scale
connectivity assessment and planning together with
stakeholders from MS administrations, EU policy bodies and
other relevant EU programs and platforms: LIFE projects,
EuropaBON, KCBD, EEA, etc.
This framework will be published in the connectivity guidelines
together with the output of task 6.2 which include a synthesis
report of the input data, statistical models and predictions of
species dispersal kernels using agent-based models.
Keywords Biodiversity policy, connectivity assessment, conservation
planning, ecological corridors, landscape fragmentation,
population models, species dispersal, species traits, user
needs.
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D6.1 Guidelines for connectivity conservation and planning in Europe with supporting web-based inventory and
databases
29.03.2024
Table of Contents
List of Figures .................................................................................................. 8
List of Tables ................................................................................................. 10
Abbreviations ................................................................................................ 11
Executive summary ....................................................................................... 12
1. Introduction................................................................................................ 17
1.1 Why is ecological connectivity important? ........................................................................ 17
1.2 Aims and target audience ..................................................................................................... 18
1.3 Summary of the content ........................................................................................................ 19
D6.1 Guidelines for connectivity conservation and planning in Europe with supporting web-based inventory and
databases
29.03.2024
D6.1 Guidelines for connectivity conservation and planning in Europe with supporting web-based inventory and
databases
29.03.2024
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Author-formatted document posted on 06/06/2024. DOI: https://doi.org/10.3897/arphapreprints.e129021
D6.1 Guidelines for connectivity conservation and planning in Europe with supporting web-based inventory and
databases
29.03.2024
List of Figures
Figure ES1: Response rates for taxonomic groups and ecosystem types for the survey on ecological
connectivity projects in Europe ............................................................................................................. 14
Figure ES2: Building blocks of the proposed framework for implementing ecological connectivity
projects .................................................................................................................................................. 16
Figure 2.1: Distribution of Natura 2000 sites and nationally designated protected areas in the European
Union, covering approx. 26% of the land surface.. ............................................................................... 22
Figure 4.1: Number of projects per country and reported project duration. .......................................... 45
Figure 4.2: Response frequencies for stated connectivity goals, thematic scope and other benefits that
a project may bring ................................................................................................................................ 46
Figure 4.3: Response rates for taxonomic groups and ecosystem types. ............................................ 47
Figure 4.4: Response frequencies for the questions concerning policy context, target users and funding
sources. ................................................................................................................................................. 48
Figure 4.5: Response frequencies for questions related to projects’ spatial scope and the
biogeographical regions where they took place. ................................................................................... 49
Figure 4.6: Response frequencies for questions concerning selected approaches and what kind of
spatially explicit information projects produced..................................................................................... 49
Figure 4.7: Number of projects that did and did not implement monitoring. ......................................... 50
Figure 4.8: Response frequencies for potential negative effects caused by increasing ecological
connectivity. .......................................................................................................................................... 51
Figure 4.9: Type of organisation and distribution (% of total) by country of the participants in the online
workshop “Assessing Ecological Connectivity in Europe: Conservation goals and information gaps”. 52
Figure 4.10: Miro board produced in the breakout group on “Enhancing Connectivity for Endangered
Species and Habitats” from the workshop “Assessing Ecological Connectivity in Europe: Conservation
goals and information gaps”.. ................................................................................................................ 54
Figure 4.11: Word response frequency retrieved from the Miro boards for Day Two of the “Assessing
Ecological Connectivity in Europe” workshop.. ..................................................................................... 57
Figure 5.1: This simple illustration (left) shows nodes in the white pixels and the movement of electrical
current through each “resistor” (i.e., pixel). ........................................................................................... 71
Figure 5.2: Illustration of the moving window (left) that moves over the source weight and resistance
surfaces ................................................................................................................................................. 72
Figure 5.3: Map of Europe showing the result of using the moving window approach in conjunction with
the Effective mesh size structural connectivity metric .......................................................................... 80
Figure 6.1: A schematic representation of a framework for connectivity network design. .................... 83
Figure 6.2: Habitat preferences (percentage of species) of European tetrapods using three coarse
classes. ................................................................................................................................................. 90
Figure 6.3: Checklist of common spatial data needs for connectivity analyses. ................................... 96
Figure S2.1: Miro board from the “Terrestrial and Freshwater Habitats” breakout group on day 1 of the
workshop. ............................................................................................................................................ 130
Figure S2.2: Miro board from the “Ecosystem Processes & Services” breakout group on day 1 of the
workshop. ............................................................................................................................................ 132
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D6.1 Guidelines for connectivity conservation and planning in Europe with supporting web-based inventory and
databases
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Figure S2.3: Miro board from the “Planning & Management of Multifunctional Corridors” breakout group
1 on day 2 of the workshop. ................................................................................................................ 134
Figure S2.4: Miro board from the “Human Infrastructure & Land Use Impacts” breakout group on day 2
of the workshop. .................................................................................................................................. 136
Figure S3.1: Aggregation of IUCN habitat classes (level 1) into natural and artificial classes... ........ 144
Figure S3.2: Habitat preferences of European tetrapods. .................................................................. 145
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D6.1 Guidelines for connectivity conservation and planning in Europe with supporting web-based inventory and
databases
29.03.2024
List of Tables
Table 4.1: Breakout group themes for the two days of the “Assessing Ecological Connectivity in Europe:
Conservation goals and information gaps” workshop.. ......................................................................... 53
Table 5.1: Outline of the most common modelling families for functional and structural connectivity. . 62
Table 5.2: Software and programming packages that can implement least-cost path and resistant kernel
analyses. ............................................................................................................................................... 67
Table 5.3: Circuit theory applications developed detailing needed data inputs and sources. .............. 74
Table 5.4: Examples of software packages commonly used to implement agent-based models. ....... 75
Table 5.5: Examples of various structural connectivity metrics, including references, all varying in
complexity, with explanation of what the respective metrics measures.. ............................................. 77
Table 6.1: Four potential connectivity problems with an example objective, target and action ............ 86
Table 6.2: Minimum and recommended corridor widths based on the review of 66 scientific studies
(Bentrup, 2008). .................................................................................................................................... 92
Table S3.1: Trait databases considered for the archetypes analysis. ................................................ 138
Table S3.2: Median values of traits used to define the five European non-volant mammals archetypes.
............................................................................................................................................................ 140
Table S3.3: Median values of traits used to define the three European bats archetypes. ................. 140
Table S3.4: Median values of traits used to define the four European birds archetypes. .................. 141
Table S3.5: Median values of traits used to define the three European frogs archetypes. ................ 141
Table S3.6: Median values of traits used to define the three European salamanders archetypes. ... 142
Table S3.7: Median values of traits used to define the two European turtles archetypes. ................. 142
Table S3.8: Median values of traits used to define the three European snakes archetypes. ............. 143
Table S3.9: Median values of traits used to define the three European lizards archetypes. .............. 143
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D6.1 Guidelines for connectivity conservation and planning in Europe with supporting web-based inventory and
databases
29.03.2024
Abbreviations
ABM Agent-Based Model
ASCII American Standard Code for Information Interchange
BGI Blue and Green Infrastructure
CMS Convention on Migratory Species
CSV Comma-separated values
EC European Commission
ES Ecosystem Services
GBF Global Biodiversity Framework
GI Green Infrastructure
GIS Geographic Information System
GUI Graphical User Interface
IUCN International Union for Conservation of Nature
LULC Land Use / Land Cover
MAES Mapping and Assessment of Ecosystems and their Services
OECM Other Effective area-based Conservation Measure
PCA Principal Component Analysis
PIT Passive Integrated Transponder
RBMP River Basin Management Plan
WFD Water Framework Directive
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D6.1 Guidelines for connectivity conservation and planning in Europe with supporting web-based inventory and
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29.03.2024
Executive summary
Ecological connectivity is key to maintaining a coherent and resilient network of protected
areas in the EU. The EU Biodiversity Strategy for 2030 has identified the unhindered
movement of species, nutrients and ecological processes across connected landscapes as a
key feature of a coherent Trans-European Nature Network (TEN-N) of protected and
conserved areas. However, to date, streamlined guidance on planning for and implementing
connectivity measures specifically at the European scale has been limited.
This report presents a coherent methodological framework and guidelines for mapping
functional and structural connectivity at the European scale, as part of the Horizon Europe
NaturaConnect project, which is supporting EU Member States in developing a coherent TEN-
N of protected and conserved areas.
It describes key ecological connectivity concepts and approaches; outlines methods and tools
for estimating connectivity; presents an overview of connectivity projects across Europe;
identifies connectivity priorities, gaps and challenges following a stakeholder consultation
process; and provides practical and operational guidelines for implementing ecological
connectivity for conservation projects ranging from regional to national and European levels.
The guidelines present a strategic blueprint aimed at enhancing ecological connectivity across
Europe, and address the specific challenges and opportunities related to planning ecological
connectivity in the European context.
This report has been written for practitioners and individuals involved in the management and
administration of protected areas and ecological connectivity projects across Europe. This
includes professionals working in TEN-N implementation at national or regional levels, others
involved in spatial planning outside protected areas, and professionals engaged in the
implementation of connectivity projects and protected area management.
The primary focus is on the terrestrial realm, although challenges in freshwater connectivity
are also addressed.
Key insights and results presented in the report include:
• Connectivity is an integral component of protected area planning in Europe. As
highlighted by the EU Biodiversity Strategy for 2030, a coherent Trans-European
Nature Network (TEN-N) depends on the setting up of ecological corridors “to prevent
genetic isolation, allow for species migration, and maintain and enhance healthy
ecosystems”.
• When planning for and implementing connectivity, understanding and
distinguishing between different connectivity concepts and approaches is vital.
For example, concepts such as the role of protected areas versus ecological corridors,
structural and functional connectivity, Green and Blue Infrastructure, spatial scale and
dispersal issues, design of corridors and stepping stones, integration of connectivity in
spatial conservation prioritization, and freshwater and cross-realm connectivity (all
defined in Chapter 2).
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• Connectivity goals are being featured prominently in several recent global and
EU policy instruments, including the post-2020 Global Biodiversity Framework, the
EU Biodiversity Strategy for 2030, the EU Forest Strategy for 2030, the Green and
Blue Infrastructure (GBI) strategy, the Nature Restoration Regulation, the Water
Framework Directive, and the EU pollinators initiative (Chapter 3).
• To date, comprehensive assessments providing a pan-European overview of
connectivity projects taking place across Europe, as well as connectivity
implementation gaps and needs, have been limited. To address this need, as part
of the NaturaConnect project, an online survey and follow up webinar were carried out
with stakeholders in the conservation community to gather this key information
(Chapter 4). Key findings from the online survey which received submissions on 80
projects across 35 European countries include:
o The most common connectivity goals of projects are connectivity between
protected areas or between specific habitat types
o Additional benefits of connectivity projects included recreation, climate
regulation and pollination services
o The most targeted taxa in projects were large carnivores, followed by
arthropods and birds
o The most targeted ecosystems were forests and grasslands (Fig. ES1)
o The main target users of project results were regional or local administrations
o The main funding sources were nature conservation funds from national and
regional administrations, and private funds
o The spatial scope of most projects was subnational
o The most common targeted biogeographical region was Continental, followed
by Alpine
o Selected approaches for estimating connectivity were mainly land cover and
expert-based
o Most projects provided spatial information on locations for ecological corridors,
stepping stones and locations for habitat restoration
o In most cases (over 70%) there was no monitoring of project effectiveness, and
the potential negative impacts of increased connectivity were often not
considered (though mentioned ones included human-wildlife conflicts and
increased spread of invasive species)
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D6.1 Guidelines for connectivity conservation and planning in Europe with supporting web-based inventory and
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Figure ES1: Response rates for taxonomic groups and ecosystem types for the survey on ecological connectivity
projects in Europe
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D6.1 Guidelines for connectivity conservation and planning in Europe with supporting web-based inventory and
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D6.1 Guidelines for connectivity conservation and planning in Europe with supporting web-based inventory and
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Figure ES2: Building blocks of the proposed framework for implementing ecological connectivity projects
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D6.1 Guidelines for connectivity conservation and planning in Europe with supporting web-based inventory and
databases
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1. Introduction
1.1 Why is ecological connectivity important?
Earth's ecosystems have undergone substantial degradation and significant loss of ecosystem
processes and functions caused by human activities. Approximately 75% of the land surface
has been significantly modified, more than 85% of wetland areas have been lost, and an
average of 25% of globally assessed animal and plant species are threatened with extinction
due to land use changes and unsustainable logging, harvesting, hunting, and fishing (IPBES,
2022). Over the next few decades, climate change is expected to play an increasingly
significant role as a direct driver of biodiversity loss (IPBES, 2022).
