# Manifesto for multidimensional ecological networks: a perspective to better account for the complexity of habitat loss and fragmentation

Romain Sordello, Jérémie Cornuau, Aurélie Coulon, Fabien Paquier, Yorick

Reyjol, Sylvie Vanpeene

► To cite this version:

Romain Sordello, Jérémie Cornuau, Aurélie Coulon, Fabien Paquier, Yorick Reyjol, et al.. Manifesto for multidimensional ecological networks: a perspective to better account for the complexity of habitat loss and fragmentation. Landscape Ecology, 2025, 40 (6), pp.110. 10.1007/s10980-025-02123-y.
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# Manifesto for multidimensional ecological networks: a perspective to better account for the complexity of habitat loss and fragmentation

Romain Sordello · Jérémie Cornuau · Aurélie Coulon · Fabien Paquier · Yorick Reyjol · Sylvie Vanpeene

Received: 21 January 2025 / Accepted: 5 May 2025 / Published online: 27 May 2025
© The Author(s) 2025

## Abstract

*Context* Ecological networks (i.e. sets of connected natural habitats) emerged in the 1980s and have been implemented worldwide. They have been developed as a land-use planning tool that enables to ensure both habitat amount and landscape connectivity in response to ground based physical anthropogenic drivers (e.g. forest cuttings, roads, dams).

*Objective* We wanted to highlight some gaps in this conception of habitat loss and fragmentation that led to potential lacks in the current methods and approaches to design and manage ecological networks, with a view to suggest a way of improving the

situation for a more comprehensive consideration of biodiversity.

*Methods* We compared new scientific knowledge about habitat loss and fragmentation with a critical assessment of feedback from ecological networks, in order to highlight bottlenecks. On this basis we formulated proposals for a new framework and discussed the operational implications for designing and managing broader ecological networks.

*Results* Airscape or soils—key compartments for activities and movements of species—appear to be rarely considered in ecological networks, as well as sensory pollutants—i.e. artificial light at night, man-induced sounds and odors—while they have been demonstrated to disturb nightscapes, soundscapes and odorscapes.

**Supplementary Information** The online version contains supplementary material available at https://doi.org/10.1007/s10980-025-02123-y.

R. Sordello (✉) · Y. Reyjol
PatriNat (OFB-MNHN), Museum national d'Histoire naturelle, Chez Le CBNBP, 61 Rue Buffon CP53, 75005 Paris, France
e-mail: romain.sordello@mnhn.fr

J. Cornuau
OïkoLab, TerrOïko, 2 Place Dom Devic, 81540 Soreze, France

A. Coulon
Centre d'Ecologie et des Sciences de la Conservation (CESCO), Museum national d'Histoire naturelle, Centre National de la Recherche Scientifique, Sorbonne Université, CP 135, 57 Rue Cuvier, 75005 Paris, France

A. Coulon
CEFE, Univ Montpellier, CNRS, EPHE, IRD, Montpellier, France

F. Paquier
Office Français de la Biodiversité. Service Mobilisation Et Accompagnement Des Entreprises Et Des Territoires - Direction Acteurs Et Citoyens, Pôle de Montpellier-Le Tabella, 125 Impasse Adam Smith, 34470 Pérols, France

S. Vanpeene
INRAE - Aix Marseille Université, UMR RECOVER, 3275 route Cezanne, CS, 40061. 13182 Aix-en-Provence, France
**Conclusion** We invite landscape designers and decision-makers to transition to multidimensional ecological networks, developing specific infrastructure against habitat loss and fragmentation, in particular an aerial infrastructure for flying species, a brown infrastructure for soil wildlife, a dark infrastructure for darkness at night, a noise-free infrastructure and an olfactory infrastructure to deal with anthropogenic odors.

**Keywords** Corridors · Light pollution · Noise · Olfactory · Underground · Windfarms

## Introduction

For decades, territory planning—whether through urban, intensive agriculture or transport infrastructure spreading—has led to a deep transformation of land-cover, resulting in a vast decrease of natural habitats (Kleinschroth et al. 2019; Simkin et al. 2022). Habitat loss reduces species ability to complete their life cycle in habitat patches, affecting foraging (Buchmann et al. 2012), breeding (Homan et al. 2004) and survival (Kotze et al. 2021). Habitat degradation, rather than loss, can also be detrimental (Fontúrbel et al. 2022), as it impacts demography (Moreno-Opo 2020), distribution (Botello et al. 2015) and viability (Zhang et al. 2018) of populations and causes damages at the ecosystem level, e.g. on species diversity (Vellend 2003) or species interactions (Spiesman and Inouye 2013).

In addition to being reduced in area, relictual habitat patches are becoming disconnected from each other, generating an anthropogenic habitat fragmentation (Fahrig 2003). Habitat fragmentation may also be due to specific barriers with no significant loss of habitat amount—such as fences or walls in farmland or gardens (McInturff et al. 2020). Habitat fragmentation has many consequences for species and ecosystems (Saunders et al. 1991). It can generate direct or indirect attrition of individuals (e.g. roadkills) which have to move through inhospitable environments between habitat patches (Medrano-Vizcaíno et al. 2022) and then affect demography (Glista et al. 2008). Habitat fragmentation has also long-term effects by limiting gene flow between populations, for animals (Keyghobadi et al. 2005) or plants (Cristóbal-Pérez et al. 2021). This may lead to isolated

populations, eventually causing their extinction due to inbreeding depression and stochastic demographic factors (Andersen et al. 2004). Lastly, fragmentation can impede the spatial adaptation of species to changing climatic conditions (Opdam and Wascher 2004) and condemn certain populations under unfavorable climates (Holyoak and Heath 2016).

Habitat loss and fragmentation have been identified as major threats to biodiversity (Moulatlet et al. 2021; Kuipers et al. 2021). As a consequence, maintaining substantial areas of optimal habitats and improving landscape connectivity (i.e. the degree to which a landscape allows or impedes movements among resource patches (Taylor et al. 1993)) have been regarded as key measures to enhance biodiversity (Hyseni et al. 2021; Timmers et al. 2022; Cerda-Peña and Rau 2023). Connectivity appears to be a factor that facilitates dispersal, migration and gene flow in fauna—for various taxa: mammals (Coulon et al. 2004), amphibians (Todd et al. 2009), birds (Radvan et al. 2023), insects (Ouin et al. 2008)—and in flora (Sork and Smouse 2006).

To address these concerns, a new concept based on landscape ecology principles, has emerged in the 1980s: ecological networks (ENs), defined as sets of natural or semi-natural habitats whose diversity, composition and configuration enable to ensure both activities and movements of species within the landscape (Jongman 1995; Bennett 2004). ENs are composed of cores closely-spaced or physically linked by linear habitats named corridors (such as hedgerows, watercourses, riparian forests) (Hilty et al. 2006). Role of corridors in facilitating the movement of species in the landscape has been observed or demonstrated by numerous studies (Beier and Noss 1998; Bolger et al. 2001; Gutiérrez-Chacón et al. 2020).

Ecological networks has brought two major innovations in nature conservation while allowing a degree of human use of the landscape: 1/ More than just a network of protected areas: they go beyond the preservation of habitat amount to include landscape connectivity as a goal (Battisti 2003); 2/ A generalized land management tool: they can be used in both natural or urban contexts (Samways and Pryke 2016; Hamid and Tan 2017). Moreover, in the range of adaptation measures to be implemented in the face of climate change, ENs are among those most recommended by scientists (Heller and Zavaleta 2009; Prober et al. 2019). Indeed, they enable species to
adjust their range in altitude or latitude (Rüter et al. 2014; Su et al. 2021).

Given their benefits, ENs have long been promoted by nature conservation policies worldwide against habitat loss and fragmentation (Hilty et al. 2019; Keeley et al. 2019). Following the Convention on Biological Diversity of Rio 1992, the EU Member States have committed themselves to the constitution of a pan-European ecological network, to enable the conservation of ecosystems and habitats in Europe, to be implemented at the national, regional or local levels. In 2002, the Johannesburg Summit provided an action plan which aim is to « promote the establishment of ecological networks and corridors at the national and regional levels », to ensure the conservation and sustainable use of biodiversity (target 44.g).<fnref n="1" /> Then, Aichi Biodiversity Targets 2011–2020 included the following objective: « By 2020, [...] degradation and fragmentation is significantly reduced. » (target 5).<fnref n="2" /> To meet these objectives, many states of the world have implemented public policies for ENs (Jongman et al. 2004) that can take various names—such as ‘Trame verte et bleue’ (France), ‘Biotopverbund’ (Germany), ‘Wlaams Ecologisch Netwerk’ (Belgium Flanders), ‘Green Network’ (Estonia), ‘National Ecological Network’ (Hungary). These public policies often result in maps of natural and semi-natural habitats and connections to be preserved and restored. For instance, in France 15,7 million hectares of core areas of habitats (29% of French continental territory) and 50,717 km of linear corridors have been identified (Billon and Sordello 2017). Therefore, ENs provide a decision-making tool for land-use planning that makes it possible to take biodiversity into account in political trade-offs at various levels, from local to international (Wickham et al. 2010; Pietsch 2018), particularly in countries where landscapes are constrained by all types of artificialisation.

EN is a flexible concept and approaches for EN design and management are manifold. The theoretical basis of ENs (i.e. including habitat amount and connectivity) offers the greatest opportunities to design multidimensional and comprehensive ENs,

considering all types of environments and all types of anthropogenic disturbance. However in practice, several vital dimensions to biodiversity functioning—such as aerial/soil habitats or sensory disturbance—seem to be regularly omitted. In the context of biodiversity shrinking on Earth, immediate action is urgently needed to deal with these issues—and ENs implementation make part of them—given that a quarter of terrestrial species is estimated to live in soils (Guerra et al. 2021), that 64% of invertebrates organisms are active at night (Holker et al. 2010) and that Earth’s natural soundscapes are vanishing (e.g. 70% of the soundscapes recorded by Bernie Krause since the 1960s would have disappeared today). Even if scientific knowledge on these topics remains incomplete, we are sufficiently informed to take action now.

