0% found this document useful (0 votes)
23 views149 pages

Guidelines Connect Conserv Plann EU

The document outlines the 'Guidelines for connectivity conservation and planning in Europe' as part of the NaturaConnect project, which aims to design a resilient Trans-European Network for Nature and People. It includes a framework for multi-scale connectivity assessment and planning, developed in collaboration with various stakeholders, and emphasizes the importance of ecological connectivity for biodiversity conservation. The guidelines also provide tools and data sources for implementing connectivity projects across Europe.

Uploaded by

RM Miau
Copyright
© © All Rights Reserved
We take content rights seriously. If you suspect this is your content, claim it here.
Available Formats
Download as PDF, TXT or read online on Scribd
0% found this document useful (0 votes)
23 views149 pages

Guidelines Connect Conserv Plann EU

The document outlines the 'Guidelines for connectivity conservation and planning in Europe' as part of the NaturaConnect project, which aims to design a resilient Trans-European Network for Nature and People. It includes a framework for multi-scale connectivity assessment and planning, developed in collaboration with various stakeholders, and emphasizes the importance of ecological connectivity for biodiversity conservation. The guidelines also provide tools and data sources for implementing connectivity projects across Europe.

Uploaded by

RM Miau
Copyright
© © All Rights Reserved
We take content rights seriously. If you suspect this is your content, claim it here.
Available Formats
Download as PDF, TXT or read online on Scribd
You are on page 1/ 149

Project Report

Author-formatted document posted on 06/06/2024

Published in a RIO article collection by decision of the collection editors.

DOI: https://doi.org/10.3897/arphapreprints.e129021

Guidelines for connectivity conservation and planning in


Europe
Francisco Moreira, Filipe S. Dias, Jeremy Dertien, Ana Ceia Hasse, Luis Borda-de-Água, Silvia
Carvalho, Miguel Porto, Francesca Cosentino, Luigi Maiorano, Andrea Sacchi, Luca Santini,
Florian Borgwardt, Georg Gruber, Nikolaj Poulsen, Rafaela Schinegger, Carina Seliger, Néstor
Fernández
Author-formatted document posted on 06/06/2024. DOI: https://doi.org/10.3897/arphapreprints.e129021

Guidelines for connectivity


conservation and planning in Europe
D6.1 Guidelines for connectivity conservation and planning in Europe with supporting web-
based inventory and databases

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.

Prepared under contract from the European Commission.

Project acronym: NaturaConnect


Project full title: NaturaConnect - Designing a resilient and coherent Trans-
European Network for Nature and People

Grant agreement 101060429


number:
Start of the project: 1 July 2022
Duration: 48 months
Project coordinators: International Institute for Applied Systems Analysis (IIASA)
and
Martin-Luther Universität Halle-Wittenberg (MLU)

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.

Front cover: Laarderhoogt Ecoduct, Netherlands. © Getty Images/Artur Debat/WWF-US

2
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

Project reference 101060429


number
Project title NATURACONNECT - DESIGNING A RESILIENT AND
COHERENT TRANS- EUROPEAN NETWORK FOR NATURE
AND PEOPLE

Deliverable title Guidelines for connectivity conservation and planning in Europe


with supporting web-based inventory and databases
Deliverable number D6.1
Contractual date of 31.03.2024
delivery
Actual date of delivery 29.03.2024
Type of deliverable R - Document, Report
Dissemination level PU - Public
Work package number WP6
Institution leading work MLU
package
Task number 6.1
Institution leading task CIBIO
Author(s) Francisco Moreira, Filipe S. Dias, Jeremy Dertien, Ana Ceia-Hasse,
Luís Borda-de-Água, Silvia Carvalho, Miguel Porto, Francesca
Cosentino, Luigi Maiorano, Andrea Sacchi, Luca Santini, Florian
Borgwardt, Georg Gruber, Nikolaj Poulsen, Rafaela Schinegger,
Carina Seliger, Néstor Fernández
EC project officer Christophe Coudun

3
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.

4
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

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

Part I: Connectivity in Europe: Key concepts, policy context, and


implementation ................................................................................... 21
2. Connectivity concepts and approaches ................................................. 22
2.1 Protected areas and ecological corridors ........................................................................... 22
2.2 Structural and functional connectivity ................................................................................ 23
2.3 Connectivity in the context of Green and Blue Infrastructure .......................................... 26
2.4 Spatial scale issues and dispersal ...................................................................................... 27
2.5 Corridors and stepping stone design.................................................................................. 29
2.6 Freshwater and cross-realm connectivity........................................................................... 32
2.6.1 Four dimensions of connectivity in rivers ..................................................................................... 33

2.6.2 Connectivity between freshwater and other realms ..................................................................... 35

2.6.3 Tools and approaches to assess connectivity across realms ...................................................... 36


2.7 Integration of connectivity in the process of area-based planning ................................. 36
2.8 Caveats of corridor design ................................................................................................... 38
2.9 Do ecological corridors work? ............................................................................................. 40

3. Global and EU policy instruments addressing connectivity................. 41


3.1 Connectivity in the post-2020 Global Biodiversity Framework ........................................ 41
3.2. Connectivity in the EU Biodiversity Strategy .................................................................... 41
3.3 Connectivity in the EU Forest Strategy ............................................................................... 42
3.4 Connectivity in the Green and Blue infrastructure strategy ............................................. 42
3.5 Connectivity in the Water Framework Directive ................................................................. 43
3.6 Connectivity in the EU Pollinators Initiative ....................................................................... 43
5
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

4. Connectivity projects in Europe and information needs ....................... 44


4.1 Survey of connectivity projects in Europe.......................................................................... 44
4.1.1 Project information, location and duration.................................................................................... 45

4.1.2 Goals and scope .......................................................................................................................... 45

4.1.3 Taxa and ecosystems .................................................................................................................. 46

4.1.4 Policy context, target users and funding ...................................................................................... 47

4.1.5 Spatial scope ................................................................................................................................ 48

4.1.6 Selected approaches and outputs ............................................................................................... 49

4.1.7 Assessing project effectiveness ................................................................................................... 50

4.1.8 Potential negative effects ............................................................................................................. 50


4.2 Priorities, gaps, and challenges in European connectivity planning .............................. 51
4.2.1 Stakeholders’ priorities for connectivity planning ......................................................................... 54

4.2.1.1 Long-term ecological resilience .................................................................................... 54

4.2.1.2 Connecting across realms and patch sizes .................................................................. 55

4.2.1.3 On the health and wellbeing of humans........................................................................ 55

4.2.1.4 Freshwater and coastal areas ....................................................................................... 55

4.2.1.5 Policies and actions that support enhanced connectivity ............................................. 55

4.2.2 Technical challenges for connectivity planning ............................................................................ 56

4.2.2.1 Data gaps for implementing connectivity projects ........................................................ 56

4.2.2.2 Critical information gaps................................................................................................ 57

4.2.2.3 Technology and capacity constraints ............................................................................ 58

4.2.3 Solutions to overcome challenges and needs ............................................................................ 59

4.2.3.1 Repositories for data and capacity building resources ................................................. 59

4.2.3.2 Collaboration and engagement at the forefront ............................................................ 59

4.2.3.3 Policies, regulations, and funding streams ................................................................... 59

4.2.3.4 Planning for global change ........................................................................................... 60

Part II: Tools and guidelines for implementation of connectivity projects


in Europe ............................................................................................ 61
5. Tools and data sources for modelling connectivity .............................. 62
5.1 Introduction ............................................................................................................................ 62
5.2 Least-cost path and Resistant Kernels ............................................................................... 65
5.3 Graph Theory.......................................................................................................................... 68
6
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

5.4 Circuit Theory ......................................................................................................................... 70


5.5 Agent-based models.............................................................................................................. 74
5.6 Structural Connectivity Metrics and moving window-analysis ........................................ 77
5.7 Assessing ecosystem services ............................................................................................ 80

6. A framework for connectivity conservation and planning .................... 82


6.1 Introduction to the framework .............................................................................................. 82
6.2 Scoping and problem assessment ...................................................................................... 83
6.3 Setting of Objectives ............................................................................................................. 86
6.3.1 Focal & archetype species for assessing connectivity in Europe ................................................ 87

6.3.2 Corridor width ............................................................................................................................... 90

6.3.3 Final Spatial Extent and Resolution ............................................................................................. 92


6.4 Analysis Selection & Data Preparation ............................................................................... 94
6.5 Assessment of Connectivity................................................................................................. 97
6.5.1 Prioritisation and restoration for connectivity objectives .............................................................. 98
6.6 Implementation, Monitoring & Evaluation......................................................................... 100

7. References ............................................................................................... 103


Annex S1. Survey of connectivity projects .............................................. 119
S1.1 Questions ........................................................................................................................... 119
S1.2 Structure ............................................................................................................................. 127
S1.3 Distribution and response rates ...................................................................................... 127
S1.4 Response processing ....................................................................................................... 128

Annex S2. Connectivity workshop Miro boards examples ..................... 129


Annex S3. Archetypes ................................................................................ 137
S3.1 Archetypes definition ........................................................................................................ 137
S3.2 Habitat preferences ........................................................................................................... 144

Annex S4: Geospatial Data Sources for Europe ...................................... 146

7
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

8
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

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

9
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 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

10
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

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

11
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

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).

12
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

• 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)

13
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

Figure ES1: Response rates for taxonomic groups and ecosystem types for the survey on ecological connectivity
projects in Europe

o A companion online database presenting all 80 projects submitted to the online


survey is publicly available online at https://naturaconnect.idiv.de/projects/. Its
aim is to serve as a dynamic resource for researchers, policymakers,
conservationists, and the public interested in connectivity conservation projects
conducted in Europe.
• Stakeholder priorities for future multi-scale connectivity planning and
implementation across Europe are wide ranging and target multiple aspects and
stages of connectivity planning (Chapter 4). Main priorities identified by
stakeholders during a two-day NaturaConnect follow up webinar included:
o Identification of climate and evolutionary refugia
o Setting of stepping stones for long distance migrants (mainly birds)
o Promoting connecting across realms
o A focus on human well-being in the planning of multi-functional corridors,
mainly in urban and peri-urban contexts
o A focus on rivers as backbones of connectivity planning
o Setting of policies and incentives targeting the promotion of connectivity, mainly
on private land
o Dam removal and the restoration of natural rivers
o Increase the permeability of linear infrastructures as roads and railway
• Connectivity planning and implementation challenges identified by stakeholder
included those on gaps in data availability, lack of streamlined guidance on
connectivity planning and implementation, and lack of opportunities for
technical training (Chapter 4). Main challenges identified by stakeholders during a
two-day NaturaConnect follow up webinar included:

14
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

o Gaps in data, mainly in what concerns species movement and dispersal


o Species requirements during migration
o The incorporation of management information in current maps of land cover
o Lack of socio-economic data to better frame people’s perceptions of corridors
and connectivity
o Lack of guidance and rules needed for future connectivity modelling and on-
the-ground implementation
o Land ownership issues
o How to assign economic value to connectivity
o Lack of a multi-level governance structure for the planning and management of
connectivity networks
o Lack of technical training opportunities
• To adequately plan for and implement connectivity measures, it is often required
to use models to estimate connectivity across regions. A portfolio of publicly
accessible tools and data sources are already available to help practitioners with this
and are summarised in this report (Chapter 5).
• The need exists for a dedicated framework for connectivity conservation and
planning in Europe, which presents clear steps to consider when designing a
connectivity project (Fig. ES2; Chapter 6). The NaturaConnect connectivity network
design framework presented in this report aims to address this need and outlines a set
of five key steps for practitioners:
(1) Scoping and Problem Assessment: Conduct a comprehensive analysis of the
entire landscape to identify potential threats, connectivity actions, and impact of
those actions, identify all relevant stakeholders and build an interdisciplinary
collaboration team for connectivity analysis, communication, and implementation.
Establish the general spatial extent at which your study will take place.
(2) Setting of Objectives: Use the assessment of the connectivity problem to
establish spatial and temporally explicit objectives and targets that will help mitigate
the identified problem. Determine the appropriate width and characteristics of
corridors and stepping stones based on the target species and landscape
characteristics. Finalize the spatial extent and needed data resolution.
(3) Analysis Selection and Data Preparation: Determine the correct model or
models to analyse ecological connectivity. Given the model and your objectives
collect and produce all the necessary data and spatial layers necessary to run the
spatial analysis.
(4) Assessment of connectivity: Use connectivity metrics and models to determine
the most effective design for a connectivity network that integrates with the current
network of protected areas. Present draft results to stakeholders, iterate new
models, and prioritise corridors and stepping stones; and
(5) Implementation, Monitoring and Evaluation: Develop a comprehensive
management and monitoring plan for the ecological corridor and/or stepping
stones. This includes activities such as habitat restoration, invasive species

15
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

control, monitoring of species movement, and assessing the corridor's


effectiveness in achieving the connectivity objectives.

Figure ES2: Building blocks of the proposed framework for implementing ecological connectivity projects

16
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

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
17
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

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.

1.2 Aims and target audience


This document aims to provide base knowledge and guidance for planning ecological
connectivity conservation and restoration. It is designed to address the specific needs,
opportunities, and challenges for connectivity planning, with a specific focus on supporting the
implementation of policy commitments for biodiversity conservation and restoration. The
report provides a thorough review of available methods, tools and data sources for
connectivity planning, and proposes recommendations for designing connectivity projects
supporting a coherent and resilient Trans-European Network of Protected Areas
(TEN-N).
18
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

We also present an analysis of the characteristics of connectivity projects in Europe based on


an extensive survey, and the identification of data and information gaps and needs for
supporting connectivity assessments, conservation, and restoration. These analyses included
the input of multiple stakeholders through online surveys and workshops.
The IUCN (International Union for Conservation of Nature) “Guidelines for conserving
connectivity through ecological networks and corridors” (Hilty et al., 2020) introduced a set of
recommendations for building ecological networks and for implementing ecological
connectivity between protected areas and other conservation areas. They showcase different
approaches for conserving ecological corridors by presenting case studies aimed at protecting
or restoring ecological connectivity from around the world and in terrestrial, freshwater and
marine ecosystems. The present report complements these guidelines and other existing
literature by providing a thorough review of approaches, the information needs as identified
by stakeholders, and practical recommendations.
This document is primarily intended for analysts, practitioners, and scientists involved in the
design and management of nature conservation and restoration projects, e.g., from public
National and regional administrations, environmental planners and managers within and
outside protected areas, as well as private initiatives, foundations, etc. interested in
connectivity planning. It is tailored to meet the needs of those responsible for developing and
implementing strategies, policies, and management plans.
The document is focused on the terrestrial realm, although some aspects of freshwater
connectivity are also addressed. The marine realm is only partly tackled, in the context of
cross-realm connectivity (Section 2.6).

1.3 Summary of the content


Part I: Connectivity in Europe: Key concepts, policy context, and implementation
Chapter 2 discusses connectivity concepts and approaches. We contrast the different
objectives of protected areas versus ecological corridors, that do not necessarily need to
provide key habitats for in situ conservation. Then the distinction between structural
connectivity (focusing on land use patterns) and functional connectivity (focusing on species
traits, including dispersal movements) is made. Connectivity is also presented in the context
of Green and Blue Infrastructure (GBI), based on multi-functionality to incorporate both
biodiversity and ecosystem services targets. The multi-functional perspective of GBI spans
many policy sectors and raises some challenges to connectivity design, namely through
defining the relative weight of the biodiversity versus ecosystem services objectives. Spatial
scale and dispersal issues are also addressed. In fact, projects addressing the restoration and
protection of connectivity may be implemented at various spatial scales, ranging from local to
regional or even continental, depending on the specific objectives and the species being
considered. The scale of analyses must be linked to the ecological traits of the species being
considered, particularly on dispersal capacity. The key principles for the design of corridors
and stepping stones is also presented, as well as the integration of connectivity in spatial
conservation prioritization. This Chapter also includes a Section on freshwater and cross-
realm connectivity.

19
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

Chapter 3 focuses on policy instruments addressing connectivity. An overview is made


of how connectivity is tackled in seven major policy instruments, 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 Law, the Water
Framework Directive, and the EU pollinators initiative.
Chapter 4 focuses on current connectivity projects and practices in Europe. It presents
the results from a survey conducted to gather information on ecological connectivity projects
in Europe carried out at regional, national, and Pan-European levels. The survey consisted of
27 questions covering project information, scope, participants, and selected approaches. The
survey was conducted between May 2023 and January 2024, and gathered information on 80
projects conducted in 35 European countries, regarding: goals and scope; targeted taxa and
ecosystems; policy context, target users and funding; spatial scope; selected approaches and
outputs; assessing effectiveness; and potential negative effects. The companion online
connectivity projects database, containing all the projects included in the survey, is also
presented.
Chapter 4 also includes a synthesis of identified stakeholder priorities and challenges for
future multi-scale connectivity planning across Europe, identified during a two-day online
NaturaConnect workshop in October 2023 on “Assessing Ecological Connectivity in Europe:
Conservation goals and information gaps” which gathered ~ 90 experts and stakeholders.
Part II: Tools and guidelines for implementation of connectivity projects in Europe
Chapter 5 provides a comprehensive review of available tools and data sources for
modelling connectivity. Here, we provide an overview of existing software and applications
based on three major types of model families: least-cost path and resistant kernels, graph
theory and circuit theory. The use of agent-based models is also addressed, as well as the
assessment of ecosystem services in the case of multi-functional connectivity projects.
Chapter 6 proposes a framework for connectivity conservation and planning, where the steps
to consider when designing a connectivity project are presented.

20
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

Part I: Connectivity in
Europe: Key concepts,
policy context, and
implementation

21
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

2. Connectivity concepts and approaches


The term ecological connectivity has been defined in several ways, each of them with
different implications for the management applications of the concept and the associated
analytical and mapping approaches. In this Section, we give an overview of concepts
underpinning different perspectives on connectivity used in nature conservation and
assessments.

2.1 Protected areas and ecological corridors


Protected areas have long been the foundation of nature conservation strategies worldwide.
Between 2010 and 2023 the global coverage of terrestrial protected areas (including inland
waters) grew from 14.1% to 17.19%. In the European Union, the percentage of protected
areas has been on a similar trajectory, with the Natura 2000 network currently covering 18.6%
of the land area (Fig. 2.1).

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.

22
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

However, the current network of protected areas alone is insufficient to safeguard


biodiversity. First, the location of historically designated protected areas is often biased
towards least productive and least threatened areas, leaving several types of ecosystems
under-represented (Joppa and Pfaff, 2009). Second, climate change can alter temperature
and precipitation patterns, and cause changes in the distribution of ecosystems, habitats and
species. Many species will require shifting their distribution ranges to more suitable climatic
conditions to persist, and therefore the current distribution of protected areas may not be
sufficient to protect these species (Dobrowski et al., 2021). Third, many protected areas are
too small (Monaco and Genovesi, 2014) and they are insufficently connected to be effective
(Santini et al., 2016; Ward et al., 2020). According to recent estimates, only 9.7% of
Earth’s terrestrial protected network can be considered structurally connected (Ward et
al. 2020). In the European Union's Natura 2000 network, connectivity estimates seem to be
more optimistic. Approximately 80% of Natura 2000 sites dominated by woodland and forest
are considered connected by natural and semi-natural features in the wider landscape
(outside of the Natura 2000 sites) across the 27 EU Member States. Among these sites, over
50% are linked by continuous patches of unprotected forest and woodland ecosystems
(Carrao et al., 2020).
Creating ecological corridors between protected areas is of the utmost importance. Corridors
work as a pathway for the movement of plants, animals, and other organisms, allowing them
to migrate, disperse, and interact. Ecological corridors are usually defined as strips of natural
habitat that connect two or more areas of similar habitat that are surrounded by a nonhabitat
matrix (Beier and Noss, 1998) and that are managed over the long term to maintain or restore
connectivity (Hilty et al., 2019). Corridors may include landscape features such as hedgerows,
tree lines, riparian strips, or managed agricultural land, offering a blend of open space and
more sheltered passageways for species (Travers et al., 2021). However, corridors do not
need to be narrow strips of habitat. They can also consist of a combination of natural and
semi-natural habitats able to increase the permeability to dispersal and other movements of
organisms (Eggers et al., 2010; Travers et al., 2021). This is particularly relevant in highly
transformed landscapes, where protected areas often include heterogeneous mixtures of
different ecosystem types under different management regimes and intensity. In Europe,
many protected areas are surrounded by a matrix of intensive land use with low natural
connectivity potential (EEA, 2024).
The main purpose of ecological corridors is to maintain connectivity (Hilty et al., 2020).
They may also contribute to conserve in situ biodiversity but this is not a strict requirement. In
contrast, the main objective of protected areas is in situ conservation, although they may
conserve connectivity as well. Each corridor should be designed with specific objectives in
mind and be managed accordingly. Regardless of the types of areas that compose them,
ecological corridors should always be distinguishable from non-designated areas based on
the specific activities that are permitted or prohibited within them (Hilty et al., 2019, 2020).

2.2 Structural and functional connectivity


The concept of ecological connectivity encompasses both structural connectivity and
functional connectivity (Hilty et al., 2019; Taylor et al., 2006).

23
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

Structural connectivity focuses on the physical arrangement and configuration of


habitat patches and corridors. It primarily considers the spatial contiguity of species
habitats, ecosystem types, etc., and how this contiguity is affected by fragmenting features.
When applied to specific species, structural connectivity captures the spatial arrangement and
configuration of habitat patches within a landscape, hence, it depends on the size, shape, and
location of habitat patches (Fahrig, 2003). Different species perceive the landscape differently
(in terms of seeing distinct land covers as resources) and possess distinct habitat
requirements, movement behaviours, dispersal propensity and distances. Therefore, a given
level of structural connectivity is likely to translate into different levels of functional connectivity
for different species (Taylor et al., 2006). Structural species connectivity can be seen as a
simplified alternative (compared to functional approaches) that focuses solely on the spatial
arrangement and configuration of species habitats within a landscape, where one ignores, on
a first approach, the details of individual species biology. Examples include classifying
different land cover types in term of suitable habitats for species movement of (e.g. forests,
wetlands, extensive agriculture) or reducing these movements (e.g. urban areas,
infrastructures, intensive agriculture). Assessments based exclusively on structural landscape
analyses can provide a cost-effective means of assessing connectivity for conservation
purposes, although at the cost of ecological detail (Saura et al., 2011). Structural connectivity
approaches can also integrate multiple ecological features when assessing the spatial
contiguity of focal patches, such as land use types, species composition, and human
pressures, therefore providing more holistic perspectives on the ecological connectivity of
natural habitats (e.g., Fernández et al. 2020).

©OlaJenner/WWF Sweden

24
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

Functional connectivity, on the other


hand, emphasises the demographic,
genetic, and community processes
affected by dispersal and movement.
Functional connectivity quantifies the
(potential) movement of genes,
gametes, propagules or individuals
move through landscapes, and their
ecological function. Naturally, the two
concepts are interconnected, as
functional connectivity often depends on
the presence and configuration of
structurally connected habitat patches.
The functional connectivity of a
© Andreas Beckmann/WWF landscape for a particular species (or a
group of species) is assumed to be
determined by both movement potential through different habitat patches and local
subpopulation dynamics including demographic effects (Doerr et al., 2011). Assessments of
the functional connectivity of populations inform on the effects of dispersal across the
landscape on the distribution, abundance, dynamics, and genetics of populations
(Fernández et al. 2016; Bruggeman et al. 2010).
Functional connectivity is therefore specific for a given species or community assemblage
occurring in a given region. In fact, species are predicted to change their behaviour and
experience variations in their fitness based on the type, shape, and spatial arrangement of
habitat patches and ecotones (Bélisle, 2005). The pattern of movements of a given species
can be strongly influenced by biological characteristics such as sex, age, and individual
behaviour (Maiorano et al., 2017).
Functional connectivity has been often measured at the level of a single species. However,
the idea of finding functional corridors for multiple species is certainly appealing. Two possible
approaches are commonly considered:
1. the use of “connectivity umbrella” or indicator species: a species that has a large
body size, home range, charisma and conservation status, and habitat requirements
that overlap with other species. These are typically large and mobile, with large
requirements of space and resources. The postulate is that a landscape that ensures
connectivity for an umbrella species can potentially serve also other (smaller or less
mobile) organisms (Dutta et al., 2023). However, the umbrella species approach is
limited by the ecological similarity and the level of interactions with co-occurring
species (Natsukawa & Sergio 2022).
2. Consider multiple species explicitly, measuring functional connectivity for all of them
and obtaining integrated or consensus connectivity maps. This should allow for the
identification of critical areas for facilitating the movement of multiple focal species
accounting for their specific ecological requirements, such as for multiple threatened
species, community guilds, or particular interacting species.

