https://orcid.org/0000-0003-4342-5044
Figures
Abstract
Global narratives emphasise an increasingly urban world, yet the emerging parallel dynamics of counter-urbanisation, the population redistribution from higher- to lower-density settlements, is reshaping human–ecological relations. However, existing evidence on the ecological consequences associated with counter-urbanisation remains fragmented and lacks a unified framework to reconcile the diverse risks and opportunities. This review develops a framework for assessing these effects through two primary mechanisms: landscape pathways, involving the physical modification of landscapes through settlements and infrastructure development, and lifestyle pathways, reflecting changes in human presence, mobility, recreation, and consumption that do not directly alter land cover. By linking these pathways to different landscape contexts and relocation motivations, the review provides an ecological interpretation of counter-urbanisation. The framework can thereby inform policies that manage trade-offs, realise potential benefits and limit emerging pressures in an increasingly mobile world.
Author summary
It is widely recognised that the world is becoming more urban, yet growing numbers of people are also moving from cities to rural and suburban areas, a trend commonly described as counter-urbanisation. While the ecological impacts of city growth are well documented, the ecological consequences of this outward population movement and redistribution remain poorly understood. In this review, we show that these effects are neither consistently beneficial nor harmful, but arise through connected pathways: physical landscape transformations, such as new housing, roads, and farming changes, and lifestyle shifts, including commuting, recreation, and consumption. The outcomes depend strongly on who moves, why they move and the characteristics of the environment that receive them. We bring this evidence together in a framework that links relocation motivations and the local landscape context to changes in habitats, biodiversity and ecosystem services. Our work provides an ecological interpretation of counter-urbanisation and a practical framework for assessing and managing its complex ecological consequences.
Citation: Duan Q, Lai S, Sorichetta A, Eigenbrod F (2026) A review of ecological consequences of emerging counter-urbanisation. PLOS Ecosyst 1(2): e0000030. https://doi.org/10.1371/journal.pesy.0000030
Editor: Carolyn J. Lundquist, Central Queensland University - Gladstone Marina Campus: Central Queensland University - Gladstone Campus, AUSTRALIA
Received: February 4, 2026; Accepted: August 10, 2026; Published: October 1, 2026
Copyright: © 2026 Duan et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Funding: This work was supported by the ESRC South Coast Doctoral Training Partnership (ES/P000673/1 to QD). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
Competing interests: The authors have declared that no competing interests exist.
Introduction
Global narratives emphasise an increasingly urban world [1]. Over the past six decades, the world has undergone rapid urbanisation, with the urban population having grown from one to 4.4 billion, and more than half of the global population now living in urban areas [2] (Fig 1). Alongside this dominant pattern of urban population concentration, however, a contrasting pattern of population redistribution from larger urban centres to smaller and less densely populated places has emerged, particularly in the Global North [3,4] (Fig 1). This is commonly described as counter-urbanisation. Although the term has been used inconsistently in the literature, it is fundamentally a demographic concept, referring to relocation down the settlement hierarchy and, more broadly, to a deconcentrated settlement pattern [3]. It encompasses heterogeneous relocation flows driven by amenity seeking, retirement and housing affordability [5,6], and may overlap with suburbanisation and exurbanisation depending on their destination. As a demographic process, counter-urbanisation is distinct from low-density urban dispersal, which refers more directly to physical landscape transformation. Counter-urbanisation may be accompanied by such transformations, but they are not inherent to its definition. Nevertheless, this review examines the associated land-use changes as potential spatial expressions of population redistribution. The COVID-19 pandemic further reshaped relocation patterns, triggering an unprecedented global shock [7–9]. Lockdowns, concerns about disease transmission, and the widespread adoption of remote work prompted many to seek greener and more spacious living environments [10,11]. Although still modest globally [12], counter-urbanisation may signal a long-term shift in settlement and lifestyle preferences as development progresses worldwide [13].
a Data sources: World Bank [24]. b Changes in weekly night-time population during workdays in the UK during 2020−2022, as measured from Facebook data [13,25]. The basemap shapefile used to plot the data was obtained from the Global Administrative Areas (GADM) spatial database (version 4.1, https://gadm.org/). Zoomed-in views of major UK cities: (b-1) London; (b-2) Birmingham; (b-3) Liverpool, Manchester, Leeds & Sheffield; (b-4) Glasgow. Black polygons represent cities from the GHS – Urban Centre Database R2024A [26].
This population distribution shift creates new social and ecological interdependencies between urban, suburban, and rural regions, raising complex questions about ecological sustainability. Unlike urbanisation, whose ecological impacts are often closely associated with the direct expansion of built-up land [14,15], landscape changes associated with counter-urbanisation are more complex. Even when people leave cities, buildings, roads and other infrastructure usually remain, and ecological recovery occurs gradually. Urban depopulation may therefore create socioeconomic challenges through underused infrastructure and urban decay [16,17], while reduced anthropogenic pressure allows vacant land to develop into habitats supporting plants, insects, and birds [18]. Conversely, population growth in receiving rural areas may increase land consumption, infrastructure demand, and habitat loss [19]. Where this growth occurs around metropolitan fringes, the associated landscape changes may overlap with peri-urbanisation and other physical expressions of the broader urbanisation process, reproducing some of the ecological pressures associated with urban expansion. However, recent studies suggest the impacts may be more subtle [20], with new residents seeking to preserve the unique appeal of their new homes [20,21]. As a result, counter-urbanisation has been recognised as one potential pathway for forest transition, where forest cover shifts from net loss to net gain [22].
Beyond these gradually emerging landscape changes, counter-urbanisation also alters how people travel, consume resources and interact with their surroundings, thereby reshaping their ecological footprint. The shift from compact urban living to more dispersed settlement patterns may increase household energy and water demand, dependence on private vehicles, resource consumption and greenhouse gas emissions [17,23]. At the same time, new patterns of recreation and agriculture engagement may create both additional ecological pressures and new opportunities for environmental stewardship.
Despite ongoing discussions, evidence on the ecological consequences of counter-urbanisation remains fragmented and often contradictory, lacking a unified framework to reconcile the diverse risks, opportunities and trade-offs. Existing studies have focused mainly on physical changes in land use and landscape structure, giving less attention to changes in human mobility, resource use and everyday activities associated with population relocation. Furthermore, while ecological outcomes vary according to relocation motives and receiving-area contexts, existing literature has rarely synthesised these impacts cohesively.
Here, this review synthesises existing evidence and proposes a framework (Fig 2) for assessing the ecological consequences of counter-urbanisation. The framework distinguishes two complementary pathways: a landscape footprint arising from changes in land use, built form and habitat structure, and a lifestyle footprint arising from changes in mobility, resource consumption, recreation and agricultural engagement. This review draws predominantly on evidence from the Global North, where counter-urbanisation has been most extensively studied, while including relevant cases from other regions, such as China. The framework further recognises that ecological outcomes depend on diverse relocation motives and the landscape contexts of receiving areas. Finally, practical methods for evaluating these impacts are discussed.