In this article, we want to highlight gaps both in the conceptualization and the implementation of ENs in various aspects. Our aim is to address both scientists and practitioners communities:

- researchers (e.g. landscape ecology, sensory ecology): identifying overlooked characteristics in the conceptualisation of habitat loss and fragmentation and highlighting key research gaps that would require further primary studies.

- landscape designers and decision-makers: inviting them to transition to more comprehensive ENs implementation, by providing them with operational guidance and examples of projects already completed.

First, we will demonstrate the limitations of the current implementation of ENs worldwide, that are based on the usual conception of habitat loss and fragmentation. Second, we will organize the argument on the requirement to overcome current bottlenecks in EN implementations, stressing the need to consider new forms of habitat loss/fragmentation in the light of scientific knowledge development. Third, we will propose to implement multidimensional ecological networks by suggesting that the concept of *Green infrastructure* could be a solution and by naming six specific infrastructure to be developed: aerial, soil, dark, man-induced noise-free, and anthropogenic odour-free. Fourth, we will discuss the operational implications for designing and managing ENs within the broader conception we recommend and we recognized a framework for this process. In this article, France is the starting point for our reflection because it is the country we know best, and we use this case

<footnote n="1">¹ https://ise.unige.ch/isdd/IMG/pdf/plan-action_johannesburg.pdf</footnote>

<footnote n="2">² https://www.cbd.int/doc/strategic-plan/targets/T5-quick-guide-en.pdf</footnote>
study to generalise, because this proposal to extend ENs is valid for many other countries, particularly all urbanised and industrialised countries.

## Limitations of the current implementations of ecological networks

Considering the various methodological guides for practitioners to design ENs and the feedback reports (e.g. Bennett et al. Bennett and Mulongoy 2006; Nau-mann et al. 2011; Science for Environment Policy 2012; Sordello et al. 2017; European Commission 2019; Hilty et al. 2020), some potential limits can be addressed about how ENs are currently designed: ENs seem to mostly focus on above ground ecosystems and mainly consider the physical drivers of hab-itat loss and fragmentation.

First, ENs are often implemented at various spatial levels, from the municipality to an entire continent, including regions or states. This multiscale process enables to take into account a great diversity of spe-cies' life history traits (e.g. dispersal distances, home range sizes). It allows to cover a wide range of spatial variability in ecological processes, from large move-ments (e.g. migration of a big mammal over several kilometers) to very short movements (e.g. a small mammal that moves over a few dozen square meters). In contrast, ENs usually forget the temporal axis of ecological processes; e.g. circadian rhythm based on day/night cycle.

Second, ENs generally include several sub-net-works according to different types of habitats which are often described in terms of terrestrial/aquatic compartments and patterns of vegetation; e.g. for-ests, hedges, wetlands, grasslands. ENs also deal with watercourses, to preserve or restore aquatic and sedimentary flows. As a result, ENs are mainly interested in what happens on the ground while aer-ial and underground habitats are more ignored. To take two examples: 1/ Voigt et al. (2018) underline that conservation strategies for bats focus almost exclusively on ground-based habitats (habitats for foraging, hibernation or reproduction) while aero-sphere is under-represented; 2/ Mony et al. (2022) highlight that microorganisms—closely linked to soils—form the dominant proportion of biodiversity

on earth while they are systematically forgotten in ENs policies.

ENs have mostly emerged to respond to habitat fragmentation caused by linear transportation infra-structure. The latter have been considered for sev-eral decades as the most evident landscape barrier (Forman and Alexander 1998; Coffin 2007), maybe because of their linear features that clearly generate a structural landscape cutting. Urbanization has also been identified as a major source of habitat loss and fragmentation for a long time (e.g. Dickman 1987), making ENs a regularly used tool in urban plan-ning to create a spatial coherence in urban green-ing (Voghera and La Riccia 2019). Farming and for-estry intensification can also lead to ruptures in the landscape (e.g. destruction of hedges, clear-cutting) (e.g. Van Apeldoorn et al. 1992; Grashof-Bokdam 1997) and ENs are used to dealing with these issues (Schmidt and Hauck 2018). Dams, weirs and other obstacles on watercourses were also very early the object of particular attention (e.g. Gehrke et al. 2002). The main drivers considered in ENs are hence currently ‘physical’ landscape features while sensory pollutants are barely included.

Sensory pollutants include anthropogenic distur-bances caused by waves or volatil compounds, such as artificial light at night (ALAN), man-induced noise and odors. There are very few international conservation strategies to reduce ALAN at the land-scape level (Schroer et al. 2020; Jägerbrand and Bouroussis 2021). For example, an assessment of French regional ENs performed in 2015 showed that they did not include ALAN (Sordello 2015). Regarding noise pollution the European Commis-sion has recommended the identification of qui-tness areas since 2014 (European Environment Agency 2014) but a real promotion of a noise-free EN is still lacking. Even less actions exist concern-ing odorscapes. Indeed, attention paid to odors is currently limited to ecological engineering meas-ures, for instance to reduce wildlife damage; manip-ulating odours to protect crops from unwanted ani-mals (Baker et al. 2008; Hansen et al. 2017).
Various dimensions that ecological networks
should embed

Air and soil compartments

Many species fly, especially birds, bats, insects and all aerial planktons (Fig. 1). Some of them may follow well identified ‘flight routes’ (Barchi et al. 2013) and some of these flyways can concentrate thousands of individuals (Fülöp et al. 2018). Yet, the aerosphere is full of obstacles (Drewitt and Langston 2008; Loss et al. 2015). Buildings such as telecommunication towers, maritime lighthouses and skyscrapers are a considerable source of mortality for flying species; this has been assessed especially for avifauna (Loss et al. 2014). Wind farms cause casualties for bats and birds (Cabrera-Cruz et al. 2020), via collisions or barotrauma (i.e. trauma due to the extreme speed of the blades which generates a sudden drop in atmospheric pressure in their vicinity) (Baerwald et al. 2008). This issue is becoming more and more concerning regarding wind farms expansion in the context of climate change and renewable energy deployment. Powerlines can lead to damage, by collisions or electrocutions (Sergio et al. 2004). Aircraft are a factor of mortality for birds (Desholm 2009). Other aerial sources of collisions may exist although they are not well assessed yet, e.g. all types of cables (wood longshoring cables, ski lift, telecommunication

cable). For instance, a study on 252 skiable domains between 2000 and 2004 in France counted close to 240 birds killed by ski-tow, ski lifts and cable cars (Maillard 2009). Therefore, it seems obvious that airscape fragmentation requires a dedicated strategy (Sordello et al. 2022b).

The soil is also an habitat for many taxa (Fig. 1), such as bacteria, fungi, invertebrates (nematodes, springtails, earthworms, ...), mammals (rodents, insectivores, carnivores and other burrow builders, galleries) or plants (root system) (Bardgett and Cook 1998; Wolters 2001). For all these organisms, the soil is both a place to live and to move (Auclerc et al. 2009; Eijsackers 2011). Some species live exclusively in the soil and have only this compartment to move. For instance, some springtails are repelled by light and live completely in the soil at depths of around thirty centimetres (eu-edaphic species) (Eisenbeis and Wichard 1987). Trees may also need to exchange nutrients among each other via their roots (Klein et al. 2016). Yet, soil wildlife may encounter many man-induced obstacles: housing foundations, transportation networks for energy (e.g. gas pipelines) or people (e.g. subway). In cities, many green areas—for example, on streetsides—are limited to large, unconnected bins of soil. Some human activities, such as material extraction or agriculture, threaten soil quality (Menta et al. 2014; Falco et al. 2015) and may generate barriers to wildlife movements, either physical (e.g. compacted soil due to intensive agricultural practices) or chemical (e.g. virtual

<figure><img src="image_2.png" /><figcaption>**Fig. 1** Aerosphere and soil are key compartments for biodiversity to live (habitats) and move (connectivity)—with specific man-induced threats—while ecological networks generally focus on above ground issues (road, dams, land-use)</figcaption></figure>
“wall” of toxic substances within a polluted soil). The soil is not just a surface but a compartment made up of different strata which—when the soil is excavated and moved elsewhere—are mixed together, destroying the way the soil functions.

Aerosphere is a compartment where take place ecological and biochemical processes that are essen-
tial to life on Earth, providing a vital resource,
including for humans: pure and oxygenated air (Kho-
nyongwa et al. 2023). Soil performs numerous eco-
logical functions (e.g. nutrient and water cycles, car-
bon storage, degradation of organic matter) and also
provides crucial benefits for our societies (e.g. cli-
mate regulation, recycling, food provisioning) (Adhi-
kari and Hartemink 2016; Banwart et al. 2019). As
a result, habitat loss and fragmentation in airscapes
and soil should be considered in ecological networks.
For the moment this consideration remains limited,
and restricted to roads; e.g. crossing passages for soil
wildlife (worm bridges) and aerial fauna (bat bridges
and gantries) (Berthinussen and Altringham 2012;
Claireau et al. 2021).

Sensory pollutants

Species are sensitive to visual, auditory or olfactory
stimuli which play a key role to seek habitats (e.g.
finding refugia, identifying food resources, detecting
an available partner (Conover 2007; Goerlitz et al.
2008; Jiang et al. 2015)) and to move (e.g. pursu-
ing prey, fleeing predators, memorising its way (Bee
2007; Reddy et al. 2022; Freas and Spetch 2023)).
Nowadays, human activities emit artificial stimuli
which interfer with natural ones (Dominoni et al.
2020) and that can combine with each other (Willems
et al. 2022; Ichikawa and Kuriwada 2023).