As climate change and direct human pressures increase in severity, ecosystems with high
biodiversity value become smaller and increasingly isolated (Fahrig, 2019, 2003). Smaller and
isolated species habitats harbour smaller populations with lower genetic diversity and higher
threats of extinction (Frankham et al., 2010; Schlaepfer et al., 2018). These habitats are also
vulnerable to edge effects (i.e., pressures on species populations at their habitat boundaries)
(e.g., Weathers et al., 2001) and to the simplification of species community composition and
interactions (e.g., Razafindratsima et al., 2018; Valladares et al., 2006). Synergistic effects of
habitat loss and isolation interfere with critical ecological processes sustaining the ecosystem
integrity, such as pollination, seed dispersal and the nutrients flow, which, in turn, has
cascading effects on both ecosystem structure and functions (Haddad et al., 2015; Laurance
and Bierregaard, 1997).
© Olla Jennersten/WWF-Sweden
Connectivity conservation and restoration are key to counteract the detrimental effects of
ecosystem degradation, habitat loss, and fragmentation. Ecological connectivity
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encompasses the functional and physical connections between different habitats and
ecosystems that enable the movement of species, nutrients, and ecological processes
across landscapes (Crooks et al., 2011; Crooks and Sanjayan, 2006; Hilty et al., 2020).
Ecological connectivity plays a pivotal role in preserving biodiversity, ensuring the long-term
persistence and adaptability of species populations and communities (van Rees et al., 2021).
Individuals are often compelled to move and disperse due to various ecological drivers,
including reproduction, access to food sources, seasonal habitat and climate change,
intraspecific competition, evasion from predators and competitors, and temporal or permanent
habitat degradation and destruction. The seamless movement facilitated by ecological
connectivity is fundamental for their survival and reproductive success (Crooks and Sanjayan,
2006). When populations are interconnected, the flow of individuals and genes are fostered,
enhancing adaptability and resilience to environmental change and stochastic events.
Moreover, as the climate changes, species may need to track spatial changes in the habitat
quality. Therefore, connected landscapes are key to species adaptability since it can facilitate
distribution shifts in response to climate and land-use changes (Heller and Zavaleta, 2009;
Opdam and Wascher, 2004).
Ecological connectivity has an even broader impact on ecosystem-level processes. Critical
ecological phenomena, such as nutrient cycling, pollination, and predator-prey interactions,
hinge on the unhindered movement of species (Crooks et al., 2011; Razafindratsima et al.,
2018). Furthermore, the interlinking of habitats provides a safety net against antropogenic
disturbances. For example, when a particular habitat patch undergoes a catastrophic event
like a fire or a disease outbreak, species can seek refuge in neighbouring, connected habitats.
This not only allows for immediate survival but also facilitates recolonization processes and
passive ecological restoration.
Recent international agreements have placed connectivity at the core of the pathway
towards nature recovery. Goal A of the United Nations Global Biodiversity Framework aims
to maintain, enhance and restore the integrity, connectivity and resilience of all ecosystems
by 2050. Associated targets include reducing threats to biodiversity through enhancing
connectivity with ecosystem restoration (Target 2), designing well-connected protected areas
(Target 3) and increasing the connectivity of green and blue spaces in urban areas,
contributing to the provision of ecosystem services (Target 12). The European Union
Biodiversity Strategy for 2030 aims to enhance connectivity among habitats, protected areas
and green and blue infrastructure, for instance, with the designation of additional protected
areas and the creation of high diversity landscape features and ecological corridors.
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D6.1 Guidelines for connectivity conservation and planning in Europe with supporting web-based inventory and
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Part I: Connectivity in
Europe: Key concepts,
policy context, and
implementation
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Figure 2.1: Distribution of Natura 2000 sites and nationally designated protected areas in the European Union,
covering approx. 26% of the land surface. Data source: European Environmental Agency, 2023.
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©OlaJenner/WWF Sweden
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D6.1 Guidelines for connectivity conservation and planning in Europe with supporting web-based inventory and
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therefore depends on the system under consideration and on the goal of the conservation
strategy.
At the local scale (Noss, 1991), several
studies have demonstrated the value of
field margins and other small, linear
patches of natural habitats in agricultural
landscapes. Fencerows may act as local
corridors between woodlots, allowing for
metapopulation persistence in
micromammals (Fahrig and Merriam,
1985), antipredator strategies in birds
(Ausprey et al., 2023), and movement in
reptiles and amphibians (Noss, 1991). A
hedgerow approach focusing on local
connectivity in a human-dominated
landscape (e.g., agriculture
conservation) can be extremely
important for animals and plants with
limited dispersal capabilities (most
herptiles, many small mammals, non- © Călin Ardelean/WWF România
flying invertebrates, etc.). A clear
disadvantage of hedgerows (and of linear corridors in general) is that they are often narrow
strips of habitat and therefore not all species can use them (e.g., forest interior species will
not, and edge effects will increase exposure to threats).
At the scale of a landscape, we often deal with landscape mosaics, including habitat patches
and corridors. At this scale, any wide-ranging mammals (e.g., a bear) require corridors to
move from one habitat patch to the other to meet their daily needs for food, water, and shelter,
often spanning tens of kilometres in a single day. At the same scale, but on a different time
frame, ungulates use landscape connectivity for seasonal movements, while amphibians
migrate between wintering grounds and breeding ponds.
At the regional to continental scale, in many parts of the world, there is an ambitious strategy
to connect nature reserves into regional networks (e.g., the Natura 2000 network in Europe or
the Yukon to Yellowstone National Parks in North America). This strategy is particularly
important when we enlarge our temporal view up to centuries and millennia. In the past, huge
biogeographical corridors have been critical in permitting the shift of floras and faunas in
response to global changes (e.g., the Bering land bridge, and the isthmus of Panama). Future
climate changes, probably occurring over the next few decades, also require a large-scale
vision. In this framework, human-related habitat fragmentation has greatly increased the
number of barriers for most native species and at a large scale, therefore regional networks
of nature reserves will play a fundamental role. A Europe-wide approach (focusing on
transboundary conservation/management) is potentially vital for species having large-scale
requirements and long dispersal distances, for example for the Italian/French wolf population
(Ciucci et al., 2009), for most large carnivores in western Europe (Chapron and Arlettaz, 2006),
or anadromous sturgeons (Friedrichs et al., 2018). Focusing particularly on the large scale of
analyses, connectivity is central to many global change analyses, in which the response of
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single species or entire communities is related to ongoing and future climate changes (Tesson
and Edelaar, 2013). For example, many Mediterranean species will relocate their distributions
to temperate or boreal regions (Maiorano et al., 2011).
The scale of analyses must be linked to the ecological traits of the species being
considered, focusing particularly on dispersal capacity, and its commonly used proxies such
as body size and home range size. Dispersal, or better natal dispersal, can be defined as the
movement of an organism (animal or plant) from its natal place to the place where it will
reproduce. In some cases, natal dispersal information is not necessarily useful for connectivity
projects. Take long distance migratory birds as an example, where the distance between natal
and breeding locations are in reality separated by thousands of kilometres spent in autumn
and spring migration. In this case, using natal dispersal as an indicator of dispersal capacity
is not useful.
The effects of dispersal can be seen at all spatial scales, from intraspecific genetic diversity
(Suárez et al., 2022) to species geographic ranges (Gaston, 2003). Furthermore, there is a
growing understanding of its importance in a global change context (Anderson et al., 2012).
Dispersal plays a central role in the response of populations and species to global changes,
including climate change, habitat loss and fragmentation, as well as invasive species (Tesson
and Edelaar, 2013). Distance between patches of suitable habitat, or distance among
protected areas must consider the dispersal capability of the species. In general, spatial
requirements may help define stepping stones both in their size and in their distance (e.g.,
Parks et al., 2023).
However, data availability on dispersal is limited in many taxonomic groups (Nathan,
2001). The reasons for this gap of knowledge go from inconsistencies in both the
measurement and the definition of dispersal to difficulties in collecting field data, and the
existence of unpredictable long-distance dispersal events (Bowman et al., 2002).
Furthermore, a dispersal event can take many forms, going from a gradual shift (e.g., typical
of philopatric mammals like brown bears) to a one-way movement over great distance (Sarkar
et al., 2021).
Often dispersal distances have been estimated using correlative models with ecological and
physiological traits as covariates. Multiple traits have been demonstrated as important
correlates for dispersal. For example, gestation length and maximum life span have been
identified as important to explaining the distance moved by species ranges during the North
American glacial/interglacial cycles (Lyons et al., 2010). A suite of demographic traits (e.g.
fecundity) has been correlated with dispersal abilities in butterflies (Stevens et al., 2012).
Although no single model outperformed all others in the literature (Whitmee and Orme, 2013),
body mass and home range consistently emerged as important predictors of dispersal ability
(see also Section 6.3).
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protected areas that act as refugia for species to maintain genetic connections and move
between the protected areas. Quite often a complex connectivity design will contain a
combination of corridors and stepping stones to connect dozens of protected areas
across a broad region, and including across borders (e.g., multi-country).
Corridors are more heavily relied upon as they are
intended to maintain an unbroken connection
through a landscape that may already be heavily
modified by humans. Examples of large
landscape corridors include the connections of
undeveloped habitat for mountain lions (Puma
concolor) in heavily urbanised southern
California, unbroken protected forests for the
movement of tigers (Panthera tigris) between
reserves in India, and the Mount Kenya Elephant
Corridor for the migration of African savanna
elephants (Loxodonta africana), or the initiative to
set cross-border connectivity corridors in the
Carpathians and Danube basin, in Europe.
In the European context, intensive land
management and the mixture of habitat types
in and around protected areas bring added
complexity to the placement and
configuration of corridors. These complexities
have implications that need to be considered
when designing corridors, e.g.: (i) protected
areas are often a mixture of habitats, raising the ©Jeremy Dertien
question of which habitat types should be
included in the corridor; (ii) should we design different corridors among protected areas for
each target group?; (iii) if two nearby Natura 2000 areas do not share any habitat type (e.g.
one is a forest and the other is a wetland), is a corridor justified?; (iv) what are the socio-
economic challenges of corridor management in these humanized landscapes?
Appropriate corridor width is important to ensure that species of concern will
successfully move through the corridor and reduce the chances that wildlife will come into
conflict with humans. If a corridor is too narrow, then the edge effects from human presence
likely affect successful movement and the habitat may be too degraded for species movement.
For example, forest birds are impacted 50-70 metres from human recreation and
development, thus a corridor that is less than 150 metres wide contains very little core habitat
that is not in some way influenced by human pressure. Large mammals are impacted at much
greaterdistance thresholds, thus corridors that are greater than 1 km in width can reduce the
impacts of humans while reducing the chances of human-wildlife interactions (Dertien et al.,
2021).
Stepping stones facilitate the movement of species and help maintaining connection between
two or more larger habitat areas without direct structural connection. Just like a chain of small
islands between two large land masses, a stepping stone design can be seen as islands of
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habitat surrounded by an ocean of unprotected land. Stepping stone designs can be more
feasible to implement in more highly populated areas where land protection for a fully
connected corridor may be much more difficult to implement (Lynch 2019). Also, the protection
of stepping stones can be the first step in the planning for the restoration of permanent
corridors between stepping stones. While stepping stones are often considered secondary
substitutes for corridors, studies have shown the importance that even small habitat patches
can have on maintaining ecological connectivity for some species (Herrera et al., 2017). Birds,
plants and aerial insects are known to benefit the most from stepping stones given their
dispersal abilities, and in fact can be key to a species range expansion (for example Saura et
al., 2014). For example, wetlands are important stepping stones along migratory flyways for
aquatic birds (Merken et al., 2015).