This figure was created using resources from Flaticon.com, with specific author attributions provided in the Acknowledgments.
Landscape footprint: Physical modification of land use and land cover
Although counter-urbanisation is defined primarily in demographic terms, sustained population redistribution may also alter land use, land cover and built form. These physical modifications can directly affect habitats and associated ecological outcomes, even though such changes often emerge gradually. The relocation of people may reshape physical landscapes in both receiving and origin areas (Fig 3). In receiving areas, the form and ecological consequences of these changes depend on the landscape context. In rural and suburban settings, demographic growth alters competition for space among built-up land, cropland, and natural land. In origin cities, by contrast, population decline may lead to vacant plots and underused infrastructure.
These are simplified schematics that provide examples of potential consequences and do not capture more complex interactions. Photographs are from the public domain and free for public use. Icons are from Flaticon.com, with specific author attributions provided in the Acknowledgments.
Built development and settlement restructuring
The influx of new residents into rural areas often reshapes land use patterns, typically resulting in low-density, diffuse settlements [27]. To accommodate the growing population, built-up land, including roads, housing and other infrastructure, may expand [20,28]. For example, in Turkey, the designation of a national park has spurred a second-home and amenity real estate market, intensifying demand for housing in and around rural villages [29]. This physical expansion replaces natural or semi-natural vegetation, causing habitat loss, which remains a primary threat to biodiversity [30,31], while also reducing habitat integrity and connectivity. Mammals relying on large habitats [32], amphibians requiring connected aquatic and terrestrial habitats [33], and reptiles dependent on structurally complex vegetation [34] are often particularly sensitive, whereas disturbance-tolerant generalists and some non-native species may persist or increase [35]. The expansion of built environments can therefore contribute to biotic homogenisation, as invasive species increasingly dominating on the habitats [36,37]. In particular, the expansion of road networks outside of cities creates physical barriers that subdivide populations, restrict movement between feeding and breeding habitats [38], and also contribute to broader ecological degradation, including altered hydrology and increased erosion [39,40].
Although such expansion at the expense of natural land mirrors urbanisation patterns, its physical character differs. Low-density rural housing often retains a patchwork of gardens, hedgerows, and green spaces. Urban–rural gradient studies [41–43] indicate that species richness may be relatively high at low or moderate development intensity because these mixed habitats create structural heterogeneity and provide microclimatic refugia during environmental perturbations, as well as water, food and nest sites [44,45]. As a result, they can support higher species richness than surrounding monocultural farmland [46] and even intensive forest plantations [47]. Landscape perforation, although reducing the continuity of natural habitat, may also create new microhabitats or corridors that benefit some species relatively adapted to built environments, including birds and butterflies [48,49], and may enhance community recovery after local perturbations [44].
These effects depend on the surrounding landscape context. For example, settlement-associated gardens and hedgerows may increase habitat heterogeneity within intensively farmed landscapes in Europe [45], and dryland built patches supported higher bird richness than natural desert sites [50]. By contrast, similar low-density development in Australian woodland fringes may reduce biodiversity by fragmenting native vegetation and simplifying habitat structure [51]. Similar effects arise from the construction of recreational facilities with tourism development. Although golf courses and tennis courts can transform land into homogeneous, simplified and fragile landscapes compared with natural vegetation, they may maintain greater vegetative cover and soil stability than the intensive agricultural land they often replace, and support higher biodiversity [52].
Overall, biodiversity outcomes are shaped by settlement density, impervious surface cover, the quality, area and connectivity of retained habitats, and the landscape context [35,43,53]. However, much of this evidence derives from built expansion examined in the context of urbanisation. Although counter-urbanisation may generate comparable forms of built development, the extent, density and spatial configuration of this expansion remain poorly understood and require longer-term observation. Furthermore, navigating the ecological opportunities and challenges of these low-density built environments requires deliberate management and design to coordinate production, vegetation, and infrastructure, as illustrated by archaeological evidence of past low-density settlements [54–56].
Agricultural restructuring
The arrival of new residents can restructure rural economies in different ways, depending on their motivations and resources. In amenity and retirement destinations, demand may shift away from traditional agriculture toward tourism-oriented services and specialised high-value cultivation [20,21]. As land-use priorities shift toward recreational and commercial functions, the maintenance of traditional agriculture can be jeopardised [57]. For example, in the Alicante region of Spain, a popular destination for retirement relocation, local agriculture has shifted from traditional crops to more intensive, high-value cultivation (e.g., irrigated fruits and vineyards). Where these systems retain natural vegetation, field margins or vegetated inter-rows, they may support local biodiversity [58]. At the same time, the shift towards non-agricultural services to fulfil the demands of tourists and retirees and open up further income sources [19,59] can accelerate agricultural abandonment and enable natural regeneration (rewilding) on some marginal lands. Counter-urbanisation can therefore contribute to forest transition (i.e., forest recovery from net loss to net gain) through the agricultural dis-intensification pathway [22], benefiting many forest- and shrubland-associated bird and mammal species [60].
Similar restructuring may occur among return migrants. Some returnees use accumulated capital, skills and market connections to shift towards commercial, specialised or capital-intensive agriculture [61]. Others may restore household labour to smallholdings and sustain traditional farming practices, potentially helping to maintain agro-biodiversity [62]. Amenity-oriented migrants may follow a different pathway by prioritising landscape aesthetics, recreation and conservation over agricultural production [63]. These preferences can increase the provision of ecosystem services, such as habitat conservation, scenic value, and soil restoration, or create opportunities for small-scale, niche agro-ecological activities. However, limited agricultural knowledge, equipment or time may lead to intermittent management or partial abandonment, altering vegetation structure and landscape connectivity, for example, through woody plant encroachment in rangelands [63].
Furthermore, local rewilding gains may be partly offset by indirect land-use change. The reduction of agriculture in areas receiving new residents, which are often located on relatively productive land, can displace food production to other regions. This may trigger cropland expansion or intensification elsewhere to meet existing demand, causing habitat loss and biodiversity pressure [64,65].
Stewardship and landscaping of natural and amenity land
Since the pursuit of natural amenities is one of the major drivers of counter-urbanisation, new rural residents may contribute to natural land recovery and conservation to protect the scenic views they value [21,66]. Drawn by the aesthetic and ecological appeal of rural landscapes, these new residents may engage in environmental activism and land stewardship, opposing the conversion of natural land into hardened surfaces and prompting the recovery of green lands [67]. For example, in one village near Shanghai, newcomers opposed the full-scale hardening of riverbanks [21]. Similarly, in Canada, new residents seeking a rural lifestyle have restored native riparian vegetation and reallocated water from irrigation to instream flows to sustain trout populations [68].