Nighttime lighting has expanded substantially
worldwide over the past decades, exposing all living
organisms to light pollution (Falchi et al. 2016). Arti-
ficial light at night (ALAN) has been demonstrated to
cause many damage on fauna, flora and ecosystems
(Falcón et al. 2020; Sanders et al. 2021). It can alter
habitat quality (Picchi et al. 2013; Ciach and Fröhlich
2019) and act as a barrier in the nightscape (Zeale
et al. 2018; Vowles and Kemp 2021; Korpach et al.
2022).

Human activities also generate sounds whose impacts
on biodiversity are extensively documented (Sordello

et al. 2020); for instance industry (Mason et al. 2016),
roads (Grade and Sieving 2016), aircraft (Injaian et al.
2021), wind farms (Keehn and Feldman 2018), recrea-
tion activities (Payne et al. 2012). This noise pollution
affects animal communication—whether in predator/
prey or mating interactions (Roca et al. 2016; Duquette
et al. 2021)—behavior or foraging (Klett-Mingo et al.
2016; Mancera et al. 2017). It alters species occurrence
(Luscier et al. 2023) and ultimately ecosystem compo-
sition (McClure et al. 2017; Sangermano 2022). In the
same way as ALAN does, anthropogenic noise degrades
habitat quality (Francis et al. 2011; Ware et al. 2015; Luo
et al. 2018) and is likely to fragment natural soundscapes.
For example, traffic noise represents an acoustic barrier
for owl hunting (Senzaki et al. 2016).

Less attention has been paid to date to the impacts
on wildlife of odors emitted by human activities such
as transportation, industry or agriculture. However, we
know that odors also contribute to sensory landscapes in
the form of an odorscape within which many taxa inter-
act (Conchou et al. 2019; Finnerty et al. 2022). Thus,
anthropogenic odors are likely to interfere with the olfac-
tory communication of living organisms, by decreas-
ing the intensity of signals or by luring individuals with
wrong information (Lurling and Scheffer 2007; David
et al. 2022; Jiang et al. 2023). Potentially, these man-
induced odors can degrade the quality of natural habitats,
or even constitute areas of disruption, for example in the
context of linear infrastructure (Girling et al. 2013; Jür-
gens and Bischoff 2017).

Therefore, nature conservation would benefit
from considering the sensory perception of species
(Madliger 2012; Elmer et al. 2021), including at the
landscape level (Little et al. 2022). This raises the
need for ecological networks to include sensory pol-
lutants, at least starting with artificial light at night
(Sordello 2017), man-induced noise (Moulherat
et al. 2023) and anthropogenic odors (Renou et al.
2021). This means we have to stop considering land-
scape features solely from a tangible point of view.
For instance, a road is not only a physical issue (e.g.
impervious surface, traffic, fences) but also a multi-
sensory polluting infrastructure (Fig. 2). Healthy nat-
ural sensory landscapes are also essential for human
beings; e.g. artificial light at night disturbs human
health (Cupertino et al. 2023) and five million peo-
ple in the European Union suffer from major chronic
sleep disorders caused by noise according to Euro-
pean Environment Agency.
**Caption:** A road combines both physical—such as impervious ground, fencing, verge management (Ouédraogo et al. 2020)—and sensory threats—such as artificial light at night (Davies et al. 2012), man-induced noise (Bautista et al. 2004; Bee and Swanson 2007) and odors (Girling et al. 2013). These sensory pollutants are likely to reduce the effectiveness of road crossing structures (Bliss-Ketchum et al. 2016; Sołowczuk 2019; Bhardwaj et al. 2020) and widen the road ecological footprint. For instance, ALAN was shown to act on bats up to 50 m (Azam et al. 2018) and noise effect on decreasing bird density was true up to 100 m (Wiącek 2023).

**Green infrastructure: a solution to implement multidimensional ecological networks?**

For decades, several tools have been developed for ecological planning, using the term ‘*infrastructure*’.

‘Ecological infrastructure’ was firstly introduced in 1984 in UNESCO’s Man and Biosphere Programme to promote « *a natural landscape and natural areas as the framework for spatially organising the city*» (UNESCO 1984). Then, during the 1990s, ‘Ecological infrastructure’ relatively equated with ‘Ecological Networks’—both terms focusing on nature conservation (Ahern 1995). In recent decades, new terms have been flowering, such as ‘Urban Ecological Infrastructure’ (UEI) or ‘Green

<figure><img src="image_3.png" /><figcaption>The same road as considered from a sensory point of view</figcaption></figure>
Infrastructure' (GI) (Childers et al. 2019; Sun et al. 2020). Today, while GI spontaneously refers to 'green' components of ecosystems, it has emerged as the more inclusive and commonly used name, whatever the environement (terrestrial or aquatic) and the context (natural or urban), among natural resource professionals (Benedict and MacMahon 2002). In recent years, GI have been spreading on a variety of territory levels, including urban contexts (e.g. in London (Jones and Somper 2014), Madrid (Velázquez Melero and Rodríguez-Espinosa 2023), New-York (Meerow 2020), Tokyo (Aleksejeva et al. 2022)). On a broader level, the European Commission has been developing a GI strategy since 2013,<fnref n="3" /> which is now part of the EU Biodiversity Strategy for 2030.

However, since GI originate from a variety of disciplines including ecology, urban planning or civil engineering (Ying et al. 2022), a kind of confusion exists about what they really aim at (Silva and Wheeler 2017). Some conceptions emphasise the more anthropogenic objective—i.e. improving human well-being (Li et al. 2017)—while other definitions focus on biodiversity persistence (Science for Environment Policy 2012; Mell 2014). Some definitions are hybdrid but give priority to human benefits; for instance, European Commission defines GI as « *A strategically planned network of natural and semi-natural areas with other environmental features, designed and managed to deliver a wide range of ecosystem services, while also enhancing biodiversity*».<fnref n="4" />

Here, we refer to *Green Infrastructure* assuming a conception focused primarily on biodiversity while benefiting humans, taking the definition of Benedict & MacMahon (2002): « *an interconnected network of green space that conserves natural ecosystem values and functions and provides associated benefits to human populations* ». Hence, we consider GI as the practical implementations of ENs. As a result, we preconize that GI also transition to encompass multidimensional ENs.

In France, a process has already been started to expand GI to other ecosystem dimensions. Especially, designers are now invited to take into account light pollution to ensure that ecosystems function

properly at night, leading to the emergence of a Dark infrastructure (Sordello et al. 2022a). Recently, the European Commission promoted the extension of GI to night-time environment (European Commission 2023).

To continue in this direction, we suggest to decline GI according to various dimensions that affect ecological networks. Namely, an aerial infrastructure for flying species, an underground infrastructure for soil wildlife (that could be named ‘brown infrastructure’ in reference to soil color), a dark infrastructure for darkness at night, a noise-free infrastructure (that could be named ‘white infrastructure’ in reference to ‘white noise’; i.e. sound emitted to mask surrounding noise) and an olfactory infrastructure to deal with anthropogenic odors (see Fig. 3). In France, such new infrastructure have begun to be implemented. Indeed, dark infrastructure are proliferating across the country (Sordello et al. 2021), a noise-free infrastructure has been undertaken in a French regional nature Park (Le Bougnec 2018) and the French metropolises of Limoges and Nancy have been involved in a brown infrastructure (Cluzeau et al. 2022; Vincent et al. 2022). Moreover, projects that combine these several dimensions to achieve a more comprehensive Green infrastructure are emerging. Details of such exploratory projects are given in Additional File 1.

## Exploratory practical concerns

Multidimensional ecological networks and their implementation in GI are a practical challenge: how can this complexity be taken into account while remaining operational and readable? We suggest here two practical outlets: designing and management—and we preconize a framework that could be operationalized by managers and decision-makers to process multidimensional GI (Fig. 4). We also highlight key knowledge gaps that would need further research, detailed in Additional File 2.

Designing from extended ecological networks to multidimensional GI

The design of ecological networks (i.e. identifying cores, corridors and barriers) is an essential first step to identify which areas are still preserved and which

<footnote n="3">³ https://environment.ec.europa.eu/topics/nature-and-biodiversity/green-infrastructure_en</footnote>

<footnote n="4">⁴ https://biodiversity.europa.eu/green-infrastructure</footnote>
<figure><img src="image_4.png" /><figcaption>**Fig. 3** The green (and blue) infrastructure could be a relevant lever to encompass multidimensional ecological networks, including specific infrastructure concerning aerial and underground compartments and sensory pollutants</figcaption></figure>

ones are degraded (i.e. have to be restored) and then to achieve in the end to a planning and decision-making tool, the multidimensional GI (see Fig. 4—Part A/Desiging). In a perspective of more comprehensive ENs, such mapping should involve airscapes and soils and include sensory drivers of habitat loss and fragmentation—artificial light, noise and olfactory disturbances (see Fig. 4—Step 1). Preserved areas should benefit from protection to prevent the spread of pressures in all these dimensions in the future, for example to avoid the implantation of wind farms on aerial migratory corridors (wind power is growing every year and will continue to do so (GWEC 2023)) or the extension of night lighting in areas of relict darkness (ALAN is estimated to grow by 2% a year (Kyba et al. 2017)).

Ecological networks are often designed based on habitat amount (ensuring population viability) and landscape connectivity (ensuring ability of species

to move). This step involves species to be selected as biological model to represent a diversity of ecology and environment. Hence, the aim will be to diversify the species guilds—e.g. to include soil fauna, flying species, species sensitive to ALAN, with different traits (dispersal distance, etc.) (see Fig. 4—Step 2). Next, population viability and landscape permeability can be assessed, for instance by including new drivers (ALAN, noise, odors) as additional parameters in spatial models through the coefficients determining habitat quality and land cover resistance (Hale et al. 2015; Pauwels et al. 2019; Moulherat et al. 2023) (but see the issue about data availabity, below).