25
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

Theoretically, multispecies strategies should provide positive conservation outcomes more


effectively than single-species strategies. However, they have a potential downside for single
species spatial planning (Brodie et al., 2015). Several studies have explored and compared
the two options (e.g., Meurant et al., 2018; Wang et al., 2018) highlighting how the use of a
multi-species approach could be more effective in the development of conservation actions,
such as the creation of corridors, because it allows for the protection of a wide range of
different habitat types and also to meet the different spatial and structural requirements of the
several species considered (Cushman and Landguth, 2012).
There is still a third approach, based on identifying archetypal species. Archetypical species
are stylized “virtual species” obtained by the combination of species with similar
ecological and physiological traits. Theoretically, if properly selected, an archetypal
species may be representative of groups of species that are expected to have fairly similar
characteristics in the context of connectivity, sharing similar spatial requirements and habitat
selection and perception.

2.3 Connectivity in the context of Green and Blue Infrastructure


In 2013, the European Commission (EC) adopted a strategy to develop green and blue
infrastructure (GBI) in the EU (see Section 3.4) in the scope of the Action Plan for Nature,
People and the Economy. GBI is broadly defined as a network of natural and semi-natural
areas, together with other environmental features designed to deliver a wide range of
ecosystem services to people, while enhancing biodiversity. GBI, with its green (terrestrial)
and blue (if aquatic systems are concerned) components, is a promising approach for land-
use planning (Houet et al., 2022) as it is multi-functional to incorporate both biodiversity
(although not related to specific targets or species) and ecosystem services targets.
Connectivity is one of the principles of GBI design, along with multi-functionality and spatial
planning (Estreguil et al., 2019). The social and ecological benefits of GBI depend to a large
degree on connectivity (Benedict and MacMahon, 2002; Ignatieva et al., 2011; Petrisor et al.,
2021).
The multi-functional perspective of GBI spans many policy sectors and raises some
challenges to connectivity design, namely through defining the relative weight of the
biodiversity versus ecosystem services objectives. As an example, when designing
connectivity corridors, the fact that vegetation is native or exotic may make no difference for
some specific ecosystem services (e.g., carbon sequestration), but it might have strong
implications for biodiversity potential as non-native vegetation may not serve as suitable
habitat for many species. However, the prevalent view is that there should be a
complementary perspective assuming that GBI is composed of biodiversity-rich areas
that also provide multiple ecosystem services to people (Estreguil et al., 2019). In any
case, the involvement of stakeholders from different sectors is crucial to determining priorities,
costs, and benefits for GBI, and consequently how to approach connectivity to deliver benefits
for specific species but also to society.

26
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

Approaches for using GBI in spatial


planning are based on two components:
(1) the mapping of existing GBI
components (protected areas and
ecological networks), focusing on the
identification and physical delineation of
landscape features consisting of green
and blue elements; and (2) an ecosystem
service-based mapping, based on
assessing the capacity of the current land
cover to provide ecosystem services. In
contrast to the physical mapping
approach, which refers to the delineation
of physical landscape elements, the
ecosystem
© Bárbara Pais
service-based mapping
approach further adds a function to the
physical element by expressing the
ecosystem services they deliver (Estreguil et al., 2019), for example the potential for increased
pollination supply due to existing hedgerows.

2.4 Spatial scale issues and dispersal


Projects addressing the restoration and protection of connectivity may be implemented
at various spatial scales, ranging from local to regional or even continental, depending
on the specific objectives and the species being considered. For example, migratory birds
may be extremely philopatric (return to the same locations) in their breeding ranges while
migrating for thousands of kilometres (from Europe to south Saharan Africa) twice a year (e.g.,
the dunlin Calidris alpina has a natal dispersal distance below 5 km and a migratory distance
which can be over 4,000 km, with peaks of 1,200 km per day during migration. On the opposite
extreme, saproxylic beetles (e.g., the hermit beetle Osmoderma eremita) never move from
their natal tree (Ranius 2006) and cave salamanders (e.g., Speleomantes strinatii) do not
move more than 20 metres, being sedentary throughout their entire life. Large carnivores are
somewhat in the middle, although with variability, even at the species level. For example,
female bears are extremely philopatric (with females often taking over part of their mother’s
home range) while male bears make relatively long dispersal movements (Maiorano et al.,
2017). Migratory fish species, particularly susceptible to fragmentation, include endangered
medium-distance migrants (e.g., Acipenser ruthenus, Hucho hucho) and large-distance
migratory species (e.g., Acipenser stellatus, Huso huso) which became extinct in the upper
Danube catchment as a consequence of the closure of the Iron Gate dams (Jungwirthh et al.,
2003).
These differences must be considered in any management and/or conservation planning with
each scale of approach having different requirements and challenges. Both habitat selection
and connectivity are very sensitive to the spatial (and temporal) scale of analysis (Ashrafzadeh
et al., 2020). An incorrect scale can result in incorrect inferences that may lead to inefficient
conservation actions. The optimal scale for planning and managing habitat connectivity
27
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

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
28
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

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).

2.5 Corridors and stepping stone design


Corridors and stepping stones are the two main landscape designs targeting the conservation
or restoration of ecological connectivity. Corridors are more traditionally utilised in
connectivity conservation as they are continuous linear connections between two or more
habitat patches. Stepping-stones are a network of smaller habitat patches between large
29
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

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
30
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

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

1. Proximity and Alignment: Stepping stones should be strategically placed to minimise


the distance between them and larger habitat areas. They should form a series of rest stops
that align with the natural movement patterns of species (e.g. North-South migrations)
(Bennet, 2003, Saura et al., 2014).
2. Habitat Quality and Diversity: Each stepping stone should offer a variety of
microhabitats and resources (such as food, water, and shelter) to cater to the needs of
different species. The quality of these habitats is just as important as their presence (Bennet,
2003)
3. Landscape Context: The surrounding landscape should be considered in the design of
stepping stones. This includes understanding the matrix of agricultural, urban, and natural
areas to optimise the placement and composition of stepping stones for the target species
(Bennet, 2003, Hilty et al., 2006).
4. Size and Shape: While stepping stones are inherently smaller than core habitats, their
size and shape should be optimised for the species of interest. Larger stepping stones can
support more species and provide more extended stays, while the shape can influence the
edge effects and the internal microclimate of the habitat (Saura et al., 2014).
5. Target Species Needs: The specific needs of target species or groups of species should
guide the design of stepping stones. This includes considerations of the species' dispersal
abilities, habitat preferences, and threats they face (Bennet, 2003).
6. Monitoring and Management: Once established, stepping stones require monitoring
and management to ensure they continue to serve their intended function. This might
include measures to control invasive species, maintain habitat structures, and adapt to
changing conditions or new scientific knowledge (Bennet, 2003, Hilty et al., 2006, Saura et
al., 2014).

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
31
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

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.

2.6 Freshwater and cross-realm connectivity


Connectivity analyses of terrestrial habitats have often included rivers as two-dimensional
elements but neglected their internal structure and heterogeneity that in turn represent a
‘riverscape’ (Wiens, 2002). Although knowledge and approaches from terrestrial assessments
can also be transferred to aquatic ecosystems, rivers exhibit certain characteristics, which
should grant them a special position in connectivity conservation e.g., (1) rivers
represent both a habitat and migration corridor (Ward, 1989; Wiens, 2002), (2) connectivity
acts on four (one temporal and three spatial) dimensions, whereby the importance of each
dimension changes along the river course (Vannote et al., 1980; Ward and Stanford, 1995),
(3) hydrologic connectivity allows the passive downstream transport of matter and energy
(Pringle, 2006; Ward and Stanford, 1995), but enables a multidimensional dispersal of
organisms (Branco et al., 2014; Ward and Stanford, 1995), and (4) while terrestrial analyses
often focus on the connectivity of specific habitat types (e.g. grasslands or forests), the
connection of different habitats may be more important in the freshwater realm where certain
species and life stages require diverse habitat patches to complete their life cycle, e.g.
sturgeons of salmons (Jungwirth et al., 2003).
Freshwater ecosystems host remarkable biodiversity and provide substantial ecosystem
services such as flood retention, water purification or recreation (Hanna et al., 2018; Kaval,
2019; Tickner et al., 2020). However, the decline of freshwater species populations is
happening globally faster than declines in marine and terrestrial realms (Reid et al., 2019).
This decline results from a set of threats, including fragmentation, and others such as flow
regulation, pollution, habitat loss and overexploitation of biological resources, and invasive
species (Dudgeon, 2019; Haase et al., 2023; Strayer and Dudgeon, 2010).

32
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

©Wild Wonders of Europe/Diego Lopez/WWF

Connectivity plays a key role in the conservation, restoration and management of


freshwater ecosystems because of their connected nature, which facilitates the
maintenance of ecological processes (including the movement of freshwater organisms, the
hydrological transport of energy, solutes, pollutants and sediments), but also allows for the
propagation of threats (such as pollution and invasive species) (Linke et al., 2011; Pringle,
2003; van Rees et al., 2021). However, human pressures constrain freshwater connectivity
worldwide, particularly river fragmentation, water abstraction and flow regulation,
sedimentation, water consumption, and urbanisation (Grill et al., 2019).
Due to limited migration opportunities, fragmentation is particularly damaging in stream
networks, since it is more difficult or even impossible for fish to avoid disconnections (Fagan,
2002). Even though weirs represent the most obvious way of fragmentation in riverine
habitats, dams may also be associated with other hydromorphological changes, which then
alter the spatial and temporal patch composition and, consequently, connectivity patterns
(Wiens, 2002). In this context, residual flow sections and impoundments may not only alter
habitats but can also contribute to habitat fragmentation by preventing fish migrations through
sections with limited water depths and flow velocities (Schmutz and Sendzimir, 2018).

2.6.1 Four dimensions of connectivity in rivers


Ward (1989) defined freshwater ecosystem connectivity across four dimensions: longitudinal,
lateral, vertical, and temporal. Firstly, the longitudinal dimension refers to the connection
between upstream and downstream regions, which facilitates the life cycle of migratory
species and species dispersal (Vannote et al., 1980). Freshwater longitudinal connectivity is
distinct from general terrestrial connectivity because it has a strong directional component
(Moilanen et al., 2008). Some studies focused on maximising linear connectivity throughout
the river catchment. For example, Moilanen et al. (2008) suggested measuring freshwater
connectivity by considering its hierarchical network topology and creating functions to describe
upstream and downstream connectivity, reflecting the water flow. These functions can be
33
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

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.

34
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

© Ante Gugić/WWF

2.6.2 Connectivity between freshwater and other realms


In addition to addressing the four dimensions of connectivity in rivers, there has been
increased attention to considering connectivity among terrestrial, freshwater and
marine ecosystems (Adams et al., 2014; Alvarez-Romero et al., 2011; Álvarez-Romero et
al., 2015; Beger et al., 2010a). The interactions between these ecosystems are necessary for
species persistence and for the maintenance of ecosystem services through the flows of
energy, materials and organisms (Giakoumi et al., 2019; Hermoso et al., 2021a; Soininen et
al., 2015). Hermoso et al. (2021b). However, there are several approaches to assess
connectivity: Tsang et al. (2019) measured the connectivity between inland and marine
habitats as a function of distance. Leontiou et al. (2022) measured connectivity for species
that breed inland but forage on the sea as the inverse distance between these areas. Tulloch
et al. (2021) used models of land-sea runoff and ocean dispersal to estimate threats to the
marine ecosystem and Devlin et al. (2012) identified inshore ecosystems exposed to surface
plume pollutants.
However, an increase in connectivity within and between realms may foster the propagation
of threats (see Section 6.7), such as the dispersion of pollutants, the spread of invasive
species, and exposure to diseases (Adams et al., 2014; Alvarez-Romero et al., 2011; Tulloch
et al., 2021). Identifying, measuring and actively managing these connections is vital, but such
practice is complex, given the broad spectrum of connectivity concepts, particularly when
focusing on realms and accounting for spatial-temporal dynamics (Beger et al., 2022). These
complexities require integrative planning that assesses the co-benefits and trade-offs of
various links to make cross-realm connectivity more informative and to highlight advantages
and disadvantages (Álvarez-Romero et al., 2015).

35
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

2.6.3 Tools and approaches to assess connectivity across realms


Although connections between the different realms may be difficult to measure, some
tools are available to examine proxies for connectivity patterns from large to short spatial
and temporal scales (Beger et al., 2022). A common approach is to identify connections
between important conservation areas in the different realms. For instance, Naia et al. (2021)
measured connectivity between desert water pools (gueltas) and their upstream contributing
sub-catchments based Euclidean distances between them. This connectivity can be very
important to relict populations which use gueltas as refugia during dry periods and disperse
during the wet season, such as the Crocodylus niloticus in Mauritania (Brito et al., 2011; Vale
et al., 2015). For large spatial scales, remotely sensed metrics such as the Normalised
Difference Vegetation Index (NDVI) provide proxies for carbon, nitrogen and phosphorus
eventually entering the freshwater ecosystems (Dahlin et al., 2021; Soininen et al., 2015). This
information can be used to approximate biodiversity-ecosystem functioning linkages,
particularly interactions, feedback, and synergies between terrestrial and freshwater
ecosystems, and across trophic levels. Remote sensing can also aid in assessing types of
vegetation that function as freshwater corridors, such as riparian vegetation and floodplain
seasonality (Reis et al., 2019). Riparian corridors provide and regulate a wide range of
ecosystem services (Atkinson and Lake, 2020), including contributing to functional
connectivity. For instance, they enable the connection of forest fragments widespread over
the landscapes and allow movement and dispersal of several species (Keten et al., 2020;
Larsen-Gray and Loehle, 2022). Remote sensing techniques can also track the propagation
of some pollutants (Gholizadeh et al., 2016), which is important to assess the potential impacts
on biodiversity patterns and processes to which they are connected through water flow
(Álvarez-Romero et al., 2015).
Investigating aquatic animal movements has become widespread and popular with the use of
radio or acoustic telemetry as well as Passive Integrated Transponder (PIT) tags (Burnett et
al., 2021). This technology can aid in assessing connectivity directly for anadromous and
catadromous fishes based on their migration pathways including the assessment of the
effectiveness of fish passages that aim to restore functional connectivity for fish. Furthermore,
genetic tools and landscape genetics have also been used to assess connectivity indirectly,
as barriers and fragmentation affect gene flows between populations (Keller et al., 2015). This
can be important to species with terrestrial and aquatic habitat requirements such as the
Eurasian otter (Leoncini et al., 2023), or for wetland-breeding amphibians (Gutiérrez-
Rodríguez et al., 2017; Watts et al., 2015).

2.7 Integration of connectivity in the process of area-based planning


Spatial conservation prioritisation is a process concerned with the identification of priority
areas accounting for quantitative or qualitative conservation targets to achieve specified
benefits for biodiversity, while allowing to account for social, economic and political constraints
(Margules and Pressey, 2000; Moilanen et al., 2009). Conservation prioritisation can optimize
and integrate multiple conservation features - e.g. ecosystems, species, populations, genetic
lineages, multiple facets of biodiversity or targeting multiple objectives (Carvalho et al., 2017;
Jung et al., 2021; Margules and Sarkar, 2007; Pollock et al., 2017), over space and time.
Connectivity can be an important aspect in deciding where to allocate future
36
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

conservation efforts and the inclusion of connectivity in conservation prioritisations


can bring advantages in the long term. However, it can also facilitate the propagation of
threats, such as diseases, pollution, invasive species and fires.
One common approach for incorporating connectivity in the prioritization process is
that more connected areas should have a higher priority, e.g. are more likely to be
selected as candidates for expanding a network of sites. However, benefits for species
will likely rely on a set of trade-offs considering size, shape and habitat quality of sites, and
connectivity between sites. Previous research shows that spatial priorities differ significantly
when connectivity is not integrated into the conservation optimization (Hanson et al., 2022;
Makino et al., 2013), with conventional methods approximating connectivity often fail reflect
functional connectivity that supports greater gene flow between sites (Hanson et al. 2019).
There has been a proliferation of mathematical and analytical methods to solve complex
decision-making problems in various aspects of conservation efforts, from heuristic algorithms
to more sophisticated optimization methods using operation research (Alagador and Cerdeira,
2022; Pressey, 2002). These include identifying priority conservation areas space (Allan et
al., 2022; Montesino Pouzols et al., 2014; Pollock et al., 2017; Ward et al., 2020), optimising
surveys and monitoring networks (Carvalho et al., 2016; Hanson et al., 2023), and allocating
management actions (Adams et al., 2014; Cattarino et al., 2015).
Different algorithms and tools have been developed to support decision-making in spatial
conservation prioritisation optimization, including Marxan (Ball and Possingham, 2000),
Marxan Connect (Daigle et al., 2020), Zonation (Moilanen et al., 2014), and the prioritizr R
package (Hanson et al., 2019). These tools vary in their algorithms, optimization approaches,
flexibility in objectives and constraints, and output types. For instance, Marxan uses a
minimum-set approach to heuristically identify near-optimal sets of planning units that achieve
a set of conservation targets at a near-minimal cost, while Zonation uses a Maximum coverage
approach, aiming at ranking gridded planning units for their biodiversity benefits and
considering a specified cost. Prioritizr, on the other hand, allows the use of multiple problem
formulations (including both minimum set, minimum shortfall and maximum coverage
formulations) and relies on exact algorithm that guarantees optimality, e.g. the areas selected
are the best possible given supplied data (Beyer et al., 2016). All these algorithms enable the
consideration of connectivity between chosen priority conservation areas. However, different
tools vary in the connectivity concepts used and respective methodologies.
In mathematical terms, connectivity can be integrated in the prioritization problem as a goal or
as a constraint (Daigle et al., 2020). Traditionally, the consideration of connectivity in spatial
planning focused on preferring compact and contiguous configurations. To achieve larger
conservation areas and a less fragmented network, connectivity is used as a constraint in the
objective function, which penalises solutions with a higher total length of edges between
selected areas, thus preferring areas in the ranking that are more compact and contiguous
rather than isolated in space. The data required for this procedure is a relation table specifying
the shared edges between each planning unit and its direct neighbours. However, connectivity
concepts go beyond achieving network compactness and, in many cases, it may require an
extended relational table specifying a connectivity measure between each pair of planning
units. This relation matrix can be obtained with different methods, for instance, using
Euclidean distances, least cost paths, resistance matrices attained with circuit theory,
biophysical models, etc. (see Chapter 4). This is the case, for example, when the goal is
37
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

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).

2.8 Caveats of corridor design


Corridors design must also consider possible drawbacks which, under some
circumstances, might entail unexpected or negative consequences for biodiversity
conservation.
Ecological corridors are often planned with long and narrow shapes, which create extensive
boundaries. Edge effects along these boundaries can negatively influence the corridor
effectiveness. Furthermore, abrupt transitions between corridors and surrounding habitats
may alter the microclimatic conditions (Bernaschini et al., 2019; Hofmeister et al., 2019;
Laurance et al., 2002) and the distribution and behaviour of species within the corridor.
Research also indicates that edge effects can cause certain species to perceive corridors as
unfavourable habitats or ecological traps and avoid them. Ecological corridors may also
increase predation rates, as they create a narrow pathway that predators can exploit. By
connecting populations or habitats that were previously isolated, corridors may also increase
the likelihood of disease dispersal and facilitate the spread of invasive species (Resasco et
al., 2014). Similarly, increased connectivity might induce gene flow interfering with population
distinctiveness between formerly isolated populations.

38
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

Ecological corridors can


also entail conservation
conflicts. The
consequences of
species dispersal
through corridors include
potential damages to
agriculture, e.g.,
ungulate species can
cause damage to crops
during raids. Corridor
use by large carnivores
can also result in attacks
to livestock (Hilty et al.,
2012).

Wild Wonders of Europe. © Cornelia Doerr/WWF

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.

39
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

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

2.9 Do ecological corridors work?


Ecological corridors play a crucial role in conserving biodiversity. However, their
implementation can be costly and may have significant economic and social implications.
Therefore, it is essential to assess their effectiveness in promoting connectivity between
habitats.
The effectiveness of ecological corridors has been assessed in two large meta-analyses. In
2010, Gilbert-Norton et al. (2010) conducted a meta-analysis of 78 experiments from 35
studies conducted between 1985 and 2008 to investigate the impact of corridors on increasing
movement between habitat patches. The results showed that corridors had a significant effect
on movement, increasing it by approximately 50% compared to patches that were not
connected with corridors. The study also found that corridors were particularly important for
the movement of invertebrates, non-avian vertebrates, and plants, rather than birds. These
findings suggest that corridors can be an effective strategy for enhancing movement
between habitat patches, which in turn can help maintain and restore populations of
plants and animals in fragmented landscapes. In 2019, Resasco (2019) conducted a meta-
analysis involving 32 additional studies conducted between 2008 and 2018, utilizing the
selection criteria established by Gilbert-Norton et al. (2010), and arrived at comparable
conclusions.
Overall, the effectiveness of ecological corridors is well founded on scientific evidence.
However, the effectiveness of corridors will vary depending on factors including the species
involved, the characteristics and quality of the corridor, the landscape context, and the
pressures.

40
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

3. Global and EU policy instruments


addressing connectivity
Connectivity is addressed in different policy instruments related to biodiversity conservation,
with different approaches and targets. A short overview is made below for the key biodiversity
policy instruments in the EU.

3.1 Connectivity in the post-2020 Global Biodiversity Framework


In the post-2020 Global Biodiversity Framework (GBF) (CBD, 2022a), the overall aim of
maintaining, enhancing or restoring natural ecosystem connectivity is mentioned in its first
goal (Goal A). Global targets for 2030 also include connectivity objectives in the scope of
restoration of degraded areas (target 2), implementing a well-connected system of
protected areas (target 3), and improving ecological connectivity in green and blue spaces
in urban and densely populated areas (target 12). Proposed headline connectivity
indicators for the GBF (CBD, 2022b) include a Protected Area Connectedness index, a
dendritic connectivity index, and a Convention on Migratory Species (CMS) connectivity index.
These are expected to be discussed and operationalized by an ad hoc technical expert group
on indicators, with a time-bound mandate until the sixteenth meeting of the Conference of the
Parties.

3.2 Connectivity in the EU Biodiversity Strategy


The EU Biodiversity Strategy for 2030 (EC, 2020) includes the desing of a coherent Trans-
European Nature Network including ecological corridors “to prevent genetic isolation,
allow for species migration, and maintain and enhance healthy ecosystems”. In this
context, the protection and restoration of ecological corridors, investments in green and blue
infrastructure, and cooperation between Member States across borders is promoted.
Preserving and restoring habitat connectivity is also key for bringing back at least 10% of
farmland under high-diversity landscape features (including e.g. buffer strips hedges,
terrace walls, fallow land). Within the scope of urban greening plans, connections between
green spaces should also be promoted. Lastly, the Strategy sets out a pledge to plant at least
three billion additional trees by 2030 in full respect of ecological principles. The respective
Commission staff working document (EC, 2021b) acknowledges the need to ensure
connectivity benefits of afforestation at the landscape level, including in forests,
agricultural landscapes and in actions related to the impact of infrastructures on habitat
fragmentation. Such actions must combine large-scale conservation and restoration planning
with the creation of new connecting infrastructures, i.e., from the creation of corridor habitats
to strategically placed green bridges and tunnels for wildlife movement.
The goals and targets of the Nature Restoration Regulation proposed by the European
Commission is central to the implementation of the EU Biodiversity Strategy (EC, 2022). The
regulation places special regard to the connectivity between the habitats of species listed
under the Habitats Directive and the Birds Directive. It also requires improving the connectivity
41
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

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.

3.3 Connectivity in the EU Forest Strategy


In the EU Forest Strategy for 2030 (EC, 2021a) connectivity is not a central topic. However,
there is a mention of establishing ecological corridors in agricultural areas, as an
expected outcome of reforestation or afforestation of biodiverse forests.