Beyond protection, new residents may actively modify the landscape to enhance diversity and scenic appeal. Motivated by leisure, aesthetic preference or the desire to attract tourists, they often aim to diversify and enhance scenic views by introducing a broad variety of plants [21,22]. For example, homestay operators in China invest substantial resources in landscaping to create environments that serve both tourism and personal enjoyment [21]. These plantings include both native [69] and non-native species [70]. In rural tourism villages in tropical China, ornamental cultivation was the principal source of non-native garden plants [71]. Similar preferences are also reflected in their housing renovations. One study in Australia found that over 60% of surveyed new residents chose to plant either a mix of native and non-native species or predominantly non-natives when establishing gardens or landscaping around their homes [72].
While the intention to enhance scenic views can theoretically support biodiversity by introducing a wider variety of plant species and creating habitats for birds and wildlife [45], human landscaping preferences often exacerbate biodiversity pressure. The pursuit of a ‘tidy’ aesthetic can drive biotic homogenisation, and some introduced non-native plants may become invasive and outcompete native flora [70]. For example, Norway maple (Acer platanoides), a shade tree introduced to eastern deciduous forests of North America, has been found to outcompete native maples and beeches [73]. Such aesthetic-driven modifications can also disrupt local pollinator networks through phenological mismatches between exotic blooms and native insect activity [74], while non-native plants may reduce habitat quality for insectivorous birds where they support fewer native insects [75]. Furthermore, introduced exotics often host generalist soil-borne pathogens which can cross-infect and degrade the health of surrounding native vegetation [76], potentially causing unpredictable disruptions to local ecosystems.
Origin urban vacancy and ecological succession
The landscape transformations associated with counter-urbanisation are not confined to rural receiving regions. Where sustained population dispersal contributes to urban population decline, source cities may experience housing abandonment, disused facilities and the formation of vacant land [77]. Counter-urbanisation is not the sole cause of these changes; rather, urban vacancy represents a linked but distinct process that may generate ecological change in source cities.
These vacant sites present opportunities for reuse by reinforcing green infrastructure [78], establishing urban agriculture [79], or creating ‘voids’ that can serve as biodiversity sanctuaries [80]. Whether through the creation of gardens and greenways or simply allowing spontaneous vegetation to colonise vacant lots, these spaces may contribute to biodiversity recovery by providing new habitats and ecological niches [77,81]. Many studies have shown that wastelands can harbour more species than other urban green spaces such as managed parks, lawns and urban forests [82–84]. For example, in the Hauts-de-Seine département near Paris, plant richness in urban wastelands represented 58% of the total recorded for the entire area [85]. In Detroit, researchers discovered that despite the urban setting, vacant lots hosting wildflowers supported a diversity and abundance of bumblebees comparable to rural nature reserves [18]. Although unmanaged regeneration raises concerns regarding invasiveness [86], these non-native species provide critical ecosystem services. A study of vacant lots in Cleveland found that trees on these lands provided roughly triple the value in ecosystem services per hectare compared to occupied residential lots, with exotic species, such as the tree-of-heaven, contributing approximately half of this value [18,87].
Importantly, the biodiversity value of urban vacant land depends on its origin and land-use legacy [88]. Former residential lots may retain garden vegetation and mature trees [89], whereas industrial and transport sites can develop heterogeneous open habitats on rubble or nutrient-poor substrates [90]. It also varies with the surrounding landscape context. For example, nearby green spaces can provide habitat and facilitate species movement, while surrounding buildings may increase isolation and restrict seed dispersal [88]. Site size and age are also key determinants of ecological succession and species richness [88]. These factors should therefore be considered together when evaluating the ecological effects of urban vacancy associated with counter-urbanisation.
Lifestyle footprint: Changes in human presence and activities
Unlike urbanisation, which is typically measured by the direct physical transformation of landscapes, the impacts of counter-urbanisation are more subtle and complex to quantify. While counter-urbanisation may involve built-up expansion, new rural residents often modify existing housing stock and maintain the low-density character of the environment. Furthermore, land abandonment is a long-term process that does not easily revert to its original natural state. Consequently, a primary driver of environmental pressure in counter-urbanisation is not just where people live, but how they live, specifically through three primary dimensions: daily living requirements, cultural tourism engagement, and agricultural stewardship (Fig 4).
These are simplified schematics that provide examples of potential consequences and do not capture more complex interactions. Photographs are from the public domain and free for public use. Icons are from Flaticon.com, with specific author attributions provided in the Acknowledgments.
Daily living
The shifts in lifestyle to low-density living [17,91] fundamentally alters the metabolic profile of the household. In the United States, for example, household carbon footprints are typically higher in outer suburbs compared to urban cores [92]. A major contributor to this difference is the energy required to heat, cool and power homes. Housing stock in outer-suburban and rural areas is generally dominated by detached single-family homes, which tend to be larger and consume more energy than urban apartments [93,94]. This is also reflected in conversion factors from the US Department of Energy, which show that larger units in lower-density areas consume more energy and create higher emissions, in both construction and operation, compared to smaller units in multi-family buildings [95,96]. Additionally, a comprehensive life-cycle assessment (LCA) accounting for both operational energy (e.g., heating, electricity) and embodied energy (e.g., materials for construction) found that suburban neighbourhoods dominated by detached homes consume up to 320% more embodied energy, 150% more operational energy, and about 160% more total life-cycle energy per capita than densely developed neighbourhoods composed mainly of low-rise apartments and duplexes [97].
Similarly, water demand often rises because low-density residential development tends to increase overall water use due to the irrigation of lawns and gardens [98]. Some newcomers, even with reduced agricultural needs, cite the personal satisfaction they derive from irrigating their land [99]. Compared to the low irrigation needs of city living, this can place considerable strain on local water resources, particularly in water-scarce regions [19]. However, in these dry regions, supplement irrigation may increase vegetation cover and plant diversity [50].
Beyond the home, rural and outer suburban areas often lack adequate public transportation infrastructure, leading to increased reliance on private vehicles [100]. Data from the U.S. National Household Travel Survey shows that 97% of rural households are more car-dependent than urban ones, with higher rate of car ownership (97% vs. 92%) and car use for trips (91% vs. 86%) [101]. Similar urban-rural gradients are evident in England. Only around 5% of households in rural villages have no car, compared with 33% in urban conurbations. Consistent with this, car travel accounted for 75% of trips in the most rural areas in 2021, versus 51% in urban conurbations [102]. Furthermore, many residents relocating to outer suburbs, due to high urban housing costs in core urban areas, continue to rely on urban-based workplaces and schools [3]. As a result, they often have a higher ecological footprint from commuting compared to residents in more centrally located neighbourhoods [103–105]. Crucially, this increased traffic volume has ecological consequences beyond emissions. The rise in vehicle frequency on rural roads, which are traditionally characterised by low traffic, increases the risk of wildlife roadkill [23] and triggers avoidance behaviours due to traffic noise and vehicle presence that subdivide animal populations into more vulnerable fractions [39,106]. Increase traffic intensity may also facilitate seed dispersal and elevate the risk of plant invasion, as exemplified by the spread of common ragweed (Ambrosia artemisiifolia L.) across road networks [107].