Another question is how to use existing maps of ecological networks supported by policies to go further. This is an important issue to avoid giving the impression, particularly to local elected representatives, that the works they have undertaken to date are now obsolete. In this respect, it is possible to start
1/ DIAGNOSIS OF
HUMAN PRESSURES

BARRIERS AND HABITAT QUALITY IN LIFE COMPARTMENTS

<figure><img src="image_5.png" /></figure>

e.g. electric lines

<figure><img src="image_6.png" /></figure>

<figure><img src="image_7.png" /></figure>

e.g. woodlands

<figure><img src="image_8.png" /></figure>

e.g. water quality

<figure><img src="image_9.png" /></figure>

<figure><img src="image_10.png" /></figure>

e.g. soil quality

<figure><img src="image_11.png" /></figure>

SENSORY DISTURBANCES

<figure><img src="image_12.png" /></figure>

artificial light at night

<figure><img src="image_13.png" /></figure>

<figure><img src="image_14.png" /></figure>

e.g. road noise

<figure><img src="image_15.png" /></figure>

<figure><img src="image_16.png" /></figure>

e.g. industrial odors

<figure><img src="image_17.png" /></figure>

2/ MAPPING THE MULTI-DIMENSIONAL
ECOLOGICAL NETWORK

SELECTION OF SPECIES GUILDS

Species sensitive to several pressures, representative of several habitats/environments, with different ecology traits

<figure><img src="image_18.png" /></figure>

<figure><img src="image_19.png" /></figure>

<figure><img src="image_20.png" /></figure>

<figure><img src="image_21.png" /></figure>

<figure><img src="image_22.png" /></figure>

<figure><img src="image_23.png" /></figure>

<figure><img src="image_24.png" /></figure>

HABITAT SUITABILITY AND LANDSCAPE CONNECTIVITY

Modelling the ability of species to complete their life cycle and to move within the landscape

<figure><img src="image_25.png" /></figure>

<figure><img src="image_26.png" /></figure>

<figure><img src="image_27.png" /></figure>

<figure><img src="image_28.png" /></figure>

3/ TRANSFORMATION INTO A DECISION-MAKING TOOL

Consultations, Territory project, Political choice, Involvement of stakeholders
Human needs and human benefits

MULTI-DIMENSIONAL INFRASTRUCTURE
(generic term: *Green infrastructure*)

<figure><img src="image_29.png" /></figure>

AERIAL
INFRASTRUCTURE

<figure><img src="image_30.png" /></figure>

UNDERGROUND
INFRASTRUCTURE

<figure><img src="image_31.png" /></figure>

GREEN AND BLUE
INFRASTRUCTURE

<figure><img src="image_32.png" /></figure>

NOISE-FREE
INFRASTRUCTURE

DARK
INFRASTRUCTURE

<figure><img src="image_33.png" /></figure>

<figure><img src="image_34.png" /></figure>

ODOUR-FREE
INFRASTRUCTURE

PRESERVATION

planning schemes
(e.g. wind power, transportation)

protected areas

quiet zones

zero net artificialisation

project location orientation

low emission zones

darksky reserves

sustainable industrial processes

reduction in inputs

RESTORATION

slowing down wind turbine speed

odor neutralization

river remeandering

lighting on demand

wildlife crossings

buffer trees

power line marking

pond creation

planting hedges

technosoils
<caption>►Fig. 4 Framework from diagnosis to action to design and manage multidimensional GI</caption>

from these already designed ecological networks by coupling them with pressure maps in order to identify the parts of the ecological networks that also meet the additional criteria; e.g. regarding night-time darkness (Ranzoni et al. 2019) or acoustic environment (Le Bougnec 2018).

Regardless the method, data (e.g. pressure, occurrence) and knowledge (species sensitivity thresholds, life traits) may not all be available today. For instance, the design of an ‘aerial infrastructure’ requires first of all a good knowledge of the migratory axes and the behavior of flying species (flight heights, frequency of movements, etc.), which is only partially known today (Dokter et al. 2011; Péron et al. 2017). The integration of sensory drivers would require a good knowledge of the sensitivity thresholds of species to light, noise and olfactory pollutants. However, these thresholds are barely known for artificial light at night (Gaston et al. 2013), much less for anthropogenic noise (Kunc and Schmidt 2019) and totally unexplored for man-made odors. Research is hence required on these topics (see Additional File 2). Identifying such thresholds (i.e. operational quantified knowledge) may require primary studies and/or secondary research (systematic reviews, meta-analyses) to synthesize available knowledge. Knowledge should evolve rapidly in certain areas; for example, soundscape ecology, which is currently undergoing strong development, offers very promising prospects based on landscape acoustic indices (Farina and Gage 2017).

Finally, the ecological network—a scientific result—will have to be transformed into Green infrastructure—a planning map based on a political project (see Fig. 4—Step 3). This implies to involve various stakeholders through consultations in order to develop a territorial project that attracts support. Human needs and human benefits are taken into account here. This stage requires education, but it also needs a strong political will to ensure that the GI is as close as possible to the EN in order to maintain the same ambition to protect biodiversity.

## Management to reduce disturbances

Mitigation measures to reduce pressures, disturbances and landscape barriers must be implemented every where it is possible. However, GI offer a guidance which allows to prioritize actions within crucial areas (cores and corridors) (see Fig. 4—Part B/Managing). These measures will contribute to improve the quality of habitats, re-establish or create corridors, remove barriers or reduce the hostility of the landscape matrix, resulting in the enhancement of ecological network functionality. Several measures have already been applied for a long time in the context of GI (e.g. planting hedges, river remeandering, wildlife crossings). Such measures are relevant but in multidimensional GI new aspects have to be considered (e.g. aerial obstacles, soil disturbance, light pollution). Given the wide range of possible actions, we will only introduce some solutions for each topic:

1/ **Aerial infrastructure:** The blades of wind turbines can be lowered or stopped to reduce mortality during specific periods (migration, agricultural works likely to attract birds, etc.) (Smallwood and Bell 2020). Concerning aircraft, since most collisions occur at less than 150 m above ground (Dolbeer 2006), a strategy is to reduce the attractiveness of airport verges to wildlife or to introduce scare tactics as a preventive measure (Villers 2019).

2/ **Brown infrastructure:** Reducing the impervious areas and increasing open ground in urban green spaces is an essential step to enhance soil biodiversity (Maher et al. 2022) and soil functions such as carbon storage (Yan et al. 2023) or water flows (Strohbach et al. 2019). Moreover, constructed Technosols are a very promising field of ecological engineering to rebuild soils by recycling mineral waste (e.g. rubble from demolition, sediments from dredging, waste from quarries) (Ivashchenko et al. 2021; Minixhofer et al. 2022; Ouédraogo et al. 2024). In rural landscapes, shift farming practices towards a more extensive agriculture (e.g. agroforestry, permaculture) may contribute to restore a brown infrastructure (Domínguez et al. 2023).

3/ **Dark infrastructure:** A lot of practices can improve night quality (Gaston et al. 2012; Sordello 2018). Lighting should be reduced to the strict minimum, which may involve eliminating unnecessary light sources or reducing lighting time; e.g. part-night lighting (Azam et al. 2015), motion-activated lighting
(Bolliger et al. 2020), dimming protocols (Merckx et al. 2023). Yellow/amber/red closed light spectrum, considered less damaging for wildlife, should be preferred (Longcore et al. 2018).

4/ Noise free infrastructure: Reducing man-made sources of noise appears to be the first response (e.g. developing silent engines). Anti-noise barriers, including vegetative, may be a solution (Halim et al. 2015) and quiet zones can be defined, both in rural or urban context (Cerwén and Mossberg 2019).

5/ Olfactory infrastructure: The best strategy seems to be a reduction of anthropogenic odors at source (e.g. reducing the use of agricultural inputs and fossil fuels). When volatile compounds are in the environment, it is difficult to neutralize them (i.e. destroy them, transform them, cover them up). Biofilters, odor-neutralizing materials (e.g. activated carbon), chemical scrubbers, essential oils are interesting ideas but very little feedback are available. For the moment, they are being tested in industry, to reduce olfactory disturbances inside buildings, and in agriculture, to reduce odour nuisance for neighbours. For instance in France, following a complaint from neighbours, a farmer installed raspberry essential oil foggers to cover up the odours caused by his duck farm. The feasibility and effectiveness of transposing such techniques to odorscape restoration for nature conservation has yet to be fully assessed. Landscape management (e.g. tree buffers, hedgerows) might also help to reduce propagation of volatile compounds (Tyndall and Colletti 2007) but assessments show a limited effect (Hernandez et al. 2012). Scent manipulation could become a promising avenue to counterbalance the negative effects of man-made odours, to enhance landscape restoration (e.g. wildlife crossings) or to prevent roadkills (Bíl et al. 2018).

It should be noted that the panel of available measures is uneven between topics (for example, mitigating odors remains entirely exploratory to date). Moreover, measures have varying degrees of efficiency, sometimes not assessed, and can lead to undesirable consequences (e.g. introducing noise to avoid a collision; exposing trees to sensory pollutants to build ‘natural barriers’). Research is therefore needed on these emerging topics (see Additional File 2) (Madliger et al. 2016). However, the need for more knowledge must not be paralizing since reducing pressures can only go in the right direction for biodiversity protection; then wide range of actions can be

implemented without delay and without regret (no-regret strategies; i.e. measures that can be started now without being certain of everything, because they have no detrimental effect) such as reducing nighttime lighting, limiting noise or decreasing the amount of impervious surfaces.