To maximise connectivity through ecological stepping stones, several key principles
should be considered in their design (Box 2.1):
Box 2.1 Key principles to consider when designing ecological stepping stones
In urban settings, the difficulties of land acquisition and city design may require a
stepping-stone approach (Lynch, 2019), though there is evidence that corridors rather than
stepping stones are more effective at supporting urban biodiversity (Beninde et al., 2015). In
the rural or peri-urban (exurban) landscape, stepping stones may be forest patches, riparian
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zones, or highly heterogeneous agricultural areas. Moving into the urban matrix these
stepping stones can become even more human-dominated such as parks, brownfields,
greenways and even rooftop gardens. As the urban matrix becomes more developed and
difficult to traverse, stepping stones must be closer together to maintain linkages. However, if
the urban matrix becomes entirely too hostile to the movement of wild species (e.g., urban
centres), stepping stones are not effective and a continuous protected corridor is the only
option to maintain ecological connectivity. Difficulties may also persist with planning and
managing stepping stones since many are available opportunistically (rather than pre-
planned) and may be owned privately rather than publicly.
Ultimately, the use of corridors or stepping stones as features for maintaining connectivity is
reliant on the geographic situation, the species or ecological processes that one is trying to
conserve, and the political will for the creation of a connectivity network.
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adjusted based on the ecological needs of different species. Hermoso et al. (2011) modified
the hierarchically based approach by measuring longitudinal connectivity between each sub-
catchment and each upstream sub-catchment as the inverse of the distance (measured
through the river) between them.
Secondly, the lateral dimension plays a key role too in maintaining the flow of matter and
energy and the daily routine of semi-aquatic species by connecting freshwater ecosystems
with nearby floodplains and wetlands. This connection can be established through river floods
during the wet season, or by dispersal of semi-aquatic organisms, such as some insects,
turtles, birds and mammals. The lateral dimension and connectivity to riparian areas play a
vital role in connecting aquatic and terrestrial habitats. Hermoso et al. (2012a) and Reis et al.
(2019) proposed a set of new inter-sub-catchment metrics to account for connectivity between
the river network and adjacent wetlands not connected by the river network. The aim was to
account for the ecological requirements of species that move across drainage divides, such
as waterbirds.
Thirdly, the vertical dimension includes interactions between the surface and groundwater,
comprising gradients in habitat stratification, temperature, light and oxygen levels, which
together condition the vertical migrations of species. This dimension of connectivity has
received less attention than the others. Notably, Nel et al. (2011) measured vertical
connectivity by developing a predictive model to map the probability of groundwater interacting
with surface water. This method was used to identify areas most critical to maintaining
seasonal refuge pools.
The fourth dimension is time, which affects freshwater ecosystems through changes in river
flow (conditioning drought and floods), climatic conditions and life cycle dynamics. Freshwater
connectivity is limited by water flow, which seasonally fluctuates in permanent rivers and is
even not permanent in a high proportion of rivers (known as temporary or intermittent rivers).
Freshwater habitats in these systems may become restricted to a reduced and disconnected
set of pools, which become ecological refugia, vital to recolonization after the dry period.
These types of rivers are more likely to occur in climatic regions with pronounced wet-to-dry
seasonality, such as the Mediterranean, which is predicted to become increasingly subjected
to droughts (Estrela-Segrelles et al., 2023; Naumann et al., 2018), and in arid regions where
in most of the year the water is restricted to water pools. In addition, water availability can also
fluctuate inter-annually, given that extreme drought events are predicted with global climate
warming (Naumann et al., 2018). Examples to address temporal connectivity include Hermoso
et al., 2012a), who used water residency time as a proxy.
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© Ante Gugić/WWF
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facilitating movement or migrations of a particular species (Mazor et al., 2016), account for
directional connectivity within and across freshwater and marine ecosystems (Beger et al.,
2010b; Bode et al., 2008; Hermoso et al., 2021a), optimise gene flow (Hanson et al., 2019),
deal with horizontally and vertically connectivity in a three-dimensional space (Venegas-Li et
al., 2018) and when accounting for species’ migratory needs derived from ongoing and future
climate change (Sonntag and Fourcade, 2022).
In general, available algorithms are restricted to a single connectivity constraint, making it
difficult to address multiple connectivity objectives. An alternative is to use a composite index
of connectivity, which can combine multiple connectivity metrics (Magris et al., 2014) and
provide a quantitative estimate of how each planning unit contributes to maintaining or
enhancing connectivity. In this case, connectivity can be treated as a conservation feature in
the optimization algorithm, rather than as a constraint (D’Aloia et al., 2017). While this
approach can be computationally practical, it can have the caveat that through aggregation it
becomes impossible to differentiate which connectivity aspect is driving the solution.
Additionally, another challenge in planning for connectivity in a network of protected
areas is the assumption that all connections between protected areas (and the areas
themselves) will persist through time once they are established, but it is likely that land
use, habitats and species ranges will shift and change under climate change, which can
disrupt the functionality of the network (Nuñez et al., 2013).
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However, scientific evidence does not support widespread negative corridor effects. An
analysis of thirty-three corridor studies found no evidence that corridors increase species
invasions or disturbances; edge effects can have either positive or negative impact on species
abundances; and effects on antagonistic species effects or population synchrony were mixed.
Whether or not the potential negative effects counteract the benefit of increasing connectivity
strongly depends on the local ecological context and therefore no generalization should be
made. Therefore, connectivity planning should acknowledge, identify, anticipate and monitor
the potential these potential unwanted effects.
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Five major factors have been proposed as potential negative ecological consequences of
corridors (Haddad et al. 2014):
1. Edge effects particularly affecting of long and narrow corridors
2. Colonization or population reinforcement of species that are antagonistic to
conservation targets, such as pathogen hosts or competitor species
3. Proliferation and increased abundance of invasive species
4. Propagation of disturbances like fires
5. Synchronisation of population dynamics between connected habitats
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of the habitats listed in Annex I to the Habitats Directive. These provisions require that nature
restoration measures contribute with ecological corridors and other measures enhancing
connectivity, with the aim of improving the habitat quality and conservation status of species
and the ecological condition of habitats. In freshwater ecosystems, restoration measures
should include the removal of artificial barriers in rivers, lakes and alluvial habitats to achieve
significant increases in the longitudinal, vertical and lateral connectivity (such as restoring the
natural functions of floodplains). Specific emphasis is also placed in restoring the connectivity
of forest ecosystems. Other obligations will similarly require increasing connectivity to restore
pollinator populations, such as implementing “buzz lines” where insect pollinators could move
across landscapes, and harshening high-diversity landscape features with great potential to
increase connectivity for species and habitats across agricultural ecosystems.
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comply with the following criteria: (i) contribute to the conservation and/or enhancement of
multiple ecosystem services at a significant scale, (ii) contribute to the goals of Nature
Directives, (iii) have a strategic approach with EU-level impact, with at least a national or
regional level approach. An overview of scientific and technical tools for GI mapping, including
the European Mapping and Assessment of Ecosystems and their Services (MAES) initiative;
and geospatial methods, data and tools (e.g., CORINE, LUCAS, Copernicus), are detailed in
a technical report (Estreguil et al., 2019).
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Figure 4.1: Number of projects per country and reported project duration (inset, top left).
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Regarding the thematic scope of the projects, which refers to what they aim to enhance,
"Ecological corridors (continuous corridors or stepping stones)" and "Ecosystem restoration"
accounted for 54% of the responses.
Respondents were also asked to identify other benefits their projects may deliver, in addition
to promoting biodiversity conservation. The three most frequently mentioned benefits were
"None", "Recreation", and "Climate regulation", which comprised 52% of all replies.
Figure 4.2: Response frequencies for stated connectivity goals, thematic scope and other benefits that a project
may bring
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Figure 4.3: Response rates for taxonomic groups and ecosystem types.
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Figure 4.4: Response frequencies for the questions concerning policy context, target users and funding sources.
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Figure 4.5: Response frequencies for questions related to projects’ spatial scope and the biogeographical regions
where they took place.
Figure 4.6: Response frequencies for questions concerning selected approaches and what kind of spatially explicit
information projects produced.
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Figure 4.7: Number of projects that did and did not implement monitoring.
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Figure 4.8: Response frequencies for potential negative effects caused by increasing ecological connectivity.
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Figure 4.9: Type of organisation and distribution (% of total) by country of the participants in the online workshop
“Assessing Ecological Connectivity in Europe: Conservation goals and information gaps”.
The workshop consisted of two sessions, each divided into several breakout groups and
aimed at answering a set of questions provided at the beginning of the sessions for guidance.
Within both sessions participants were divided into different thematic breakout group
discussions (Table 4.1); each discussion centred on a virtual whiteboard (“Miro Board”) with
multiple prompts where participants could leave responses (Fig. 4.10). The first day focused
on identifying priorities for connectivity planning in Europe from five different subjects. The
questions tackled ranged from the desired outcomes of connectivity across scales, to which
areas should be better connected, where to establish corridors, and what measures have
been, or should be, implemented to maintain and enhance connectivity. The second day
focused on the technical challenges, data needs and potential solutions for connectivity
planning. The questions addressed related to the data needed to implement connectivity
projects, critical information gaps perceived, technical challenges and guidance constraining
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connectivity design, how to plan for connectivity in the face of climate change, and other
challenges, needs, or solutions for connectivity planning and corridor design (Fig. 4.10; see
Annex S2 for additional examples).
All responses from the Miro Boards were gathered, and a two-stage qualitative thematic
analysis was conducted across all the groups. This form of analysis aims to synthesise
responses into major groups to classify and categorise survey or discussion responses.
There were multiple repeating themes in the responses provided by the participants for both
priorities (Day 1) and challenges and solutions (Day 2) (see breakout group themes in Table
4.1), which are summarised in the following Sections.
Table 4.1: Breakout group themes for the two days of the “Assessing Ecological Connectivity in Europe:
Conservation goals and information gaps” workshop. Stakeholders participating in the workshop self-selected their
group and then interactively added responses to prompts derived from their general theme.
Day 1: “Identifying priorities for Day 2: “Technical challenges and gaps for
connectivity planning in Europe” connectivity analyses and planning”
Protected Areas & Natura2000 Human Infrastructure & Land Use Impacts
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Figure 4.10: Miro board produced in the breakout group on “Enhancing Connectivity for Endangered Species and
Habitats” from the workshop “Assessing Ecological Connectivity in Europe: Conservation goals and information
gaps”. The red circles indicate the issues that participants identified during the session as the most pressing ones
and the arrows indicate some connections between the questions being addressed.
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Figure 4.11: Word response frequency retrieved from the Miro boards for Day Two of the “Assessing Ecological
Connectivity in Europe” workshop. Day Two was concerned with identifying the technical challenges, data needs,
and solutions for connectivity planning in Europe. Data on species spatial distribution, abundance and movement
was the most highly discussed response.
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multiple corridors to add redundancies into the protected areas network or when one corridor
will likely suffice.
Designing and implementing multifunctional corridors was seen as a means for stacking
multiple goals into one conservation action (e.g., integrating biodiversity conservation, human
recreation, and provisioning ecosystem services - amongst other goals - into one coherent
design). However, the existing frameworks for complex problems and multiple objectives and
constraints have not been widely applied yet to connectivity related problems.
Other challenges to implement connectivity projects relate to land ownership issues; creating
corridors on private land requires working closely with landowners, and setting participatory
approaches where they have the required knowledge and are involved in decision making.
Finally, the issue of assigning economic value for connectivity was also raised to better argue
for the need for restoration/connectivity planning, namely compensation schemes for
conservation actions including features that increase connectivity.
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Table 5.1: Outline of the most common modelling families for functional and structural connectivity. Lists of
applications and software are not exhaustive but demonstrate some of the possibilities with these models.
Resistance and source weight surfaces are rasters that are inputted typically in ASCII format or another raster
format depending on the software package (see Dutta et al. 2022 for an extensive listing of software).
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Analysis of landscape
structure and potential
Focal nodes & connection file Conefor, ArcPro Network
functionality, prioritisation of
Graph Theory of attributes between node Analyst, R packages
patches and connections,
pairs (‘iGraph’, ‘riverconn’)
long-term population
persistence
The habitat patches or “focal nodes” where species or processes of interest occur and where
movement is expected to originate and/or terminate are often protected areas, but this
depends on the research question as, for example, the analysis could revolve around the
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inclusion of hypothetical unidentified nodes that could improve ecological connectivity. This
can be represented as single or clusters of pixels derived from a raster or polygon shapefiles.