Finally, daily routines introduce direct biotic disturbances. Indeed, the lifestyle of the new residents often includes domestic pets, especially free-roaming cats and dogs, which act as introduced predators with severe impacts on bird and small mammal populations [108]. One estimate suggests that free-ranging domestic cats kill 1.3–4.0 billion birds and 6.3–22.3 billion mammals annually in the United States [109]. These impacts are often more pronounced in rural areas, where pets have greater access to wildlife than those in suburban or urban environments [108]. Furthermore, the daily mobility of rural residents, including activities such as dog-walking and outdoor exercise, increases the frequency of human-wildlife encounters [110], creating disturbances that may prompt wildlife to alter their temporal and spatial behaviours [110–112]. Simultaneously, this increased proximity facilitates pathogen pollution, enabling diseases to spill back into wildlife populations (reverse zoonosis) and imposing a biological pressure that can devastate susceptible native species [113]. Additionally, increased foot traffic can lead to the physical trampling of vegetation, which compacts soil and damages sensitive understory flora [114].
Cultural engagement and tourism
The counter-urbanisation lifestyle is heavily influenced by the aesthetic appeal of the countryside, propelling local economic transformation and development [115,116]. New residents often act as catalysts for nature-based cultural activities, actively shifting the local economy from traditional production to service-oriented sectors such as birdwatching, photography tours, and eco-tourism [117,118]. The distinctive natural landscapes and cultural activities attract tourists, fostering a unique regional identity that draws substantial funding and investment toward conservation initiatives [119].
However, these activities inevitably increase the spatial and temporal overlap between humans and wildlife [110–112]. Higher visitor frequencies in sensitive habitats can disrupt foraging, breeding, and resting behaviours, potentially stressing local species [120–122]. For example, recreational activities in coastal areas can reduce the feeding success and survival of plover chicks [121]. Thus, it is essential to balance the economic benefits of the new nature-based economy, which generates the necessary funds to protect the ecosystem, with the risk that increased human presence may inadvertently degrade the very biodiversity it seeks to preserve.
Agricultural stewardship and practices
The lifestyle preferences of new residents can transform local agricultural systems through both consumption demands and active participation. First, the specific need for green and organic agricultural products among this demographic acts as a market force, prompting the local farming industry to pivot toward higher environmental standards [123,124]. Simultaneously, new residents who often possess scientific knowledge may introduce advanced sustainable farming techniques to local farmers, thereby upgrading regional production capabilities [123,125]. Beyond influencing the industry, these residents also personally maintain or revive agricultural production, motivated not by economic returns but by a desire to experience ‘farming culture’ and rural identity [49,126]. This form of production differs from conventional productivist activities, as it typically relies on organic methods and avoids the heavy use of fertilisers, pesticides, and other agrochemicals that are harmful to the environment [21,49,127]. This may benefit soil organisms, pollinators and other farmland species [128], although the magnitude of these benefits varies with farming intensity [129], and surrounding landscape context [130].
From pathways to ecological consequences
The ecological consequences of counter-urbanisation are complex and highly variable, shaped by the interaction of landscape and lifestyle pathways that can both alleviate and intensify ecological pressures. As shown above, their direction and magnitude depend on the relocation motivations, the landscape contexts of source and receiving regions, the baseline against which change is assessed, and the timescale of evaluation.
To interpret the diversity of ecological responses triggered by counter-urbanisation, it is important to distinguish between where relocations occur and who is relocating, while recognising that these dimensions are interdependent. In terms of context, rural and peri-urban landscapes can be grouped into categories such as peri-urban production belts under strong metropolitan influence, intensive agricultural regions and high-nature-value areas [131]. In terms of motivations, new residents can be grouped by dominant motivation and life stage, including amenity seekers [5], remote workers [132], affordability-driven movers [133], crisis-induced movers [134,135] or tourism entrepreneurs [21,136]. In practice, local housing markets, labour-market links and environmental qualities filter which mover groups may arrive in each setting [137,138]. Thus, specific context-motivation combinations drive distinct ecological pathways and consequences. Three illustrative combinations are discussed below:
1. Affordability-driven movers in peri-urban or suburban areas.
Affordability-driven relocation, prompted by high housing and living costs in cities, is often directed towards peri-urban or suburban areas under strong metropolitan influence [133]. These relocators often prioritise housing affordability over environmental quality while remaining employed in urban centres, reinforcing energy-intensive housing and car-dependent sprawl that increases household energy use and commuting emissions [93,94]. Regular commuting can also increase wildlife roadkill [23] and facilitate the spread of invasive plants across the urban-rural gradient [107], contributing to biotic homogenisation as sensitive native species are replaced by urban-adapted generalists. Furthermore, the expansion of road networks and impervious surfaces to fulfil increasing demand can fragment habitats, subdividing contiguous natural areas into isolated units. These changes can also degrade ecosystem services such as pollination, carbon storage and water regulation.
2. Tourism entrepreneurs in traditional agricultural landscapes.
Tourism entrepreneurs may be attracted to traditional agricultural landscapes where natural amenities, farming heritage and cultural character support rural tourism. They may drive a shift from conventional farming to high-value cultivations, such as fruit production linked to fruit-picking and other experience-based activities, or contribute to agricultural abandonment as land and labour move into the service economy [21,22,136]. Such abandonment can trigger passive rewilding and benefit various native species [60]. To enhance aesthetic appeal and attract visitors, these relocators often landscape their properties with ornamental, non-native plants, which can provide habitat but may also increase invasion risk and disrupt local ecosystems [71]. Meanwhile, the expansion of accommodation, transport networks, and recreational facilities directly converts and fragments habitats [30], while seasonal visitor influxes further disturb wildlife and reduce the functional availability of feeding, breeding and resting habitats [110–112].
3. Amenity seekers and remote workers in high-nature-value landscapes.
Nature-amenity seekers [5] and remote workers [132] may be drawn to high-nature-value landscapes. Compared with tourism entrepreneurs, they may be more likely to seek quieter locations outside tourism hotspots and to integrate into the existing local environment, rather than substantially transforming it. Some of these relocators may have relatively high environmental awareness and disposable income, increasing their willingness and capacity to support conservation initiatives, oppose environmentally damaging development, and landscape their private gardens [21,66]. These activities can improve habitat condition, support native biodiversity and strengthen ecosystem services such as pollination and local climate regulation. However, preferences for particular landscaping aesthetics may introduce non-native plants, potentially disrupting local ecological communities [72]. Amenity seekers may also engage in small-scale, low-intensity agriculture to experience farming and rural culture, helping to maintain habitat heterogeneity and agrobiodiversity, while partial abandonment may promote vegetation succession and benefit some species [60,115]. At the same time, closer engagement with the surrounding environment may increase recreational pressure through activities such as running and dog walking, disturbing native wildlife and degrading sensitive vegetation through trampling [114] (Fig 5).