The implementation of these actions requires a cross-sectoral coordination by the Green infrastructure project manager. Long-term monitoring protocols are needed to assess the efficiency of GI implementation by measuring changes in both human pressures and biodiversity responses (i.e. habitat amount, quality and connectivity). In the light of this monitoring, it is recommended that the Green infrastructure (design and/or the action plan) be updated periodically (every 5 years, for example).

## Conclusion

Scientific knowledge about human impacts on wildlife activities and movements is constantly increasing. New threats to species have been identified as diverse as windfarms, man-induced sounds or light pollution. This increase in the awareness of anthropogenic disturbances on biodiversity raises the number of parameters that should be accounted for when implementing ecological networks. To be more comprehensive, we encourage ecological network public policies to cover different living compartments—from soil to air—and more drivers—both physics and sensory—that lead to habitat loss and fragmentation. To this end, extended Green infrastructure could be a relevant solution. They should include several specific infrastructure, especially an aerial infrastructure for flying species, a brown infrastructure for soil wildlife, a dark infrastructure for darkness at night, a noise-free infrastructure to deal with anthropogenic sounds and an olfactory infrastructure to deal with anthropogenic odors. Transitioning to multidimensional ecological networks is an enormous challenge for societies that will involve more stakeholders in the process of design and management of GI. A lot of actors have now to feel concerned by such public policies, far from transportation managers, farmers or water stakeholders, such as the aviation or wind power sectors, technosol engineers, lighting engineers or noise and odor emitting industries.
Although knowledge is still incomplete, there is an urgent need for action on these issues without delay, for biodiversity and for us. Indeed, this challenge is essential for our societies since a vast literature highlights the benefits of a dark night environment (Wang et al. 2023), healthy soils (Hassan et al. 2022) and quiet spaces (Hahad et al. 2024) for our well-being. Then, the gain of a healthy environment for humans and non-humans go in the same direction, which is underlined by the « *One health* » concept currently developed by the United Nations (ONU 2022).

**Acknowledgements** We thank Aleksandra Delcourt for the drawing (Figure 3).

**Author contributions** All authors collectively conceptualized the problem addressed by the article. R.S. wrote the main manuscript text and prepared the figures and table. All authors reviewed the manuscript.

**Data availability** No datasets were generated or analysed during the current study.

**Declarations**

**Competing interests** The authors declare no competing interests.

**Open Access** This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or other third party material in this article are included in the article's Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article's Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/.

**References**

Adhikari K, Hartemink AE (2016) Linking soils to ecosystem services — a global review. Geoderma 262:101–111

Ahern J (1995) Greenways as a planning strategy. Landsc Urban Plan 33:131–155

Aleksejeva J, Voulgaris G, Gasparatos A (2022) Assessing the potential of strategic green roof implementation for green infrastructure: insights from Sumida ward. Tokyo Urban Forestry Urban Greening 74:127632

Andersen LW, Fog K, Damgaard C (2004) Habitat fragmentation causes bottlenecks and inbreeding in the European tree frog (*Hyla arborea*). Proc R Soc Lond B 271:1293–1302

Auclerc A, Ponge JF, Barot S, Dubs F (2009) Experimental assessment of habitat preference and dispersal ability of soil springtails. Soil Biol Biochem 41:1596–1604

Azam C, Kerbiriou C, Vernet A et al (2015) Is part-night lighting an effective measure to limit the impacts of artificial lighting on bats? Glob Change Biol 21:4333–4341

Azam C, Le Viol I, Bas Y et al (2018) Evidence for distance and illuminance thresholds in the effects of artificial lighting on bat activity. Landsc Urban Plan 175:123–135

Baerwald EF, D'Amours GH, Klug BJ, Barclay RMR (2008) Barotrauma is a significant cause of bat fatalities at wind turbines. Current Biol. https://doi.org/10.1016/j.cub.2008.06.029

Baker SE, Ellwood SA, Slater D et al (2008) Food aversion plus odor cue protects crop from wild mammals. J Wildl Manag 72:785–791

Banwart SA, Nikolaidis NP, Zhu Y-G et al (2019) Soil Functions: connecting earth's critical zone. Annu Rev Earth Planet Sci 47:333–359

Barchi JR, Knowles JM, Simmons JA (2013) Spatial memory and stereotypy of flight paths by big brown bats in clutered surroundings. J Exp Biol 216:1053–1063

Bardgett RD, Cook R (1998) Functional aspects of soil animal diversity in agricultural grasslands. Appl Soil Ecol 10:263–276

Battisti C (2003) Habitat fragmentation, fauna and ecological network planning: toward a theoretical conceptual framework. Italian J Zool 70:241–247

Bautista LM, García JT, Calmaestra RG et al (2004) Effect of weekend road traffic on the use of space by raptors. Conserv Biol 18:726–732

Bee MA (2007) Selective phonotaxis by male wood frogs (*Rana sylvatica*) to the sound of a chorus. Behav Ecol Sociobiol 61:955–966

Bee MA, Swanson EM (2007) Auditory masking of anuran advertisement calls by road traffic noise. Anim Behav 74:1765–1776

Beier P, Noss RF (1998) Do habitat corridors provide connectivity? Conserv Biol 12:1241–1252

Benedict M, MacMahon ET (2002) Green infrastructure: smart conservation for the 21st Century. Renew Res J 20:12–17

Bennett G (2004) Integrating biodiversity conservation and sustainable use: lessons learned from ecological networks. IUCN, World Conservation Union, Gland, Switzerland

Bennett G, Mulongoy KJ, Secretariat of the Convention on Biological Diversity (2006) Review of experience with ecological networks, corridors and buffer zones

Berthinussen A, Altringham J (2012) Do bat gantries and underpasses help bats cross roads safely? PLoS ONE 7:e38775

Bhardwaj M, Soanes K, Lahoz-Monfort JJ et al (2020) Artificial lighting reduces the effectiveness of wildlife-crossing structures for insectivorous bats. J Environ Manage 262:110313
Bíl M, Andrášik R, Bartonička T et al (2018) An evaluation of odor repellent effectiveness in prevention of wild-life-vehicle collisions. J Environ Manage 205:209–214

Billon L, Sordello R (2017) Schémas régionaux de cohérence écologique : Chiffres de synthèse à l'échelle nationale. Service du patrimoine naturel, Muséum national d'Histoire naturelle, Paris

Bliss-Ketchum LL, de Rivera CE, Turner BC, Weisbaum DM (2016) The effect of artificial light on wildlife use of a passage structure. Biol Conservat 199:25–28

Bolger DT, Scott TA, Rotenberry JT (2001) Use of corridor-like landscape structures by bird and small mammal species. Biol Cons 102:213–224

Bolliger J, Hennet T, Wermelinger B et al (2020) Effects of traffic-regulated street lighting on nocturnal insect abundance and bat activity. Basic Appl Ecol 47:44–56

Botello F, Sarkar S, Sánchez-Cordero V (2015) Impact of habitat loss on distributions of terrestrial vertebrates in a high-biodiversity region in Mexico. Biol Cons 184:59–65

Buchmann CM, Schurr FM, Nathan R, Jeltsch F (2012) Movement upscaled – the importance of individual foraging movement for community response to habitat loss. Ecography 35:436–445

Cabrera-Cruz SA, Cervantes-Pasqualli J, Franquesa-Soler M et al (2020) Estimates of aerial vertebrate mortality at wind farms in a bird migration corridor and bat diversity hotspot. Global Ecol Conservat 22:e00966

Cerda-Peña C, Rau JR (2023) The importance of wetland habitat area for waterbird species-richness. Ibis 165:739–752

Cerwén G, Mossberg F (2019) Implementation of quiet areas in Sweden. IJERPH 16:134

Childers DL, Bois P, Hartnett HE, et al (2019) Urban Ecological Infrastructure: An inclusive concept for the non-built urban environment. Elementa: Science of the Anthropocene 7:46.

Ciach M, Fröhlich A (2019) Ungulates in the city: light pollution and open habitats predict the probability of roe deer occurring in an urban environment. Urban Ecosyst 22:513–523

Claireau F, Kerbiriou C, Charton F et al (2021) Bat overpasses help bats to cross roads safely by increasing their flight height. Acta 23:189–198

Cluzeau D, Leprisé A, Maréchal J, et al (2022) Trame brune de limoges métropole. développement expérimental d'une trame brune sur le territoire d'un EPCI. Phase 1 : Approche méthodologique

Coffin AW (2007) From roadkill to road ecology: a review of the ecological effects of roads. J Trans Geogr 15:396–406

Commission E (2019) Review of progress on implementation of the EU green infrastructure strategy. Publ. Off. of the Europ, Union, Luxembourg

Conchou L, Lucas P, Meslin C et al (2019) Insect odorscapes: from plant volatiles to natural olfactory scenes. Front Physiol 10:972.