3.4 Connectivity in the Green and Blue infrastructure strategy


In parallel to the strictly biodiversity-focused approach, the EC drafted a strategy to develop
green and blue infrastructure (GBI) in the EU (EC, 2013, 2019), in the scope of the Action
Plan for Nature, People and the Economy. Here, green infrastructure (GI) is broadly
defined as a network of natural and semi-natural areas, together with other
environmental features designed to deliver a wide range of ecosystem services. The
blue component (BI) includes freshwater and marine realms. The EC encourages the
preservation, restoration and enhancement of green infrastructure to halt the loss of
biodiversity and enable the provision of ecosystem services. So, although GI is assumed as
a tool for biodiversity objectives (Natura 2000 is assumed as the backbone of the EU GI), it
includes a strong component of ecosystem service delivery objectives (it is assumed as
“services-oriented”), therefore aiming for ecological, economic and social benefits through
natural solutions (assumed as more sustainable than conventional civil engineering solutions,
the so-called “grey infrastructure”). GI has a strong focus on urban settings and includes a
wide range of objectives including climate change, health and disaster risk management.
Green and blue infrastructure is expected to positively contribute to the sustainability of
several EU policies, including regional development, social cohesion, agriculture, transport,
energy production and transmission, disaster risk management, fisheries and maritime
policies. Because of the multifunctional perspective of GBI, projects may have very
different scopes, including increasing landscape permeability for wildlife through better road
and railway planning (e.g., TRANSGREEN project), large-scale corridors for large mammals,
using power lines rights-of-way as ecological corridors, river and dune restoration, green
infrastructure in cities, or setting regional networks of protected areas.
The EC drafted a guidance document focused on supporting tools and instruments to support
investment, as well as good practices for GBI (EC, 2019). EU-level GI should cumulatively

42
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

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).

3.5 Connectivity in the Water Framework Directive


The EU Water Framework Directive (WFD) targets connectivity indirectly by recognizing the
interconnected nature of water bodies and emphasizing the need to maintain and enhance
connectivity within and between them. The directive aims to achieve "good ecological status"
for all European water bodies and acknowledges that the ecological health and functioning of
aquatic ecosystems depend on the free movement of water, sediment, and biota across
different habitats. The WFD identifies threats to connectivity, such as dams, weirs, and
other physical obstacles in its River Basin Management Plans (RBMPs). For at least 20%
of EU river water bodies, barriers are considered a significant pressure under the WFD,
contributing to the non-achievement of a good ecological status (EEA, 2021) and a decline of
93% in migratory freshwater fish in Europe since 1970 (Deinet et al., 2020). Consequently,
Member States are encouraged to adopt measures that promote connectivity and
integrated water management approaches. However, no direct measures to improve
connectivity are foreseen in the EU Water Framework Directive.

3.6 Connectivity in the EU Pollinators Initiative


Connectivity is addressed in the EU Pollinators Initiative (EC, 2023) through the promotion
of well-connected, high-quality habitats for pollinators. These ecological corridors
(named “buzz lines”) are expected to enable species movement in search of food, shelter, and
nesting and breeding sites, as well as acting as migration routes for species impacted by
climate change. Connectivity for pollinators is particularly important in farmland, but also in
urban areas, with the expectation that GI expansion can result in benefits for pollinators in
those environments as well. By 2027, the Commission and Member States should devise a
blueprint for a network of ecological corridors for pollinators and develop a plan of
measures for implementing it.

43
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

4. Connectivity projects in Europe and


information needs
4.1 Survey of connectivity projects in Europe
To obtain a comprehensive, pan-European snapshot of connectivity initiatives taking place
across Europe, an online survey was developed and disseminated, gathering information on
ecological connectivity projects being implemented by actors at regional, national, and pan-
European levels. The breadth of projects submitted included public and private conservation
action plans and strategies, connectivity conservation projects, restoration projects, and
research and innovation projects. The survey consisted of 27 questions covering project
information, scope, participants, and selected approaches (see Annex S1.1). The survey was
conducted between May 2023 and January 2024, using Google Forms and distributed via
email and social media to project stakeholders and members of the wider conservation
community and public. For more information on survey design, dissemination and response
processing please see Annex S1.2.
This Section presents key results and findings from the survey, which help to build a more
complete picture of ecological connectivity efforts across Europe. The insights derived from
this exercise contributed to developing the framework for connectivity conservation and
planning outlined in Section 6.
A companion online database has also been developed for public use, which contains all the
projects included in the survey (80 projects). Its aim is to serve as a dynamic resource for
researchers, policymakers, conservationists and members of the public interested in
connectivity conservation projects conducted in Europe. To utilize this database, users can

44
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

navigate through a user-friendly interface to access detailed information on a project, including


its scope, participants, and methodologies. The results are systematically organized to enable
intuitive exploration and comparison of projects at regional, national, and Pan-European
levels. The database is accessible at https://naturaconnect.idiv.de/projects/:

4.1.1 Project information, location and duration


The final project list comprised 80 projects conducted in 35 European countries, along with
Russia. The top five countries where connectivity projects were most frequently reported were
Spain, France, Italy, Germany, and Austria (Fig. 4.1). Most (~75%) of the projects were
completed within two to six years.

Connectivity projects online database - Connectivity projects online database –


homepage project page

Figure 4.1: Number of projects per country and reported project duration (inset, top left).

4.1.2 Goals and scope


In reference to the connectivity goals of projects, the three most frequently stated goals were
“Connectivity between protected areas", "Connectivity between specific habitat types", and
"Protection of multiple species", which represented 72% of all responses (Fig. 4.2).

45
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

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

4.1.3 Taxa and ecosystems


Respondents were asked to identify the taxonomic groups that their project focused on. The
results showed that large carnivores make up 30% of the reported taxa, while arthropods and
birds make up 19% and 13%, respectively. In terms of ecosystem types, forests account for
20% of the responses, grasslands 17% and agricultural areas 15% (Fig. 4.3).

46
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

Figure 4.3: Response rates for taxonomic groups and ecosystem types.

4.1.4 Policy context, target users and funding


To evaluate the policy context of the projects, respondents were asked to indicate if their
project was commissioned by an administration. We found that this was the case for only 38%
of projects (Fig. 4.4).
In terms of the target users of the projects, "Regional and/or local administration(s)" and
"National administration" accounted for 50% of reported target users. Regarding the policies
that the projects aim to support, the most common responses were "Biodiversity conservation
policy and strategies", "Spatial planning of protected areas", and "Green and Blue
Infrastructure policies", which together accounted for 64% of the replies.
Finally, in relation to projects’ funding sources, “Nature conservation funds from National
and/or Regional administrations", “European funds associated with sustainability policies" and
“Private funds" were identified as funding sources most frequently, accounting for 58% of
answers.

47
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

Figure 4.4: Response frequencies for the questions concerning policy context, target users and funding sources.

4.1.5 Spatial scope


With regards to the spatial scope of projects, “Sub-national (spatially comprehensive for one
or several administrative regions)", “Transboundary (connecting across 2 or more countries)"
and “Local (e.g., covering one or several municipalities or a specific infrastructure)"
accounted for 73% of responses (Fig. 4.5).
For the biogeographical region where projects took place, the continental region was reported
most frequently and accounted for 23% of the reported regions, while the alpine and
Mediterranean regions accounted for 20% and 18%, respectively.

48
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

Figure 4.5: Response frequencies for questions related to projects’ spatial scope and the biogeographical regions
where they took place.

4.1.6 Selected approaches and outputs


For approaches used in projects for assessing connectivity, “Land cover and land use
analyses", “Expert-based" and “Analysis of infrastructures (e.g., roads and railway) and urban
sprawl" were the most frequent answers, accounting for 55% of the reported approaches.
Regarding the kinds of spatially explicit information projects produced, “Locations for
ecological corridors" and “Locations for stepping stones" accounted for 49% of the responses
(Fig. 4.6).

Figure 4.6: Response frequencies for questions concerning selected approaches and what kind of spatially explicit
information projects produced.

49
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

4.1.7 Assessing project effectiveness


Respondents were asked to determine if their projects included connectivity monitoring. Only
28% of projects reported implementing monitoring (Fig. 4.7), which suggests two possibilities.
The first is that many projects did not propose testable approaches for enhancing connectivity.
The second is that they did not consider or had insufficient funding for conducting monitoring.

Figure 4.7: Number of projects that did and did not implement monitoring.

4.1.8 Potential negative effects


Regarding any potential negative impacts associated with increased connectivity, the most
common response, accounting for 41% of the answers, was “None" (Fig. 4.8). The answers
“Increased human-wildlife conflicts” and “Increased spread of invasive species” accounted for
17% and 14% of the responses, respectively.

50
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

Figure 4.8: Response frequencies for potential negative effects caused by increasing ecological connectivity.

4.2 Priorities, gaps, and challenges in European connectivity planning


Maintaining and enhancing connectivity requires careful design, implementation, monitoring
and the involvement of different stakeholder groups, including experts and practitioners from
multiple sectors. Following the past and current state of European connectivity projects
through the online survey, further insights on stakeholder priorities and challenges for future
multi-scale connectivity planning across the continent were gathered via an online workshop.
Overcoming gaps and challenges is critical for meeting the long-term success of a resilient
ecological network across Europe for nature and people, and the online workshop served a
crucial role in collecting stakeholders about the difficulties they face with connectivity
planning, and potential actions that can help with remediating those challenges.
The two-day online workshop was organised in October 2023 and titled “Assessing
Ecological Connectivity in Europe: Conservation goals and information gaps”. It gathered ~70
experts and stakeholders across the two days, from multiple sectors including EU/EC
institutions, national and sub-national governmental administration and authorities, non-
governmental organisations (NGOs), the private sector, and research institutes and
universities (Fig. 4.9).

51
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

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

52
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

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”

Enhancing Connectivity for Endangered Species &


Ecosystem Processes & Services
Habitats

Protected Areas & Natura2000 Human Infrastructure & Land Use Impacts

Species Conservation Planning & Management of Multifunctional Corridors

Terrestrial & Freshwater Habitat Socio-Cultural Barriers & Opportunities

Urban & Peri-urban Biodiversity

53
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

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.

4.2.1 Stakeholders’ priorities for connectivity planning


4.2.1.1 Long-term ecological resilience
Overarching all the priorities identified by stakeholders across Europe who participated in the
online workshop, was the need for ecological connectivity to maintain ecosystem resilience
into the future. Given the impacts of fragmentation on the landscape and ongoing climate
change, the increased and efficient planning of connections for species and ecosystem
services is of paramount concern. Multi-functional areas with high redundancies in species
communities and in ecosystem services and multi-scale planning to incorporate different
species needs at different scales were specific needs repeatedly highlighted. So was the need
for stepping stones (in addition to corridors), especially as stopover sites for migratory species
such as waterbirds.
A high level of future uncertainty also requires the identification of probable climate and
evolutionary refugia in multiple locations to create redundancies in the system. This will aid in
the relative short-term for resilience of species sub-populations and for the maintenance of
extant populations of the species in the long-term.
54
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

4.2.1.2 Connecting across realms and patch sizes


Connectivity planning across major ecological realms (e.g., terrestrial and freshwater) has
been historically less of a focus in connectivity planning, however, stakeholders view it as very
important for creating a truly holistic network of protected areas. Determining how to best
create connections across ecosystem types as well as including smaller potentially suboptimal
patches within the highly fragmented European landscape was seen as a high priority. There
was some disagreement on whether to prioritise connections of the big intact habitats or a
broader system of connections between a mixture of larger “source” patches and smaller
potentially “sink” patches. This extended to the need for creating connections for ecosystem
services especially between patches of “surplus” services to those with increased “demand”
or deficit for those services.

4.2.1.3 On the health and wellbeing of humans


Increased ecological connectivity can have multiple benefits for human health and wellbeing.
Stakeholders included considerations of human wellbeing as one of the many facets in the
planning of multi-functional corridors with a focus on recreation, prioritising those ecosystem
services that ultimately benefit humans, such as provisioning services that promote clean air
and water, carbon sequestration, aid for organic farming, etc. Such multi-functional corridors
can benefit the local economy through recreation and ecotourism and can preserve cultural
services within the region, maintaining or building a ‘sense-of-place’ for the residents. A better
sense-of-place then has the potential to aid in the maintenance of conservation initiatives
within that area. These principles are particularly relevant in urban and peri-urban settings,
where connectivity should deliver ecosystem services in addition to bringing biodiversity into
the city (rather than aiming to have major ecological corridors crossing urban areas). Peri-
urban agricultural areas and nature parks are particularly important to link to important
biodiversity (protected) areas surrounding cities.

4.2.1.4 Freshwater and coastal areas


Freshwater and coastal ecosystems connectivity is often overlooked in many connectivity
studies, but is key for the conservation of biodiversity, the movement of nutrients, and for a
plethora of other ecosystem services. Existing knowledge within freshwater and coastal
systems is sparse compared to larger terrestrial systems. Connectivity planning that is
inclusive of riverscapes and in coastal systems was often noted as a priority concern for
stakeholders as they have seen little research and guidance concerning the connectivity
management of these ecosystems. Rivers, in particular, were considered as backbones of
connectivity planning in several contexts, recognising their critical role as linear habitats.

4.2.1.5 Policies and actions that support enhanced connectivity


There were a wide assortment of policies and actions suggested as priorities for connectivity,
from the local to continental scale. Promoting the creation of more natural and artificial
connections on public and private lands, increased funding, and a common EU to local level
policy framework were frequently cited as overarching priorities for connectivity conservation.
Policies that assist landowners/managers to maintain connectivity through their property, such
as forest grants and agricultural subsidies for farmers to maintain small-scale landscape
55
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

elements, should be supported and expanded upon, according to workshop participants. In


addition, legal actions are needed to expand the size of current protected areas and assign
Other Effective area-based Conservation Measures (OECMs) to corridors that may not
receive traditional protected area status. Additionally, some comments focused on the need
to increase legal protection for specific habitat types and features such as creating effective
measures to protect riparian galleries or incentives to protect unique wetland pockets that
could act as stopover locations for migrating waterbirds.
Management actions should target mainly the local level, where they could potentially have
the greatest impact. Actions such as the creation of fish ladders, dam removal and the
restoration of natural rivers, and remodelling existing bridges to include a parallel “green strip”,
were some of the ideas given as priority actions. Other priorities included temporal actions
that reduce landscape resistance and wildlife mortality, such as the promotion of “dark
passages” for bat movement and halting wind turbines during bird migrations. The need to
increase permeability of linear infrastructures (roads and railway) featured as an
acknowledged priority.

4.2.2 Technical challenges for connectivity planning


4.2.2.1 Data gaps for implementing connectivity projects
Data availability gaps were a definitive challenge identified by most participants in the
workshop. Data needs varied from the fine-scale species data to land cover and
socioeconomics. The availability of species data was particularly a prominent concern (Fig.
4.11) with movement and dispersal data, for all taxa, appearing to be the most in-demand
given its utility in models to identify corridors and stepping-stones. There is also the need for
information on species requirements during migrations and the magnitude of impact from
different anthropogenic barriers. In addition, the stakeholders identified population
demography, abundance, habitat use and the relationship between habitat suitability and
demographics as important data needs.
Without accurate land-use / land-cover (LULC) data, projections of potential movement
pathways and the spatial configuration of connectivity networks could be wildly inaccurate.
Having accurate LULC data that are continually updated at fine resolution (<100m) can assist
with reducing error in predicted relationships with all species and processes of connectivity
concern. Historical land cover photos and rasters would also assist in understanding current
land dynamics and, perhaps more importantly, could be used to parameterize LULC models
and contribute to better predict future land system configurations under human global change.
Finally, data on current habitat management information such as timber management and
sales, would be in the next level of land use detail beyond just categorical information on the
land class. Incorporating finer land management detail would add a needed temporal
component to predictions and assist in large landscapes that appear homogeneous from
satellite derived LULC layers but are in fact temporally dynamic in their structure (e.g., forests
of central and northern Finland).
Another major data gap is spatial and non-spatial social and socio-economic data. Items such
as peoples’ perceptions of environmental management actions, values or socio-economic
interests are lacking according to workshop participants. Information on perceptions and

56
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

knowledge of ecosystem services and valuation of protected areas or for recreation


opportunities would assist in planning multi-functional corridors.

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.

4.2.2.2 Critical information gaps


Compared to data gaps, information gaps included those guidance and rules needed for future
connectivity modelling and on-the-ground implementation. Primarily, this included aspects of
the correct parameters of corridor configuration and overall connectivity network design.
Among these parameters, minimum corridor width is often discussed especially in areas of
high human development. While there is no one simple answer, participants noted the need
for clear guidance on how much land needs to be protected to enable free movement of wildlife
through a corridor, or the maximum distance between connectivity stepping-stones (see
Section 6.5). There is also the need for better guidance on what situations warrant the use of
57
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

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.

4.2.2.3 Technology and capacity constraints


Connectivity analyses rely heavily on geographic information systems (GIS) to capture
landscape dynamics. However, the resolution of the data (i.e., grain size) may not be fine
enough to realistically capture many of the ecological dynamics of the system, if at all.
Therefore, many participants in the workshop noted that there is a mismatch between the
grain size and smaller scale dynamics which may lead to the wrong predictions in modelling
species preferences and likely movement patterns, amongst other conclusions. In addition,
there is still difficulty and lack of guidance, in the opinion of many of the participants, on the
complexities of integrating social, economic, and political factors into spatial planning. While
some frameworks have been introduced (Ogletree et al. 2019), some data is difficult to
represent in a spatial manner and even, when possible, how do you relate that to its impacts
on the ecological dynamics?
Another major challenge is the need to simultaneously account for horizontal (i.e., across
space) and vertical (i.e., through time) connectivity. This challenge is due in part to the static
nature of most GIS data, the difficulties of integrating the management of species with different
life histories, and the complexities of creating multi-functional corridors with multiple, often
competing, goals. Furthermore, these difficulties will likely only increase with the predicted
volatility in climatic patterns and the increase in extreme stochastic weather events.
Many workshop participants also focused on the multitude of other capacity deficiencies that
can hinder effective connectivity planning. Planning and management of connectivity networks
require the organisation of a multi-level governance structure which is, first, difficult to
construct and second, difficult to maintain given the perceived lack of funding and time for its
maintenance. There is also often the lack of capacity to implement a participatory spatial
planning process, given deficiencies in funding, staff time, and other resources necessary for
proper communication with multiple landowners. Built into this is the need for better
coordination between public and private organisations to increase engagement across user
groups and regions. Increased technical training opportunities are also required.

58
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

4.2.3 Solutions to overcome challenges and needs


4.2.3.1 Repositories for data and capacity building resources
Building upon many of the challenges identified by participants, there were also many
potential solutions. Participants saw better availability of relevant data and capacity building
resources as a paramount solution to several of the challenges. A more centralised repository
of the different data types identified in Section 4.2.2 could allow for more efficient and
accurate connectivity modelling and planning. In addition, well documented case studies and
methodologies that can be used as examples for further planning and management as well
as a harmonisation of methods across scales would promote and streamline other
connectivity planning projects. Capacity building resources should include technical training
for different connectivity tools, training materials for the facilitation of stakeholder
engagements, and education materials aimed at teaching the general public about the
importance of connectivity for the conservation of nature.

4.2.3.2 Collaboration and engagement at the forefront


Connectivity planning is inherently a multidisciplinary endeavour; early, continual
collaboration and engagement across public and private stakeholders was seen as one of
the most important aspects of a successful connectivity planning exercise. Engaging with
decision makers could help “mainstream” the importance of connectivity measures and
cement it as a factor in local and regional spatial planning. Continual engagement with NGOs
and community stakeholders will assist in identifying the multitude of goals from different
community groups and assist in garnering support for the implementation of the project's final
recommendations.

4.2.3.3 Policies, regulations, and funding streams


As noted in the priorities above, a unified set of policies and regulations that can be
implemented from the EU to the local level would be a major step in improving and promoting
connectivity planning projects. Incentives for private landowners to increase permeability on
their lands and regulations that can reduce wildlife mortality are both general solutions that
can have large landscape impacts. There should also be a focus on establishing basic
requirements for ecological corridor protection including expanded use of legal land
protections, including OECMs, for corridors and stepping stones. Other important factors
highlighted by participants included providing resources for connectivity planning best
practices, exhibiting the importance of connectivity planning to policymakers, and finally
lobbying for higher funding availability from multiple levels of government agencies.
One of the key findings from the survey of connectivity projects (see Section 4.1) is that only
38% of the projects were commissioned by an administration, suggesting that promoting
ecological connectivity has not been a priority for government agencies. Another significant
finding was that only 26.8% of the projects reported conducting any monitoring or evaluation
of effectiveness. This suggests two possibilities. The first is that many projects did not propose
testable approaches for enhancing connectivity. The second is that they did not consider or
had insufficient funding for conducting monitoring.
59
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

4.2.3.4 Planning for global change


Further connectivity planning needs to be inclusive of shifting landscapes due to global
change. A focus on the temporal aspects of connectivity is increasingly important given the
increasing changes to climatic and habitat patterns altering nutrient cycling and species
distributions. Identifying habitat sensitivities to these changes, then projecting species
distributions through time will assist in the accuracy of connectivity planning outcomes. Going
further, including adaptive planning that contains multifunctionality and corridor or stepping
stone redundancies can fortify a network against increasing unexpected weather events. One
must be careful though when considering public opinion, as some workshop participants
noted, to balance both reactive approaches such as those to decreasing species abundance
or improving water connectivity and proactive approaches such as rewildling corridors for
species that do not currently occupy an area. Finally, the workshop participants came back
to the need for environmental education for key stakeholders (landowners, city councils,
agricultural associations, etc.) to explain how climate change will alter historical patterns and
why the need to fortify the connections across a fragmented landscape is now more important
than ever given an increasingly dynamic system.

60
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

Part II: Tools and guidelines


for implementation of
connectivity projects in
Europe

61
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

5. Tools and data sources for modelling


connectivity
5.1 Introduction
Connectivity analyses are being used for a broad array of subjects from water conservation to
megafauna movement. However, what currently works best, for example, to identify key
corridors to conserve European bison (Bison bonasus) movement likely is not the best model
to predict structural connectivity for dead wood beetles (Rocca et al., 2017). Therefore,
understanding the strengths and appropriate applications of different connectivity modelling
approaches is vital to producing the most accurate results to address your problem.
Modelling connectivity is an important step in the design and implementation of a connectivity
project, and should take place after initial steps of scoping and problem assessment, and
setting of connectivity objectives (see the connectivity network design framework in Chapter
6 for more details).
In the following subsections, different frameworks as families of models are introduced, and
the applications of these methods, and their strengths and weaknesses are discussed.

A summary of modelling families


Data inputs can vary across the different types of connectivity analyses. Central to nearly all
connectivity modelling families, however, is some form of information on the habitat patches
or “focal nodes” where species or processes of interest occur and where movement is
expected to originate and/or terminate (Table 5.1).

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).

Model Family Data Needs Applications Software & Packages

Focal species corridors,


LCP: ArcGIS Tools,
Least-cost population dispersal
Resistance surface, focal QGIS plugin, R
Path & potential, area of potential
nodes, and species dispersal packages ('gdistance',
Resistant use, pollinator movement,
data (RK) 'leastcostpath'); RK:
Kernels probability of human
UNICOR
movement

62
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

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

Focal species connectivity


Resistance surface & focal Circuitscape, GFlow,
and pinch points, water
Circuit Theory nodes (Circuitscape) or Omniscape, Linkage
flow, pollinator movement,
source weight surface (Omni) Mapper (multi-family)
invasive species control

Model specific: Focal node


and network data, survival Long-term population
MetaIPM, HexSIM,
Agent-based rate, population growth rate, persistence, patch and
NetLogo, R packages (‘p
models fecundity, node transition connection importance,
SiMRiv’)
probabilities, resistance source-sink analysis
surface, etc.

(Will differ depending on the


Assessing connectivity of
used metric(s)) Number of
select components in the Conefor, ArcPro Network
patches, patch size(s),
Structural physical landscape, e.g. Analyst, R packages
boundaries and perimeter,
Connectivity protected areas, specific (‘iGraph’, ‘riverconn’,
distance between patches,
Metrics habitats and/or corridors. 'gdistance')
focal nodes,
Both in relation to intra- and
presence/absence of links,
inter-patch connectivity.
number of paths.

Study area planning units Identifying structural


Spatial layer, biodiversity connectivity via
Marxan, Zonation, R
prioritisation distributions, land cover/land prioritisation of landscape
packages (‘prioritizr’)
tools use, current protected areas, elements and systematic
etc. conservation planning

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
63
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

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.