Dashed boxes indicate examples of datasets that may be used to quantify these complex dynamics.
Landscape context can be characterised using land-cover composition, infrastructure maps, landscape configuration, nature value and human accessibility. Broader geographical conditions, including climate and topography, should also be considered because they influence settlement suitability, species distributions and ecosystem sensitivity. These characteristics could be synthesised using classifications such as land-system archetypes [139,140] to support further analysis of the context dependency of counter-urbanisation impacts. Although relocation motivations are difficult to capture directly, they can be identified through structured household surveys and inferred from demographic characteristics, such as age and household composition, and socio-economic metrics, including local house prices, income or deprivation, employment patterns and housing tenure. Because movers within the same landscape context often have diverse motivations, their relative prevalence can be represented as proportions. The timescale of evaluation is critical. In the short term, ecological effects may arise primarily through lifestyle changes, which can be assessed using mobility traces [141], tourism statistics and household energy/utility data [142]. Over longer periods, cumulative settlement expansion, infrastructure development, agricultural restructuring and land abandonment transform landscapes more fundamentally; these changes can be tracked by high-temporal-resolution land cover products [143], building footprints [144], and AI-driven geospatial foundation models capable of recognising subtle landscape changes (e.g., Alpha Earth [145]). Measurable ecological outcomes can then be assessed across three dimensions: habitat structure, including habitat extent, patch size, fragmentation and connectivity; biodiversity, including species richness, distribution, community composition and invasive-species occurrence; and ecosystem services, including carbon storage, pollination and water regulation.
Finally, observed outcomes can be compared with clearly defined counterfactual baselines representing the most plausible alternative trajectory for each context–relocation combination. For example, in peri-urban areas favoured by counter-urban movers, short-term changes in traffic volume and household energy use, together with longer-term settlement development, could be compared with a continued urban-densification trajectory. This would help determine whether counter-urbanisation shifts development towards more dispersed peri-urban settlement, changes the amount of land conversion and increases habitat fragmentation relative to a more compact growth pattern. In intensive agricultural regions, tourism-related restructuring and changes in cropland patterns could be compared with the continuation of existing farming practices or ongoing rural depopulation. Where these regions are close to cities, they could also be compared with the trajectory expected under continued urban expansion. Depending on data availability, counterfactual comparisons can employ longitudinal to track ecological indicators before and after relocation- or redistribution-driven change, matched control areas selected for similar environmental and socio-economic conditions but contrasting population redistribution patterns, or scenario-based modelling that projects how land use and ecological conditions would have evolved without counter-urban relocation. Comparing the same indicators across observed and counterfactual trajectories would help isolate the net ecological effects attributable to counter-urbanisation.
Summary and future perspectives
This review shows that counter-urbanisation is reshaping human-ecology relations in ways that are distinct from the classic narrative of urbanisation. Rather than a dominant process, it consists of heterogeneous flows operating through both landscape and lifestyle pathways that can either alleviate or intensity ecological pressure. A central contribution of this review is to reinterpret counter-urbanisation within an ecological context. Although fundamentally a demographic process, it can be understood ecologically as the spatial redistribution of human metabolic demands, land-use pressures and everyday environmental impacts from urban centres towards small settlements and rural areas. The proposed framework therefore links the motivations for relocation and the characteristics of receiving landscape contexts to the landscape and lifestyle pathways through which these redistributed pressures generate ecological consequences.
Future research should combine this framework with various datasets and complementary quantitative and qualitative evidence to jointly track population redistribution, land-use change, human activity and ecological indicators. Developing transferable typologies of landscape contexts and relocating groups will be essential for synthesising case-study findings across regions. This integrated approach will also allow researchers to explore how alternative policy choices steer rural trajectories under different scenarios, informing proactive planning for nature recovery and climate resilience, rather than necessitating reactive, crisis-driven interventions.
Acknowledgments
For the purpose of open access, the author has applied a Creative Commons attribution license (CC BY) to any Author Accepted Manuscript version arising from this submission. The icons used in figures were designed by srip, Icongeek26, Mehwish, Magnific, Nuricon, kornkun, ekays.dsgn, BZZRINCANTATION, Iconjam, Smashicons and Prashanth Rapolu 15 from https://www.flaticon.com.
References
- 1. Wahba Tadros SN, Wellenstein A, Das MB, Palmarini N, D’Aoust OS, Singh G. Demographic Trends and Urbanization. 158009. Washington, D.C.: World Bank Group. 2021. A vailable from: http://documents.worldbank.org/curated/en/260581617988607640/Demographic-Trends-and-Urbanization
- 2.
World Bank. World Bank Open Data. 2022. Available from: https://data.worldbank.org
- 3. Mitchell CJA. Making sense of counterurbanization. J Rural Stud. 2004;20(1):15–34.
- 4. Rowe F, Bell M, Bernard A, Charles-Edwards E, Ueffing P. Impact of internal migration on population redistribution in Europe: Urbanisation, counterurbanisation or spatial equilibrium?. CPoS. 2019;44.
- 5. Halfacree K. To revitalise counterurbanisation research? Recognising an international and fuller picture. Popul Space Place. 2008;14(6):479–95.
- 6. Halfacree K. Heterolocal identities? Counter-urbanisation, second homes, and rural consumption in the era of mobilities. Popul Space Place. 2012;18:209–24.
- 7. Gamlen A. Migration isn’t increasing, border restrictions don’t reduce crossings - and other home truths. Nature. 2023;623(7988):683–4. pmid:37985836
- 8. Wolff M, Mykhnenko V. COVID-19 as a game-changer? The impact of the pandemic on urban trajectories. Cities. 2023;134:104162. pmid:36593903
- 9. Halfacree K. Counterurbanisation in post-covid-19 times. Signifier of resurgent interest in rural space across the global North?. J Rural Stud. 2024;110:103378.
- 10. Stawarz N, Rosenbaum-Feldbrügge M, Sander N, Sulak H, Knobloch V. The impact of the COVID-19 pandemic on internal migration in Germany: A descriptive analysis. Popul Space Place. 2022;28(6):e2566. pmid:35601664
- 11. Ilham MA, Fonzone A, Fountas G, Mora L. To move or not to move: a review of residential relocation trends after COVID-19. Cities. 2024;151:105078.