Data requirements on these nodes can vary from a simple text (i.e., ASCII (American Standard
Code for Information Interchange) format), CSV (comma-separated values) file with the
distances between pairs of pixels, to spatial data formats (e.g., shapefiles) that contain
georeferenced characteristics about each patch.
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connectivity models to determine an optimal resistance surface (Dutta et al., 2022; Peterman,
2018).
Spatial prioritisation
Finally, spatial prioritisation software can serve as a means of identifying and prioritising
structural connectivity between habitat patches (Beger et al., 2022; Daigle et al., 2020). Spatial
prioritisation software is typically used within a broader systematic conservation planning
(SCP) process to identify efficient means of conserving a broad suite of natural and cultural
features (Table 5.1; see also Section 2.7). Generally, the user gives the program a spatial
layer of the study area divided into “planning units”, each unit serving as the base-level
decision-making unit. The user also provides spatial features of interest such as endangered
species presence, key biodiversity areas, culturally sensitive landscapes, important
watersheds, etc. In addition, output from the other connectivity modelling families (e.g., least-
cost corridors, high current density areas) that explicitly measure levels of structural or
functional connectivity can then be used as a conservation feature in the spatial prioritisation
software (Hanson et al., 2022). The program can then attempt to meet the conservation targets
maximizing the occurrence of considered spatial features, while creating a connected and
relatively compact protected area network. An advantage of including connectivity in
prioritizations (see Section 2.7) is that any area selections (e.g., protected area expansions)
are not considered in isolation from connectivity within a wider land- or seascape.
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resistance. It is therefore considered the path that a hypothetical animal would most likely take
as it would exert the least amount of energy. Given the methodology, the path it identifies is
only one pixel-wide, leaving the user with theoretically the single most effective corridor for
conservation between those two points. This method was extended to the factorial least cost
path, which considers multiple source points or locations where an animal may be originating
and ending its movement (Cushman et al., 2013, 2009). Factorial models predict all the
possible combinations of source to endpoint paths cumulating the resulting paths together.
This creates a quasi-prioritization of path areas, as the higher the cumulative weight on the
landscape the more overlapping pathways you are conserving. This is somewhat more
realistic, especially for larger protected areas where it is much more difficult to predict where
on the PA border an animal is likely to enter or leave.
However, there are a couple of key assumptions and issues when relying on a least-cost path
analysis. First, we are assuming an animal has the knowledge and foresight of the landscape
matrix to choose the path that gives it the least resistance (Unnithan Kumar and Cushman,
2022). Depending on the situation this could be a troublesome assumption as most individuals
will likely not have this foresight for the landscape, especially for dispersing individuals that
have potentially never been in that area. Second, as with many of the connectivity analyses,
we are assuming that the cost layer that we are using captures the true resistance that a
species faces when it attempts to move through that land area. Third, we are identifying only
one “best” corridor that is the width of a pixel, rather than multiple solutions that take into
account a buffer area around the corridor. Highlighting the one best connection is only the
starting point in determining if conserving that area is sufficient to ensure first structural and
then functional connectivity. Most likely, a much wider corridor that possibly incorporates even
wider stepping stones is necessary to achieve the functional connectivity needed for
conservation success. Some of this can be solved by factorial least-cost path analysis, but
there can still be heavy overlap in the identified paths leading to very narrow corridors.
Resistant kernels are the last member of this family. It is a unique adaptation of least-cost path
analysis that seeks to broaden the interpreted area of the least resistant path by using a hull
or moving window estimator and information on the dispersal potential of the animal. This
moving window approach can assist in identifying the corridor buffer area needed by that
species, especially when constrained to the output from a least-cost path analysis (Cushman
et al., 2013). In addition, there is no information needed on the end point of movement, as the
model does not assume that the focal species has prior or precise knowledge of its final
destination, an assumption that must be made in least-cost path analyses.
Least-cost path and factorial least-cost path have been broadly applied in connectivity
analyses but are primarily used for modelling the likely movement of a single wildlife species
between protected areas including mammals, reptiles, amphibians and insects. There is also
application in simulating possible downstream water flow and flooding routes and routes for
pollinator movement (over short and long periods). The applications are the same for resistant
kernels but with the added benefit of including the dispersal ability of the species. This can be
expanded to include the scattered dispersal of insect or animal movements from hives or
colonies. Given the functional connectivity aspects of resistant kernels, they could also
hypothetically be used to simulate likely sediment flow and potential plant dispersal under
climate change.
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Some of the software and programming packages that can implement least-cost path and
resistant kernel analyses include (Table 5.2):
Table 5.2: Software and programming packages that can implement least-cost path and resistant kernel analyses.
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Resistant UNICOR Universal Corridor This is the primary software used for
kernels network simulator resistant kernels estimation. It was created
(https://github.com/Co by the developers of the resistant kernels
methodology. It is a Python-based program
mputationalEcologyLa
and can be used through the Python
b/UNICOR) command line or in a graphical user
interface (GUI), which can be launched
following the instructions in the UNICOR
manual.
Ultimately, these methods are still used and in the case of least-cost path analysis is a simpler
and quicker way to identify the potentially important movement corridors in a landscape. This
is especially useful for large and high-resolution datasets given the heavy computing capacity
needed for some of the other connectivity methods. However, while the more information and
preparation may be required, resistant kernels can give a better and more realistic output as
to the best overall areas to be conserved for connectivity (Sumar & Cushman, 2022). Also,
with the addition of dispersal information into the model, resistant kernel output can be seen
as measuring functional rather than just structural connectivity.
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and
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between vertices, more sophisticated methods can estimate distance as a least-cost path or
similar approaches.
Connectivity analyses using graph theory have incorporated asymmetric connectivity, which
considers differential probabilities of dispersal from one patch (e.g., patch A) to another (e.g.,
patch B) compared to the reverse direction (from B to A). This development has applications
in source-sink dynamics or the modelling of passive dispersal, such as wind or water-mediated
dispersal.
The R package "iGraph" is commonly used for computing graph theory analyses and
visualising graphs (Csárdi et al., 2023). However, ecological connectivity analysis using graph
theory is often performed using the software Conefor, which includes a wide array of metrics
commonly used in connectivity analysis.
Conefor
Conefor (originally ‘Conefor Sensinode’) is an open-source software implemented in C,
providing computational efficiency compared to R. It offers a user-friendly graphical user
interface (GUI) and can also be run from the command line (Saura and Torné, 2009). Both
versions of the software can be downloaded from the official Conefor website
(http://www.conefor.org/index.html), which also provides a comprehensive manual and
reference list. Several GIS software have also developed extensions to apply connectivity
analyses using Conefor (e.g. QGIS: https://github.com/ricardogsilva/qgisconefor; ArcGIS:
http://www.jennessent.com/arcgis/conefor_inputs.htm).
Conefor requires a node file and a connection file as inputs. The connection file can contain
attributes represented as Booleans (0 or 1), distances (which are converted to probabilities of
connection based on an average dispersal distance and assuming a negative exponential
kernel for probabilistic metrics), or direct probabilities of connection. Users can select specific
connectivity indices and define a maximum dispersal distance to limit connections between
nodes beyond a certain biological threshold. The software also allows users to compute
metrics for the overall network (faster) or for each vertex and edge (slower). Additionally,
Conefor provides the capability to assess scenarios involving the addition of vertices to the
network, as well as improvements or deteriorations in the existing connections between
nodes.
For publications using Conefor on landscape planning and monitoring case studies see
http://www.conefor.org/applications.html, which includes a map allowing to find publications
for specific parts of the world, including Europe.
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especially when we are trying to identify corridors and pinch points (i.e., areas where the
movement of species is highly concentrated into a very narrow path). This has similarities to
factorial least-cost path analysis; however, circuit theory models can provide a continuous
raster surface of “current densities” which can be viewed as an index depicting the likely
concentration of species movement over each pixel (Fig. 5.1). The higher the current density
the greater the electric current is being concentrated at that point.
Figure 5.1: This simple illustration (left) shows nodes in the white pixels and the movement of electrical current
through each “resistor” (i.e., pixel). The black pixel is a permanent barrier that blocks any current flow. Once applied
to a real-world landscape (right) one can see how the current densities between the four focal nodes concentrate
along certain paths or diffuse (northern border) when no clear paths are present. This example is of North American
river otter connectivity through an agri/silviculture landscape surrounding forested wetlands. The model
concentrated current flow into riparian areas and the forested wetlands in the centre of the map (Sources: McRae
et al. 2013, Dertien and Baldwin 2023).
Models in the circuit theory family can be divided into directional and omnidirectional circuit
models. Directional circuit models use focal nodes or patches similar to the least-cost path;
however, in circuit theory, these node pairs are designated as either a source (origin of
electrical current) or ground (termination of electrical current). Hence, your source node is the
point where your species or process of interest is starting from, moving in the direction of and
terminating at the ground node. You can model movement in both directions by switching
these designations. The resulting raster output covers your entire study area and can present
one or multiple options for the most important movement corridors. Directional circuit models
are best applied to problems involving multiple patches, such as a system of separated
protected areas surrounded by unprotected “matrix” habitats. Additionally, directional circuit
theory may be applied to problems such as known migration routes or predicted directional
movement under climate change.
Newer omnidirectional circuit models run through the program Omniscape allow you to model
the connectivity potential of the landscape from all directions. Therefore, this model does not
require the selection of focal nodes, but it does require a new form of data input, a source
weight surface. The source weight surface depicts the potential for each pixel to, in essence,
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be a source node (Fig. 5.2). The model then uses a moving window calculation where the
centre pixel at each stop of the moving window is considered the ground node (McRae et al.,
2016). The model then calculates the connectivity from all the remaining pixels (i.e., source
nodes) within the moving window to the central ground node. Given the limited use so far of
this method there is little information on the correct moving window size, however, it should
be in part informed by the process or wildlife species that you are trying to model and your
computational resources.
Figure 5.2: Illustration of the moving window (left) that moves over the source weight and resistance surfaces.The
green pixels are coded as potential source pixels of varying strengths while the open white pixels are coded as not
a potential source (e.g., urban core). Current movement is then calculated between all potential sources to the
target (right). Cumulative current density for the landscape is calculated once all moving window calculations for
the landscape are complete. Adapted from Landau et al. (2021) and McRae et al. (2016).
Omni-directional models have great potential to solve many of the pressing landscape
conservation questions, especially for large landscapes where there are limited protected
areas, where there is extensive use by wildlife in the unprotected matrix, or where it is not
clear where wildlife or ecosystem processes may originate. However, given the cumulative
moving window approach taken by the model, the computational demand is significantly
higher than in directional models (i.e., Circuitscape). Running models for even smaller regions
can take days if using a single computer and may not be possible without access to high-
throughput computing or a supercomputer. Reducing the size of the moving window or
instructing the model to create a multi-pixel target node in the centre of the moving window
are two ways of reducing the computational demand for a model.
Like least-cost path analyses and resistant kernels, circuit theory models have been used for
a plethora of different applications. Dickson et al. (2019) found hundreds of peer-reviewed
studies over a ten-year period that applied circuit theory. These primarily focused on
mammalian movement but also included studies on birds, amphibians, reptiles, arthropods,
and fish. Additional applications have included the potential for wildfire movement, water flow,
and ecosystem services (e.g., pollinators and seed dispersers). Other studies include
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Table 5.3: Circuit theory applications developed detailing needed data inputs and sources.
Circuitscape 4 This older version of circuitscape Inputs for rasters layers https://circuitscape
is restricted to smaller regional- are in ASCII format like .org/downloads/
scale landscapes and has a Circuitscape 5
slower processing speed.
However, it still uses an easily
navigable GUI that is especially
helpful for first-time users and
contains many of the same
advanced options.
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patterns and processes that arise from the interaction of the agents among themselves and
with their environment. Not surprisingly, ABMs have found application in a wide variety of
fields, such as social sciences (Silverman, 2018), in particular economics (Farmer and Foley,
2009; Hamill and Gilbert, 2015), and in biology and ecology (Grimm and Railsback, 2005;
Railsback and Grimm, 2019; Zhang, 2018).