Creating resistance surfaces


Also, central to the least-cost path, resistant kernels (Section 5.2), and circuit theory models
(Section 5.4) is a measurement of resistance of movement between and possibly within
habitat patches (Table 5.1). Resistance is generally defined as the inherent difficulty that an
animal, plant or abiotic process will have while trying to cross a section of land or water. This
resistance is input into the models as a resistance surface, a raster of spatial data where each
pixel is the energetic cost, risk, or force exerted to move across that pixel. The cumulative
resistance to movement can thus be calculated as the summation of all the pixels crossed on
a path between two habitat patches. The easiest conceptualization of this cumulative
resistance to movement may be the difficulties faced by a large mammal attempting to cross
a landscape over hours or days. However, the temporal scale of this cumulative resistance
can vary depending on the focal problem. Other examples could include the varying seasonal
forces acting on the seed dispersal of particular plant species, the resistance met by
subsequent generations of a grouse species as it expands its range, or the seasonal and
anthropogenic forces acting on the flow of water in a regional watershed.
There are a variety of methods to create resistance surfaces, depending on the type of data
available for the species of interest (Zeller et al., 2012). Species distribution models (SDM)
that correlate environmental variables to species habitat use are currently the most common
method for the creation of resistance surfaces. These models may use presence data (e.g.,
GLM, random forest, Maxent, etc.) or detection-nondetection data (e.g., occupancy models)
to estimate habitat suitability. Results from the SDM are often then inverted so that the areas
with the greatest habitat suitability values have the lowest values of resistance. A major
assumption in this is that habitat suitability values are completely and inversely correlated with
the resistance to movement that land cover exerts on that species (i.e., the same magnitude
of force dictating habitat preference for an area dictates the ability to move across that area).
Data transformations that decrease the level of resistance of moderately suitable habitats
have been proposed and used in recent studies to compensate for a part of this distinction
(Keeley et al., 2016). Other methods include the use of telemetry or GPS point and tracking
data from collared wildlife individuals. Point data can be applied to resource selection functions
to generate the relative probability of resource use and track data is used in path-selection
functions, which generate a relative probability of movement raster (Zeller et al., 2018). Finally,
resistance surfaces can be generated from landscape genetic data collected from across the
study area. The genetics of a population differentiates with increasing distance between
subpopulations and obstacles to individual movement, therefore, differences in the spatial
patterns of pairwise genetics can be used as a proxy for the spatial distribution of resistance
values. While this can be seen as a measure of functional connectivity, optimization of
resistance surfaces using landscape genetic data is much more complex and requires running
a genetic algorithm in conjunction with calculations of pairwise effective distances from
64
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

connectivity models to determine an optimal resistance surface (Dutta et al., 2022; Peterman,
2018).

More complex modelling frameworks


Data types and needs change as we move further into more complex modelling frameworks
that attempt to simulate individual movement or metapopulation dynamics (Table 5.1). Models
in this family are especially useful for analysing population viability over time, therefore, inputs
can include data on population survival rates, abundance and fecundity, probability of
dispersal from patches, and multiple characteristics of the focal patches. Like graph theory
(Section 5.3), one can use these models to prioritise focal patches, with the added benefit of
determining long-term processes such as identifying potential sources and sinks amongst the
different patches. Source-sink dynamics analyses how population growth is affected by
variation in habitat quality (Pulliam 1988). In this approach, sources correspond to high quality
habitat allowing populations to increase, while sinks correspond to low quality habitat that on
its own would not support a population. However, sink populations may persist indefinitely if
surplus individuals move from sources to the sinks. Thus, considering source-sink dynamics
can inform conservation decisions.

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.

5.2 Least-cost path and Resistant Kernels


Least-cost path analysis is the form of connectivity modelling that has been applied the
longest. These models find the most cost-effective path between two points across a
resistance surface taking into consideration the distance travelled and the cumulative

65
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

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.

66
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

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.

Model Software & Functions & tools Details


Family Packages
Least-cost ArcGIS toolbox Distance Accumulation Beyond the basic functionality of
Path function calculating an individual least-cost path
(ArcPro:Spatial this tool can calculate the equivalent of
Analyst) factorial least-cost. The function also can
(https://pro.arcgis.com/ consider true surface distance and a raster
en/pro- of landscape barriers as input.
app/latest/help/analysi
s/raster- This function will automatically attempt to
functions/distance- parallelize the analysis across half the
accumulation-global- available cores in your computer, which
function.htm) should increase processing time.

Cost Path tool Can calculate least-cost paths between 2


(ArcMap:Spatial or more sources, however, typically
Analyst) & Optimal requires the output from one of the other
Regions Connection weighted cost tools in the ArcGIS toolbox
(ArcPro) (e.g., Cost Distance) to create the cost
(https://pro.arcgis.com/ distance raster.
en/pro-app/latest/tool-
reference/spatial- Note: ArcMap will no longer be updated
analyst/optimal-region- post-February 2024 and will lose technical
connections.htm) support March 2026.
QGIS Least-Cost Path plugin Capable of calculating the least cost
(https://plugins.qgis.or path(s) when given a cost raster, start
g/plugins/leastcostpath points, and end points as inputs.
/)
https://github.com/Gooong/LeastCostPath
for more information.
R packages leastcostpath Fully integrated package with functions to
(https://cran.r- create different resistance/cost layers and
project.org/web/packa then run least-cost path analysis.
ges/leastcostpath/inde
x.html) Includes functionality for different forms of
least-cost path analyses, though primarily
oriented to barriers and slope costs.
gdistance Greater functionality than ‘leastcostpath’
(https://cran.r- which makes it more complicated to
project.org/web/packa operate.
ges/gdistance/index.ht
ml) Includes abilities beyond least-cost path
including constrained and non-constrained
random walk functions with similarities to
resistant kernels and circuit theory.

67
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

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.

Grid/raster input data is in ASCII format.


Additional input information includes
distance of movement thresholds and
parameters of the kernel density
estimator.

An older but more in-depth manual can be


found at this web address:
https://www.fs.usda.gov/rm/pubs_other/rmr
s_2011_landguth_e002.pdf

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.

5.3 Graph Theory


Definitions and applicable situations
Graph Theory is a mathematical discipline that finds wide application in various fields,
including computer science, linguistics, social network analysis, transportation network
analysis, and ecological connectivity. Graph Theory focuses on structures known as graphs,
which consist of vertices (nodes) and edges (connections).
In the context of ecological connectivity, graph theory models allow us to prioritise habitat
patches and links between patches relative to one another. Vertices represent patches of
habitat (or any other relevant features such as protected areas), while edges represent the
connectivity between these patches (Bunn et al., 2000). Vertices can be assigned weights that
influence their connectedness with neighbouring nodes and their overall importance in the
network. In ecological connectivity analysis, the weight is typically based on some property of
the habitat patch, such as its area, overall quality, population size, estimated number of
propagules produced, etc. Similarly, edges can be modelled in different ways. In a simple
binary representation, an edge indicates the presence of a connection between a pair of
vertices, for example, when two vertices are closer than the species' dispersal distance. In
68
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

more sophisticated versions, edges are represented as probabilities of connection between


vertices, which can be a function of dispersal probability at a given distance, based on
dispersal kernels.
The application of graph theory to connectivity analysis enables the examination of network
connectivity, allow us to compare between different networks. It also allows for the analysis of
adding or subtracting individual vertices or edges to the overall network connectivity (i.e., if
habitat patch X is added how much better is the network connected). Various metrics have
been developed to describe the degree of connectivity in a network, some of the more
common metrics used in connectivity modelling is the Integral Index of Connectivity [IIC],
Probability of Connectivity [PC], and Equivalent Connected Area [ECA] (Pascual-Hortal and
Saura, 2006; Saura et al., 2011; Saura and Pascual-Hortal, 2007). For individual vertices
metrics such as the generalised betweenness centrality metric measures how centralised or
well-connected a node is within the network (Bodin and Saura, 2010).
To estimate the relative contribution (i.e., importance) and non-redundancy of vertices or
edges to the network, these metrics can be applied to evaluate network connectivity with and
without specific vertices or edges, leading to the above indices IIC and PC to be expressed
as the percentage of variation: dIIC, dPC, respectively. This can be interpreted as the
importance of the node or edge according to the index. The relative contribution of a patch
can be further analysed by estimating its contribution in terms of habitat area (dIICintra,
dPCintra), its connectivity to neighbouring patches (dIICflux, dPCflux), and its role as a
stepping stone (e.g., dIICconnector, dPCconnector), resulting in (Saura and Rubio, 2010):
dIIC = dIICintra + dIICflux + dIICconnector

and

dPC = dPCintra + dPCflux + dPCconnector

Data inputs, packages and software


Ecological connectivity analyses using graph theory typically require two types of input data:
a list of vertices with their attributes (typically a list of protected areas), and a list of edges with
their attributes. The vertex dataset includes all vertices present in the network and generally
consists of two columns: a vertex ID and its weight. The edge dataset represents realised
connections in the network and is typically reported with three columns: two columns indicating
the connected nodes and one column indicating the measure of connectivity between them.
Connectivity between patches can be expressed as a probability, which can be estimated
based on the probability of dispersal to a given distance. To obtain such a probability,
knowledge of the average or median dispersal distance of a species is necessary, along with
information about the distribution of dispersal distances in a population. The commonly
assumed distribution is a negative exponential distribution, but other distributions are also
possible such as for instance Weibull for plants (García and Borda-de-Água, 2017).
Additionally, the probability of connection between vertices can be influenced by the
permeability of the landscape. While the simplest approach assumes Euclidean distance

69
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

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.

5.4 Circuit Theory


Circuit theory connectivity models are graph theoretical methods which use principles of
electrical current to simulate the potential movement of an animal or other entity across a
resistance surface. We know that electrical current will attempt to flow through areas of least
resistance and that in those areas where there is high resistance current density can be
compressed through lower resistance areas (Fig. 5.1; McRae et al., 2008). This becomes very
useful as an allegory for the movement of living species as well as abiotic processes,
70
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

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,
71
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

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
72
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

mapping connectivity and conservation opportunities on agricultural landscapes (Suraci et al.,


2023) and prioritizing multispecies habitat networks that are robust to climate and land-use
change (Albert et al., 2017). Different circuit theory applications and versions developed are
detailed in Table 5.3.

73
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

Table 5.3: Circuit theory applications developed detailing needed data inputs and sources.

Application Details Inputs URL

Circuitscape 5 The current version of Data inputs include ASCII https://github.com/


Circuitscape is developed in the text files for focal nodes Circuitscape/Circui
Julia programming language, and resistance surfaces, tscape.jl
allowing the program to run much and a .ini file that gives
faster and model much larger the program instructions
landscapes. on how the model should
The only coding needed is when calculate currents
loading the program and to input between the nodes, and
the needed data files (only 6-7 what type of outputs the
lines of code). Preparation of data model should produce
inputs is however much more (see instructions and
time-consuming. adaptable templates for
the .ini file in the link
provided).

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.

Omniscape Omniscape is written in Julia by Data inputs include ASCII https://github.com/


the same developers as text files for the Circuitscape/Omni
Circuitscape. resistance and source scape.jl
Like Circuitscape 5 there are only weight surfaces, and a .ini
a few lines of code necessary to file that gives the program
run these models in Julia, and instructions on moving
formatting the data correctly is window size, target pixel
much more time-consuming. block size, what type of
Computational times can take outputs you would like the
days or weeks depending on model to produce, and
landscape extent, data resolution, several other advanced
moving window size, and if target options.
pixels are clumped in the moving
window.
It is advised to start with a much
smaller extent to adjustments
these options.

5.5 Agent-based models


Agent-based models (ABM) or, for our purposes, more aptly called individual-based models,
simulate the behaviour of individual agents, such as animals, in their environment. They
provide a flexible approach by allowing the simulation of each agent as a distinct entity, with
a set of rules and behaviours. They are powerful tools to understand and explore the emergent
74
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

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.

Software packages Details Source


moveHMM Based on Hidden Markov Models and Maximum Michelot et al. (2016)
Likelihood estimation.
momentuHMM McClintock and
Allow fitting individual multistate movement
Michelot (2018)
models to real location data.
SiMRiv Uses optimization to fit a multistate model to Quaglietta and Porto
location data. (2019)

75
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

Can be used to simulate the movement of aquatic


species and predict road mortality hotspots.
samc Connectivity modelling with spatial absorbing Marx et al. (2020)
Markov chains.
rangeShifter Built using object-oriented C++ providing Bocedi et al. (2020)
computationally efficient simulation of complex
individual-based, eco-evolutionary models and
species responses to environmental changes.
NetLogo Flexible, programmable simulation environments, Wilenski
greatly simplify the task of translating a movement (1999)
HexSim model into a computer program and simulating the Schumaker and
dynamics and analysing the outcomes of an entire Brookes (2018)
system based on its agents

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
76
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

a trial-and-error process, to achieve a realistic movement pattern in light of what is known


about the species. Although this manual approach may achieve reasonable results (Quaglietta
and Porto, 2019), it is plagued by the typical problems raised by a priori and arbitrary
decisions, and, naturally, should be avoided if there is real tracking data available. However,
manual parameterization may be the only option when a complex or customised movement
model is needed, as statistical estimation procedures are not available for all kinds of models
and do not cover all particular cases.
After the user has a parameterized movement model (or models, if there are multiple species
of interest), they can be used to simulate a large number of individuals in landscapes, for a
given time frame. The emergent patterns of these simulations are of paramount utility in
addressing questions of connectivity, landscape fragmentation, and landscape change. By
combining all the simulated movements along the simulated years, it is possible to derive, for
example, metrics of connectivity (Whittington et al., 2022) and infer the best location for
corridors. For instance, areas that are more intensively used by the simulated individuals may
be seen as areas that are important as corridors. Further, the impacts of changes in the
landscape can be explicitly addressed, by simulating individuals in different landscape
configurations, that may be dynamic according to different scenarios, and assessing the
differences in connectivity that result (Whittington et al., 2022). This ultimately allows
assessing the consequences and effectiveness of different planning and management
approaches on population spread (migration) or persistence (extinction). In theory, there are
no limitations to the potential applications of such models in landscape connectivity problems,
the foremost practical limitation is that the higher the model complexity, the more difficult it is
to parameterize it in an ecologically meaningful way. Validating the outputs of such models
will always require expert knowledge of the system at hand.

5.6 Structural Connectivity Metrics and moving window-analysis


When assessing the structural connectivity of landscapes, applying one or more of a larger
collection of connectivity metrics, all varying in complexity can often be a direct and
straightforward methodology. Connectivity metrics are formulas that use various components
and/or characteristics of the landscape, to provide the user with distinct values of connectivity
estimations. The values can either be between different patches of habitats or across larger
landscapes (Keeley et al., 2021). Different metrics can vary a lot both in terms of input data
and parameters needed for their implementation as well as in their resulting estimation and
interpretation. Most structural connectivity metrics (Table 5.5) use physical components of the
study area to provide its assessment, this could include characteristics such as but not limited
to: the area-size of focal patches, the distance between patches, the threshold distance or
perimeter of patches (Yang et al., 2024). Other metrics can require graph components based
on graph theory, such as nodes/centroids or links and path (see Section 5.3).

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.

Metric Explanation Reference

77
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

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.

Area-weighted mean radius An Area-weighted summarization of the mean McGarigal, 1995


of gyration* radius of gyration

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

78
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

(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.

79
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

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.

5.7 Assessing ecosystem services


There are several approaches and tools to map ecosystem supply service and demand. For
details, Burkhard and Maes (2017) provide a relevant overview and source of information.
Most of these tools are openly available and are constantly evolving. The selection of an
appropriate tool depends on multiple factors including the questions to be addressed, the
spatial scale, data availability and so on.

80
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

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).

81
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

6. A framework for connectivity


conservation and planning
6.1 Introduction to the framework
Designing and implementing an ecological connectivity network either at the local, regional,
or continental scale involves considering several fundamental steps to ensure its effectiveness
in promoting ecological connectivity. In this Chapter we present a general framework for
planning and implementing a connectivity project (Fig. 6.1) and walk through five major steps.
While the framework is presented linearly, it is often a very iterative process in part due to the
continuous need to engage and collaborate with area stakeholders to ensure the production
of the most accurate and useful plan possible. The framework covers these five steps:
(1) Scoping and Problem Assessment: Conduct a comprehensive analysis of the entire
landscape to identify potential threats, connectivity actions, and impact of those actions,
identify all relevant stakeholders and build an interdisciplinary collaboration team for
connectivity analysis, communication, and implementation. Establish the general spatial
extent at which your study will take place;
(2) Setting of Objectives: Use the assessment of the connectivity problem to establish spatial
and temporally explicit objectives and targets that will help mitigate the identified problem.
Determine the appropriate width and characteristics of corridors and stepping stones based
on the target species and landscape characteristics. Finalize the spatial extent and needed
data resolution;
(3) Analysis Selection & Data Preparation: Determine the correct model or models to analyse
ecological connectivity. Given the model and your objectives collect and produce all the
necessary data and spatial layers necessary to run the spatial analysis;
(4) Assessment of connectivity: Use connectivity metrics and models to determine the most
effective design for a connectivity network that integrates with the current network of protected
areas. Present draft results to stakeholders, iterate new models, and prioritize corridors and
stepping stones;
(5) Implementation, Monitoring & Evaluation: Develop a comprehensive management and
monitoring plan for the ecological corridor and/or stepping stones. This includes activities such
as habitat restoration, invasive species control, monitoring of species movement, and
assessing the corridor's effectiveness in achieving the connectivity objectives.
While the landscape characteristics, focal species or conservation objectives of each
connectivity project can be different, these steps, in conjunction with the information provided
thus far, provide a foundation for designing a connectivity network that may effectively facilitate
ecological connectivity.

82
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

Figure 6.1: A schematic representation of a framework for connectivity network design.

6.2 Scoping and problem assessment

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
83
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

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.

©PA Images / Alamy Stock Photo

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

84
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

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

85
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

6.3 Setting of Objectives

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.

Problem Objective Target Action

86
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

Implement the protection


of a forested
Low genetic diversity and Four or more breeding
corridor >1km wide
a decreasing abundance Increase brown bear females establish new
between a source
of brown bear in an movement to isolated PA territories in the isolated
population of brown
isolated PA PA
bears and the isolated
PA.

Ensure spatial and Return pollinator Seasonal ban on


Rapidly declining
temporal connectivity capture numbers to mowing of key stepping
pollinator presence in
between grassland 110% of baseline before stone grasslands and
grassland habitat
patches for pollinators rapid decline fields

Restrict grazing within


Increasing instances of Identify and restore new Reduce livestock and expand the
livestock mortality due to corridors to decrease mortality by 40% within minimum width of two
wolf dispersal human-wolf conflict 5 years corridors prioritized by
connectivity models

Work with local


Establish 5 new
environmental ministries
Increasing mortality of Define new stepping migratory stopover
to implement other
waterbirds during stone habitats for areas AND increase
effective area-based
migration migrating waterbirds annual waterbird
conservation measures
survival by 3%
on identified wetlands.

6.3.1. Focal & archetype species for assessing connectivity in Europe


Connectivity studies often focus on a single or small suite of focal species to represent other
species or the broader ecosystem. Historically, these were charismatic species threatened by
habitat fragmentation and were thought of as an umbrella species whose conservation would
hopefully benefit a broad suite of species and ecosystem processes. These focal species
approaches are still frequently used given the special attention still paid to large charismatic
mammals, the scarcity of presence or movement data for many species, and the relative
complexity of attempting to model the connectivity of dozens of different species at the same
time.
While the focal species approach can be effective in promoting and protecting some areas for
connectivity, identifying a small grouping of species with differing habitat preferences has been
shown to be more effective at capturing the needs of a broader species pool for a given habitat
type (Meurant et al., 2018). Similarly, projects may target groups of species sharing some
fundamental traits and with similar conservation needs. A typical example is large carnivores,
which have large area requirements and might be particularly vulnerable to fragmentation. In
these cases, 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
(see Box 6.1 for plant and vertebrate species archetypes in Europe).

Box 6.1 Description of archetype tetrapods.

87
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

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.

• Salamanders was represented by three archetypes: (1) medium-sized newts with


high clutch size and presence of larval stages, (2) big-sized viviparous salamanders
with high clutch size, and (3) small- to medium-sized salamanders with low clutch
size and presence of direct development.

• 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.

88
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

• 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.

89
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

Figure 6.2: Habitat preferences (percentage of species) of European tetrapods using three coarse classes.

6.3.2 Corridor width


The width of ecological corridors is a critical factor for determining their effectiveness. Narrow
corridors may not provide sufficient habitat or protection for wildlife, especially for species with
wide-ranging territories. Conversely, overly wide corridors may be impractical or economically
unfeasible, especially in densely populated or agriculturally intensive regions. Corridor width
should be sufficient to accommodate two general groups of species based on their mobility,
passage species and corridor dwellers. Passage species refer to those species in which
an individual animal can traverse the length of the corridor in a single event, typically in a few
hours or days. Corridor dwellers require more than a generation to move individuals and/or
genes across the corridor (Beier and Loe, 1992). For corridor dwellers, it is important that the
corridor is sufficiently wide to allow for overlapping home ranges allowing animals to live, find
mates and reproduce in the corridor (Beier and Loe, 1992).
90
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

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).

Box 6.2 Debate and Guidance on Corridor Width

Beier’s rule of thumb


Beier (2019) suggests that ecological corridors connecting habitat patches that are 8–80
km apart should have a minimum width of 2 kilometres, except for unavoidable bottlenecks
such as highway crossings. This recommendation is based on the notion that a width of this
magnitude would be sufficient to accommodate home ranges of up to approximately 8 km².
This coverage would meet the needs of 345 species of mammals that are probable corridor
dwellers, selected from a list of 429 terrestrial mammals provided by Tucker et al. (2014).
Concerning edge effects, Beier (2019) notes that in North America negative edge effects
were considered biologically significant at distances of up to 300 m (Kennedy et al., 2003),
therefore a corridor that is 2 km wide would have at least 1700 m free of edge effects.
Ford’s et al. approach
Ford et al. (2020) proposal is based on the concepts of zone of influence (Boulanger et al.,
2012) and flight initiation distance (FID), that is, the distance at which animals flee from an
approaching person. Specifically, their approach is based on determining how both these
distances reduce effective corridor widths and use that knowledge to make
recommendations on corridor width in different landscape settings. The authors searched
the literature for case studies documenting the zone of influence and the FID from
recreational trails and residential development for four North American carnivore species,
black bears, grizzly bears, grey wolves, and cougars. From this analysis, they concluded
that the effective corridor width should vary from 3000 to 6000 m close to residential areas
and 400 to 1000 m in areas containing recreational trails, depending on the species.
USDA’s recommendations
The United States Department of Agriculture (USDA) conducted an extensive review of 66
studies, including movement ecology studies, observational studies, and habitat
management studies conducted in North America, encompassing a diverse range of
species groups that are also present in Europe (Bentrup, 2008). While the USDA
acknowledged that many of the studies did not encompass a wide enough range of corridor
widths to definitively determine optimal sizes, they did provide general recommendations.

91
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

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).

Group Minimum corridor width Upper end of recommended


(m) width (m)

Plants 30 101

Invertebrates 30 61

Aquatic species 30 61

Reptiles & Amphibians 30 183

Birds: interior species 61 1609

Birds: edge species 30 101

Small mammals 101 101

Large mammals 101 2414

Large predator mammals 101 > 4828

6.3.3 Final Spatial Extent and Resolution


During the scoping process a general spatial extent was identified at which the project will be
operating. Now at this step once specific objectives are defined and potentially the targets
associated with those objectives, there can be greater concentration on the precise spatial
extent and resolution of an analysis that will assist decision makers in an impactful
conservation outcome.
Two main factors should be of focus when determining the spatial extent (i.e., total study area)
and resolution (i.e., grain) of your connectivity analysis. First, what is the extent and resolution
that will most accurately represent and capture the ecological process which you are
attempting to model and conserve? Second, what is the extent and resolution that will be
effective for practitioners and policymakers to implement the findings? For example,
attempting to model the corridors that are important for a seasonally migratory large ungulate
will likely require a spatial extent that is orders of magnitude larger than that for the connectivity
for a local ground beetle.
92
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

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.

93
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

6.4 Analysis Selection & Data Preparation

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

94
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

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.

95
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

Checklist of Spatial Data:


Data layers frequently needed during a connectivity analysis with some
more specific features.

▢ Protected Area Boundaries


▢ Political Boundaries
▢ Land Cover/Land Use
▢ Land use projections
▢ Land management practices
▢ Elevation Data
▢ Rivers & Lakes
▢ Hydrologic features (vector data)
▢ Wetness level* (raster data)
▢ Linear Infrastructure
▢ Roads & Trails
▢ Border Walls & Fences
▢ Dams
▢ Wildlife passages
▢ Species Presence Data
▢ Climate Data
▢ Historical trends
▢ Future climate scenarios
* Includes information on permanent vs. temporary wet areas, snow, etc.

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.

96
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

6.5 Assessment of Connectivity

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
97
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

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.