- 12. Niva V, Horton A, Virkki V, Heino M, Kosonen M, Kallio M, et al. World’s human migration patterns in 2000-2019 unveiled by high-resolution data. Nat Hum Behav. 2023;7(11):2023–37. pmid:37679443
- 13. Duan Q, Lai S, Sorichetta A, Tatem AJ, Steele J, Eigenbrod F. COVID-19 and urban exodus: diverging population redistribution patterns across countries from 2020 to 2022. npj Urban Sustain. 2026;6: 59.
- 14. Guetté A, Gaüzère P, Devictor V, Jiguet F, Godet L. Measuring the synanthropy of species and communities to monitor the effects of urbanization on biodiversity. Ecol Indic. 2017;79:139–54.
- 15. Kondratyeva A, Knapp S, Durka W, Kühn I, Vallet J, Machon N. Urbanization effects on biodiversity revealed by a two-scale analysis of species functional uniqueness vs. redundancy. Front Ecol Evol. 2020;8.
- 16. Faust KM, Abraham DM, DeLaurentis D. Coupled human and water infrastructure systems sector interdependencies: framework evaluating the impact of cities experiencing urban decline. J Water Resour Plann Manage. 2017;143(8).
- 17. Sutradhar U, Spearing L, Derrible S. Depopulation and associated challenges for US cities by 2100. Nat Cities. 2024;1(1):51–61.
- 18. Kwok R. Accidental urban oases. Proc Natl Acad Sci USA. 2018;115(19):4800–4.
- 19. Zasada I, Alves S, Müller FC, Piorr A, Berges R, Bell S. International retirement migration in the Alicante region, Spain: process, spatial pattern and environmental impacts. J Environ Plan Manage. 2010;53(1):125–41.
- 20. Herrero-Jáuregui C, Concepción ED. Effects of counter-urbanization on Mediterranean rural landscapes. Landsc Ecol. 2023;38: 3695–711.
- 21. Yu L, Wang Y, Li M. The emergence of counter-urbanisation in China: can it be a pathway for rural revitalisation?. Habitat Int. 2024;144:102998.
- 22. Jimenez YG, Aráoz E, Fernandez RD, Nanni S, Ovejero R, Paolini L, et al. Counterurbanization: a neglected pathway of forest transition. Ambio. 2022;51(4):823–35. pmid:34677811
- 23. Denneboom D, Bar-Massada A, Shwartz A. Wildlife mortality risk posed by high and low traffic roads. Conserv Biol. 2024;38(2):e14159. pmid:37551769
- 24.
World Bank. Urban population (% of total population). 2023. Available from: https://data.worldbank.org/indicator/SP.URB.TOTL.IN.ZS
- 25. Duan Q, Steele J, Cheng Z, Cleary E, Ruktanonchai N, Voepel H, et al. Identifying counter-urbanisation using Facebook’s user count data. Habitat Int. 2024;150:103113.
- 26. Mari Rivero I, Melchiorri M, Florio P, Schiavina M, Goch K, Politis P. GHS-UCDB R2024A - GHS Urban Centre Database 2025. European Commission, Joint Research Centre (JRC). 2024.
- 27. Cuadrado-Ciuraneta S, Durà-Guimerà A, Salvati L. Not only tourism: unravelling suburbanization, second-home expansion and “rural” sprawl in Catalonia, Spain. Urban Geogr. 2016;38(1):66–89.
- 28. Woods M. Rural Geography: Processes, Responses and Experiences in Rural Restructuring. London: SAGE Publications; 2011.
- 29. Hurley PT, Arı Y. Saying “No” to (the) Oxygen Capital? Amenity migration, counter-territorialization, and uneven rural landscape change in the Kaz Dağları (Ida Mountains) of western Turkey. J Rural Stud. 2018;62:195–208.
- 30. Newbold T, Hudson LN, Hill SLL, Contu S, Lysenko I, Senior RA, et al. Global effects of land use on local terrestrial biodiversity. Nature. 2015;520(7545):45–50. pmid:25832402
- 31. Jaureguiberry P, Titeux N, Wiemers M, Bowler DE, Coscieme L, Golden AS, et al. The direct drivers of recent global anthropogenic biodiversity loss. Sci Adv. 2022;8(45):eabm9982. pmid:36351024
- 32. Milda D, Ashish K, Ramesh T, Kalle R, Thanikodi M. Evaluation of anthropogenic pressure on the occupancy patterns of large mammals in the Western and Eastern Ghats. Landsc Ecol. 2023;38:409–22.
- 33. Caballero-Díaz C, Tarroso P, Sánchez-Montes G, Reyes-Moya I, Polo-Cavia N, Martínez-Solano Í. Connecting the dots: regional assessment of landscape connectivity in amphibian communities in Central Spain. Landsc Ecol. 2025;40:155.
- 34. Wright LJ, Nimmo DG, Green MC, Simpson REL, Wassens S, Michael DR. Remnant Habitat Patches Have Lower Reptile Richness and Diversity Compared to Areas of Continuous Habitat. Austral Ecology. 2025;50(3).
- 35. Müller N, Ignatieva M, Nilon CH, Werner P, Zipperer WC. Patterns and trends in urban biodiversity and landscape design. In: Elmqvist T, Fragkias M, Goodness J, Güneralp B, Marcotullio PJ, McDonald RI, editors. Urbanization, biodiversity and ecosystem services: challenges and opportunities: a global assessment. Dordrecht: Springer Netherlands; 2013. p. 123–74.
- 36. van Rensburg BJ, Peacock DS, Robertson MP. Biotic homogenization and alien bird species along an urban gradient in South Africa. Landsc Urban Plan. 2009;92(3–4):233–41.
- 37. McKinney ML. Urbanization as a major cause of biotic homogenization. Biol Conserv. 2006;127(3):247–60.
- 38. Dániel‐Ferreira J, Berggren Å, Wissman J, Öckinger E. Road verges are corridors and roads barriers for the movement of flower‐visiting insects. Ecography. 2021;2022(2).
- 39. van Strien MJ, Grêt-Regamey A. Global expansion of the ecological impact of extra-urban road traffic. Nat Sustain. 2025;8:1294–303.
- 40. Forman RTT, Alexander LE. Roads and their major ecological effects. Annu Rev Ecol Syst. 1998;29(1):207–31.
- 41. McKinney ML. Effects of urbanization on species richness: a review of plants and animals. Urban Ecosyst. 2008;11(2):161–76.
- 42. Čiliak M, Čejka T, Tej B, Oboňa J, Manko P. Species richness patterns and community structure of land snail communities along an urban-rural gradient in river floodplains. Urban Ecosyst. 2024;27(3):953–63.
- 43. Kowarik I. Novel urban ecosystems, biodiversity, and conservation. Environ Pollut. 2011;159(8–9):1974–83. pmid:21435761
- 44. Oliver TH, Heard MS, Isaac NJB, Roy DB, Procter D, Eigenbrod F. Biodiversity and resilience of ecosystem functions. Trends Ecol Evol. 2015;30:673–84.