The main advantage of ABMs relative to other mathematical approaches, such as those with
differential equations, is that the latter are restricted by mathematical tractability. This means
that the only mathematical models that could be analytically developed were studied, but these
were often too simplified, to the point that one could doubt whether the relevant characteristics
of the system at hand were being properly modelled. Such doubts often arose when (i)
nonlinearities were present, (ii) the inclusion of interaction among individuals, or these with
the environments, were essential to understanding emergent patterns, (iii) modelling the
space explicitly, and its heterogeneity was required, (iv) agents/individuals were different, and
(v) agents/individuals exhibit complex and adaptive behaviour, such as learning (Zhang,
2018). Therefore, an important characteristic of an ABM, though necessarily a simplification
of a real system, is that it still retains enough processes and interactions so that simulations
include some of the most prominent processes of the real system that it attempts to model,
and thus an ABM can more closely emulate natural systems. Moreover, experiments that
cannot be feasibly carried out from a practical (or even ethical) perspective, e.g., the impact
of fragmentation on the persistence of populations, can be simulated with ABMs and the
consequences of different actions (and policies) evaluated.
The development of an ABM requires a multidisciplinary approach. At its core is the
development of a model that describes the movement patterns of individuals and how they
are affected by the environment, by the presence of other individuals, and eventually any other
factors that are known to influence movement. This model, which is a set of mathematical
rules that connect the movement properties (e.g. the distributions of step lengths and turning
angles of different movement states) to all the factors that influence them, is characteristic of
a species (or a species archetype) and can be developed either from expert knowledge alone,
estimated from real movement location data, or a combination of the two (see Table 5.4 for
examples of Software packages). Other, more complex or customised types of models may
be developed by writing a specific computer program that accounts for all the processes of
interest with no limitations, but in that case, there are no packages to allow estimating model
parameters from real data, and the model must be parameterized from expert knowledge.
Table 5.4: Table 5.4. Examples of software packages commonly used to implement agent-based models.
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As said, there are two pathways to parameterize an individual-based model: either estimating
the parameters from real location data or using expert knowledge. The first approach needs
real tracking data (e.g., telemetry data) over a time series, and, depending on the complexity
of the model, may have hard requirements as to the time resolution and period of the data. A
complex movement model (e.g., multistate movement in a heterogeneous landscape) may
require higher resolution data (e.g., location in every minute) and longer periods, to properly
fit the model. Models that account for landscape heterogeneity additionally require the
landscape structure to be provided as inputs. This can be in the form of a single landscape
resistance raster, as in SiMRiv (Quaglietta and Porto, 2019), or in the form of spatial covariates
that are known to influence movement, like forest cover, wind velocity, etc. (McClintock and
Michelot, 2018). In the case of landscape resistance data, it usually requires expert knowledge
about the species in question to score land cover types in terms of “resistance”, from the prism
of that species.
momentuHMM (McClintock and Michelot, 2018) and SiMRiv (Quaglietta and Porto, 2019) are
noteworthy models in that they allow integration of the influence of landscape structure in the
movement model, and thus can be used to simulate realistic movements constrained by real
or simulated landscapes, for example, as derived from scenarios of landscape change.
rangeShifter (Bocedi et al., 2020) can be used to explore the concept of evolving connectivity
in response to land-use modification, by examining how movement rules come under selection
over landscapes of different structure and composition.
If real tracking data is not available, or not sufficiently detailed to allow estimating model
parameters, these can be set manually from expert knowledge. For example, a given animal
may be known to have two movement states, and to spend most of the time in “state one”.
Then, the state transition matrix (i.e. the probabilities of changing from one state to another)
may be parameterized manually to respect this known behaviour. Similarly, the type of
movement in each state may also be set manually by defining basic movement properties like
the step length distribution and the amount of correlation between successive step angles, in
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Table 5.5: Table 5.5 Examples of various structural connectivity metrics, including references, all varying in
complexity, with explanation of what the respective metrics measures. “*” indicates metrics requiring only physical
components. “**” indicates metrics requiring both physical and graph components.
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Distance to nearest Edge distance to the nearest neighbouring patch. Prugh, 2009
neighbour*
Effective mesh size* The probability of two randomly placed points in the Jaeger, 2000
landscape being connected, converted to area.
Habitat within buffer* A measure of how isolated or aggregated focal Prugh, 2009
patches are in the landscape
Patch cohesion index* Standardized area-weighted mean of ratio between Schumaker, 1996
the perimeter and area
Mean radius of gyration* A measure of how far-reaching a patch is across McGarigal, 1995
the landscape.
Proximity index* A measure of the size and proximity of all focal McGarigal, 1995
areas within a user-defined buffer around focal
patch
Betweenness centrality** The degree for which a patch can serve as a Albert et al. 2017
stepping inside a network.
Clustering coefficient** The average fraction of a node’s neighbours that Jordán et al. 2003
are also neighbours with each other.
Compartmentalization** The relationship between the degree of focal nodes Minor et al. 2008
and the average degree of neighbouring nodes.
Integral index of The connectivity of focal areas based on habitat Pascual-Hortal and
connectivity** size and the number of stepping stone patches Saura, 2006
separating focal areas.
The advantages of using structural connectivity metrics are that they are often easier to apply
since the input data needed are often more widely available or can be acquired
computationally since most metrics do not require any ecological of species-specific data. The
downside of these metrics is the sometimes lack of spatial explicitness. Some metrics only
result in a single value of connectivity for the focal area, and they are therefore most useful for
quicker assessments of connectivity in smaller areas or for comparisons of regions/countries.
However, on larger spatial scales structural connectivity metrics can become less informative,
as some metrics do not take the geographical placement of areas into account in the final
results.
An approach to attempt to make structural connectivity metrics more spatially explicit, could
be the implementation of a moving window analysis, also referred to as focal analysis or
neighbourhood analysis. This type of analysis is a common technique when it comes to data
analysis, image processing and feature extraction, particularly in relation to multi-scale data
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(Hagen-Zanker 2016). This method consists of a window, with a fixed size, moving sequentially
along a dataset, where computations are then performed within each window, with the result
often applied to the focal area or individual raster pixels. By combining a structural connectivity
metric into a moving window approach, the metric is effectively to applied to the individual
pixels and the whole study area at the same time, rather than providing the user with only a
single connectivity value. This results in a gradient map of connectivity values; this provides
more spatial explicitness to the metric as changes in connectivity across the landscape can
be observed both at the local scale as well as a larger country-wide or even continental scale
(Fig. 5.3).
It is worth noting that with the implementation of this moving window approach, the resulting
ranges for which some connectivity metrics operate, will change, due to the metrics now being
confined to a pre-determined size of the moving window. This is only applicable for metrics
where the results can range from [0, + ∞) (e.g. Distance to nearest neighbour, Effective mesh
size, Habitat within buffer, Mean radius of gyration, Area-weighted mean radius of gyration,
Proximity index, etc.) (Yang et al., 2024). However, as mentioned, this approach allows for
structural connectivity metrics to provide a better overview of connectivity in the landscape by
being more spatially informative on larger and more local scales at the same time.
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Figure 5.3: Map of Europe showing the result of using the moving window approach in conjunction with the Effective
mesh size structural connectivity metric. In this case the connectivity metrics was applied to all forest patches
extracted from the CORINE Land Cover. Darker brown colours indicate a larger effective mesh size value, further
indicating higher connectivity.
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Ecosystem Services (ES) mapping approaches can broadly be classified into five categories
(Burkhard and Maes, 2017; Honeck et al., 2020; Martínez-Harms and Balvanera, 2012).
The "lookup table" approach is a commonly employed and straightforward method connecting
ES with geographic information, predominantly relying on land cover data. In this method, land
cover data serves as proxies representing the supply or demand of various ES. The lookup
table incorporates ES information often derived from statistics like crop yield in the context of
agricultural production.
1. “Expert knowledge approaches” mainly rely on specialists to rank land cover classes
based on their potential to provide services; experts estimate ES values in lookup
tables but also use other methods such as Delphi surveys.
2. The "causal relationship" approach involves estimating ES by leveraging established
connections between ES and spatial information extracted from literature or statistical
sources. As an illustration, the estimation of timber production utilises harvesting
statistics specific to various areas, elevations, and forest types as documented in a
national forest inventory.
3. The “extrapolation of primary data” method associates weighted field data with land
cover and other cartographical data; Approaches that estimate ES extrapolated from
primary data such as field surveys linked to spatial information.
4. The “Regression models” method combines biophysical information from field data and
the literature into a quantitative ecological system model.
The use of GIS in ES mapping can take three general approaches: (1) analysis tools built into
GIS software packages; (2) disciplinary biophysical models for ES assessment (e.g.,
hydrological models such as the Soil and Water Assessment Tool or Variable Infiltration
Capacity model for water-related ES); and (3) integrated modelling tools designed specifically
for ES assessment (e.g., InVEST, ARIES) (Burkhard and Maes 2017). The initial method is
suitable for straightforward analyses based on land cover and indicator mapping of Ecosystem
Services (ES), as demonstrated, for instance, in Mapping and Assessment of Ecosystems and
their Services (MAES). The second approach is suited for intricate model-based analyses of
services, integrating expertise from distinct disciplines (e.g., ecology for crop pollination or
hydrology for flood regulation mapping). The third approach builds upon the second by
employing modelling tools capable of evaluating trade-offs and scenarios for multiple services
(Burkhard and Maes 2017).
An overview of scientific and technical tools for GI mapping, including the European Mapping
and Assessment of Ecosystems and their Services (MAES) initiative; and geospatial methods,
data and tools (e.g. CORINE, LUCAS, Copernicus), are detailed in Estreguil et al. (2019).
Relevant maps and data have been produced by the European Environment Agency and the
Joint Research Centre (https://data.jrc.ec.europa.eu/collection/MAES).
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Assessing and developing a clear understanding of the connectivity problem is the first critical
step in a connectivity planning exercise. In the context of connectivity such issues could include
declining species population numbers, decreased migratory movement, or habitat loss due to
climate change. Scoping the problem at this stage includes i) assessing the threats in the
system (e.g., road mortality, encroaching development, dams), the possible actions that can
be taken to mitigate these threats (e.g., protected corridors, underpasses, seasonal closures),
and the likely impact from such actions. When assessing the likely impacts of any action, it
would also be appropriate to consider how they will be evaluated.
A part of scoping will involve identifying all the key stakeholders involved with the problem
and with the targeted region. Identifying as many stakeholders as possible and involving key
individuals in the decision process from the beginning is vitally important to produce the best
possible outcome. This steady involvement from the beginning will produce a more accurate
pre-implementation plan, increase the chances of the plan being implemented, and lay the
groundwork for management and monitoring post-implementation. Decades of collaborative
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conservation projects have shown the importance of such sustained stakeholder involvement,
and this was confirmed in our stakeholder webinar (Section 4.2) where participants frequently
identified the need for comprehensive stakeholder engagement from the beginning of any
conservation action or initiative. If producing deliverables informing landscape-scale planning
is an ultimate goal, one should think about what products need to be produced to inform private
citizens and public decision-makers about the important areas for corridor conservation.
Before determining the final objectives of the study and while still assessing the overall problem
a collaborative team should be set. Given the ecological and socio-political complexities of
these projects an interdisciplinary team should include natural resources practitioners,
communication specialists, scientists, policymakers and ideally project administrators. This will
assist in producing robust scientific results which are more representative of the on-the-ground
situation and increase the chances that such a plan will garner support for implementation.
Defining the scope of the connectivity planning study will also include identifying the study’s
basic ecological realm (e.g., terrestrial, freshwater, etc.) and general spatial extent (i.e., total
study area). The most effective spatial extent for successful implementation of connectivity
and conservation planning is often context dependent; however, the spatial extents should
ideally be informed by an intersection of the biologically relevant area and the socio-political
conditions that support the implementation. Therefore, it is important to consider if the project
is operating at a spatial extent that captures important ecological and abiotic processes while
also being contained within a governance structure that can maintain support for the
implementation of project aims.
A focus on species’ natural history and especially dispersal capabilities is important when
deciding upon the spatial scale of your study (see Ch. 2.4). This is especially true for certain
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groups such as semi-aquatic species including river otters, salamanders, and turtles, which
are cross-realm species especially threatened by linear infrastructure either blocking
movement or causing mortality during dispersal (e.g., roadkill). Connectivity assessments in
riverine ecosystems should ideally take place at large spatial extents (e.g., catchment or sub-
catchment). Especially for long-distance migratory species, large-scale assessments can
support the identification of bottlenecks and priority areas for restoration. For potamodromous
species (i.e., species limited to freshwater ecosystems), smaller scales covering the
distribution range of respective species might be sufficient. At the least, the selected spatial
scale should include all required habitats for the target species to complete their life cycle (i.e.,
habitats for spawning, feeding, and wintering).