6.5.1 Prioritisation and restoration for connectivity objectives


Not all corridors or stepping stones are equally important to maintain the overall connectivity
of the system. Beyond identifying potential corridors or stepping stones, connectivity analysis
can be used as a decision tool to prioritise those identified areas, either to improve existing
ones or to establish new ones through restoration activities, by helping to identify areas that
hold the greatest potential for overall connectivity in the region (Rudnick et al., 2012). One
way of getting this information is to evaluate the importance of removing each of the corridors
(linkages) in the overall connectivity of the region, using different methods such as program
Conefor or Linkage Mapper (see Chapter 4). For example, de la Fuente et al. (2018) used a
graph-based approach to identify the conservation importance and the restoration importance
of the corridors linking Natura 2000 sites across Spain. The latter was quantified as the
increase in the connectivity of the network of Natura 2000 sites that would occur if the current
conditions in each corridor were improved so that the land cover was fully composed of the
most favourable habitat for species movements.
In the process of conservation area selection, there is a tendency to select areas with better
conditions for movement by animals and plants and therefore higher connectivity. However, if
these corridors are not maintained, e.g. due to land use changes, their function might be lost.
This is an important aspect to consider when using connectivity information in spatial
conservation planning (e.g., Daigle et al., 2018; see also Section 2.7), as the optimization
algorithm will influence protected area selection assuming a priori connectivity estimates that
might be changed in the future if the land cover types responsible for the high values are
changed.
Restoration for connectivity is not restricted to habitat restoration but can include interventions
on human infrastructures that act as barriers to movement. In the terrestrial realm,
underpasses or overpasses in roads and railways have proved effective, though evaluating
their effectiveness is still not a common practice and needs to be generalized (Soanes et al.,
2024). Mitigating the impacts of such linear infrastructure on wildlife populations also includes
preventing mortality due to wildlife-vehicle collisions. Under that perspective, methods to
prioritise the placement of mitigation measures such as under or overpasses in roads or
railways can include the use of connectivity models to identify crossing areas (Lee et al., 2023)
and the identification of road mortality hotspots based on carcass surveys. The implementation
98
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

of mitigation measures such as crossing structures should consider species-specific


ecological requirements and follow best-practice guidelines (Soanes et al., 2024). For
example, Gurrutxaga and Saura (2014) used habitat network analysis to evaluate how the
location of road permeabilization measures for highway defragmentation would restore
landscape connectivity in a forest-protected area network in the Basque country in Spain.
More recently, Lee et al. (2023) used a combination of connectivity modelling (based on
empirical data of migratory routes and using least-cost pathways and corridor prioritisation)
and road mortality hotspot identification together with citizen science data to prioritise road
mitigation opportunities for pronghorn in Canada.
In the freshwater realm and especially in rivers, dams and weirs as well as other obstacles
represent significant impacts on connectivity thus affecting biodiversity. While large dams
inherently create barriers, even smaller infrastructural elements, like small dams, act as
(micro-)barriers. These smaller barriers, although often overlooked, can cumulatively affect
larger areas and hence are seen as priority zones for restoration to enhance connectivity.
Although possibilities exist to enhance partial connectivity in rivers, such as migration facilities
for fish, full connectivity (i.e. enabling not only the movement of fish but also a free flow of
water and materials such as sediments) can only be established in rivers by removing the
barrier. Dam removal has gained considerable importance through the last years and will gain
further importance as the EU Nature Restoration Law aims for at least 25 000 km of free-
flowing rivers, compared to 2020. Thus, approaches to prioritise the removal of dams and
barriers can help to efficiently reach this target. The prioritisation of barrier removal should first
increase the connectivity of habitats that are relevant for species to fulfil their life cycle (e.g.
habitats for spawning, juveniles, adults for migratory fish). However, trade-offs between costs
(e.g. reduced energy production) and conservation benefits must be considered too (ideally
this should first be identified during step 1 of the decision framework) (McKay et al., 2017).
Overall, the connectivity of the river network can be optimised in regard of different aspects
(Branco et al., 2014; Hermoso et al., 2021a).
Another option to mitigate the effects of barriers in rivers is migration facilities, such as fish
ladders at hydropower plants. These facilities function as an alternative corridor for organisms
for up- and downstream connectivity restoration. To be used by the different species of
interest, their design must meet the requirements of the most demanding migratory species
concerning, e.g., swimming capabilities, preferred migration corridors, and space
requirements (Seliger and Zeiringer, 2018). However, a migration facility does not restore the
‘full’ connectivity of a corridor (e.g. sediments are still trapped upstream of the barrier).
Moreover, migration facilities also provide corridors for invasive species or pathogens, which
has also to be assessed compared to the benefits for a species in need. See McKay et al.
(2020a) for guidelines on criteria to identify priority areas for setting these facilities. Prioritising
hydropower plants where such migration facilities should be built is mostly related to the
species that should benefit from such a facility. For example, the ICPDR developed an
‘Ecological Prioritisation Approach River and Habitat Continuity Restoration’ approach that
combines structural as well as functional connectivity aspects where the functional aspects
are addressed by different migratory guilds of fish (mainly medium- and long-distance
migratory).
Overall, rivers, wetlands and riparian land have been identified as priority areas for restoration
as corridors, increasing connectivity for multiple purposes as they contain both blue and green
99
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

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.

6.6 Implementation, Monitoring & Evaluation

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).

100
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

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.

101
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

102
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

7. References
Abbott, I.M., Butler, F., Harrison, S., 2012. When flyways meet highways – The relative permeability of different
motorway crossing sites to functionally diverse bat species. Landsc. Urban Plan. 106, 293–302.
https://doi.org/10.1016/j.landurbplan.2012.03.015
Adams, V.M., Alvarez-Romero, J.G., Carwardine, J., Cattarino, L., Hermoso, V., Kennard, M.J., Linke, S., Pressey,
R.L., Stoeckl, N., 2014. Planning Across Freshwater and Terrestrial Realms: Cobenefits and Tradeoffs
Between Conservation Actions. Conserv. Lett. 7, 425–440. https://doi.org/10.1111/conl.12080
Alagador, D., Cerdeira, J.O., 2022. Operations research applicability in spatial conservation planning. J. Environ.
Manage. 315, 115172. https://doi.org/10.1016/j.jenvman.2022.115172
Albert, C.H., Rayfield, B., Dumitru, M. and Gonzalez, A. (2017), Applying network theory to prioritize multispecies
habitat networks that are robust to climate and land-use change. Conservation Biology, 31: 1383-1396.
https://doi.org/10.1111/cobi.12943
Alencar, L.R.V., Quental, T.B., 2023. Geographical and ecological drivers of coexistence dynamics in squamate
reptiles. Glob. Ecol. Biogeogr. 32, 1937–1951. https://doi.org/10.1111/geb.13745
Allan, J.R., Possingham, H.P., Atkinson, S.C., Waldron, A., Di Marco, M., Butchart, S.H.M., Adams, V.M., Kissling,
W.D., Worsdell, T., Sandbrook, C., Gibbon, G., Kumar, K., Mehta, P., Maron, M., Williams, B.A., Jones, K.R.,
Wintle, B.A., Reside, A.E., Watson, J.E.M., 2022. The minimum land area requiring conservation attention to
safeguard biodiversity. Science 376, 1094–1101. https://doi.org/10.1126/science.abl9127
Álvarez-Romero, J.G., Pressey, R.L., Ban, N.C., Brodie, J., 2015. Advancing Land-Sea Conservation Planning:
Integrating Modelling of Catchments, Land-Use Change, and River Plumes to Prioritise Catchment
Management and Protection. PLOS ONE 10, e0145574. https://doi.org/10.1371/journal.pone.0145574
Alvarez-Romero, J.G., Pressey, R.L., Ban, N.C., Vance-Borland, K., Willer, C., Klein, C.J., Gaines, S.D., 2011.
Integrated land-sea conservation planning: The missing links, in: Futuyma, D., Shaffer, H., Simberloff, D.
(Eds.), Annual Review of Ecology, Evolution, and Systematics, Vol 42, pp. 381–409.
https://doi.org/10.1146/annurev-ecolsys-102209-144702
Anderson, A.S., Reside, A.E., VanDerWal, J.J., Shoo, L.P., Pearson, R.G., Williams, S.E., 2012. Immigrants and
refugees: the importance of dispersal in mediating biotic attrition under climate change. Glob. Change Biol.
18, 2126–2134. https://doi.org/10.1111/j.1365-2486.2012.02683.x
Arthington, A.H., 2021. Grand Challenges to Support the Freshwater Biodiversity Emergency Recovery Plan. Front.
Environ. Sci. 9. https://doi.org/10.3389/fenvs.2021.664313
Ashrafzadeh, M.R., Khosravi, R., Adibi, M.A., Taktehrani, A., Wan, H.Y., Cushman, S.A., 2020. A multi-scale, multi-
species approach for assessing effectiveness of habitat and connectivity conservation for endangered felids.
Biol. Conserv. 245, 108523. https://doi.org/10.1016/j.biocon.2020.108523
Atkinson, S.F., Lake, M.C., 2020. Prioritizing riparian corridors for ecosystem restoration in urbanizing watersheds.
PeerJ 8, e8174. https://doi.org/10.7717/peerj.8174
Ausprey, I.J., Newell, F.L., Robinson, S.K., 2023. Sensitivity of tropical montane birds to anthropogenic disturbance
and management strategies for their conservation in agricultural landscapes. Conserv. Biol. e14136.
https://doi.org/10.1111/cobi.14136
Ball, I.R., Possingham, H.P., 2000. Marxan (v 1.8.6): Marine reserve design using spatially explicit annealing. A
manual prepared for the Great Barrier Reef Marine Park Authority. (phd).
Bastianelli, M.L., Premier, J., Herrmann, M., Anile, S., Monterroso, P., Kuemmerle, T., Dormann, C.F., Streif, S.,
Jerosch, S., Götz, M., Simon, O., Moleón, M., Gil-Sánchez, J.M., Biró, Z., Dekker, J., Severon, A., Krannich,
A., Hupe, K., Germain, E., Pontier, D., Janssen, R., Ferreras, P., Díaz-Ruiz, F., López-Martín, J.M., Urra, F.,
Bizzarri, L., Bertos-Martín, E., Dietz, M., Trinzen, M., Ballesteros-Duperón, E., Barea-Azcón, J.M., Sforzi, A.,
Poulle, M.-L., Heurich, M., 2021. Survival and cause-specific mortality of European wildcat (Felis silvestris)
across Europe. Biol. Conserv. 261, 109239. https://doi.org/10.1016/j.biocon.2021.109239
103
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

Beger, M., Grantham, H.S., Pressey, R.L., Wilson, K.A., Peterson, E.L., Dorfman, D., Mumby, P.J., Lourival, R.,
Brumbaugh, D.R., Possingham, H.P., 2010a. Conservation planning for connectivity across marine,
freshwater, and terrestrial realms. Biol. Conserv. 143, 565–575. https://doi.org/10.1016/j.biocon.2009.11.006
Beger, M., Linke, S., Watts, M., Game, E., Treml, E., Ball, I., Possingham, H.P., 2010b. Incorporating asymmetric
connectivity into spatial decision making for conservation. Conserv. Lett. 3, 359–368.
https://doi.org/10.1111/j.1755-263X.2010.00123.x
Beger, M., Metaxas, A., Balbar, A.C., McGowan, J.A., Daigle, R., Kuempel, C.D., Treml, E.A., Possingham, H.P.,
2022. Demystifying ecological connectivity for actionable spatial conservation planning. Trends Ecol. Evol.
37, 1079–1091. https://doi.org/10.1016/j.tree.2022.09.002
Beier, P., 2019. A rule of thumb for widths of conservation corridors. Conserv. Biol. 33, 976–978.
https://doi.org/10.1111/cobi.13256
Beier, P., Loe, S., 1992. In My Experience: A Checklist for Evaluating Impacts to Wildlife Movement Corridors.
Wildl. Soc. Bull. 1973-2006 20, 434–440.
Beier, P., Majka, D.R., Spencer, W.D., 2008. Forks in the Road: Choices in Procedures for Designing Wildland
Linkages. Conservation Biology 22, 836–851. https://doi.org/10.1111/j.1523-1739.2008.00942.x
Beier, P., Majka, D.R., Newell, S.L., 2009. Uncertainty analysis of least-cost modeling for designing wildlife
linkages. Ecol. Appl. 19, 2067–2077. https://doi.org/10.1890/08-1898.1
Beier, P., Noss, R., 1998. Do Habitat Corridors Provide Connectivity? Conserv. Biol. 12, 1241–1252.
https://doi.org/10.1111/j.1523-1739.1998.98036.x
Bélisle, M., 2005. Measuring landscape connectivity: the challenge of behavioral landscape ecology. Ecology 86,
1988–1995. https://doi.org/10.1890/04-0923
Benedict, M., MacMahon, E., 2002. Green Infrastructure: Smart Conservation for the 21st Century, Renewable
Resources Journal.
Beninde, J., Veith, M., Hochkirch, A., 2015. Biodiversity in cities needs space: a meta-analysis of factors
determining intra-urban biodiversity variation. Ecol. Lett. 18, 581–592.
Bennett, A.F. (2003). Linkages in the Landscape: The Role of Corridors and Connectivity in Wildlife Conservation.
IUCN, Gland, Switzerland and Cambridge, UK. xiv + 254 pp. https://doi.org/10.2305/IUCN.CH.2004.FR.1.en
Bentrup, G., 2008. Conservation Buffers Design Guidelines for Buffers, Corridors, and Greenways.
https://doi.org/10.2737/SRS-GTR-109
Bernaschini, M. L., Trumper, E., Valladares, G., & Salvo, A. (2019). Are all edges equal? Microclimatic conditions,
geographical orientation and biological implications in a fragmented forest. Agriculture, Ecosystems &
Environment, 280, 142–151. https://doi.org/10.1016/j.agee.2019.04.035
Beyer, H. L., Dujardin, Y., Watts, M. E., & Possingham, H. P. (2016). Solving conservation planning problems with
integer linear programming. Ecological Modelling, 328, 14–22.
https://doi.org/10.1016/j.ecolmodel.2016.02.005
Birds of the World (2022). Edited by S. M. Billerman, B. K. Keeney, P. G. Rodewald, and T. S. Schulenberg. Cornell
Laboratory of Ornithology, Ithaca, NY, USA.
Bocedi, G., Palmer, S.C.F., Malchow, A.-K., Zurell, D., Watts, K. and Travis, J.M.J. (2021), RangeShifter 2.0: an
extended and enhanced platform for modelling spatial eco-evolutionary dynamics and species' responses to
environmental changes. Ecography, 44: 1453-1462. https://doi.org/10.1111/ecog.05687
Bode, M., Burrage, K., Possingham, H.P., 2008. Using complex network metrics to predict the persistence of
metapopulations with asymmetric connectivity patterns. Ecol. Model. 214, 201–209.
https://doi.org/10.1016/j.ecolmodel.2008.02.040
Bodin, Ö., Saura, S., 2010. Ranking individual habitat patches as connectivity providers: integrating network
analysis and patch removal experiments. Ecol. Model. 221, 2393–2405.
https://doi.org/10.1016/j.ecolmodel.2010.06.017

104
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

Borda-de-Água, L., Ascensão, F., Sapage, M., Barrientos, R., Pereira, H.M., 2019. On the identification of mortality
hotspots in linear infrastructures. Basic Appl. Ecol. 34, 25–35. https://doi.org/10.1016/j.baae.2018.11.001
Boulanger, J., Poole, K.G., Gunn, A., Wierzchowski, J., 2012. Estimating the zone of influence of industrial
developments on wildlife: a migratory caribou Rangifer tarandus groenlandicus and diamond mine case study.
Wildl. Biol. 18, 164–179. https://doi.org/10.2981/11-045
Bowman, J., Jaeger, J.A.G., Fahrig, L., 2002. Dispersal distance of mammals is proportional to home range size.
Ecology 83, 2049–2055. https://doi.org/10.1890/0012-9658
Branco, P., Segurado, P., Santos, J.M., Ferreira, M.T., 2014. Prioritizing barrier removal to improve functional
connectivity of rivers. J. Appl. Ecol. 51, 1197–1206. https://doi.org/10.1111/1365-2664.12317
Brito, J.C., Martínez-Freiría, F., Sierra, P., Sillero, N., Tarroso, P., 2011. Crocodiles in the Sahara Desert: An
Update of Distribution, Habitats and Population Status for Conservation Planning in Mauritania. PLOS ONE
6, e14734. https://doi.org/10.1371/journal.pone.0014734
Brodie, J.F., Giordano, A.J., Dickson, B., Hebblewhite, M., Bernard, H., Mohd-Azlan, J., Anderson, J., Ambu, L.,
2015. Evaluating multispecies landscape connectivity in a threatened tropical mammal community:
Multispecies Habitat Corridors. Conserv. Biol. 29, 122–132. https://doi.org/10.1111/cobi.12337
Bruggeman, D. J., Wiegand, T., & Fernández, N. (2010). The relative effects of habitat loss and fragmentation on
population genetic variation in the red‐cockaded woodpecker (Picoides borealis). Molecular Ecology, 19(17),
3679-3691.
Bunn, A.G., Urban, D.L., Keitt, T.H., 2000. Landscape connectivity: a conservation application of graph theory. J.
Environ. Manage. 59, 265–278.
Burkhard, B., Maes, J., 2017. Mapping ecosystem services. Pensoft Publishers, Sofia.
Burnett, M.J., O’Brien, G.C., Jacobs, F.J., Jewitt, G., Downs, C.T., 2021. Fish telemetry in African inland waters
and its use in management: a review. Rev. Fish Biol. Fish. 31, 337–357. https://doi.org/10.1007/s11160-021-
09650-2
Campos, P., Caparrós, A., Oviedo, J.L., Ovando, P., Álvarez-Farizo, B., Díaz-Balteiro, L., Carranza, J., Beguería,
S., Díaz, M., Herruzo, A.C., Martínez-Peña, F., Soliño, M., Álvarez, A., Martínez-Jauregui, M., Pasalodos-
Tato, M., de Frutos, P., Aldea, J., Almazán, E., Concepción, E.D., Mesa, B., Romero, C., Serrano-Notivoli,
R., Fernández, C., Torres-Porras, J., Montero, G., 2019. Bridging the Gap Between National and Ecosystem
Accounting Application in Andalusian Forests, Spain. Ecol. Econ. 157, 218–236.
https://doi.org/10.1016/j.ecolecon.2018.11.017
Carrao, H., Kleeschulte, S., Naumann, S., Davis, M., Schröder, C., Abdul Malak, D., Condé, S., Erhard, M., & Dige,
G. (2020). Contributions to building a coherent Trans-European Nature Network. (p. 38). Environment Agency
Austria.
Caro, T., 2010. Conservation by Proxy: Indicator, Umbrella, Keystone, Flagship, and Other Surrogate Species.
Island Press.
Carvalho, S.B., Gonçalves, J., Guisan, A., Honrado, J., 2016. Systematic site selection for multi-species monitoring
networks. J. Appl. Ecol. 53, 1305–1316. https://doi.org/10.1111/1365-2664.12505
Carvalho, S.B., Velo-Antón, G., Tarroso, P., Portela, A.P., Barata, M., Carranza, S., Moritz, C., Possingham, H.P.,
2017. Spatial conservation prioritization of biodiversity spanning the evolutionary continuum. Nat. Ecol. Evol.
1, 0151. https://doi.org/10.1038/s41559-017-0151
Cattarino, L., Hermoso, V., Carwardine, J., Kennard, M.J., Linke, S., 2015. Multi-Action Planning for Threat
Management: A Novel Approach for the Spatial Prioritization of Conservation Actions. PLOS ONE 10,
e0128027. https://doi.org/10.1371/journal.pone.0128027
CBD, 2022a. 15/4. Kunming-Montreal Global Biodiversity Framework.
CBD, 2022b. 15/5. Monitoring framework for the Kunming-Montreal Global Biodiversity Framework.
Chapron, G., Arlettaz, R., 2006. Using Models to Manage Carnivores. Science 314, 1682–1683.
https://doi.org/10.1126/science.314.5806.1682c
105
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

Cisneros-Araujo, P., Ramirez-Lopez, M., Juffe-Bignoli, D., Fensholt, R., Muro, J., Mateo-Sánchez, M.C., Burgess,
N.D., 2021. Remote sensing of wildlife connectivity networks and priority locations for conservation in the
Southern Agricultural Growth Corridor (SAGCOT) in Tanzania. Remote Sens. Ecol. Conserv. 7, 430–444.
https://doi.org/10.1002/rse2.199
Ciucci, P., Reggioni, W., Maiorano, L., Boitani, L., 2009. Long-Distance Dispersal of a Rescued Wolf From the
Northern Apennines to the Western Alps. J. Wildl. Manag. 73, 1300–1306. https://doi.org/10.2193/2008-510
Crooks, K.R., Burdett, C.L., Theobald, D.M., Rondinini, C., Boitani, L., 2011. Global patterns of fragmentation and
connectivity of mammalian carnivore habitat. Philos. Trans. R. Soc. B Biol. Sci. 366, 2642–2651.
https://doi.org/10.1098/rstb.2011.0120
Crooks, K.R., Sanjayan, M., 2006. Connectivity conservation: maintaining connections for nature, in: Crooks, K.R.,
Sanjayan, M. (Eds.), Connectivity Conservation, Conservation Biology. Cambridge University Press,
Cambridge, pp. 1–20. https://doi.org/10.1017/CBO9780511754821.001
Csárdi, G., Nepusz, T., Müller, K., Horvát, S., Traag, V., Zanini, F., Noom, D., 2023. igraph for R: R interface of the
igraph library for graph theory and network analysis. https://doi.org/10.5281/ZENODO.7682609
Cushman, S.A., Landguth, E.L., 2012. Multi-taxa population connectivity in the Northern Rocky Mountains. Ecol.
Model. 231, 101–112. https://doi.org/10.1016/j.ecolmodel.2012.02.011
Cushman, S.A., Lewis, J.S., Landguth, E.L., 2013. Evaluating the intersection of a regional wildlife connectivity
network with highways. Mov. Ecol. 1, 12. https://doi.org/10.1186/2051-3933-1-12
Cushman, S.A., McKELVEY, K.S., Schwartz, M.K., 2009. Use of Empirically Derived Source-Destination Models
to Map Regional Conservation Corridors. Conserv. Biol. 23, 368–376. https://doi.org/10.1111/j.1523-
1739.2008.01111.x
Dahlin, K.M., Zarnetske, P.L., Read, Q.D., Twardochleb, L.A., Kamoske, A.G., Cheruvelil, K.S., Soranno, P.A.,
2021. Linking Terrestrial and Aquatic Biodiversity to Ecosystem Function Across Scales, Trophic Levels, and
Realms. Front. Environ. Sci. 9. https://doi.org/10.3389/fenvs.2021.692401
Daigle, R., Metaxas, A., Balbar, A., Mcgowan, J., Treml, E., Kuempel, C., Possingham, H., Beger, M., 2018.
Operationalizing ecological connectivity in spatial conservation planning with Marxan Connect.
https://doi.org/10.1101/315424
Daigle, R.M., Metaxas, A., Balbar, A.C., McGowan, J., Treml, E.A., Kuempel, C.D., Possingham, H.P., Beger, M.,
2020. Operationalizing ecological connectivity in spatial conservation planning with Marxan Connect. Methods
Ecol. Evol. 11, 570–579. https://doi.org/10.1111/2041-210X.13349
D’Aloia, C.C., Daigle, R.M., Côté, I.M., Curtis, J.M.R., Guichard, F., Fortin, M.-J., 2017. A multiple-species
framework for integrating movement processes across life stages into the design of marine protected areas.
Biol. Conserv. 216, 93–100. https://doi.org/10.1016/j.biocon.2017.10.012
de la Fuente, B., Mateo-Sánchez, M.C., Rodríguez, G., Gastón, A., Pérez de Ayala, R., Colomina-Pérez, D.,
Melero, M., Saura, S., 2018. Natura 2000 sites, public forests and riparian corridors: The connectivity
backbone of forest green infrastructure. Land Use Policy 75, 429–441.
https://doi.org/10.1016/j.landusepol.2018.04.002
Deinet, S., Scott-Gatty, K., Rotton, H., Twardek, W.M., Marconi, V., McRae, L., Baumgartner, L.J., Brink, K.,
Claussen, J.E., Cooke, S.J., Darwall, W., Eriksson, B.K., Garcia de Leaniz, C., Hogan, Z., Royte, J., Silva,
L.G.M., Thieme, M.L., Tickner, D., Waldman, J., Wanningen, H., Weyl, O.L.F., Berkhuysen, A., 2020. The
Living Planet Index (LPI) for migratory freshwater fish: Technical Report. World Fish Migration Foundation,
Groningen.
Dertien, J.S., Larson, C.L., Reed, S.E., 2021. Recreation effects on wildlife: A review of potential quantitative
thresholds. Nat. Conserv. 44, 51–68.
Dertien, J.S., Baldwin, R.F., 2023. Does Scale or Method Matter for Conservation? Application of Directional and
Omnidirectional Connectivity Models in Spatial Prioritizations. Front. Conserv. Sci. 4.
https://doi.org/10.3389/fcosc.2023.976914.