- 45. Clark K, Mauchline AL, Blanuša T, Felton M, Potts S, Garratt MPD. The multiple benefits delivered by hedgerows: where is the evidence and does it meet current knowledge needs?. People Nat. 2025;7(10):2437–50.
- 46. Phillips M, Page S, Saratsi E, Tansey K, Moore K. Diversity, scale and green landscapes in the gentrification process: traversing ecological and social science perspectives. Appl Geogr. 2008;28(1):54–76.
- 47. Haskell DG, Evans JP, Pelkey NW. Depauperate avifauna in plantations compared to forests and exurban areas. PLoS One. 2006;1(1):e63. pmid:17183694
- 48. Bock CE, Bock JH. Biodiversity and residential development beyond the urban fringe. In: Esparza AX, McPherson G, editors. The planner’s guide to natural resource conservation: the science of land development beyond the metropolitan fringe. New York, NY: Springer; 2009. p. 59–84.
- 49. Abrams J, Gosnell H, Gill N, Klepeis P. Re-creating the Rural, Reconstructing Nature: An International Literature Review of the Environmental Implications of Amenity Migration. Conservat Soc. 2012;10(3):270.
- 50. Diamant ES, Oswald KN, Awoyemi AG, Gaston KJ, MacGregor-Fors I, Berger-Tal O, et al. The importance of biome in shaping urban biodiversity. Trends Ecol Evol. 2025;40(6):601–12. pmid:40254468
- 51. Suárez-Castro AF, Maron M, Mitchell MGE, Rhodes JR. Disentangling direct and indirect effects of landscape structure on urban bird richness and functional diversity. Ecol Appl. 2022;32(8):e2713. pmid:36196040
- 52. Petrosillo I, Valente D, Pasimeni MR, Aretano R, Semeraro T, Zurlini G. Can a golf course support biodiversity and ecosystem services? The landscape context matter. Landscape Ecol. 2019;34(10):2213–28.
- 53. Kowarik I. Urban biodiversity, ecosystems and the city. Insights from 50 years of the Berlin School of urban ecology. Landsc Urban Plan. 2023;240: 104877.
- 54. Isendahl C, Dunning NP, Grazioso L, Hawken S, Lentz DL, Scarborough VL. Growth and decline of a sustainable city: a multitemporal perspective on blue-black-green infrastructures at the pre-Columbian Lowland Maya city of Tikal. Urban Stud. 2024;62(3):487–506.
- 55. Hawken S, Fletcher R. A Long-term archaeological reappraisal of low-density urbanism: implications for contemporary cities. J Urban Archaeol. 2021;3:29–50.
- 56. Hawken S, Isendahl C. Patch urbanism: towards an integrated theoretical framework for examining spatial and temporal dynamics in the Asian rice belt. Urban Stud. 2025;63(3):484–504.
- 57. Akgün AA, Baycan-Levent T, Nijkamp P, Poot J. Roles of local and newcomer entrepreneurs in rural development: a comparative meta-analytic study. Reg Stud. 2011;45(9):1207–23.
- 58. Paiola A, Assandri G, Brambilla M, Zottini M, Pedrini P, Nascimbene J. Exploring the potential of vineyards for biodiversity conservation and delivery of biodiversity-mediated ecosystem services: a global-scale systematic review. Sci Total Environ. 2020;706:135839. pmid:31846877
- 59. Buciega A, Pitarch M-D, Esparcia J. The context of rural–urban relationships in Finland, France, Hungary, The Netherlands and Spain. J Environ Policy Plan. 2009;11(1):9–27.
- 60. Crawford CL, Wiebe RA, Yin H, Radeloff VC, Wilcove DS. Biodiversity consequences of cropland abandonment. Nat Sustain. 2024;7:1596–607.
- 61. Qian W, Wang D, Zheng L. The impact of migration on agricultural restructuring: evidence from Jiangxi Province in China. J Rural Stud. 2016;47:542–51.
- 62. Yaregal Y, Sime G. Traditional home garden agro‐biodiversity dynamics, agro‐ecosystem services, and management practices in smallholder farmers setting, South‐Central Ethiopia. Food Energy Secur. 2024;13(4).
- 63. Matarrita-Cascante D. More than a population movement: Post-COVID amenity migration and the contemporary social-ecological transformation of rural working lands. Land Use Policy. 2026;169:108136.
- 64. van Vliet J. Direct and indirect loss of natural area from urban expansion. Nat Sustain. 2019;2:755–63.
- 65. Li F, Wu S, Liu H, Yan D. Biodiversity loss through cropland displacement for urban expansion in China. Sci Total Environ. 2024;907:167988. pmid:37875196
- 66. Zhang Y, Yasin SM, Sun L. The implications of urban-to-rural migration on rural development: A systematic literature review. GeoJournal. 2025;90(3).
- 67. Halfacree K. ‘Glow worms show the path we have to tread’: the counterurbanisation of Vashti Bunyan. Soc Cult Geogr. 2009;10(7):771–89.
- 68. Gosnell H, Haggerty JH, Byorth PA. Ranch ownership change and new approaches to water resource management in southwestern Montana: implications for fisheries1. J Am Water Resour Assoc. 2007;43(4):990–1003.
- 69. Jimenez YG, Aráoz E, Grau HR, Paolini L. Linking forest transition, plant invasion and forest succession theories: socioeconomic drivers and composition of new subtropical andean forests. Landsc Ecol. 2021;36(4):1161–76.
- 70. Gill N, Klepeis P, Chisholm L. Stewardship among lifestyle oriented rural landowners. J Environ Plan Manage. 2010;53(3):317–34.
- 71. Yang M, Zhang D, Geng Y. Non-native flora changes in rural gardens of China: the role of tourism. Discov Sustain. 2024;5:129.
- 72. Kurz T, Baudains C. Biodiversity in the front yard: an investigation of landscape preference in a domestic urban context. Environ Behav. 2012;44:166–96.
- 73. Webb SL, Pendergast TH IV, Dwyer ME. Response of native and exotic maple seedling banks to removal of the exotic, invasive norway maple (Acer platanoides). J Torrey Bot Soc. 2001;128(2):141.
- 74. Garbuzov M, Alton K, Ratnieks FLW. Most ornamental plants on sale in garden centres are unattractive to flower-visiting insects. PeerJ. 2017;5:e3066. pmid:28286716
- 75. Narango DL, Tallamy DW, Marra PP. Nonnative plants reduce population growth of an insectivorous bird. Proc Natl Acad Sci U S A. 2018;115(45):11549–54. pmid:30348792
- 76. Waller LP, Allen WJ, Black A, Condron L, Tonkin JD, Tylianakis JM, et al. Asymmetric sharing of generalist pathogens between exotic and native plants correlates with exotic impact in communities. J Ecol. 2024;112(10):2264–76.