A final consideration while scoping the project is the potential impact improving ecological
connectivity could have on the provisioning of ecosystem services. Connectivity can directly
or indirectly affect ecosystem service provisioning, making these services valuable targets
themselves. Furthermore, many ecosystem services are strictly linked to the movement of
certain species. Examples are seed dispersal, pollination, nutrient cycling, and cultural and
recreational activities. However, priority areas that improve connectivity to increase ecosystem
services supply might not be the same as for the aim to mitigate threats for specific species
to improve their conservation status. Aiming to increase ecosystem services is particularly
important in locations near urban areas, where the multifunctionality of the green infrastructure
is more relevant. By enhancing the connectivity for the species or functional groups that
provide these services, connectivity interventions indirectly contribute to the provision of
ecosystem services.
Once full scoping of the problem is complete and it appears that an action taken to promote
the creation or restoration of a connectivity network will help mitigate the problem, the project
team should decide on specific project objective
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The planning of connectivity requires careful consideration of various factors, and it is essential
to define the specific goals and beneficiaries of these efforts. Connectivity can focus on whole
ecosystems, specific habitat types, species, functional groups or ecosystem services. This
choice entails trade-offs between generality and specificity. Taking the findings from project
scoping in step 1, it is then essential to establish one or more clear and achievable
objectives that will guide the analysis and actions taken by the project (Table 6.1). Objectives
can be customized towards the conservation of one species or wide-ranging to include multiple
overlapping ecological functions that would benefit from connectivity conservation.
Objectives can include conserving the daily movement of a species, maintaining seasonal
migratory pathways, increasing genetic exchange between populations, dispersal of seeds or
pollen, or movement of nutrients (Hilty et al., 2019). Spatial and temporal scale are most
important to consider when developing objectives and indicators as the planning decisions
made henceforth will vary dramatically if, for example, you are interested in the dispersal
capability of an individual animal or plant versus the long-term population viability of those
species. Examples for general objectives could include: i) biodiversity conservation (e.g.,
daily movement of species; multi-species movement over decades; yearly fish migrations;
wildlife and plant dispersal corridors due to climate change; invertebrate connectivity across
intensive farmland or forestry); ii) ecosystem services (e.g., sediment capture and reduction
of erosion; pollinator connectivity; water filtration; nutrient cycling and carbon sequestration;
recreation); iii) reduction of impacts from infrastructure (e.g., locations for green bridges or
underpasses; prioritisation of dam or road removal; establishment of best land zoning
regulations); iv) connectivity planning in urban/peri-urban areas (e.g., placement of
greenbelts; restoring water flow between urban wetlands; v) multi-functional corridors (e.g.,
combination of objectives for species conservation, ecosystem services, human recreation,
and/or other goals).
Depending on the connectivity project certain objectives may also include specific quantitative
targets to be met to achieve that objective. For example, while the objective may be to
establish a corridor to an isolated protected area to conserve movement of brown bears, a
target could include a quantitative measure of the number of individuals or breeding females
desired to disperse via the corridor (Table 6.1).
Table 6.1: Four potential connectivity problems with an example objective, target and action. Note that actions are
not always just implementing and restoring a protected area (PA) or corridor but can include seasonal changes of
human disturbance or alterations in permissible land use.
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When projects target groups of species sharing some fundamental traits and with similar
conservation needs, a possible approach is focusing on species archetypes, i.e. generic sets
of traits representing groups of species that are functionally similar from a connectivity
perspective. Connectivity between populations results from a combination of their physical
distance, landscape resistance to movements, species reproductive potential (timing and
output), and dispersal ability. Among species traits, reproductive potential determines the
number of propagules that are produced over time, whereas dispersal determines the ability
of each propagule to reach a given distance.
This approach has been explored for European plants by Lososová and colleagues (2023),
while for terrestrial vertebrates no similar analysis has been published up to now. Following
an approach like that published for plants, it has been identified a total of 27 archetypes in
European tetrapods (see below) considering reproductive traits and other variables that can
relate to spatial requirements and energy consumption (e.g., body mass, home range size;
detailed methods are reported in Annex S3). Median trait values are reported in Annex S3
(Tables S3.2-S3.9).
• Non-volant mammals were divided in (1) large mammals with long life span and
long dispersal distances, (2) large mammals with long life span and medium
dispersal distances, (3) medium mammals with long life span and medium dispersal
distances, (4) small-medium mammals with medium life span and short dispersal
distances, (5) small mammals with short life span and short dispersal distances.
• Bats were divided in (1) bats with small home ranges and short dispersal distances,
(2) bats with large home ranges and long dispersal distances, (3) bats with medium
home ranges and short dispersal distances.
• Birds were represented by (1) large birds with long lifespan, medium dispersal and
wide home range, (2) medium-sized birds with medium longevity, medium dispersal
and medium home range, (3) small-medium birds with medium-short lifespan, long
dispersal and small home range, (4) small birds, with short lifespan, short dispersal
and small home range.
• Frogs were divided in (1) large frogs with a high clutch size, (2) small-medium toads
with a low clutch size, and (3) small frogs with a medium clutch size.
• Turtles were divided in two archetypes: (1) small-medium turtles with high clutch
size, and (2) large tortoises with medium clutch size.
• Snakes were characterised by three archetypes: (1) big-size snakes with high
clutch size, (2) small-medium snakes with high clutch size, and (3) small-size
snakes with low clutch size.
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• Lizards were represented by three archetypes: (1) big-size terrestrial lizards, (2)
small-medium size saxicolous lizards, and (3) small-medium size terrestrial lizards.
Understanding species habitat preferences (Fig. 6.2) is crucial to establish a connection
between archetypes and a specific environmental context highlighting the critical role of
landscape resistance in conducting effective connectivity analyses. However, most species
have species-specific combinations of habitat requirements (Figs. S3.1 and S3.2 in Annex
S3), which prevents generalizations, even considering coarse habitat classes. In this context,
a major challenge in connectivity modelling is represented by the consideration of geographic
barriers. The modification of landscapes through the development of roads, railways, fences,
and canals increases habitat fragmentation by reducing species movement and increasing
mortality (Bastianelli et al., 2021). For example, birds that live on the boundary between
roads and forest or pastureland are more susceptible to being injured or killed by vehicles or
noise barriers; similarly, birds in anthropogenic environments have been reported to collide
with wind turbines, power lines, and building windows (Medrano‐Vizcaíno et al., 2022). In
addition, traffic and roadkill represent a main threat for mammals across Europe, because of
their movement ecology and the large amount of space that they require for dispersal and
home range (Klar et al., 2009). In the European scenario, overcoming or eliminating
geographical barriers could be crucial to enhance spatial connectivity for species of
conservation concern. However, when analysing species-specific connectivity, it is essential
to account for the resistance posed by such infrastructures to prevent overestimating or
underestimating the species’ movement abilities in connectivity modelling.
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Figure 6.2: Habitat preferences (percentage of species) of European tetrapods using three coarse classes.
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Corridor design should also consider edge effects and the zone of influence of human
activities which changes ecological processes inside corridors and reduces their effective
width (Boulanger et al., 2012), that is, the residual space that occurs beyond the zone of
influence of human activity. For instance, if a 400 m wide ecological corridor is adjacent to a
residential area and the influence from that development is 100 m, then the effective corridor
width is reduced to 300 m.
The current body of scientific research does not provide fully comprehensive evidence to
determine how wide ecological corridors should be to attain all conservation goals (Gilbert-
Norton et al., 2010; Haddad et al., 2011; Sawyer et al., 2011). Given the complexity,
confounding effects, and time-consuming nature of assessing the effectiveness of ecological
corridors and determining the optimal width (Beier, 2019; Gregory and Beier, 2014), the
available evidence is relatively scattered (Beier, 2019); however, there are some key
resources that provide valuable recommendations (Box 6.2).
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Concerning the general recommendations, they suggest that 1) larger species need wider
corridors to facilitate movement and provide potential habitat, 2) longer corridors should be
wider than shorter corridors and 3) shorter corridors are more likely to provide connectivity
than longer corridors.
As for the specific recommendations, they proposed minimum corridor widths and
recommended corridor widths for several groups of species ranging from plants to large
predator mammals (Table 6.2).
Table 6.2: Minimum and recommended corridor widths based on the review of 66 scientific studies (Bentrup,
2008).
Plants 30 101
Invertebrates 30 61
Aquatic species 30 61
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Likewise, data resolution must be much finer (i.e., smaller pixel size) for an analysis of ground
beetle connectivity (<10 m) compared to the ungulate population (30 m - 1 km) given the
differing scales at which the species respond to their environment. Local and regional scales
of assessment are frequently the scale management actions are implemented at to increase
connectivity and thus require fine-grained, accurate and detailed data. The large-scale
(transborder or continental) mapping of corridors needs less detail and is adequate for projects
related to the planning of transportation infrastructure or trans-boundary coherence for the
connectivity of protected areas.
The primary concern is to produce analysis products that are at fine enough resolution to give
meaningful and operational information for practitioners and decision-makers, at a spatial
extent which will be implementable. Producing coarse grain connectivity assessments may not
necessarily provide any new information to local resource managers to aid them in
conservation decision-making.
In addition, to produce more accurate results, the
impacts of study area boundaries on different
models should be considered when determining
the entire spatial extent to conduct the
connectivity analysis. This is true for circuity
theory models (see Section 5.4) and other models
influenced by boundary edge effects where results
near boundaries can be significantly biased and
are less trustworthy (Koen et al., 2010). Therefore,
depending on the model, analyses should include
buffer areas beyond the agreed-upon study area
to negate any boundary impacts within the area of
interest. These buffer areas can be simply clipped
off from the final connectivity results and
deliverables.
Finally, a third factor to consider for large-scale
projects is the computational power needed to run
analyses as they approach large extents and finer
spatial resolutions. This may not be a factor for
relatively simpler analyses such as least-cost
path, but with analyses such as resistant kernels,
circuit theory models and certain movement
models fine spatial resolution at large spatial Overlay of satellite imagery and Circuitscape results
extents may require access to a high-throughput
computing cluster or supercomputer to complete analyses over multiple days.
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Having established your objectives and before extensive data collection, it is important to
determine the analysis that will achieve them. As was introduced, there are many tools to
choose from to address the project question or objective (Chapter 4). When the objective is
connectivity of habitat types (e.g., connectivity of forests) or to enhance the connectivity of
particularly intact natural ecosystem systems, structural connectivity metrics or graph theory
models that only consider the basic structure of the landscape could suffice (e.g., iGraph,
riverconn, Conefor; see Table 4.1). Conversely, functional connectivity (e.g., agent-based,
circuit theory, and resistant kernels models) should be prioritised when project objectives
centre on a focal species or a specific group of species.
Structural connectivity analysis via graph theory (e.g., Bunn et al., 2000) has been applied
for analysing landscape structure and functionality, prioritising patches and connections, and
for assessing long-term population persistence. Applying graph theory to connectivity analysis
allows for examining network connectivity as a whole, which facilitates comparative analyses
among different networks. Data needs include focal nodes and connection attributes between
node pairs. The emphasis is put on the configuration of a network, including the isolation, size,
and shape of the patches, connecting elements (e.g. corridors, stepping stones), and elements
that can act as barriers (e.g. anthropogenic such as roads or natural such as rivers). Such
simplified assessments are likely to be easier to communicate to stakeholders and
policymakers (Saura et al., 2011).
If structural connectivity is being used as an index for multiple species, this approach assumes
that enhancing connectivity at a structural level can facilitate the movement of various species,
reducing the risk of population isolation and contributing to genetic diversity. This might be
particularly sensible when species of these systems face similar connectivity challenges (e.g.
riverine ecosystems where species face similar movement barriers). For instance,
interventions to enhance connectivity in river networks, despite hosting diverse species, can
equally benefit numerous freshwater species. Overall, enhancing the structural connectivity of
certain habitat types is expected to yield benefits to plant and animal species dependent on
these habitats.