Conover MR (2007) Predator-prey dynamics: the role of olfaction, CRC Press, Taylor and Francis. New York, USA

Coulon A, Cosson JF, Angibault JM et al (2004) Landscape connectivity influences gene flow in a roe deer

population inhabiting a fragmented landscape: an individual-based approach. Mol Ecol 13:2841–2850

Cristóbal-Pérez EJ, Fuchs EJ, Martén-Rodríguez S, Quesada M (2021) Habitat fragmentation negatively affects effective gene flow via pollen, and male and female fitness in the dioecious tree, Spondias purpurea (Anacardiaceae). Biol Cons 256:109007

Cupertino MDC, Guimarães BT, Pimenta JFG et al (2023) Light pollution: a systematic review about the impacts of artificial light on human health. Biol Rhythm Res 54:263–275

da Silva JMC, Wheeler E (2017) Ecosystems as infrastructure. Perspect Ecol Conservat 15:32–35

David NF, Henry TJ, Sprayberry JDH (2022) Odor-pollution from fungicides disrupts learning and recognition of a common floral scent in bumblebees (*Bombus impatiens*). Front Ecol Evol 10:765388

Davies TW, Bennie J, Gaston KJ (2012) Street lighting changes the composition of invertebrate communities. Biol Lett 8:764–767

Desholm M (2009) Avian sensitivity to mortality: Prioritising migratory bird species for assessment at proposed wind farms. J Environ Manage 90:2672–2679

Dickman CR (1987) Habitat fragmentation and vertebrate species richness in an urban environment. J Appl Ecol 24:337

Dokter AM, Liechti F, Stark H et al (2011) Bird migration flight altitudes studied by a network of operational weather radars. J R Soc Interface 8:30–43

Dolbeer RA (2006) Height distribution of birds recorded by collisions with civil aircraft. J Wildl Manag 70:1345–1350

Domínguez A, Escudero HJ, Rodríguez MP et al (2023) Agroecology and organic farming foster soil health by promoting soil fauna. Environ Dev Sustain. https://doi.org/10.1007/s10668-022-02885-4

Dominoni DM, Halfwerk W, Baird E et al (2020) Why conservation biology can benefit from sensory ecology. Nat Ecol Evol 4:502–511

Drewitt AL, Langston RHW (2008) Collision effects of wind-power generators and other obstacles on birds. Ann New York Acad Sci 1134:233–266

Duquette CA, Loss SR, Hovick TJ (2021) A meta-analysis of the influence of anthropogenic noise on terrestrial wildlife communication strategies. J Appl Ecol 58:1112–1121

Eijsackers H (2011) Earthworms as colonizers of natural and cultivated soil environments. Appl Soil Ecol 50:1–13

Eisenbeis G, Wichard W (1987) Atlas on the Biology of Soil Arthropods. Springer, Berlin Heidelberg, Berlin, Heidelberg

Elmer LK, Madliger CL, Blumstein DT et al (2021) Exploiting common senses: sensory ecology meets wildlife conservation and management. Conservat Phys. https://doi.org/10.1093/conphys/coab002

European Commission (2023) Science for environment policy future brief 28: light pollution

European Environment Agency (2014) Quiet areas in Europe — the environment unaffected by noise pollution

Fahrig L (2003) Effects of habitat fragmentation on biodiversity. Annu Rev Ecol Evol Syst 34:487–515
Falchi F, Cinzano P, Duriscoe D et al (2016) The new world atlas of artificial night sky brightness. Sci Adv. https://doi.org/10.1126/sciadv.1600377

Falco L, Sandler R, Momo F et al (2015) Earthworm assemblages in different intensity of agricultural uses and their relation to edaphic variables. PeerJ 3:e979

Falcón J, Torriglia A, Attia D et al (2020) Exposure to artificial light at night and the consequences for flora, fauna, and ecosystems. Front Neurosci 14:602796

Farina A, Gage SH (eds) (2017) Ecoacoustics: the ecological role of sounds. Wiley, Hoboken (N.J.), Etats-Unis d'Amérique

Finnerty PB, McArthur C, Banks P et al (2022) The Olfactory landscape concept: a key source of past, present, and future information driving animal movement and decision-making. Bioscience 72:745–752

Fontúrbel FE, Betancurt-Grisales JF, Vargas-Daza AM, Castaño-Villa GJ (2022) Effects of habitat degradation on bird functional diversity: a field test in the Valdivian rainforest. For Ecol Manage 522:120466

Forman RTT, Alexander LE (1998) Roads and their major ecological effects. Annu Rev Ecol Syst 29:207–231

Francis CD, Paritsis J, Ortega CP, Cruz A (2011) Landscape patterns of avian habitat use and nest success are affected by chronic gas well compressor noise. Landsc Ecol 26:1269–1280

Freas CA, Spetch ML (2023) Varieties of visual navigation in insects. Anim Cogn 26:319–342

Fülöp A, Daróczi SJ, Dehelean AS et al (2018) Autumn passage of soaring birds over dobrogea (Romania): a migration corridor in Southeast Europe. Ardea 106:61

Gaston KJ, Davies TW, Bennie J, Hopkins J (2012) REVIEW: reducing the ecological consequences of night-time light pollution: options and developments. J Appl Ecol 49:1256–1266

Gaston KJ, Bennie J, Davies TW, Hopkins J (2013) The ecological impacts of nighttime light pollution: a mechanistic appraisal. Biol Rev 88:912–927

Gehrke PC, Gilligan DM, Barwick M (2002) Changes in fish communities of the Shoalhaven River 20 years after construction of Tallowa Dam, Australia. River Res Apps 18:265–286

Girling RD, Lusebrink I, Farthing E et al (2013) Diesel exhaust rapidly degrades floral odours used by honeybees. Sci Rep 3:2779

Glista DJ, DeVault TL, DeWoody JA (2008) Vertebrate road mortality predominantly impacts amphibians. Herpetol Conserv Biol 3:77–87

Goerlitz HR, Greif S, Siemens BM (2008) Cues for acoustic detection of prey: insect rustling sounds and the influence of walking substrate. J Exp Biol 211:2799–2806

Grade AM, Sieving KE (2016) When the birds go unheard: highway noise disrupts information transfer between bird species. Biol Lett. https://doi.org/10.1098/rsbl.2016.0113

Grashof-Bokdam C (1997) Forest species in an agricultural landscape in the Netherlands: effects of habitat fragmentation. J Vegetation Science 8:21–28

Guerra CA, Bardgett RD, Caon L et al (2021) Tracking, targeting, and conserving soil biodiversity. Science 371:239–241

Gutiérrez-Chacón C, Valderrama-A C, Klein A-M (2020) Biological corridors as important habitat structures for maintaining bees in a tropical fragmented landscape. J Insect Conserv 24:187–197

GWEC (2023) Global Offshore Wind Report 2023

Hahad O, Kuntic M, Al-Kindi S et al (2024) Noise and mental health: evidence, mechanisms, and consequences. J Expo Sci Environ Epidemiol. https://doi.org/10.1038/s41370-024-00642-5

Hale JD, Fairbrass AJ, Matthews TJ et al (2015) The ecological impact of city lighting scenarios: exploring gap crossing thresholds for urban bats. Glob Change Biol 21:2467–2478

Halim H, Abdullah R, Ali AAA, MohdJMohd N (2015) Effectiveness of existing noise barriers: comparison between vegetation, concrete hollow block, and panel concrete. Procedia Environ Sci 30:217–221

Hamid AR, Tan PY (2017) Urban ecological networks for biodiversity conservation in cities. In: Tan PY, Jim CY (eds) Greening Cities: Forms and Functions. Springer, Singapore, pp 251–277

Hansen SC, Stolter C, Imholt C, Jacob J (2017) Like or dislike: response of rodents to the odor of plant secondary metabolites. Integrat Zool 12:428–436

Hassan W, Li Y, Saba T et al (2022) Improved and sustainable agroecosystem, food security and environmental resilience through zero tillage with emphasis on soils of temperate and subtropical climate regions: a review. Int Soil Water Conservat Res 10:530–545

Heller NE, Zavaleta ES (2009) Biodiversity management in the face of climate change: a review of 22 years of recommendations. Biol Cons 142:14–32

Hernandez G, Trabue S, Sauer T et al (2012) Odor mitigation with tree buffers: swine production case study. Agr Ecosyst Environ 149:154–163

Hilty JA, Lidicker WZ, Merenlender AM (2006) Corridor ecology: the science and practice of linking landscapes for biodiversity conservation. Island Press, Washington, DC

Hilty JA, Keeley ATH, Lidicker WZ et al (2019) Corridor ecology: linking landscapes for biodiversity conservation and climate adaptation, 2nd edn. Island Press, Washington, Covelo London

Hilty J, Worboys GL, Keeley A et al (2020) Guidelines for conserving connectivity through ecological networks and corridors. IUCN, International Union for Conservation of Nature

Holker F, Wolter C, Perkin EK, Tockner K (2010) Light pollution as a biodiversity threat. Trends Ecol Evol 25:681–682

Holyoak M, Heath SK (2016) The integration of climate change, spatial dynamics, and habitat fragmentation: a conceptual overview. Integrative Zoology 11:40–59

Homan RN, Windmiller BS, Reed JM (2004) Critical thresholds associated with habitat loss for two vernal pool-breeding amphibians. Ecol Appl 14:1547–1553

Hyseni C, Heino J, Bini LM et al (2021) The importance of blue and green landscape connectivity for biodiversity in urban ponds. Basic Appl Ecol 57:129–145

Ichikawa I, Kuriwada T (2023) The combined effects of artificial light at night and anthropogenic noise on life history traits in ground crickets. Ecol Res 38:446–454
Injaian AS, Lane ED, Klinck H (2021) Aircraft events cor-
respond with vocal behavior in a passerine. Sci Rep
11:1197

Ivashchenko K, Lepore E, Vasenev V et al (2021) Assessing
soil-like materials for ecosystem services provided by
constructed technosols. Land 10:1185

Jägerbrand AK, Bouroussis CA (2021) Ecological impact of artificial light at night: effective strategies and measures to deal with protected species and habitats. Sustainability 13:5991

Jiang F, Fang G, Xue F et al (2015) Male music frogs compete vocally on the basis of temporal sequence rather than spatial cues of rival calls. Asian Herpetol Res 6:305-316

Jiang N-J, Chang H, Weißflog J et al (2023) Ozone exposure disrupts insect sexual communication. Nat Commun 14:1186

Jones S, Somper C (2014) The role of green infrastruc-
ture in climate change adaptation in L ondon. Geogr J
180:191-196