106
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

Devlin, M.J., McKinna, L.W., Álvarez-Romero, J.G., Petus, C., Abott, B., Harkness, P., Brodie, J., 2012. Mapping
the pollutants in surface riverine flood plume waters in the Great Barrier Reef, Australia. Mar. Pollut. Bull.,
The Catchment to Reef Continuum: Case studies from the Great Barrier Reef 65, 224–235.
https://doi.org/10.1016/j.marpolbul.2012.03.001
Dickson, B.G., Albano, C.M., Anantharaman, R., Beier, P., Fargione, J., Graves, T.A., Gray, M.E., Hall, K.R.,
Lawler, J.J., Leonard, P.B., Littlefield, C.E., McClure, M.L., Novembre, J., Schloss, C.A., Schumaker, N.H.,
Shah, V.B., Theobald, D.M., 2019. Circuit‐theory applications to connectivity science and conservation.
Conserv. Biol. 33, 239–249. https://doi.org/10.1111/cobi.13230
Dobrowski, S.Z., Littlefield, C.E., Lyons, D.S., Hollenberg, C., Carroll, C., Parks, S.A., Abatzoglou, J.T., Hegewisch,
K., Gage, J., 2021. Protected-area targets could be undermined by climate change-driven shifts in ecoregions
and biomes. Commun. Earth Environ. 2, 1–11. https://doi.org/10.1038/s43247-021-00270-z
Doerr, V.A.J., Barrett, T., Doerr, E.D., 2011. Connectivity, dispersal behaviour and conservation under climate
change: a response to Hodgson et al.: Connectivity and dispersal behaviour. J. Appl. Ecol. 48, 143–147.
https://doi.org/10.1111/j.1365-2664.2010.01899.x
Dudgeon, D., 2019. Multiple threats imperil freshwater biodiversity in the Anthropocene. Curr. Biol. 29, R960–R967.
https://doi.org/10.1016/j.cub.2019.08.002
Dutta, T., De Barba, M., Selva, N., Fedorca, A.C., Maiorano, L., Thuiller, W., Zedrosser, A., Signer, J., Pflüger, F.,
Frank, S., Lucas, P.M., Balkenhol, N., 2023. An objective approach to select surrogate species for connectivity
conservation. Front. Ecol. Evol. 11, 1078649. https://doi.org/10.3389/fevo.2023.1078649
Dutta, T., Sharma, S., Meyer, N.F.V., Larroque, J., Balkenhol, N., 2022. An overview of computational tools for
preparing, constructing and using resistance surfaces in connectivity research. Landsc. Ecol. 37, 2195–2224.
https://doi.org/10.1007/s10980-022-01469-x
EC, 2023. Revision of the EU Pollinators Initiative.
EC, 2022. Proposal for a Regulation of the European Parliament and of the Council on nature restoration.
EC, 2021a. New EU Forest Strategy for 2030.
EC, 2021b. The 3 Billion Tree Planting Pledge For 2030.
EC, 2020. EU Biodiversity Strategy for 2030.
EC, 2019. Guidance on a strategic framework for further supporting the deployment of EU-level green and blue
infrastructure.
EC, 2013. Communication from the Commission to the European Parliament, the Council, the European Economic
and Social Committee and the Committee of the Regions — Green infrastructure (GI): enhancing Europe’s
natural capital.
EEA, 2024. Natura 2000 and land cover data viewer — European Environment Agency [WWW Document]. URL
https://www.eea.europa.eu/data-and-maps/dashboards/natura-2000-data-viewer (accessed 2.14.24).
EEA, 2021. Drivers of and pressures arising from selected key water management challenges: a European
overview. Publications Office of the European Union, LU.
Eggers, B., Matern, A., Drees, C., Eggers, J., Härdtle, W., & Assmann, T. (2010). Value of Semi-Open Corridors
for Simultaneously Connecting Open and Wooded Habitats: A Case Study with Ground Beetles. Conservation
Biology, 24(1), 256–266.
Estreguil, C., Dige, G., Kleeschulte, S., Carrao, H., Raynal, J., Teller, A., 2019. Strategic Green Infrastructure and
Ecosystem Restoration: geospatial methods, data and tools. Publications Office of the European Union,
Luxembourg.
Estreguil,C, Dige, G, Kleeschulte, S, Carrao, H, Raynal, J, Teller, A, 2019. Strategic green infrastructure and
ecosystem restoration: geospatial methods, data and tools. Publications Office, LU.
Estrela-Segrelles, C., Gómez-Martínez, G., Pérez-Martín, M.Á., 2023. Climate Change Risks on Mediterranean
River Ecosystems and Adaptation Measures (Spain). Water Resour. Manag. 37, 2757–2770.
https://doi.org/10.1007/s11269-023-03469-1
107
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

Fagan, W.F., 2002. Connectivity, Fragmentation, and Extinction Risk in Dendritic Metapopulations. Ecology 83,
3243–3249. https://doi.org/10.1890/0012-9658(2002)083[3243:CFAERI]2.0.CO;2
Fahrig, L., 2019. Habitat fragmentation: A long and tangled tale. Glob. Ecol. Biogeogr. 28, 33–41.
https://doi.org/10.1111/geb.12839
Fahrig, L., 2003. Effects of Habitat Fragmentation on Biodiversity. Annu. Rev. Ecol. Evol. Syst. 34, 487–515.
https://doi.org/10.1146/annurev.ecolsys.34.011802.132419
Fahrig, L., Merriam, G., 1985. Habitat Patch Connectivity and Population Survival: Ecological Archives E066-008.
Ecology 66, 1762–1768. https://doi.org/10.2307/2937372
Farmer, J.D., Foley, D., 2009. The economy needs agent-based modelling. Nature 460, 685–686.
https://doi.org/10.1038/460685a
Fernández, N., Román, J., & Delibes, M. (2016). Variability in primary productivity determines metapopulation
dynamics. Proceedings of the Royal Society B: Biological Sciences, 283(1828), 20152998
Fernández, N., Torres, A., Wolf, F., Quintero, L., & Pereira, H. M. (2020). Boosting Ecological Restoration for a
Wilder Europe. German Centre for Integrative Biodiversity Research (iDiv) & Martin-Luther University Halle-
Wittemberg. DOI: https://dx.doi.org/10.978.39817938/57
Field, R.D., Parrott, L., 2017. Multi-ecosystem services networks: A new perspective for assessing landscape
connectivity and resilience. Ecol. Complex. 32, 31–41. https://doi.org/10.1016/j.ecocom.2017.08.004
Ford, A.T., Sunter, E.J., Fauvelle, C., Bradshaw, J.L., Ford, B., Hutchen, J., Phillipow, N., Teichman, K.J., 2020.
Effective corridor width: linking the spatial ecology of wildlife with land use policy. Eur. J. Wildl. Res. 66, 69.
https://doi.org/10.1007/s10344-020-01385-y
Frankham, R., Ballou, J.D., Briscoe, D.A., 2010. Introduction to Conservation Genetics, 2nd ed. Cambridge
University Press, Cambridge. https://doi.org/10.1017/CBO9780511809002
Friedrichs, M., Hermoso, V., Bremerich, V., Langhans, S.D., 2018. Evaluation of habitat protection under the
European Natura 2000 conservation network – The example for Germany. PLoS ONE 13, e0208264.
https://doi.org/10.1371/journal.pone.0208264
García, C. and Borda-de-Água, L. (2017), Extended dispersal kernels in a changing world: insights from statistics
of extremes. J Ecol, 105: 63-74. https://doi.org/10.1111/1365-2745.12685
Gaston, K., 2003. The Structure and Dynamics of Geographic Ranges.
Gholizadeh, M.H., Melesse, A.M., Reddi, L., 2016. A Comprehensive Review on Water Quality Parameters
Estimation Using Remote Sensing Techniques. Sensors 16, 1298. https://doi.org/10.3390/s16081298
Giakoumi, S., Hermoso, V., Carvalho, S.B., Markantonatou, V., Dagys, M., Iwamura, T., Probst, W.N., Smith, R.J.,
Yates, K.L., Almpanidou, V., Novak, T., Ben-Moshe, N., Katsanevakis, S., Claudet, J., Coll, M., Deidun, A.,
Essl, F., Garcia-Charton, J.A., Jimenez, C., Kark, S., Mandic, M., Mazaris, A.D., Rabitsch, W., Stelzenmuller,
V., Tricarico, E., Vogiatzakis, I.N., 2019. Conserving European biodiversity across realms. Conserv. Lett. 12.
https://doi.org/10.1111/conl.12586
Gilbert-Norton, L., Wilson, R., Stevens, J.R., Beard, K.H., 2010. A Meta-Analytic Review of Corridor Effectiveness:
Corridor Meta-Analysis. Conserv. Biol. 24, 660–668. https://doi.org/10.1111/j.1523-1739.2010.01450.x
Gomes, L., Grilo, C., Silva, C., Mira, A., 2009. Identification methods and deterministic factors of owl roadkill hotspot
locations in Mediterranean landscapes. Ecol. Res. 24, 355–370. https://doi.org/10.1007/s11284-008-0515-z
Gregory, A.J., Beier, P., 2014. Response variables for evaluation of the effectiveness of conservation corridors.
Conserv. Biol. J. Soc. Conserv. Biol. 28, 689–695. https://doi.org/10.1111/cobi.12252
Grill, G., Lehner, B., Thieme, M., Geenen, B., Tickner, D., Antonelli, F., Babu, S., Borrelli, P., Cheng, L.,
Crochetiere, H., Ehalt Macedo, H., Filgueiras, R., Goichot, M., Higgins, J., Hogan, Z., Lip, B., McClain, M.E.,
Meng, J., Mulligan, M., Nilsson, C., Olden, J.D., Opperman, J.J., Petry, P., Reidy Liermann, C., Sáenz, L.,
Salinas-Rodríguez, S., Schelle, P., Schmitt, R.J.P., Snider, J., Tan, F., Tockner, K., Valdujo, P.H., van
Soesbergen, A., Zarfl, C., 2019. Mapping the world’s free-flowing rivers. Nature 569, 215–221.
https://doi.org/10.1038/s41586-019-1111-9
108
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

Grimm, V., Railsback, S.F., 2005. Individual-based Modeling and Ecology:, STU-Student edition. ed. Princeton
University Press.
Gundersen, H., Andreassen, H.P., 1998. The risk of moose Alces alces collision: A predictive logistic model for
moose-train accidents. Wildl. Biol. 4, 103–110. https://doi.org/10.2981/wlb.1998.007
Gurrutxaga, M., Saura, S., 2014. Prioritizing highway defragmentation locations for restoring landscape
connectivity. Environ. Conserv. 41, 157–164. https://doi.org/10.1017/S0376892913000325
Gutiérrez-Rodríguez, J., Gonçalves, J., Civantos, E., Martínez-Solano, I., 2017. Comparative landscape genetics
of pond-breeding amphibians in Mediterranean temporal wetlands: The positive role of structural
heterogeneity in promoting gene flow. Mol. Ecol. 26, 5407–5420. https://doi.org/10.1111/mec.14272
Haase, P., Bowler, D.E., Baker, N.J., Bonada, N., Domisch, S., Garcia Marquez, J.R., Heino, J., Hering, D., Jähnig,
S.C., Schmidt-Kloiber, A., Stubbington, R., Altermatt, F., Álvarez-Cabria, M., Amatulli, G., Angeler, D.G.,
Archambaud-Suard, G., Jorrín, I.A., Aspin, T., Azpiroz, I., Bañares, I., Ortiz, J.B., Bodin, C.L., Bonacina, L.,
Bottarin, R., Cañedo-Argüelles, M., Csabai, Z., Datry, T., de Eyto, E., Dohet, A., Dörflinger, G., Drohan, E.,
Eikland, K.A., England, J., Eriksen, T.E., Evtimova, V., Feio, M.J., Ferréol, M., Floury, M., Forcellini, M., Forio,
M.A.E., Fornaroli, R., Friberg, N., Fruget, J.-F., Georgieva, G., Goethals, P., Graça, M.A.S., Graf, W., House,
A., Huttunen, K.-L., Jensen, T.C., Johnson, R.K., Jones, J.I., Kiesel, J., Kuglerová, L., Larrañaga, A., Leitner,
P., L’Hoste, L., Lizée, M.-H., Lorenz, A.W., Maire, A., Arnaiz, J.A.M., McKie, B.G., Millán, A., Monteith, D.,
Muotka, T., Murphy, J.F., Ozolins, D., Paavola, R., Paril, P., Peñas, F.J., Pilotto, F., Polášek, M., Rasmussen,
J.J., Rubio, M., Sánchez-Fernández, D., Sandin, L., Schäfer, R.B., Scotti, A., Shen, L.Q., Skuja, A., Stoll, S.,
Straka, M., Timm, H., Tyufekchieva, V.G., Tziortzis, I., Uzunov, Y., van der Lee, G.H., Vannevel, R.,
Varadinova, E., Várbíró, G., Velle, G., Verdonschot, P.F.M., Verdonschot, R.C.M., Vidinova, Y., Wiberg-
Larsen, P., Welti, E.A.R., 2023. The recovery of European freshwater biodiversity has come to a halt. Nature
620, 582–588. https://doi.org/10.1038/s41586-023-06400-1
Haddad, N., Hudgens, B., Damschen, E., Levey, D., Orrock, J., Tewksbury, J., Weldon, A., 2011. Assessing
positive and negative ecological effects of corridors. Sources Sinks Sustain. 475–504.
https://doi.org/10.1017/CBO9780511842399.024
Haddad, N.M., Brudvig, L.A., Clobert, J., Davies, K.F., Gonzalez, A., Holt, R.D., Lovejoy, T.E., Sexton, J.O., Austin,
M.P., Collins, C.D., Cook, W.M., Damschen, E.I., Ewers, R.M., Foster, B.L., Jenkins, C.N., King, A.J.,
Laurance, W.F., Levey, D.J., Margules, C.R., Melbourne, B.A., Nicholls, A.O., Orrock, J.L., Song, D.-X.,
Townshend, J.R., 2015. Habitat fragmentation and its lasting impact on Earth’s ecosystems. Sci. Adv. 1,
e1500052. https://doi.org/10.1126/sciadv.1500052
Haddad, N.M., Brudvig, L.A., Damschen, E.I., Evans, D.M., Johnson, B.L., Levey, D.J., Orrock, J.L., Resasco, J.,
Sullivan, L.L., Tewksbury, J.J., Wagner, S.A., Weldon, A.J., 2014. Potential Negative Ecological Effects of
Corridors. Conserv. Biol. 28, 1178–1187. https://doi.org/10.1111/cobi.12323
Hagen-Zanker, A., 2016. A computational framework for generalized moving windows and its application to
landscape pattern analysis. International journal of applied earth observation and geoinformation, 44, 205-
216. https://doi.org/10.1016/j.jag.2015.09.010
Hamill, L., Gilbert, N., 2015. Agent-Based Modelling in Economics, 1st edition. ed. Wiley.
Hanna, D.E.L., Tomscha, S.A., Ouellet Dallaire, C., Bennett, E.M., 2018. A review of riverine ecosystem service
quantification: Research gaps and recommendations. J. Appl. Ecol. 55, 1299–1311.
https://doi.org/10.1111/1365-2664.13045
Hanson, Jeffrey O., Fuller, R.A., Rhodes, J.R., 2019. Conventional methods for enhancing connectivity in
conservation planning do not always maintain gene flow. J. Appl. Ecol. https://doi.org/10.1111/1365-
2664.13315
Hanson, J.O., McCune, J.L., Chadès, I., Proctor, C.A., Hudgins, E.J., Bennett, J.R., 2023. Optimizing ecological
surveys for conservation. J. Appl. Ecol. 60, 41–51. https://doi.org/10.1111/1365-2664.14309
Hanson, J. O., Schuster, R., Morrell, N., Strimas-Mackey, M., Watts, M.E., Arcese, P., Bennett, J., 2019. Prioritizr:
Systematic Conservation Prioritization in R. R Package Version 411 https://CRAN.R-
project.org/package=prioritizr.

109
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

Hanson, J.O., Vincent, J., Schuster, R., Fahrig, L., Brennan, A., Martin, A.E., Hughes, J.S., Pither, R., Bennett,
J.R., 2022. A comparison of approaches for including connectivity in systematic conservation planning. J.
Appl. Ecol. 59, 2507–2519. https://doi.org/10.1111/1365-2664.14251
Heller, N.E., Zavaleta, E.S., 2009. Biodiversity management in the face of climate change: A review of 22 years of
recommendations. Biol. Conserv. 142, 14–32. https://doi.org/10.1016/j.biocon.2008.10.006
Hermoso, V., Linke, S., Prenda, J., Possingham, H.P., 2011. Addressing longitudinal connectivity in the systematic
conservation planning of fresh waters. Freshw. Biol. 56, 57–70. https://doi.org/10.1111/j.1365-
2427.2009.02390.x
Hermoso, V., Ward, D.P., Kennard, M.J., 2012a. Using water residency time to enhance spatio-temporal
connectivity for conservation planning in seasonally dynamic freshwater ecosystems. J. Appl. Ecol. 49, 1028–
1035. https://doi.org/10.1111/j.1365-2664.2012.02191.x
Hermoso, V., Kennard, M.J., Linke, S., 2012b. Integrating multidirectional connectivity requirements in systematic
conservation planning for freshwater systems. Divers. Distrib. 18, 448–458. https://doi.org/10.1111/j.1472-
4642.2011.00879.x
Hermoso, V., Vasconcelos, R.P., Henriques, S., Filipe, A.F., Carvalho, S.B., 2021a. Conservation planning across
realms: Enhancing connectivity for multi-realm species. J. Appl. Ecol. https://doi.org/10.1111/1365-
2664.13796
Hermoso, V., Clavero, M., Filipe, A.F., 2021b. An accessible optimisation method for barrier removal planning in
stream networks. Sci. Total Environ. 752, 141943. https://doi.org/10.1016/j.scitotenv.2020.141943
Herrera, L.P., Sabatino, M.C., Jaimes, F.R., Saura, S., 2017. Landscape connectivity and the role of small habitat
patches as stepping stones: an assessment of the grassland biome in South America. Biodivers. Conserv.
26, 3465–3479. https://doi.org/10.1007/s10531-017-1416-7
Hilty, J. A., Jr, W. Z. L., Merenlender, A. M., & Dobson, A. P. (2012). Corridor Ecology: The Science and Practice
of Linking Landscapes for Biodiversity Conservation. Island Press.
Hilty, J.A., Keeley, A.T.H., Jr, W.Z.L., Merenlender, A.M., 2019. Corridor Ecology, Second Edition: Linking
Landscapes for Biodiversity Conservation and Climate Adaptation, Second Edition, New edition, Second
Edition, New. ed. Island Press, Washington.
Hilty, J., Worboys, G.L., Keeley, A., Woodley, S., Lausche, B.J., Locke, H., Carr, M., Pulsford, I., Pittock, J., White,
J.W., Theobald, D.M., Levine, J., Reuling, M., Watson, J.E.M., Ament, R., Tabor, G.M., 2020. Guidelines for
conserving connectivity through ecological networks and corridors. IUCN, International Union for
Conservation of Nature. https://doi.org/10.2305/IUCN.CH.2020.PAG.30.en
Hodgson, J.A., Thomas, C.D., Wintle, B.A., Moilanen, A., 2009. Climate change, connectivity and conservation
decision making: back to basics. J. Appl. Ecol. 46, 964–969. https://doi.org/10.1111/j.1365-
2664.2009.01695.x
Hofmeister, J., Hošek, J., Brabec, M., Střalková, R., Mýlová, P., Bouda, M., Pettit, J. L., Rydval, M., & Svoboda, M.
(2019). Microclimate edge effect in small fragments of temperate forests in the context of climate change.
Forest Ecology and Management, 448, 48–56. https://doi.org/10.1016/j.foreco.2019.05.069
Honeck, E., Sanguet, A., Schlaepfer, M.A., Wyler, N., Lehmann, A., 2020. Methods for identifying green
infrastructure. SN Appl. Sci. 2, 1916. https://doi.org/10.1007/s42452-020-03575-4
Houet, T., Palka, G., Rigo, R., Hugues, B., Baudry, J., Xavier, P., Jean-Baptiste, N., Álvarez-Martínez, J.M., Balbi,
S., Cendrine, M., Lucie, L., Johanna, B., Barquín, P., 2022. European blue and green infrastructure network
strategy vs. the common agricultural policy. Insights from an integrated case study (Couesnon, Brittany). Land
Use Policy 120, 106277. https://doi.org/10.1016/j.landusepol.2022.106277
Ignatieva, M., Stewart, G.H., Meurk, C., 2011. Planning and design of ecological networks in urban areas. Landsc.
Ecol. Eng. 7, 17–25. https://doi.org/10.1007/s11355-010-0143-y
IPBES, 2022. Global assessment report on biodiversity and ecosystem services of the Intergovernmental Science-
Policy Platform on Biodiversity and Ecosystem Services. https://doi.org/10.5281/zenodo.6417333

110
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

Joppa, L.N., Pfaff, A., 2009. High and far: biases in the location of protected areas. PLoS ONE 4, e8273.
doi:10.1371/journal.pone.0008273.
Jordán, F., Báldi, A., Orci, K.M., 2003. Characterizing the importance of habitat patches and corridors in maintaining
the landscape connectivity of a Pholidoptera transsylvanica (Orthoptera) metapopulation. Landscape Ecol.
18, 83–92. https://doi.org/10.1023/A:1022958003528
Jung, M., Arnell, A., de Lamo, X., García-Rangel, S., Lewis, M., Mark, J., Merow, C., Miles, L., Ondo, I., Pironon,
S., Ravilious, C., Rivers, M., Schepashenko, D., Tallowin, O., van Soesbergen, A., Govaerts, R., Boyle, B.L.,
Enquist, B.J., Feng, X., Gallagher, R., Maitner, B., Meiri, S., Mulligan, M., Ofer, G., Roll, U., Hanson, J.O.,
Jetz, W., Di Marco, M., McGowan, J., Rinnan, D.S., Sachs, J.D., Lesiv, M., Adams, V.M., Andrew, S.C.,
Burger, J.R., Hannah, L., Marquet, P.A., McCarthy, J.K., Morueta-Holme, N., Newman, E.A., Park, D.S.,
Roehrdanz, P.R., Svenning, J.-C., Violle, C., Wieringa, J.J., Wynne, G., Fritz, S., Strassburg, B.B.N.,
Obersteiner, M., Kapos, V., Burgess, N., Schmidt-Traub, G., Visconti, P., 2021. Areas of global importance
for conserving terrestrial biodiversity, carbon and water. Nat. Ecol. Evol. 1–11.
https://doi.org/10.1038/s41559-021-01528-7
Kaval, P., 2019. Integrated catchment management and ecosystem services: A twenty-five year overview. Ecosyst.
Serv. 37, 100912. https://doi.org/10.1016/j.ecoser.2019.100912
Keeley, A., Beier, P., Gagnon, J. W., 2016. Estimating landscape resistance from habitat suitability: effects of data
source and nonlinearities. Landsc. Ecol. 31. https://doi.org/10.1007/s10980-016-0387-5
Keeley, A.T.H., Ackerly, D.D., Cameron, D.R., Heller, N.E., Huber, P.R., Schloss, C.A., Thorne, J.H., Merenlender,
A.M., 2018. New concepts, models, and assessments of climate-wise connectivity. Environ. Res. Lett. 13,
073002. https://doi.org/10.1088/1748-9326/aacb85
Keeley, A.T.H., Beier, P., Jenness, J.S., 2021. Connectivity metrics for conservation planning and monitoring. Biol.
Conserv. 255, 109008. https://doi.org/10.1016/j.biocon.2021.109008
Keller, D., Holderegger, R., van Strien, M.J., Bolliger, J., 2015. How to make landscape genetics beneficial for
conservation management? Conserv. Genet. 16, 503–512. https://doi.org/10.1007/s10592-014-0684-y
Kennedy, C., Wilkinson, J., Balch, J., 2003. Conservation Thresholds for Land Use Planners.
Keten, A., Eroglu, E., Kaya, S., Anderson, J.T., 2020. Bird diversity along a riparian corridor in a moderate urban
landscape. Ecol. Indic. 118, 106751. https://doi.org/10.1016/j.ecolind.2020.106751
Klar, N., Herrmann, M., Kramer-Schadt, S., 2009. Effects and Mitigation of Road Impacts on Individual Movement
Behavior of Wildcats. J. Wildl. Manag. 73, 631–638. https://doi.org/10.2193/2007-574
Koen, E.L., Garroway, C.J., Wilson, P.J., Bowman, J., 2010. The effect of map boundary on estimates of landscape
resistance to animal movement. PLoS One 5. https://doi.org/10.1371/journal.pone.0011785
Kumar, S.U., Cushman, S.A., 2022. Connectivity modelling in conservation science: a comparative evaluation. Sci.
Rep. 12, 1–12. https://doi.org/10.1038/s41598-022-20370-w
Landau, V., Shah, V., Anantharaman, R., Hall, K., 2021. Omniscape.jl: Software to compute omnidirectional
landscape connectivity. J. Open Source Softw. 6, 2829. https://doi.org/10.21105/joss.02829
Larsen-Gray, A.L., Loehle, C., 2022. Relationship Between Riparian Buffers and Terrestrial Wildlife in the Eastern
United States. J. For. 120, 336–357. https://doi.org/10.1093/jofore/fvab067
Laurance, W.F., Bierregaard, R.O., 1997. Tropical Forest Remnants: Ecology, Management, and Conservation of
Fragmented Communities. University of Chicago Press, Chicago.
Laurance, W.F., Lovejoy, T.E., Vasconcelos, H.L., Bruna, E.M., Didham, R.K., Stouffer, P.C., Gascon, C.,
Bierregaard, R.O., Laurance, S.G., Sampaio, E., 2002. Ecosystem Decay of Amazonian Forest Fragments:
A 22-Year Investigation. Conserv. Biol. 16, 605–618. https://doi.org/10.1046/j.1523-1739.2002.01025.x
Lee, T.S., Jones, P.F., Jakes, A.F., Jensen, M., Sanderson, K., Duke, D., 2023. Where to invest in road mitigation?
A comparison of multiscale wildlife data to inform roadway prioritization. J. Nat. Conserv. 71, 126327.
https://doi.org/10.1016/j.jnc.2022.126327