Targeting functional connectivity must consider species-specific characteristics such as
dispersal propensity and abilities, area requirements, permeability of different landscape
elements to the species movement, behavioural response to infrastructures, etc. Movement
models or some agent-based models that can use GPS or radio-tracked animals locations
provide some of the closest information we have to true functional connectivity (Table 4.1).
Circuitscape, Omniscape and Linkage Mapper can provide quasi-functional connectivity model
estimates, especially when resistance surfaces are generated using either spatially-explicit
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genetics data of a species (i.e., allele variation between subpopulations of one species) or
from detection/non-detection wildlife data.
Resistance to movement between and within habitat patches is central to least-cost path,
resistant kernels, and circuit theory models. There are several methods to create resistance
surfaces, depending on the type of data available for the species of interest (Zeller et al.,
2012). The most common method is to invert habitat suitability values obtained with species
distribution models so that the areas with higher habitat suitability values have the lowest
values of resistance. Other methods include using telemetry, GPS point and track data.
In contrast, spatial prioritisation tools commonly used for systematic conservation planning
can be useful to identify structural connectivity, by prioritising landscape elements (Hanson et
al., 2022). Needed data inputs include a conservation planning units file, the spatial distribution
of biodiversity, land use, or protected areas. Available tools include Marxan, Zonation, or the
R package prioritizr (see Table 4.1 and Section 2.7 for details).
There are several approaches and tools to map ecosystem services (see Burkhard and Maes,
2017 and Estreguil et al., 2019 for details), whose selection depends as well on multiple factors
including the questions to be addressed, the spatial scale, and data availability (Section 4.6).
Graph theory or circuit theory models are best for problems surrounding water quality,
sediment capture, nutrient cycling, and fire. Circuitscape especially is utilised for attempts to
simulate potential fire spread. Seed dispersal and pollination services can be modelled with
resistant kernels if there is data on dispersal distances for the species or using circuit theory
models even without dispersal data. Finally, finding the best movement pathways for
recreation can be accomplished using least-cost path methods or Circuitscape.
Data requirements for many of the models are very similar (Fig. 6.3). Most require some
representation of the landscape via a land cover dataset and/or elevation model.
Understanding the current protected area network including the georeferenced boundaries of
each protected area is vital for any of the connectivity models. Spatial data should be acquired
for all the threats and potential barriers identified in step 1, such as dams, border walls,
motorways, or major water bodies. If the creation of a resistance surface is necessary, climate
data could also be important for generating species distribution models. Gathering data on the
dispersal capability of the species of interest is also important for resistant kernels, particularly.
The checklist on the proceeding page covers some of the more common data needs but is not
fully inclusive of all connectivity modelling contexts and Annex S4 provides a basic listing of
geospatial data sources for the European continent.
Finally, before conducting the connectivity analysis, stakeholders should be engaged again (if
they haven’t been continually) to inquire if there are any data pieces missing or if there are
unforeseen gaps in the data. For example, information on the protected areas network may
have been downloaded from the World Database of Protected Areas. While the coverage of
this database is quite extensive, it may not be comprehensive, especially at the local and
regional level where several parks and protected areas or OCEMs may be excluded. So too is
information on green bridges/wildlife passages whose locations may not be published or well-
advertised.
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Figure 6.3: Checklist of common spatial data needs for connectivity analyses. Data needs will vary between the
different models and research problems, so this should not be seen as an exhaustive list of what may be required.
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Having determined the analysis framework you will be using, collected all the necessary data,
and consulted with stakeholders about any gaps in the data, connectivity analysis can be
conducted. This should be a cyclic process with the collaboration team generating draft model
results and then gathering input from key stakeholders on the draft results, before running the
models again. This aids in the effectiveness of the final plans and the realistic possibilities of it
being implemented.
Depending upon the chosen approach, time for such assessments could take weeks or months
to fully process and run the models. Typically, much more time is spent processing and
preparing the data for input into the model, rather than the actual construction of code or
runtime of the model. Graph theory models with simple input information on node and link
characteristics or smaller-scale least cost path analyses can be run in minutes. Conversely,
movement models that are processing tens of thousands of GPS locations or circuit theory
models such as Omniscape can take days or weeks to complete the analysis for a single
species. The online resources highlighted in Chapter 4 can provide guidance on what to expect
in terms of runtimes and assistance in troubleshooting modelling problems.
Once you have draft results, they can either be presented to stakeholders (e.g., regional
councils, land management agencies, conservation organizations, private landowners, etc.)
for review or further analyses can be conducted to prioritise the corridors and/or stepping
stone locations before stakeholder consultation (see Section below). Prioritisation may not
always be necessary, depending on the project, or you may wish to seek stakeholder input on
draft results to assist with the ultimate prioritisation.
Stakeholder reviews can be done in-person or via webinar. In either setting, members of the
collaboration team should present the workflow followed in the project and the specific
methods used to generate the connectivity maps. Like public regional planning meetings, initial
results and maps from such assessments should be presented so stakeholders can provide
expert opinions on the preliminary results.
In-person meetings should allow stakeholders to engage in open discussion and to draw and
leave sticky notes on the maps. Ideally this will assist in identifying impractical or potentially
spurious results coming from the models, missing information not uncovered during step 3,
and opinions on prioritization of the proposed corridors or stepping stones. Online webinar
formats should follow a similar flow, but with online mapping resources provided for
stakeholders to leave georeferenced comments. Platforms that can support public
participatory GIS (PPGIS) such as ArcGIS Experience, the Google Maps interface, and the
R package ‘PPGISr’ allow stakeholders to draw points or polygon features on the map and
add comments as attribute information for the feature. In addition, online PPGIS platforms
allow participants to easily toggle between geospatial layers such as species distribution
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models or alternative connectivity scenarios, which may give the stakeholders a better
understanding of the workflow that arrived at the proposed connectivity map. A combination of
in-person and online meetings may give the most equitable coverage of stakeholders and
concerned citizens as some people may not be able to attend in-person meetings but can
leave comments remotely on a PPGIS interface.
Members of the collaborative team should synthesize all the comments and determine those
recurring or major issues that should be addressed in the next round of the connectivity
analysis. This may mean returning to previous steps in the connectivity design framework if,
for example, there were missed landscape threats or if objectives need to be altered. Once
connectivity model results are finalized, either at this round or after further consultation with
stakeholders, then there can be the final determinations of management actions and
prioritisation of corridors and stepping stones.
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elements. Thus, they are seen as a backbone of green-blue infrastructures (e.g., de la Fuente
et al., 2018). More pragmatic approaches have also been suggested by stakeholders to
prioritise restoration efforts, namely selecting the areas where restoration efforts can be more
easily implemented for technical and/or political reasons. This, however, may have the
disadvantage of not targeting the most cost-effective areas.
In intensive agricultural areas, connectivity can be increased through the introduction of the
so-called landscape features, small fragments of non-productive natural or semi-natural
vegetation including hedges, ponds, ditches, non-productive trees, field margins, terrace
walls, dry-stone or earth walls, or fallow land. These are expected to contribute to the target
of the EU Biodiversity Strategy for 2030, of achieving 10% of cover by landscape features in
the agricultural area of the EU, and will benefit a diversity of taxa including amphibians, reptiles
and farmland birds. The maintenance or uptake of low-intensity traditional agricultural
practices, which maintain a significant amount of natural or semi-natural habitats, can also be
seen as a kind of restoration effort contributing to the creation of ecological corridors breaking
the intensive agricultural landscape matrix.
The actual implementation of the now designed connectivity network will likely be the hardest
part of the process. This is where investing the time from the beginning to involve and have
buy-in from community and professional stakeholders will especially start to pay off. Compared
to traditional protected areas (PA), the corridors and stepping stones that may make a
connectivity network may not be able to achieve any level of legal protection. Therefore, these
networks can become a patchwork of PA designations, other effective area-based
conservation measures or area agreements. Therefore, having a coordinated governance
structure including members of municipal or regional councils (or equivalent governing
bodies), local environmental organizations, regional planners, and other environmental
decision-makers will assist greatly in slowly building up the protection and regulations of the
network to meet project objectives and ensure connectivity into perpetuity (McGuinn et al.
2017, WWF 2020).
During and after the network is established it will be vital to monitor and evaluate the
effectiveness of the connectivity conservation actions. Monitoring ecological corridors or
stepping stones typically involves remote sensing, field sampling, and genetic analysis,
combined with different forms of ecological modelling. Remote sensing data can be used to
monitor changes in land cover and land use patterns in and around ecological corridors and
stepping stones. For instance, it can be used to track the effects of human activities, such as
deforestation or urbanisation, on the structural and functional integrity of corridors (e.g.,
Cisneros-Araujo et al., 2021).
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Field sampling methods such as transect surveys for plants and animal sign (e.g., pellets),
camera trapping, and passive acoustic sensors can be used to in conjunction with monitor
dynamics of different species. These data in conjunction with abundance, occupancy, and
species distribution models allow researchers to more accurately estimate the seasonal or
yearly dynamics of population and habitat use. GPS and radio tracking can facilitate targeted
studies on wildlife movement, for example, to examine the effectiveness of constructed
crossing structures (i.e., green bridges). These methodologies thus allow researchers to
evaluate different ecological indicators to determine if corridors are meeting their initial project
objectives and targets.
Evaluating the effectiveness of crossing structures of roads or railways is especially prudent
given the high cost invested in their construction and the enormous potential benefit,
particularly for mammal populations (Soanes et al., 2024). Evaluating the effectiveness of such
mitigation measures must include the use of benchmarks, which can include the use of
control sites and before data (Rytwinski et al., 2015), as well as comparing unmitigated and
“no construction” sites (Soanes et al., 2013). The choice of appropriate benchmarks should
also consider that control sites should have the same qualities as mitigation sites (e.g.,
regarding habitat quality or movement paths) to ensure comparability between them (Abbott
et al., 2012). If the crossing structures are not being used effectively management actions can
be implemented such as increased fencing to better funnel individuals across the structure,
changes to the ecosystem planted on the structure, and mitigation of human activity on or near
the site.
Genetic analysis involves the use of molecular techniques to study the genetic diversity and
gene flow of species across the total landscape. By analysing DNA samples from individuals
across different habitat patches, genetic analysis can provide insights into the connectivity and
genetic exchange between populations. This approach helps identify potential barriers to gene
flow and assess the long-term effectiveness of a connectivity network (e.g., Proctor et al.,
2005). I can also assist in the generation of resistance surfaces for future connectivity model
iterations.
Integrating the data from remote sensing, field sampling, genetic analysis, and ecological
modelling, researchers and conservation practitioners can gain a better understanding of how
connectivity and protected area network is operating and its impact on biodiversity
conservation. This thus informs the adaptive management of the network allowing for new and
likely improved connectivity models.
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2. Acronym
3. Responsible institution(s) *
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6. E-mail *
Note: This information will not be public
8. Project website
9. Abstract
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LIFE programme
Cohesion funds
Biodiversity offsets
Private funds
Other:
Small landscape features (e.g. hedgerows and tree lines, riparian forest
remnants, green urban spaces, etc)
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Other:
14. In case the project is focused on selected taxa, please indicate which.
16. Regardless of who commissioned the project, who are the target users of the
results?
National administration
European institutions
National
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Pan-European
Other:
18. Will the project be used for supporting any of the following *
policies?
Climate policies
Sustainable development
Human health
Other:
Ecosystem restoration
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Other:
Alpine
Anatolian
Arctic
Atlantic
Black sea
Boreal
Continental
Macaronesia
Mediterranean
Pannonian
Steppic
Forests
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Shrublands
Grasslands
Wetlands
Coastal areas
Rivers
Agricultural areas
Analyses using genetic data and/or models (e.g., genetic diversity, gene flow, etc.)
Analyses using data and/or models for population size and demography
Expert-based
Other:
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None
Other:
24. Does (or will) the project monitor the effectiveness of * connectivity
conservation for biodiversity or ecosystem services (yes/no)? If yes, please specify
which.
Water quality
Pollination
Recreation
Soil quality
None
Other:
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Genetic homogenization
None
Other:
27. Do you know of any other connectivity projects that we may be interested in?
Please add the name of project and contact information.