Jongman RHG (1995) Nature conservation planning in Europe: developing ecological networks. Landsc Urban Plann 32:169-183

Jongman RHG, Külvik M, Kristiansen I (2004) European eco-
logical networks and greenways. Landsc Urban Plann
68:305-319

Jürgens A, Bischoff M (2017) Changing odour landscapes: the effect of anthropogenic volatile pollutants on plant-pollinator olfactory communication. Funct Ecol 31:56-64

Keehn JE, Feldman CR (2018) Disturbance affects biotic com-
munity composition at desert wind farms. Wildl Res
45:383-396

Keeley ATH, Beier P, Creech T et al (2019) Thirty years of connectivity conservation planning: an assessment of factors influencing plan implementation. Environ Res Lett 14:103001

Keyghobadi N, Roland J, Strobeck C (2005) Genetic differen-
tiation and gene flow among populations of the alpine
butterfly, Parnassius smintheus, vary with landscape con-
nectivity: LANDSCAPE CONNECTIVITY and GENE
FLOW. Mol Ecol 14:1897-1909

Khonyongwa O, Kumar S, Kumar N et al (2023) Air pollution impact on human health and its management: a review. Macromol Symp 407:2100488

Klein T, Siegwolf RTW, Körner C (2016) Belowground carbon trade among tall trees in a temperate forest. Science 352:342-344

Kleinschroth F, Laporte N, Laurance WF et al (2019) Road expansion and persistence in forests of the congo basin. Nat Sustain 2:628–634

Klett-Mingo JI, Pavón I, Gil D (2016) Great tits, Parus major, increase vigilance time and reduce feeding effort during peaks of aircraft noise. Anim Behav 115:29-34

Korpach AM, Garroway CJ, Mills AM et al (2022) Urbaniza-
tion and artificial light at night reduce the functional con-
nectivity of migratory aerial habitat. Ecography. https://
doi.org/10.1111/ecog.05581

Kotze R, Marshal JP, Winterbach CW et al (2021) Demo-
graphic consequences of habitat loss and crowding
in large carnivores: a natural experiment. Afr J Ecol
59:63-73

Kuipers KJ, Hilbers JP, Garcia-Ulloa J et al (2021) Habi-
tat fragmentation amplifies threats from habitat loss to
mammal diversity across the world's terrestrial ecoregions. One Earth 4:1505-1513

Kunc HP, Schmidt R (2019) The effects of anthropogenic noise on animals: a meta-analysis. Biol Lett 15:20190649

Kyba CCM, Kuester T, Sánchez de Miguel A et al (2017) Arti-
ficially lit surface of Earth at night increasing in radiance
and extent. Sci Adv 3:e1701528

Le Bougnec A (2018) La Trame sonore - Elaboration d'une méthode d'identification sur le territoire du Parc Naturel Régional du Golfe du Morbihan. Université Bordeaux Montaigne, Parc naturel régional du Golf du Morbihan

Li F, Liu X, Zhang X et al (2017) Urban ecological infrastruc-
ture: an integrated network for ecosystem services and
sustainable urban systems. J Clean Prod 163:S12-S18

Little CJ, Rizzuto M, Luhring TM et al (2022) Movement with meaning: integrating information into meta-ecology. Oikos 2022:e08892

Longcore T, Rodríguez A, Witherington B et al (2018) Rapid assessment of lamp spectrum to quantify ecological effects of light at night. J Exper Zool Part A 329:511-521

Loss SR, Will T, Loss SS, Marra PP (2014) Bird-building collisions in the United States: estimates of annual mortality and species vulnerability. The Condor 116:8-23

Loss SR, Will T, Marra PP (2015) Direct mortality of birds from anthropogenic causes. Annu Rev Ecol Evol Syst 46:99-120

Luo Q, Song Y, Hu X et al (2018) Effects of tourism distur-
bance on habitat quality and population size of the Chi-
nese giant salamander (Andrias davidianus). Wildl Res
45:411-420

Lurling M, Scheffer M (2007) Info-disruption: pollution and the transfer of chemical information between organisms. Trends Ecol Evol 22:374-379

Luscier JD, Christopher A, Synan H, Wilson KB (2023) Effects of light and noise pollution on occupancy of gray treefrogs (Hyla versicolor) and green frogs ( Lithobates clamitans ) in Syracuse, NY. Urban Ecosyst 26:941-953

Madliger CL (2012) Toward improved conservation manage-
ment: a consideration of sensory ecology. Biodivers Con-
serv 21:3277-3286

Madliger CL, Cooke SJ, Crespi EJ et al (2016) Success stories and emerging themes in conservation physiology. Conserv Physiol. 4:cov057

Maher GM, Johnson GA, Burdine JD (2022) Impervious sur-
face and local abiotic conditions influence arthropod
communities within urban greenspaces. PeerJ 10:e12818

Maillard D (2009) Localisation des câbles dangereux pour les oiseaux sur les domaines skiables. ONCFS

Mancera KF, Murray PJ, Lisle A et al (2017) The effects of acute exposure to mining machinery noise on the behaviour of eastern blue-tongued lizards (Tiliqua scincoides). Anim Welf 26:11-24

Mason JT, McClure CJW, Barber JR (2016) Anthropogenic noise impairs owl hunting behavior. Biol Cons 199:29-32

McClure CJW, Ware HE, Carlisle JD, Barber JR (2017) Noise from a phantom road experiment alters the age struc- ture of a community of migrating birds. Anim Conserv 20:164-172
McInturff A, Xu W, Wilkinson CE et al (2020) Fence ecology: frameworks for understanding the ecological effects of fences. *BioScience*. https://doi.org/10.1093/biosci/biaa103

Medrano-Vizcaíno P, Grilo C, Silva Pinto FA et al (2022) Roadkill patterns in Latin American birds and mammals. *Glob Ecol Biogeogr* 31:1756–1783

Meerow S (2020) The politics of multifunctional green infrastructure planning in New York City. *Cities* 100:102621

Mell IC (2014) Aligning fragmented planning structures through a green infrastructure approach to urban development in the UK and USA. *Urban Forest Urban Green* 13:612–620

Menta C, Conti FD, Pinto S et al (2014) Monitoring soil restoration in an open-pit mine in northern Italy. *Appl Soil Ecol* 83:22–29

Merckx T, Nielsen ME, Kankaanpää T et al (2023) Dim light pollution prevents diapause induction in urban and rural moths. *J Appl Ecol* 60:1022–1031

Minixhofer P, Scharf B, Hafner S et al (2022) Towards the circular soil concept: optimization of engineered soils for green infrastructure application. *Sustainability* 14:905

Mony C, Uroy L, Khalfallah F et al (2022) Landscape connectivity for the invisibles. *Ecography*. https://doi.org/10.1111/ecog.06041

Moreno-Opo R (2020) Individual and demographic responses of a marsh bird assemblage to habitat loss and subsequent restoration. *Avian Res* 11:4

Moulatlet GM, Ambriz E, Guevara J et al (2021) Multi-taxa ecological responses to habitat loss and fragmentation in western Amazonia as revealed by RAPELD biodiversity surveys. *Acta Amaz* 51:234–243

Moulherat S, Plotard C, De Roincé C, Cornuau J (2023) Pren-dre en compte la pollution sonore dans une modélisation de dynamiques de populations d'espèces. *SET*. https://doi.org/10.20870/Revue-SET.2023.43.7512

Naumann S, Davis M, Kaphengst T, et al (2011) Design, implementation and cost elements of green infrastructure projects

ONU (2022) One Health Joint Plan of Action, 2022–2026. FAO; UNEP; WHO; World Organisation for Animal Health (WOAH) (founded as OIE); Rome

Opdam P, Wascher D (2004) Climate change meets habitat fragmentation: linking landscape and biographical scale levels in research and conservation. *Biol Cons* 117:285–297

Ouédraogo D-Y, Villemey A, Vanpeene S et al (2020) Can linear transportation infrastructure verges constitute a habitat and/or a corridor for vertebrates in temperate ecosystems? *A Systemat Rev Environ Evi* 9:13

Ouédraogo D-Y, Lafitte A, Sordello R et al (2024) Existing evidence on the potential of soils constructed from mineral wastes to support biodiversity: a systematic map. *Environ Evid* 13:9

Ouin A, Martin M, Burel F (2008) Agricultural landscape connectivity for the meadow brown butterfly (*Maniola jurtina*). *Agr Ecosyst Environ* 124:193–199

Pauwels J, Le Viol I, Azam C et al (2019) Accounting for artificial light impact on bat activity for a biodiversity-friendly urban planning. *Landsc Urban Plann* 183:12–25

Payne CJ, Jessop TS, Guay P-J et al (2012) Population, behavioural and physiological responses of an urban population of black swans to an intense annual noise event. *PLoS ONE* 7:e45014

Péron G, Fleming CH, Duriez O et al (2017) The energy landscape predicts flight height and wind turbine collision hazard in three species of large soaring raptor. *J Appl Ecol* 54:1895–1906

Picchi MS, Avolio L, Azzani L et al (2013) Fireflies and land use in an urban landscape: the case of *Luciola italica* L. (*Coleoptera: Lampyridae*) in the city of Turin. *J Insect Conserv* 17:797–805

Pietsch M (2018) Contribution of connectivity metrics to the assessment of biodiversity – some methodological considerations to improve landscape planning. *Ecol Ind* 94:116–127

Prober SM, Doerr VAJ, Broadhurst LM et al (2019) Shifting the conservation paradigm: a synthesis of options for renovating nature under climate change. *Ecol Monogr* 89:e01333

Radvan M, Rendall AR, Weston MA (2023) The habitat connectivity hypothesis of escape in urban woodland birds. *Behav Ecol* 34:297–305