111
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

Leoncini, F., Semenzato, P., Di Febbraro, M., Loy, A., Ferrari, C., 2023. Come back to stay: landscape connectivity
analysis for the Eurasian otter (Lutra lutra) in the western Alps. Biodivers. Conserv. 32, 653–669.
https://doi.org/10.1007/s10531-022-02517-3
Leontiou, S., Katsanevakis, S., Vogiatzakis, I.N., 2022. Accounting for functional connectivity in cross-realm
conservation planning in a data poor context: The Cyprus case. J. Nat. Conserv. 70, 126304.
https://doi.org/10.1016/j.jnc.2022.126304
Linke, S., Turak, E., Nel, J., 2011. Freshwater conservation planning: the case for systematic approaches. Freshw.
Biol. 56, 6–20. https://doi.org/10.1111/j.1365-2427.2010.02456.x
Lososová, Z., Axmanová, I., Chytrý, M., Midolo, G., Abdulhak, S., Karger, D.N., Renaud, J., Van Es, J., Vittoz, P.,
Thuiller, W., 2023. Seed dispersal distance classes and dispersal modes for the European flora. Glob. Ecol.
Biogeogr. 32, 1485–1494. https://doi.org/10.1111/geb.13712
Lynch, A.J., 2019. Creating Effective Urban Greenways and Stepping-stones: Four Critical Gaps in Habitat
Connectivity Planning Research. J. Plan. Lit. 34, 131–155. https://doi.org/10.1177/0885412218798334
Lyons, S., Wagner, P.J., Dzikiewicz, K., 2010. Ecological correlates of range shifts of Late Pleistocene mammals.
Philos. Trans. R. Soc. B Biol. Sci. 365, 3681–3693. https://doi.org/10.1098/rstb.2010.0263
Magris, R.A., Pressey, R.L., Weeks, R., Ban, N.C., 2014. Integrating connectivity and climate change into marine
conservation planning. Biol. Conserv. 170, 207–221. https://doi.org/10.1016/j.biocon.2013.12.032
Maiorano, L., Boitani, L., Chiaverini, L., Ciucci, P., 2017. Uncertainties in the identification of potential dispersal
corridors: The importance of behaviour, sex, and algorithm. Basic Appl. Ecol. 21, 66–75.
https://doi.org/10.1016/j.baae.2017.02.005
Maiorano, L., Falcucci, A., Zimmermann, N.E., Psomas, A., Pottier, J., Baisero, D., Rondinini, C., Guisan, A.,
Boitani, L., 2011. The future of terrestrial mammals in the Mediterranean basin under climate change. Philos.
Trans. R. Soc. B Biol. Sci. 366, 2681–2692. https://doi.org/10.1098/rstb.2011.0121
Makino, A., Beger, M., Klein, C.J., Jupiter, S.D., Possingham, H.P., 2013. Integrated planning for land-sea
ecosystem connectivity to protect coral reefs. Biol. Conserv. 165, 35–42.
https://doi.org/10.1016/j.biocon.2013.05.027
Margules, C., S. Sarkar, 2007. Systematic Conservation Planning. Cambridge University Press. Cambridge.
Margules, C.R., Pressey, R.L., 2000. Systematic conservation planning. Nature 405.
https://doi.org/10.1038/35012251
Martínez-Harms, M.J., Balvanera, P., 2012. Methods for mapping ecosystem service supply: a review. Int. J.
Biodivers. Sci. Ecosyst. Serv. Manag. 8, 17–25. https://doi.org/10.1080/21513732.2012.663792
Marx, A.J., Wang, C., Sefair, J.A., Acevedo, M.A. and Fletcher, R.J., Jr. (2020), samc: an R package for connectivity
modeling with spatial absorbing Markov chains. Ecography, 43: 518-527. https://doi.org/10.1111/ecog.04891
Mazor, T., Beger, M., McGowan, J., Possingham, H.P., Kark, S., 2016. The value of migration information for
conservation prioritization of sea turtles in the Mediterranean. Glob. Ecol. Biogeogr. 25, 540–552.
https://doi.org/10.1111/geb.12434
McClintock, B.T., Michelot, T., 2018. momentuHMM: R package for generalized hidden Markov models of animal
movement. Methods Ecol. Evol. 9, 1518–1530. https://doi.org/10.1111/2041-210X.12995
McGarigal, K., Marks, B.J., 1995. FRAGSTATS: spatial pattern analysis program for quantifying landscape
structure. Gen Tech Rep PNW-GTR-351 Portland US Dep. Agric. For. Serv. Pac. Northwest Res. Stn. 122 P
351. https://doi.org/10.2737/PNW-GTR-351
McGuinn, J., Oulès, L., Bradley, H., Mcneill, A., 2017. Effective multi-level environmental governance for a better
implementation of EU environment legislation. https://doi.org/10.2863/406864
McKay, S.K., Cooper, A.R., Diebel, M.W., Elkins, D., Oldford, G., Roghair, C., Wieferich, D., 2017. Informing
Watershed Connectivity Barrier Prioritization Decisions: A Synthesis: Synthesizing Barrier Prioritization. River
Res. Appl. 33, 847–862. https://doi.org/10.1002/rra.3021

112
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

McKay, S.K., Martin, E.H., McIntyre, P.B., Milt, A.W., Moody, A.T., Neeson, T.M., 2020. A comparison of
approaches for prioritizing removal and repair of barriers to stream connectivity. River Res. Appl. 36, 1754–
1761. https://doi.org/10.1002/rra.3684
McRae, B., K. Popper, A. Jones, M. Schindel, S. Buttrick, K. Hall, R.S. Unnasch, J. Platt, 2016. Conserving Nature’s
Stage: Mapping Omnidirectional Connectivity for Resilient Terrestrial Landscapes in the Pacific Northwest.
https://doi.org/10.13140/RG.2.1.4158.6166
McRae, B.H., Dickson, B.G., Keitt, T.H., Shah, V.B., 2008. Using circuit theory to model connectivity in ecology,
evolution, and conservation. Ecology 89, 2712–2724. https://doi.org/10.1890/07-1861.1
McRae, B.H., V.B. Shah, and T.K. Mohapatra. 2013. Circuitscape 4 User Guide. The Nature Conservancy.
http://www.circuitscape.org.
Medrano‐Vizcaíno, P., Grilo, C., Silva Pinto, F.A., Carvalho, W.D., Melinski, R.D., Schultz, E.D., González‐Suárez,
M., 2022. Roadkill patterns in Latin American birds and mammals. Glob. Ecol. Biogeogr. 31, 1756–1783.
https://doi.org/10.1111/geb.13557
Merken, R., Deboelpaep, E., Teunen, J., Saura, S., Koedam, N., 2015. Wetland Suitability and Connectivity for
Trans-Saharan Migratory Waterbirds. PLOS ONE 10, e0135445.
https://doi.org/10.1371/journal.pone.0135445
Meurant, M., Gonzalez, A., Doxa, A., Albert, C.H., 2018. Selecting surrogate species for connectivity conservation.
Biol. Conserv. 227, 326–334. https://doi.org/10.1016/j.biocon.2018.09.028
Michelot, T., Langrock, R., Patterson, T.A., 2016. moveHMM: an R package for the statistical modelling of animal
movement data using hidden Markov models. Methods Ecol. Evol. 7, 1308–1315.
https://doi.org/10.1111/2041-210X.12578
Minor, E.S. and Urban, D.L., 2008. A Graph-Theory Framework for Evaluating Landscape Connectivity and
Conservation Planning. Conservation Biology, 22: 297-307. https://doi.org/10.1111/j.1523-
1739.2007.00871.x
Mitchell, M.G.E., Bennett, E.M., Gonzalez, A., 2013. Linking Landscape Connectivity and Ecosystem Service
Provision: Current Knowledge and Research Gaps. Ecosystems 16, 894–908.
Moilanen, A., Leathwick, J., Elith, J., 2008. A method for spatial freshwater conservation prioritization. Freshw. Biol.
53, 577–592. https://doi.org/10.1111/j.1365-2427.2007.01906.x
Moilanen, A., Possingham, H.P., Wilson, K.A., 2009. Spatial Conservation Prioritization: Past, Present and Future,
in: Moilanen, A., Wilson, K. A., Possingham, H. P. (Eds.), Spatial Conservation Prioritization - Quantitative
Methods and Computational Tools. Oxford University Press.
Moilanen, A., Pouzols, F.M., Meller, L., Veach, V., Arponen, A., Leppänen, J., Kujala, H., 2014. Zonation - Spatial
conservation planning methods and software. Version 4. User Manual. C-BIG Conservation Biology
Informatics Group. Department of Biosciences. University of Helsinki, Finland.
Monaco, A., Genovesi, P. (2014) European Guidelines on Protected Areas and Invasive Alien Species, Council of
Europe, Strasbourg, Regional Parks Agency – Lazio Region, Rome
Montesino Pouzols, F., Toivonen, T., Di Minin, E., Kukkala, A.S., Kullberg, P., Kuusterä, J., Lehtomäki, J.,
Tenkanen, H., Verburg, P.H., Moilanen, A., 2014. Global protected area expansion is compromised by
projected land-use and parochialism. Nature 516, 383–386. https://doi.org/10.1038/nature14032
Naia, M., Hermoso, V., Carvalho, S.B., Brito, J.C., 2021. Promoting connectivity between priority freshwater sites
for conservation in intermittent hydrological systems. Aquat. Conserv. Mar. Freshw. Ecosyst. 31, 1886–1900.
https://doi.org/10.1002/aqc.3564
Nathan, R., 2001. The challenges of studying dispersal. Trends Ecol. Evol. 16, 481–483.
https://doi.org/10.1016/S0169-5347(01)02272-8
Natsukawa, H., & Sergio, F. (2022). Top predators as biodiversity indicators: A meta‐analysis. Ecology
Letters, 25(9), 2062-2075.

113
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

Naumann, G., Alfieri, L., Wyser, K., Mentaschi, L., Betts, R.A., Carrao, H., Spinoni, J., Vogt, J., Feyen, L., 2018.
Global Changes in Drought Conditions Under Different Levels of Warming. Geophys. Res. Lett. 45, 3285–
3296. https://doi.org/10.1002/2017GL076521
Nel, J.L., Reyers, B., Roux, D.J., Dean Impson, N., Cowling, R.M., 2011. Designing a conservation area network
that supports the representation and persistence of freshwater biodiversity. Freshw. Biol. 56, 106–124.
https://doi.org/10.1111/j.1365-2427.2010.02437.x
Nguyen, T., Meurk, C., Benavidez, R., Jackson, B., Pahlow, M., 2021. The Effect of Blue-Green Infrastructure on
Habitat Connectivity and Biodiversity: A Case Study in the Ōtākaro/Avon River Catchment in Christchurch,
New Zealand. Sustainability 13, 6732. https://doi.org/10.3390/su13126732
Noss, R., 1991. Landscape connectivity: different functions at different scales 27–39.
Nuñez, T.A., Lawler, J.J., McRae, B.H., Pierce, D.J., Krosby, M.B., Kavanagh, D.M., Singleton, P.H., Tewksbury,
J.J., 2013. Connectivity Planning to Address Climate Change. Conserv. Biol. 27, 407–416.
https://doi.org/10.1111/cobi.12014
Ogletree, S.S., Powell, R.B., Baldwin, R.F., Leonard, P.B., 2019. A framework for mapping cultural resources in
landscape conservation planning. Conserv. Sci. Pract. 1, https://doi.org/10.1111/csp2.41
Opdam, P., Wascher, D., 2004. Climate change meets habitat fragmentation: linking landscape and
biogeographical scale levels in research and conservation. Biol. Conserv. 117, 285–297.
https://doi.org/10.1016/j.biocon.2003.12.008
Parks, S.A., Holsinger, L.M., Abatzoglou, J.T., Littlefield, C.E., Zeller, K.A., 2023. Protected areas not likely to serve
as steppingstones for species undergoing climate‐induced range shifts. Glob. Change Biol. 29, 2681–2696.
https://doi.org/10.1111/gcb.16629
Pascual-Hortal, L., Saura, S., 2006. Comparison and development of new graph-based landscape connectivity
indices: towards the priorization of habitat patches and corridors for conservation. Landsc. Ecol. 21, 959–967.
https://doi.org/10.1007/s10980-006-0013-z
Peterman, W.E., 2018. ResistanceGA: An R package for the optimization of resistance surfaces using genetic
algorithms. Methods Ecol. Evol. 9, 1638–1647. https://doi.org/10.1111/2041-210X.12984
Petrişor, A.-I., Andronache, I.C., Petrişor, L.E., Ciobotaru, A.-M., Peptenatu, D., 2016. Assessing the Fragmentation
of the Green Infrastructure in Romanian Cities Using Fractal Models and Numerical Taxonomy. Procedia
Environ. Sci., ECOSMART - Environment at Crossroads: Smart Approaches for a Sustainable Development
32, 110–123. https://doi.org/10.1016/j.proenv.2016.03.016
Petrisor, A.-I., Mierzejewska, L., Mitrea, A., Drachal, K., Tache, A.V., 2021. Dynamics of Open Green Areas in
Polish and Romanian Cities during 2006–2018: Insights for Spatial Planners. Remote Sens. 13, 4041.
https://doi.org/10.3390/rs13204041
Pollock, L.J., Thuiller, W., Jetz, W., 2017. Large conservation gains possible for global biodiversity facets. Nature
546, 141–144. https://doi.org/10.1038/nature22368
Popescu, V.D., Rozylowicz, L., Cogălniceanu, D., Niculae, I.M., Cucu, A.L., 2013. Moving into Protected Areas?
Setting Conservation Priorities for Romanian Reptiles and Amphibians at Risk from Climate Change. PLOS
ONE 8, e79330. https://doi.org/10.1371/journal.pone.0079330
Pressey, R.L., 2002. The first reserve selection algorithm—a retrospective on Jamie Kirkpatrick’s 1983 paper. Prog.
Phys. Geogr. 26, 434–441. https://doi.org/10.1191/0309133302pp347xx
Pringle, C., 2006. Hydrologic connectivity: a neglected dimension of conservation biology, in: Crooks, K.R.,
Sanjayan, M. (Eds.), Connectivity Conservation, Conservation Biology. Cambridge University Press,
Cambridge, pp. 233–254. https://doi.org/10.1017/CBO9780511754821.011
Pringle, C., 2003. What is hydrologic connectivity and why is it ecologically important? Hydrol. Process. 17, 2685–
2689. https://doi.org/10.1002/hyp.5145
Proctor, M.F., McLellan, B.N., Strobeck, C., Barclay, R.M.R., 2005. Genetic analysis reveals demographic
fragmentation of grizzly bears yielding vulnerable small populations. Proc. R. Soc. B Biol. Sci. 272, 2409–
2416. https://doi.org/10.1098/rspb.2005.3246
114
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

Prugh, L.R., 2009. An evaluation of patch connectivity measures. Ecological Applications, 19: 1300-1310.
https://doi.org/10.1890/08-1524.1
Pulliam, H.R. (1988). Sources, sinks, and population regulation. The American Naturalist 132 (5): 652–61.
https://doi.org/10.1086/284880
Quaglietta, L., Porto, M., 2019. SiMRiv: an R package for mechanistic simulation of individual, spatially-explicit
multistate movements in rivers, heterogeneous and homogeneous spaces incorporating landscape bias. Mov.
Ecol. 7, 11. https://doi.org/10.1186/s40462-019-0154-8
Railsback, S.F., Grimm, V., 2019. Agent-Based and Individual-Based Modeling | Princeton University Press, 2nd
ed. Princeton University Press.
Razafindratsima, O.H., Brown, K.A., Carvalho, F., Johnson, S.E., Wright, P.C., Dunham, A.E., 2018. Edge effects
on components of diversity and above-ground biomass in a tropical rainforest. J. Appl. Ecol. 55, 977–985.
https://doi.org/10.1111/1365-2664.12985
Reid, A.J., Carlson, A.K., Creed, I.F., Eliason, E.J., Gell, P.A., Johnson, P.T.J., Kidd, K.A., MacCormack, T.J.,
Olden, J.D., Ormerod, S.J., Smol, J.P., Taylor, W.W., Tockner, K., Vermaire, J.C., Dudgeon, D., Cooke, S.J.,
2019. Emerging threats and persistent conservation challenges for freshwater biodiversity. Biol. Rev. 94, 849–
873. https://doi.org/10.1111/brv.12480
Reis, V., Hermoso, V., Hamilton, S.K., Bunn, S.E., Linke, S., 2019. Conservation planning for river-wetland
mosaics: A flexible spatial approach to integrate floodplain and upstream catchment connectivity. Biol.
Conserv. 236, 356–365. https://doi.org/10.1016/j.biocon.2019.05.042
Resasco, J., 2019. Meta-analysis on a Decade of Testing Corridor Efficacy: What New Have we Learned? Curr.
Landsc. Ecol. Rep. 4, 61–69. https://doi.org/10.1007/s40823-019-00041-9
Resasco, J., Haddad, N.M., Orrock, J.L., Shoemaker, D., Brudvig, L.A., Damschen, E.I., Tewksbury, J.J., Levey,
D.J., 2014. Landscape corridors can increase invasion by an exotic species and reduce diversity of native
species. Ecology 95, 2033–2039. https://doi.org/10.1890/14-0169.1
Rocca, F.D., Bogliani, G., Milanesi, P., 2017. Patterns of distribution and landscape connectivity of the stag beetle
in a human-dominated landscape. Nat. Conserv. 19, 19–37.
https://doi.org/10.3897/natureconservation.19.12457
Rudnick, D.A., Ryan, S.J., Beier, P., Cushman, S.A., Dieffenbach, F., Epps, C.W., Gerber, L.R., Hartter, J.,
Jenness, J.S., Kintsch, J., Merenlender, A.M., Perkl, R.M., Preziosi, D.V., Trombulak, S.C., 2012. The role of
landscape connectivity in planning and implementing conservation and restoration priorities.
Rytwinski, T., van der Ree, R., Cunnington, G.M., Fahrig, L., Findlay, C.S., Houlahan, J., Jaeger, J.A.G., Soanes,
K., van der Grift, E.A., 2015. Experimental study designs to improve the evaluation of road mitigation
measures for wildlife. J. Environ. Manage. 154, 48–64. https://doi.org/10.1016/j.jenvman.2015.01.048
Santini, L., Saura, S., Rondinini, C., 2016. Connectivity of the global network of protected areas. Divers. Distrib.
22, 199–211. https://doi.org/10.1111/ddi.12390
Sarkar, M.S., Niyogi, R., Masih, R.L., Hazra, P., Maiorano, L., John, R., 2021. Long-distance dispersal and home
range establishment by a female sub-adult tiger (Panthera tigris) in the Panna landscape, central India. Eur.
J. Wildl. Res. 67, 54. https://doi.org/10.1007/s10344-021-01494-2
Saura, S., Bodin, Ö., Fortin, M.-J., 2014. EDITOR’S CHOICE: Stepping stones are crucial for species’ long-distance
dispersal and range expansion through habitat networks. J. Appl. Ecol. 51, 171–182.
https://doi.org/10.1111/1365-2664.12179
Saura, S., Estreguil, C., Mouton, C., Rodríguez-Freire, M., 2011. Network analysis to assess landscape
connectivity trends: Application to European forests (1990–2000). Ecol. Indic. 11, 407–416.
https://doi.org/10.1016/j.ecolind.2010.06.011
Saura, S., Pascual-Hortal, L., 2007. A new habitat availability index to integrate connectivity in landscape
conservation planning: Comparison with existing indices and application to a case study. Landsc. Urban Plan.
83, 91–103. https://doi.org/10.1016/j.landurbplan.2007.03.005

115
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

Saura, S., Rubio, L., 2010. A common currency for the different ways in which patches and links can contribute to
habitat availability and connectivity in the landscape. Ecography 33, 523–537.
Saura, S., Bertzky, B., Bastin, L., Battistella, L., Mandrici, A., Dubois, G., 2018. Protected area connectivity:
Shortfalls in global targets and country-level priorities. Biological Conservation 219, 53–67.
https://doi.org/10.1016/j.biocon.2017.12.020
Sawyer, S.C., Epps, C.W., Brashares, J.S., 2011. Placing linkages among fragmented habitats: do least-cost
models reflect how animals use landscapes? J. Appl. Ecol. 48, 668–678. https://doi.org/10.1111/j.1365-
2664.2011.01970.x
Schlaepfer, D.R., Braschler, B., Rusterholz, H.-P., Baur, B., 2018. Genetic effects of anthropogenic habitat
fragmentation on remnant animal and plant populations: a meta-analysis. Ecosphere 9, e02488.
https://doi.org/10.1002/ecs2.2488
Schmutz, S., Sendzimir, J. (Eds.), 2018. Riverine Ecosystem Management: Science for Governing Towards a
Sustainable Future. Springer International Publishing, Cham. https://doi.org/10.1007/978-3-319-73250-3
Schumaker, N.H., 1996. Using Landscape Indices to Predict Habitat Connectivity. Ecology, 77: 1210-1225.
https://doi.org/10.2307/2265590
Schumaker, N., Brookes, A., 2018. HexSim: a modeling environment for ecology and conservation. Landsc. Ecol.
33. https://doi.org/10.1007/s10980-017-0605-9
Seliger, C., Zeiringer, B., 2018. River connectivity, habitat fragmentation and related restoration measures. Riverine
Ecosyst. Manag. Sci. Gov. Sustain. Future 171–186.
Silverman, E., 2018. Methodological Investigations in Agent-Based Modelling. https://doi.org/10.1007/978-3-319-
72408-9
Soanes, K., Lobo, M.C., Vesk, P.A., McCarthy, M.A., Moore, J.L., van der Ree, R., 2013. Movement re-established
but not restored: Inferring the effectiveness of road-crossing mitigation for a gliding mammal by monitoring
use. Biol. Conserv. 159, 434–441. https://doi.org/10.1016/j.biocon.2012.10.016
Soanes, K., Rytwinski, T., Fahrig, L., Huijser, M.P., Jaeger, J.A.G., Teixeira, F.Z., Van Der Ree, R., Van Der Grift,
E.A., 2024. Do wildlife crossing structures mitigate the barrier effect of roads on animal movement? A global
assessment. J. Appl. Ecol. 1365-2664.14582. https://doi.org/10.1111/1365-2664.14582
Soininen, J., Bartels, P., Heino, J., Luoto, M., Hillebrand, H., 2015. Toward More Integrated Ecosystem Research
in Aquatic and Terrestrial Environments. BioScience 65, 174–182. https://doi.org/10.1093/biosci/biu216
Sonntag, S., Fourcade, Y., 2022. Where will species on the move go? Insights from climate connectivity modelling
across European terrestrial habitats. J. Nat. Conserv. 126139. https://doi.org/10.1016/j.jnc.2022.126139
Stevens, V.M., Trochet, A., Van Dyck, H., Clobert, J., Baguette, M., 2012. How is dispersal integrated in life
histories: a quantitative analysis using butterflies: Dispersal life-history correlates. Ecol. Lett. 15, 74–86.
https://doi.org/10.1111/j.1461-0248.2011.01709.x
Strayer, D.L., Dudgeon, D., 2010. Freshwater biodiversity conservation: recent progress and future challenges. J.
North Am. Benthol. Soc. 29, 344–358. https://doi.org/10.1899/08-171.1
Suárez, D., Arribas, P., Jiménez-García, E., Emerson, B.C., 2022. Dispersal ability and its consequences
for population genetic differentiation and diversification. Proc. R. Soc. B Biol. Sci. 289, 20220489.
https://doi.org/10.1098/rspb.2022.0489
Suraci, J. P., C. E. Littlefield, C. C. Nicholson, M. C. Hunter, A. Sorensen, and B. G. Dickson. 2023. Mapping
connectivity and conservation opportunity on agricultural lands across the conterminous United States.
Biological Conservation 278: 109896. https://doi.org/10.1016/j.biocon.2022.109896
Taylor, P.D., Fahrig, L., With, K.A., 2006. Landscape connectivity: a return to the basics, in: Crooks, K.R., Sanjayan,
M. (Eds.), Connectivity Conservation. Cambridge University Press, pp. 29–43.
https://doi.org/10.1017/CBO9780511754821.003
Tesson, S.V., Edelaar, P., 2013. Dispersal in a changing world: opportunities, insights and challenges. Mov. Ecol.
1, 10. https://doi.org/10.1186/2051-3933-1-10
116
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