S1.2 Structure
The survey was composed of 27 questions covering project information, scope, users and
selected approaches. The survey featured 15 open-ended questions and 12 multiple-choice
questions which also included an open-response field for additional options.
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ecological connectivity projects and encouraging recipients to share the survey with
colleagues involved in similar projects. To expand our reach, we also leveraged
NaturaConnect's social media platforms to promote the survey. This outreach resulted in 57
additional projects being submitted through the survey by entities we had not initially
contacted. The response rate, based solely on the projects we had reached out to, was
31.4%.
The survey was conducted using Google Forms (https://forms.gle/7D2kJqU83eHpGYWS9)
from May 2023 to January 2024.
Some answers needed to be processed further to facilitate the subsequent analysis. Several
multiple-choice questions provided respondents with an open field where they could add
additional information that they felt was not covered by the available options. Three team
members reviewed each of these answers and determined if they provided new information
not covered by the available options or if it could be reclassified as one of the existing options.
In the case of “Question 25 - Does the project consider other benefits in addition to biodiversity
conservation?”, we added some answers to a newly created category called “Other (e.g.
Human safety and well-being)”. In “Question 26 - Does the project identify any potential
negative impacts of connectivity?”, we added one answer to a new category called “Other
(e.g., economic costs). Concerning “Question 14 - In case the project is focused on selected
taxa, please indicate which” we reclassified the answers in group categories (e.g., large
carnivores, ungulates etc). When we found answers that were out of the scope of the question,
we added them to a new category called “Answer out of scope”.
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Figure S2.1: Miro board from the “Terrestrial and Freshwater Habitats” breakout group on day 1 of the workshop.
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Figure S2.2: Miro board from the “Ecosystem Processes & Services” breakout group on day 1 of the workshop.
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Figure S2.3: Miro board from the “Planning & Management of Multifunctional Corridors” breakout group 1 on day 2 of the workshop.
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Figure S2.4: Miro board from the “Human Infrastructure & Land Use Impacts” breakout group on day 2 of the workshop.
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In amphibians and reptiles, we could not define archetypes considering dispersal distances
due to a lack of data. The few estimates available are highly heterogeneous in terms of
summary measures provided (median, maximum, or minimum) and the approach used to
measure the dispersal distances (GPS, radio tracking, or laboratory experiment). Additionally,
while several articles claim to report “dispersal distances”, in most cases these are not
dispersal movements sensu strictu, including migration distances, pond-to-pond movements,
daily movements, movements from hibernacula, or other movements not classified. Therefore,
the representativeness of the measures provided is undermined by the large variance of this
type of data. For instance, the yellow-bellied toad (Bombina variegata) showed a mean
dispersal distance of 444 m (metres), but the longest dispersal distance is estimated at 3141
m (Cayuela et al., 2019). Similarly, the maximum dispersal distance reported for the common
wall lizard (Podarcis muralis) is 1000 m (Popescu et al., 2013), albeit Williams et al. (2021),
modelling the range expansion of this species, estimated a dispersal distance from 5 to 16
metres. In fact, given the lack of data, there are examples in the literature of modelled dispersal
abilities by using a trait-based proxy (e.g., snout to vent length or body size in Alencar and
Quental, 2023), however, the reliability of these derived data is undermined by the lack of
comparative empirical data. For these reasons, we preferred to not consider unreliable or
unrepresentative dispersal estimates in the definition of archetypes of amphibians and reptiles,
focusing only on size, locomotion mode, reproductive traits, and home range size when
available.
For each analysis group, we conducted a principal component analysis (PCA) in order to
reduce the dimensionality of the dataset while preserving most of the initial variance existing
in the dataset. Then, we performed a k-means cluster analysis considering the PCA axes that
cumulatively explained at least 80% of the variance. K-means clustering is one of the simplest
and popular unsupervised machine learning algorithms. The objective of K-means is to group
similar data points together and discover underlying patterns. The number of clusters is chosen
a priori according to the average silhouette method and on expert-based evaluation. With this
analysis, we were able, therefore, to identify homogeneous clusters within each group of
tetrapods (Tables S3.2-S3.9).
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AmphiBIO Amphibians Oliveira, B. F., São-Pedro, V. A., Santos-Barrera, G., Penone, C.,
& Costa, G. C. (2017). AmphiBIO, a global database for
amphibian ecological traits. Scientific data, 4(1), 1-7.
AnAge Amphibians, De Magalhaes, J. P., & Costa, A. J. (2009). A database of
birds, vertebrate longevity records and their relation to other life‐history
mammals, traits. Journal of evolutionary biology, 22(8), 1770-1774.
reptiles
http://genomics.senescence.info/species/
AvianBodySize Birds Lislevand, T., Figuerola, J., & Székely, T. (2007). Avian body
sizes in relation to fecundity, mating system, display behaviour,
and resource sharing: Ecological archives E088‐096. Ecology,
88(6), 1605-1605.
AVONET Birds Tobias, J. A., Sheard, C., Pigot, A. L., Devenish, A. J., Yang, J.,
Sayol, F., ... & Schleuning, M. (2022). AVONET: morphological,
ecological and geographical data for all birds. Ecology Letters,
25(3), 581-597.
Bird_behav Birds Tobias, J. A., & Pigot, A. L. (2019). Integrating behaviour and
ecology into global biodiversity conservation strategies.
Philosophical Transactions of the Royal Society B, 374(1781),
20190012.
COMBINE Mammals Soria, C. D., Pacifici, M., Di Marco, M., Stephen, S. M., &
Rondinini, C. (2021). COMBINE: a coalesced mammal database
of intrinsic and extrinsic traits.
EAmphDB Amphibians Trochet, A., Moulherat, S., Calvez, O., Stevens, V. M., Clobert, J.,
& Schmeller, D. S. (2014). A database of life-history traits of
European amphibians. Biodiversity Data Journal, (2).
EltonTraits Mammals, Wilman, H., Belmaker, J., Simpson, J., de la Rosa, C.,
birds Rivadeneira, M. M., & Jetz, W. (2014). EltonTraits 1.0: Species‐
level foraging attributes of the world's birds and mammals:
Ecological Archives E095‐178. Ecology, 95(7), 2027-2027.
Eubirds Birds Storchová, L., & Hořák, D. (2018). Life‐history characteristics of
European birds. Global Ecology and Biogeography, 27(4), 400-
406.
EuroBaTrait Bats Froidevaux, J. S., Toshkova, N., Barbaro, L., Benítez-López, A.,
Kerbiriou, C., Le Viol, I., ... & Razgour, O. (2023). A species-level
trait dataset of bats in Europe and beyond. Scientific data, 10(1),
253.
HomeRange Mammals Broekman, M. J. E., Hoeks, S., Freriks, R., Langendoen, M. M.,
Runge, K. M., Savenco, E., ... & Tucker, M. A. (2023).
HomeRange: A global database of mammalian home ranges.
Global Ecology and Biogeography, 32(2), 198-205.
https://github.com/SHoeks/HomeRange
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LHT_ERep Reptiles Grimm, A., Ramírez, A. M. P., Moulherat, S., Reynaud, J., &
Henle, K. (2014). Life-history trait database of European reptile
species. Nature Conservation, 9, 45-67.
LizardTraits Reptiles Meiri, S. (2018). Traits of lizards of the world: Variation around a
successful evolutionary design. Global ecology and
biogeography, 27(10), 1168-1172.
MammalDiet Mammals Kissling, W. D., Dalby, L., Fløjgaard, C., Lenoir, J., Sandel, B.,
Sandom, C., ... & Svenning, J. C. (2014). Establishing
macroecological trait datasets: digitalization, extrapolation, and
validation of diet preferences in terrestrial mammals worldwide.
Ecology and Evolution, 4(14), 2913-2930.
panTHERIA Mammals Jones, K. E., Bielby, J., Cardillo, M., Fritz, S. A., O'Dell, J., Orme,
C. D. L., ... & Purvis, A. (2009). PanTHERIA: a species‐level
database of life history, ecology, and geography of extant and
recently extinct mammals. Ecological Archives E090‐184.
Ecology, 90(9), 2648-2648.
https://tinyurl.com/5czp2t33
TetraDENSITY Amphibians, Santini, L., Isaac, N. J., & Ficetola, G. F. (2018). TetraDENSITY:
birds, A database of population density estimates in terrestrial
mammals, vertebrates. Global Ecology and Biogeography, 27(7), 787-791.
reptiles
Table S3.2: Median values of traits used to define the five European non-volant mammals archetypes.
1 2 3 4 5
Table S3.3: Median values of traits used to define the three European bats archetypes.
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1 2 3
Table S3.4: Median values of traits used to define the four European birds archetypes.
1 2 3 4
Table S3.5: Median values of traits used to define the three European frogs archetypes.
1 2 3
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Walker 0 1 0
Jumper 1 1 1
Climber 0 0 1
Swimmer 1 1 0
Table S3.6: Median values of traits used to define the three European salamanders archetypes.
1 2 3
Walker 1 1 0
Climber 0 0 1
Swimmer 0 1 0
Direct development 0 0 1
Larval stages 0 1 0
Viviparous 1 0 0
Table S3.7: Median values of traits used to define the two European turtles archetypes.
1 2
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Table S3.8: Median values of traits used to define the three European snakes archetypes.
1 2 3
Table S3.9: Median values of traits used to define the three European lizards archetypes.
1 2 3
Arboreal 0 0 0
Terrestrial 1 0 1
Saxicolous 0 1 0
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Figure S3.1: Aggregation of IUCN habitat classes (level 1) into natural and artificial classes.
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Copernicus-
European https://land.copernicus.eu/en/produ
CORINE Land Cover Copernicus Environment Agency cts/corine-land-cover
Copernicus- https://land.copernicus.eu/en/produ
High Resolution Layer European cts/high-resolution-layer-water-and-
Land use &
Water and Wetness Copernicus Environment Agency wetness
Land cover
Global Tree Cover Hansen, M.C. et https://glad.umd.edu/dataset/global-
2010 al. Global Forest Watch 2010-tree-cover-30-m
https://www.nature.com/articles/s41
Primary forest Sabatini et al. Nature 597-021-00988-7#Sec7
Open Street Map Open Street Map Open Street Map https://www.openstreetmap.org
Roads &
Linear Copernicus- https://land.copernicus.eu/en/produ
Features European cts/eu-hydro/eu-hydro-river-
EU Hydro Rivernet Copernicus Environment Agency network-database
https://www.eea.europa.eu/en/datah
ub/datahubitem-view/d08852bc-
European Digital 7b5f-4835-a776-
Evalutation Elevation Model (EU- European 08362e2fbf4b?activeAccordion=735
& DEM) EEA Environment Agency 550
Topography
https://spacedata.copernicus.eu/coll
Copernicus Global European Space ections/copernicus-digital-elevation-
DEM ESA Agency model
https://www.eea.europa.eu/data-
Biogeographical European and-maps/figures/biogeographical-
Boundaries regions 2016 EEA Environment Agency regions-in-europe-2
&
EEA Administrative https://sdi.eea.europa.eu/catalogue/
Bioregions
Boundaries based on srv/eng/catalog.search#/metadata/9
GISCO NUTS and European 4438969-2dd5-4ba3-b708-
EBM EEA Environment Agency e4d29a8b7699
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Global Biodiversity
Species
Information Facility
Data
(GBIF) GBIF GBIF https://www.gbif.org/
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NaturaConnect has 22 partner institutions: International Institute for Applied System Analysis (project
lead; Austria); German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig (project
co-lead; Germany); Associação Biopolis (Portugal); BirdLife Europe (Netherlands); Birdlife International
(United Kingdom); Centre National De La Recherche Scientifique (France); Doñana Research Station
- Agencia Estatal Consejo Superior De Investigaciones Cientificas (Spain); Europarc Federation
(Germany); Finnish Environment Institute (Finland); Humboldt-University of Berlin (Germany); Institute
for European Environmental Policy (Belgium); Netherlands Environmental Assessment Agency
(Netherlands); Rewilding Europe (Netherlands); University of Evora (Portugal); University of Helsinki
(Finland); University of Natural Resources and Life Sciences, Vienna (Austria); University of Rome La
Sapienza (Italy); University of Warsaw (Poland); Vrie University of Amsterdam (Netherlands); WWF
Central and Eastern Europe (Austria); WWF Romania and WWF Hungary.
www.naturaconnect.eu
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