Ranzoni J, Giuliani G, Huber L, Ray N (2019) Modelling the nocturnal ecological continuum of the State of Geneva, Switzerland, based on high-resolution nighttime imagery. *Remote Sens Appl Soc Environ* 16:100268

Reddy G, Murthy VN, Vergassola M (2022) Olfactory sensing and navigation in turbulent environments. *Annu Rev Condens Matter Phys* 13:191–213

Renou M, Sordello R, Reyjol Y (2021) La biologie de la conservation t-elle prendre en compte les paysages odorants ? *Sciences, Eaux et Territoires* https://doi.org/10.14758/SET-REVUE.2021.HS.03

Roca IT, Desrochers L, Giacomazzo M et al (2016) Shifting song frequencies in response to anthropogenic noise: a meta-analysis on birds and anurans. *Behav Ecol* 27:1269–1274

Rüter S, Vos CC, van Eupen M, Rühmkorf H (2014) Trans-boundary ecological networks as an adaptation strategy to climate change: the example of the Dutch – German border. *Basic Appl Ecol* 15:639–650

Samways MJ, Pryke JS (2016) Large-scale ecological networks do work in an ecologically complex biodiversity hotspot. *Ambio* 45:161–172

Sanders D, Frago E, Kehoe R et al (2021) A meta-analysis of biological impacts of artificial light at night. *Nat Ecol Evol* 5:74–81

Sangermano F (2022) Acoustic diversity of forested landscapes: relationships to habitat structure and anthropogenic pressure. *Landsc Urban Plan* 226:104508

Saunders DA, Hobbs RJ, Margules CR (1991) Biological consequences of ecosystem fragmentation: a review. *Conserv Biol* 5:18–32

Schmidt J, Hauck J (2018) Implementing green infrastructure policy in agricultural landscapes—scenarios for Saxony-Anhalt, Germany. *Reg Environ Change* 18:899–911

Schroer S, Huggins BJ, Azam C, Hölker F (2020) Working with inadequate tools: legislative shortcomings in protection against ecological effects of artificial light at night. *Sustainability* 12:2551
Science for Environment Policy (2012) The multifunctionality of Green Infrastructure. European Commission's Direktorate-General Environment

Senzaki M, Yamaura Y, Francis CD, Nakamura F (2016) Traffic noise reduces foraging efficiency in wild owls. *Sci Rep.* https://doi.org/10.1038/srep30602

Sergio F, Marchesi L, Pedrini P et al (2004) Electrocution alters the distribution and density of a top predator, the eagle owl Bubo bubo. *J Appl Ecol* 41:836–845

Simkin RD, Seto KC, McDonald RI, Jetz W (2022) Biodiversity impacts and conservation implications of urban land expansion projected to 2050. *Proc Natl Acad Sci USA* 119:e2117297119

Smallwood KS, Bell DA (2020) Effects of wind turbine curtailment on bird and bat fatalities. *Jour Wild Mgmt.* https://doi.org/10.1002/jwmg.21844

Sołowczuk A (2019) Determinants of the performance of bat gantries installed to carry bat commuting routes over the S3 expressway in Poland. *Symmetry* 11:1022

Sordello R (2017) Pistes méthodologiques pour prendre en compte la pollution lumineuse dans les réseaux écologiques. *VertigO - La Revue Électronique En Sciences De L'environnement.* https://doi.org/10.4000/verti go.18730

Sordello R (2018) Comment gérer la lumière artificielle dans les continuités écologiques ? *Sci Eaux Territoir* 25:86

Sordello R, Ratel O, Flamerie De Lachapelle F et al (2020) Evidence of the impact of noise pollution on biodiversity: a systematic map. *Environ Evid* 9:20

Sordello R, Busson S, Cornuau JH et al (2022a) A plea for a worldwide development of dark infrastructure for biodiversity – practical examples and ways to go forward. *Landsc Urban Plan* 219:104332

Sordello R, Reyjol Y, Amsallem J et al (2022b) Les déplacements des espèces volantes : vers la mise en œuvre d'une « Trame aérienne » dans le cadre de la politique Trame verte et bleue ? *N.* https://doi.org/10.5852/naturae2022a9

Sordello R, Billon L, Amsallem J, Vanpeene S (2017) Bilan technique et scientifique sur l'élaboration des Schémas régionaux de cohérence écologique. Méthodes d'identification des composantes de la TVB. *Centre de ressources TVB* 1:104

Sordello R, Paquier F, Daloz A (2021) Trame noire: Méthodes d'élaboration et outils pour sa mise en œuvre. *office français de la biodiversité* 116

Sordello R (2015) Première capitalisation méthodologique sur les Schémas régionaux de cohérence écologique adoptés ou en projet - Pollution lumineuse. *MNHN-SPN - CDR TVB*

Sork VL, Smouse PE (2006) Genetic analysis of landscape connectivity in tree populations. *Landsc Ecol* 21:821–836

Spiesman BJ, Inouye BD (2013) Habitat loss alters the architecture of plant—pollinator interaction networks. *Ecology* 94:2688–2696

Strohbach MW, Döring AO, Möck M et al (2019) The “hidden urbanization”: trends of impervious surface in low-density housing developments and resulting impacts on the water balance. *Front Environ Sci.* https://doi.org/10.3389/fenvs.2019.00029

Su J, Yin H, Kong F (2021) Ecological networks in response to climate change and the human footprint in the Yangtze

River Delta urban agglomeration, China. *Landsc Ecol* 36:2095–2112

Sun S, Jiang Y, Zheng S (2020) Research on ecological infrastructure from 1990 to 2018: a bibliometric analysis. *Sustainability* 12:2304

Taylor PD, Fahrig L, Henein K, Merriam G (1993) Connectivity Is a vital element of landscape structure. *Oikos* 68:571

Timmers R, Van Kuijk M, Verweij PA et al (2022) Conservation of birds in fragmented landscapes requires protected areas. *Front Ecol Environ* 20:361–369

Todd BD, Luhring TM, Rothermel BB, Gibbons JW (2009) Effects of forest removal on amphibian migrations: implications for habitat and landscape connectivity. *J Appl Ecol* 46:554–561

Tyndall J, Colletti J (2007) Mitigating swine odor with strategically designed shelterbelt systems: a review. *Agroforest Syst* 69:45–65

UNESCO (1984) International Experts Meeting on Ecological Approaches to Urban Planning, Suzdal, USSR, 1984; final report

Van Apeldoorn RC, Oostenbrink WT, Van Winden A, Van Der Zee FF (1992) Effects of habitat fragmentation on the bank vole, clethrionomys glareolus, in an agricultural landscape. *Oikos* 65:265

Velázquez Melero JC, Rodríguez-Espinosa VM (2023) Identification and assessment of green infrastructure in the community of Madrid. *Landsc Res* 48:297–312

Vellend M (2003) Habitat loss inhibits recovery of plant diversity as forests regrow. *Ecology* 84:1158–1164

Villers A (2019) Intérêt du suivi individuel pour la gestion du risque animalier : le cas de l'outarde canepetière sur la plateforme aéroportuaire de Marseille-Provence. *Faune Sauvage* 323:10–16

Vincent Q, Mangin N, Rigolot R, Blanchart A (2022) Étude de la biodiversité des sols (macrofaune épigée) urbains et péri-urbains dans le cadre de la création d'une trame brune. *Étude Et Gestion des Sols* 29:275–294

Voghera A, La Riccia L (2019) Ecological Networks in Urban Planning: Between Theoretical Approaches and Operational Measures. In: Calabro F, Della Spina L, Bevilacqua C (eds) *New Metropolitan Perspectives*. Springer International Publishing, Cham, pp 672–680

Voigt CC, Currie SE, Fritze M et al (2018) Conservation strategies for bats flying at high altitudes. *BioScience* 68:427–435

Vowles AS, Kemp PS (2021) Artificial light at night (ALAN) affects the downstream movement behaviour of the critically endangered European eel. *Anguilla Anguilla Environ Pollut* 274:116585

Wang T, Kaida N, Kaida K (2023) Effects of outdoor artificial light at night on human health and behavior: a literature review. *Environ Pollut* 323:121321

Ware HE, McClure CJW, Carlisle JD, Barber JR (2015) A phantom road experiment reveals traffic noise is an invisible source of habitat degradation. *Proc Natl Acad Sci USA* 112:12105–12109

Wiącek J (2023) How farmland birds react to traffic noise? *Belgian Journal of Zoology*. https://doi.org/10.26496/bjz.2023.108
Wickham JD, Riitters KH, Wade TG, Vogt P (2010) A national assessment of green infrastructure and change for the conterminous United States using morphological image processing. Landsc Urban Plan 94:186–195

Willems JS, Phillips JN, Francis CD (2022) Artificial light at night and anthropogenic noise alter the foraging activity and structure of vertebrate communities. Sci Total Envi-ron 805:150223

Wolters V (2001) Biodiversity of soil animals and its function. Eur J Soil Biol 37:221–227

Yan Y, Zhang W, Hu Y et al (2023) Spatially explicit subpixel-based study on the expansion of impervious surfaces and its impacts on soil organic carbon. Urban Ecosyst 26:793–806

Ying J, Zhang X, Zhang Y, Bilan S (2022) Green infrastruc-ture: systematic literature review. Econom Res Ekonom-ska Istraživanja 35:343–366

Zeale MRK, Stone EL, Zeale E et al (2018) Experimentally manipulating light spectra reveals the importance of dark corridors for commuting bats. Glob Change Biol 24:5909–5918

Zhang L, Ameca Y Juárez EI, Jiang Z (2018) Viability analy-sis of the wild sika deer (*Cervus nippon*) population in China: Threats of habitat loss and effectiveness of man-agement interventions. Journal for Nature Conservation 43:117–125

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