Tickner, D., Opperman, J.J., Abell, R., Acreman, M., Arthington, A.H., Bunn, S.E., Cooke, S.J., Dalton, J., Darwall,
W., Edwards, G., Harrison, I., Hughes, K., Jones, T., Leclère, D., Lynch, A.J., Leonard, P., McClain, M.E.,
Muruven, D., Olden, J.D., Ormerod, S.J., Robinson, J., Tharme, R.E., Thieme, M., Tockner, K., Wright, M.,
Young, L., 2020. Bending the Curve of Global Freshwater Biodiversity Loss: An Emergency Recovery Plan.
BioScience 70, 330–342. https://doi.org/10.1093/biosci/biaa002
Travers, E., Härdtle, W., & Matthies, D. (2021). Corridors as a tool for linking habitats – Shortcomings and
perspectives for plant conservation. Journal for Nature Conservation, 60, 125974.
https://doi.org/10.1016/j.jnc.2021.125974
Tsang, Y.-P., Tingley, R.W., Hsiao, J., Infante, D.M., 2019. Identifying high value areas for conservation:
Accounting for connections among terrestrial, freshwater, and marine habitats in a tropical island system. J.
Nat. Conserv. 50, 125711. https://doi.org/10.1016/j.jnc.2019.125711
Tucker, M.A., Ord, T.J., Rogers, T.L., 2014. Evolutionary predictors of mammalian home range size: body mass,
diet and the environment. Glob. Ecol. Biogeogr. 23, 1105–1114. https://doi.org/10.1111/geb.12194
Tulloch, V.J.D., Atkinson, S., Possingham, H.P., Peterson, N., Linke, S., Allan, J.R., Kaiye, A., Keako, M., Sabi, J.,
Suruman, B., Adams, V.M., 2021. Minimizing cross-realm threats from land-use change: A national-scale
conservation framework connecting land, freshwater and marine systems. Biol. Conserv.
https://doi.org/10.1016/j.biocon.2021.108954
Unnithan Kumar, S., Cushman, S.A., 2022. Connectivity modelling in conservation science: a comparative
evaluation. Sci. Rep. 12, 16680. https://doi.org/10.1038/s41598-022-20370-w
Vale, C.G., Pimm, S.L., Brito, J.C., 2015. Overlooked Mountain Rock Pools in Deserts Are Critical Local Hotspots
of Biodiversity. PLOS ONE 10, e0118367. https://doi.org/10.1371/journal.pone.0118367
Valladares, F., Sanchez-Gomez, D., Zavala, M.A., 2006. Quantitative estimation of phenotypic plasticity: bridging
the gap between the evolutionary concept and its ecological applications. J. Ecol. 94, 1103–1116.
https://doi.org/10.1111/j.1365-2745.2006.01176.x
Vannote, R.L., Minshall, G.W., Cummins, K.W., Sedell, J.R., Cushing, C.E., 1980. The River Continuum Concept.
Can. J. Fish. Aquat. Sci.
Venegas-Li, R., Levin, N., Possingham, H., Kark, S., 2018. 3D spatial conservation prioritisation: Accounting for
depth in marine environments. Methods Ecol. Evol. 9, 773–784. https://doi.org/10.1111/2041-210X.12896
Vittoz, P., Engler, R., 2007. Seed dispersal distances: a typology based on dispersal modes and plant traits. Bot.
Helvetica 117, 109–124. https://doi.org/10.1007/s00035-007-0797-8
Wang, F., McShea, W.J., Li, S., Wang, D., 2018. Does one size fit all? A multispecies approach to regional
landscape corridor planning. Divers. Distrib. 24, 415–425. https://doi.org/10.1111/ddi.12692
Wang, F., Winkler, J., Viña, A., McShea, W.J., Li, S., Connor, T., Zhao, Z., Wang, D., Yang, H., Tang, Y., Zhang,
J., Liu, J., 2021. The hidden risk of using umbrella species as conservation surrogates: A spatio-temporal
approach. Biol. Conserv. 253, 108913. https://doi.org/10.1016/j.biocon.2020.108913
Ward, J.V., 1989. The Four-Dimensional Nature of Lotic Ecosystems. J. North Am. Benthol. Soc. 8, 2–8.
https://doi.org/10.2307/1467397
Ward, J.V., Stanford, J.A., 1995. The serial discontinuity concept: Extending the model to floodplain rivers. Regul.
Rivers Res. Manag. 10, 159–168. https://doi.org/10.1002/rrr.3450100211
Ward, M., Saura, S., Williams, B., Ramírez-Delgado, J.P., Arafeh-Dalmau, N., Allan, J.R., Venter, O., Dubois, G.,
Watson, J.E.M., 2020. Just ten percent of the global terrestrial protected area network is structurally
connected via intact land. Nat. Commun. 11, 4563. https://doi.org/10.1038/s41467-020-18457-x
Watts, A., Schlichting, P., Billerman, S., Jesmer, B., Micheletti, S., Fortin, M.-J., Funk, C., Hapeman, P., Muths, E.,
Murphy, M., 2015. How spatio-temporal habitat connectivity affects amphibian genetic structure. Front. Genet.
6. https://doi.org/10.3389/fgene.2015.00275
Weathers, K.C., Cadenasso, M.L., Pickett, S.T.A., 2001. Forest Edges as Nutrient and Pollutant Concentrators:
Potential Synergisms between Fragmentation, Forest Canopies, and the Atmosphere. Conserv. Biol. 15,
1506–1514. https://doi.org/10.1046/j.1523-1739.2001.01090.x
117
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

Whitmee, S., Orme, C.D.L., 2013. Predicting dispersal distance in mammals: a trait-based approach. J. Anim. Ecol.
82, 211–221. https://doi.org/10.1111/j.1365-2656.2012.02030.x
Whittington, J., Hebblewhite, M., Baron, R.W., Ford, A.T., Paczkowski, J., 2022. Towns and trails drive carnivore
movement behaviour, resource selection, and connectivity. Mov. Ecol. 10, 17. https://doi.org/10.1186/s40462-
022-00318-5
Wiens, J.A., 2002. Riverine landscapes: taking landscape ecology into the water. Freshw. Biol. 47, 501–515.
https://doi.org/10.1046/j.1365-2427.2002.00887.x
Wilensky, U., 1999. NetLogo.
Williams, R.J., Dunn, A.M., Mendes da Costa, L., Hassall, C., 2021. Climate and habitat configuration limit range
expansion and patterns of dispersal in a non-native lizard. Ecol. Evol. 11, 3332–3346.
https://doi.org/10.1002/ece3.7284
WWF Tigers Alive, 2020. Landscape connectivity science and practice: Ways forward for large ranging species
and their landscapes. Workshop Report, WWF International.
Zeller, K.A., Jennings, M.K., Vickers, T.W., Ernest, H.B., Cushman, S.A., Boyce, W.M., 2018. Are all data types
and connectivity models created equal? Validating common connectivity approaches with dispersal data.
Divers. Distrib. 24, 868–879. https://doi.org/10.1111/ddi.12742
Zeller, K.A., McGarigal, K., Whiteley, A.R., 2012. Estimating landscape resistance to movement: a review. Landsc.
Ecol. 27, 777–797. https://doi.org/10.1007/s10980-012-9737-0
Zhang, W., 2018. Fundamentals of network biology, 1st edition. ed. World Scientific Publishing Europe Ltd, New
Jersey London Singapore Beijing Shanghai Hong Kong Taipei Chennai Tokyo.

118
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

Annex S1. Survey of connectivity projects


S1.1 Questions

Survey of ecological connectivity projects in


Europe

The NaturaConnect project (https://naturaconnect.eu) is an EU-funded Research and


Innovation action that develops knowledge, tools, and capacity- building to support
the implementation of a coherent network of protected areas across Europe - the
Trans-European nature network (TEN-N). The present survey aims to collect
information on ecological connectivity projects in Europe undertaken at Regional to
National and Pan-European levels. Projects may include public and private
conservation action plans and strategies, connectivity conservation and restoration
measures, research and innovation projects, etc. The inputs will be used to produce a
public repository of connectivity projects that can support knowledge sharing.

Contact: The NaturaConnect Project ([email protected])

* Indicates required question

Section 1: Project information

1. Name of the project *

2. Acronym

3. Responsible institution(s) *

119
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

4. Country or countries of application *

5. Name of contact person(s) *


Note: This information will not be public

6. E-mail *
Note: This information will not be public

7. Other participating institutions

8. Project website

9. Abstract

10. Start and end date (e.g. 2010-2014) *

11. Funding sources *

Check all that apply.

Nature conservation funds from National and/or Regional Administrations

National and/or regional Research & Innovation funds

Development funds from National and/or Regional administrations


120
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

LIFE programme

European research & innovation funds (E.g., Horizon, Biodiversa, etc.)

European funds associated to other sustainability policies (e.g., climate,


agriculture, Interreg, etc)

Cohesion funds

Biodiversity offsets

Private funds

Other:

12. Name of funding programme(s)

Section 2: Project scope and users

13. Connectivity goals. Please select only those categories *


explicitly addressed in the project.

Check all that apply.

Protection of one species or a particular group of species

Protection of multiple species (non-species-specific target)

Connectivity between protected areas

Connectivity between specific habitat types (forests, EU directives habitats, etc)

Small landscape features (e.g. hedgerows and tree lines, riparian forest
remnants, green urban spaces, etc)

121
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

Other:

14. In case the project is focused on selected taxa, please indicate which.

15. Has the project been commissioned by a competent *


administration (yes/no)? If yes, please specify which.

16. Regardless of who commissioned the project, who are the target users of the
results?

Check all that apply.

No specific user identified

Regional and/or local administration(s)

National administration

European institutions

Private organizations, foundations and/or NGOs

Protected area managers


Other:

17. What is the spatial scope of the project? *

Check all that apply.

Local (e.g., covering one or several municipalities, or a specific infrastructure)

Sub-national (spatially comprehensive for one or several administrative regions)

National

Transboundary (connecting across 2 or more countries)


122
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

Other multiple countries

Pan-European

Other:

18. Will the project be used for supporting any of the following *
policies?

Check all that apply.

Biodiversity conservation policy and strategies

Green and Blue Infrastructure

Spatial planning of protected areas

Restoration plans and/or actions

Climate policies

Sustainable development

Human health
Other:

19. Thematic scope: does the project aims to enhance any of *


the following?

Check all that apply.

Increase the permeability of linear infrastructures such as roads or railways

Ecosystem restoration

Ecological corridors (i.e., continuous corridors or stepping stones)

123
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

New protected areas and/or expansion of other existing protected areas

Adaptation to climate change

Urban green spaces

Other:

20. Biogeographical region(s) included

Check all that apply.

Alpine

Anatolian

Arctic

Atlantic

Black sea

Boreal

Continental

Macaronesia

Mediterranean

Pannonian

Steppic

21. Ecosystem type(s) included *

Check all that apply.

Forests

124
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

Shrublands

Grasslands

Wetlands

Coastal areas

Rivers

Agricultural areas

Urban and periurban


Other:

Section 3: Technical approaches used in the project

22. Approaches for assessing connectivity *

Check all that apply.

Analyses using species movement data (e.g., from tracked individuals)

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

Analyses using species distribution models

Land cover and land-use analyses

Analysis of infrastructures (e.g. roads, railway) and urban sprawl

Expert-based
Other:

23. Does the project provide spatially explicit information on: *

125
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

Check all that apply.

Locations for ecological corridors

Locations for stepping stones

Locations to increase permeability of linear infrastructures (e.g., wildlife passes)


Locations for habitat restoration

Locations for proposed protected areas

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.

25. Does the project consider other benefits in addition to *


biodiversity conservation?

Check all that apply.

Climate regulation (e.g. carbon sequestration, cooling effect)

Water quality

Prevention of natural hazards (e.g., fires, floods)

Pollination

Recreation

Soil quality

None
Other:

126
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

26. Does the project identify any potential negative impacts of *


connectivity?

Check all that apply.

Increased human-wildlife conflicts

Increased risk of natural hazards (e.g., fires)

Increased spread of invasive species

Increased risk of spreading diseases

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.

S1.3 Distribution and response rates

We began by compiling a comprehensive list of ecological connectivity projects by


conducting internet searches, utilizing European project databases such as Network Nature,
Biodiversa, and LIFE Public Database, and seeking suggestions from NaturaConnect's
consortium members. Our initial list comprised a total of 105 projects. Subsequently, we
distributed a survey via email to all contacts on our list, requesting additional suggestions for
127
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

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.

S1.4 Response processing

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”.

128
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

Annex S2. Connectivity workshop Miro boards examples

129
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

Figure S2.1: Miro board from the “Terrestrial and Freshwater Habitats” breakout group on day 1 of the workshop.

130
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

131
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

Figure S2.2: Miro board from the “Ecosystem Processes & Services” breakout group on day 1 of the workshop.

132
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

133
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

Figure S2.3: Miro board from the “Planning & Management of Multifunctional Corridors” breakout group 1 on day 2 of the workshop.

134
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

135
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

Figure S2.4: Miro board from the “Human Infrastructure & Land Use Impacts” breakout group on day 2 of the workshop.

136
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

Annex S3. Archetypes


S3.1. Archetypes definition
We considered all European tetrapods (amphibians, reptiles, breeding birds, mammals)
naturally occurring in Europe. We followed the taxonomy proposed by GBIF (www.gbif.org).
For each species, we gathered trait data directly from online repositories or by using the R
package ‘’traitdata” (Table S3.1) which includes most of the ecological trait datasets currently
available. For each class of tetrapods, we considered all the relevant datasets to gather as
much data as possible (raw data stored in https://zenodo.org/doi/10.5281/zenodo.10842629).
Given the ecological differences between the different classes of tetrapods, as well as the
different availability of traits in existing databases, to identify the different archetypes we
conducted separate analyses for functionally homogeneous groups, mostly selected at the
level of orders or sub-orders. Among mammals, the order Chiroptera (bats) was considered a
priori to be a cluster due to its specialisation for flight, while all other species were analysed
together in a single group. For amphibians, we considered frogs (Anura) and salamanders
(Caudata) as two separated groups, and for reptiles, we focused our analyses on three groups
(turtles, snakes, lizards). No detailed subdivision was applied to birds.
We selected ecological traits related to dispersal capacity and movement dynamics like
morphological (e.g., adult body mass, snout to vent length) and life-history traits (e.g., litter or
clutch size, maximum longevity) that can be used as proxies for each species’ spatial
requirements. We also considered traits directly related to dispersal distance and home range
size (Santini et al., 2013; Tamburello et al., 2015; Weeks et al., 2022; Alencar and Quental,
2023). As a proxy of species size, we used body mass for non-volant mammals, birds, turtles,
and snakes, the snout to vent length for frogs, salamanders, and lizards, while for bats we
considered the forearm length. For breeding birds, we considered the beak depth and width
as proxy for their trophic ecology. Among reproductive traits, we considered the litter size for
all analyzed groups except for bats. While, only for salamanders, we considered the
reproductive modalities (viviparous, larval stages, direct development). We used female
maturity as a proxy for generation length for all tetrapods except reptiles, and the number of
incubation days for turtles and lizards. Finally, we considered the maximum longevity for all
mammals, birds, turtles, and snakes. In addition, for amphibians we also used the type of
locomotion, while only for lizards we included the type of substrate used by the species
(arboreal, terrestrial, saxicolous).
Since dispersal data were not available for most species, we predicted dispersal distances
following Weeks et al. (2022) for birds, and Santini et al. (2013) for mammals. Different studies
have suggested using migration distances as a proxy for dispersal in birds. However, the link
between migration distances and dispersal distances is not clear nor direct. In fact, although
migratory species tend to disperse more, this may be related to their high flight efficiency rather
than an effect of migratory movements (Claramunt, 2021). Additionally, migrant birds often
have strong philopatric tendencies which could reduce or potentially nullify any effect of
migratory movements on dispersal distance (Chu and Claramunt, 2023). We also imputed
missing data on home range following Tamburello et al. (2015) for both birds and reptiles.

137
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

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).

Table S3.1: Trait databases considered for the archetypes analysis.

Database Taxonomic Reference


coverage
AmnioteDB Amphibians, Myhrvold, N. P., Baldridge, E., Chan, B., Sivam, D., Freeman, D.
birds, L., & Ernest, S. M. (2015). An amniote life‐history database to
mammals, perform comparative analyses with birds, mammals, and reptiles:
reptiles Ecological Archives E096‐269. Ecology, 96(11), 3109-3109.
https://esapubs.org/archive/ecol/E096/269/
Amphibian traits Amphibians Huang, N., Sun, X., Song, Y., Yuan, Z., & Zhou, W. (2023).
database Amphibian traits database: A global database on morphological
traits of amphibians. Global Ecology and Biogeography, 32(5),
633-641.

138
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

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

139
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

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

Body mass (g) 170875.0 35383.2 7518.3 234.5 23.4

Maximum longevity (d) 9909.8 8030 7300.0 4015.0 1761.1

Female maturity (d) 883.8 548 429.6 305.2 51.4

Litter size (n) 1.9 1.4 3.7 4.4 5.2

Dispersal (km) 45.8 6.5 11.6 0.8 0.1

Home Range (km2) 289.1 2.1 8.0 0.1 <0.001

Table S3.3: Median values of traits used to define the three European bats archetypes.

140
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

1 2 3

Maximum longevity (y) 29 26 21

Female maturity (d) 456 502 730

Forearm Length (cm) 37.6 40.4 47.7

Home Range (km2) 2.61 54.4 33

Dispersal (km) 0.68 50 1.46

Table S3.4: Median values of traits used to define the four European birds archetypes.

1 2 3 4

Maximum longevity (y) 45.0 22.9 16.2 13.1

Female maturity (d) 1277.9 726.9 545.2 364.2

Beak Width (mm) 18.2 12.9 4.3 4.7

Beak Depth (mm) 27.9 14.3 5.7 6.1

Kipps Distance (mm) 205.7 122.6 83.2 28.7

Body mass (g) 3846.9 866.0 132.5 36.0

Age of first breeding (y) 3 2 2 1

Long distance migrant 0 0 1 0

Dispersal (km) 26.5 23.0 38.7 15.1

Home range (km2) 147.7 12.8 0.9 0.1

Table S3.5: Median values of traits used to define the three European frogs archetypes.

1 2 3

141
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

Snout to vent length (mm) 69.1 44.7 41

Sexual maturity (y) 3 1.5 1

Number of eggs/clutch 4000 350 1400

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

Snout to vent length (mm) 102.3 63.3 54.6

Sexual maturity (y) 3 2.9 3.5

Number of eggs/clutch 28 250 10

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

142
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

Body mass (g) 1430 469.1

Incubation (d) 78 91.8

Clutch size (n) 5 9

Maximum longevity (y) 127 72.3

Home range (km2) 0.2 0.04

Table S3.8: Median values of traits used to define the three European snakes archetypes.

1 2 3

Body mass (g) 2209.5 196 104

Clutch size (n) 8 10 4

Maximum longevity (y) 5.8 19 5.9

Home range (km2) 1.2 0.06 0.03

Table S3.9: Median values of traits used to define the three European lizards archetypes.

1 2 3

Snout to vent length (cm) 14.5 6.3 6.05

Incubation period (d) 61.8 47 48

Clutch size (n) 8.9 3.5 3.7

Arboreal 0 0 0

Terrestrial 1 0 1

Saxicolous 0 1 0

143
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

S3.2. Habitat preferences


Understanding habitat composition is crucial for connectivity and spatial conservation
planning. Therefore, we used the habitat preferences of European tetrapods to relate
archetypes to a specific environmental context. Following the IUCN habitat classification
scheme (Version 3.1), we identified several natural and artificial habitat types in Europe:
Forest, Savanna, Shrublands, Grasslands, Wetlands, Rocky areas, Desert, and Artificial. For
“Artificial”, given the importance of anthropogenic factors especially upon habitat
fragmentation and connectivity, we further followed the second level of IUCN classification
distinguishing different artificial habitat types: Pastureland, Arable Land, Plantations, Rural
Garden, and Urban Areas. In our final classification, we reduced the possible number of habitat
types merging the classes Grassland, Savanna, Pastureland, Desert, and Rocky Areas in
“Open natural”; while Rural Garden was associated with “Plantations” habitat class (Fig. S3.1).
We maintained the original classification for Wetlands, Arable land, Shrublands, Urban areas,
and we defined Forest as “Close Natural”.
For each group of tetrapods, we strived to find a generalised habitat type representing most of
the species included. However, from our analyses, we found that most species had species-
specific habitat requirements with idiosyncratic combinations of habitats that cannot be
generalised without losing most of the variance (Fig. S3.2). Therefore, we were forced to
reclassify European habitats in two very broad classes: Natural and Artificial. We included in
“Natural” habitats the following classes: Close Natural, Shrublands, Wetlands, Open Natural,
and Pastureland; while Arable land, Plantations, and Urban areas were included into “Artificial”
habitats (Fig. S3.1).

Figure S3.1: Aggregation of IUCN habitat classes (level 1) into natural and artificial classes.

144
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

Figure S3.2: Habitat preferences of European tetrapods.

145
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

Annex S4: Geospatial Data Sources for


Europe
Authoring
Sector Data Source Name Author Data location
organization

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

Forest management Oostdijk, Saskia Vrije Universiteit https://dataverse.nl/dataset.xhtml?p


map for Europe et al. Amsterdam ersistentId=doi:10.34894/HQIJN5

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

146
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

Global Biodiversity
Species
Information Facility
Data
(GBIF) GBIF GBIF https://www.gbif.org/

European network of https://www.eea.europa.eu/en/datah


protected sites Natura European ub/datahubitem-view/6fc8ad2d-
2000 EEA Environment Agency 195d-40f4-bdec-576e7d1268e4
Protected
Areas https://www.eea.europa.eu/en/datah
Nationally designated European ub/datahubitem-view/f60cec02-
areas (CDDA) EEA Environment Agency 6494-4d08-b12d-17a37012cb28

147
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

More information about the project:

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.

NaturaConnect aims to design and develop a blueprint for a truly coherent


Trans-European Nature Network (TEN-N) of conserved areas that protect
at least 30% of land in the European Union, with at least one third of it under
strict protection. Our project unites universities and research institutes,
government bodies and non-governmental organizations, working together
with key stakeholders to create targeted knowledge and tools, and build the
capacity needed to support European Union Member States in realizing an
ecologically representative, resilient and well-connected network of
conserved areas across Europe.

www.naturaconnect.eu

NaturaConnect receives funding under the European Union’s


Horizon Europe research and innovation programme under
grant agreement number 101060429.

148

You might also like