- 77. Lima MF, Eischeid MR. Shrinking cities: rethinking landscape in depopulating urban contexts. Landsc Res. 2017;42(7):691–8.
- 78. Schetke S, Haase D. Multi-criteria assessment of socio-environmental aspects in shrinking cities. Experiences from eastern Germany. Environ Impact Assess Rev. 2008;28:483–503.
- 79. White MM. Sisters of the soil: urban gardening as resistance in detroit. Race/Ethnicity Multidiscipl Global Contexts. 2011;5(1):13–28.
- 80.
Langer A. Pure urban nature: Nature-park Südgelände, Berlin. Routledge; 2012.
- 81. Bonthoux S, Brun M, Di Pietro F, Greulich S, Bouché-Pillon S. How can wastelands promote biodiversity in cities? A review. Landsc Urban Plan. 2014;132:79–88.
- 82. Maurer U, Peschel T, Schmitz S. The flora of selected urban land-use types in Berlin and Potsdam with regard to nature conservation in cities. Landsc Urban Plan. 2000;46(4):209–15.
- 83. Öckinger E, Dannestam Å, Smith HG. The importance of fragmentation and habitat quality of urban grasslands for butterfly diversity. Landsc Urban Plan. 2009;93(1):31–7.
- 84. Robinson SL, Lundholm JT. Ecosystem services provided by urban spontaneous vegetation. Urban Ecosyst. 2012;15(3):545–57.
- 85. Muratet A, Machon N, Jiguet F, Moret J, Porcher E. The role of urban structures in the distribution of wasteland flora in the Greater Paris Area, France. Ecosystems. 2007;10(4):661–71.
- 86. Maurel N, Salmon S, Ponge JF, Machon N, Moret J, Muratet A. Does the invasive species Reynoutria japonica have an impact on soil and flora in urban wastelands?. Biol Invasions. 2010;12:1709–19.
- 87. Riley CB, Herms DA, Gardiner MM. Exotic trees contribute to urban forest diversity and ecosystem services in inner-city Cleveland, OH. Urban For Urban Green. 2018;29:367–76.
- 88. Cui J, Sharifi E, Bartesaghi Koc C, Yi L, Hawken S. Factors shaping biodiversity in urban voids: a systematic literature review. Land. 2025;14(4):821.
- 89. Johnson AL, Borowy D, Swan CM. Land use history and seed dispersal drive divergent plant community assembly patterns in urban vacant lots. J Appl Ecol. 2017;55(1):451–60.
- 90. Preston PD, Dunk RM, Smith GR, Cavan G. Not all brownfields are equal: a typological assessment reveals hidden green space in the city. Landsc Urban Plan. 2023;229:104590.
- 91. Ala-Mantila S, Heinonen J, Junnila S. Relationship between urbanization, direct and indirect greenhouse gas emissions, and expenditures: a multivariate analysis. Ecol Econ. 2014;104:129–39.
- 92. Jones C, Kammen DM. Spatial distribution of U.S. household carbon footprints reveals suburbanization undermines greenhouse gas benefits of urban population density. Environ Sci Technol. 2014;48(2):895–902. pmid:24328208
- 93. Heinonen J, Junnila S. Residential energy consumption patterns and the overall housing energy requirements of urban and rural households in Finland. Energy Buildings. 2014;76:295–303.
- 94.
Thorve S, Swarup S, Marathe A, Chungbaek Y, Nordberg EK, Marathe MV. Simulating residential energy demand in urban and rural areas. 2018. p. 548–59. https://doi.org/10.1109/WSC.2018.8632203
- 95.
EPA U. Emission facts: Greenhouse gas emissions from a typical passenger vehicle. Washington: US Environmental Protection Agency; 2005.
- 96. Andrews CJ. Greenhouse gas emissions along the rural-urban gradient. J Environ Plan Manage. 2008;51(6):847–70.
- 97. Nichols BG, Kockelman KM. Life-cycle energy implications of different residential settings: recognizing buildings, travel, and public infrastructure. Energy Policy. 2014;68:232–42.
- 98. Matos C, Bentes I, Pereira S, Gonçalves AM, Faria D, Briga-Sá A. Which are the factors that may explain the differences in water and energy consumptions in urban and rural environments?. Sci Total Environ. 2018;642:421–35.
- 99. Skaggs RK, Samani Z. Farm size, irrigation practices, and on-farm irrigation efficiency. Irrig Drain. 2005;54(1):43–57.
- 100. Wolny A. Are suburban commuters confined to private transport? A case study of a medium-sized functional urban area (FUA) in Poland. Cities. 2019;92:82–96.
- 101. Pucher J, Renne JL. Rural mobility and mode choice: Evidence from the 2001 National Household Travel Survey. Transportation. 2005;32(2):165–86.
- 102.
Department for Transport. National Travel Survey 2021: travel by region and rural and urban classification of residence. 2021. Available from: https://www.gov.uk/government/statistics/national-travel-survey-2021/national-travel-survey-2021-travel-by-region-and-rural-and-urban-classification-of-residence
- 103. Muñiz I, Galindo A. Urban form and the ecological footprint of commuting. The case of Barcelona. Ecol Econ. 2005;55:499–514.
- 104. McGuirk P, Argent N. Population growth and change: implications for Australia’s cities and regions. Geogr Res. 2011;49:317–35.
- 105. Stokes EC, Seto KC. Tradeoffs in environmental and equity gains from job accessibility. Proc Natl Acad Sci U S A. 2018;115(42):E9773–81. pmid:30275299
- 106. Jaeger JAG, Bowman J, Brennan J, Fahrig L, Bert D, Bouchard J, et al. Predicting when animal populations are at risk from roads: an interactive model of road avoidance behavior. Ecol Model. 2005;185(2–4):329–48.
- 107. Lemke A, Buchholz S, Kowarik I, Starfinger U, von der Lippe M. Interaction of traffic intensity and habitat features shape invasion dynamics of an invasive alien species (Ambrosia artemisiifolia) in a regional road network. NB. 2021;64:155–75.
- 108. Hansen AJ, Knight RL, Marzluff JM, Powell S, Brown K, Gude PH. Effects of exurban development on biodiversity: patterns, mechanisms, and research needs. Ecol Appl. 2005;15:1893–905.
- 109. Loss SR, Will T, Marra PP. The impact of free-ranging domestic cats on wildlife of the United States. Nat Commun. 2013;4:1396. pmid:23360987
- 110. Ellis-Soto D, Oliver RY, Brum-Bastos V, Demšar U, Jesmer B, Long JA, et al. A vision for incorporating human mobility in the study of human-wildlife interactions. Nat Ecol Evol. 2023;7(9):1362–72. pmid:37550509
- 111. Gaynor KM, Hojnowski CE, Carter NH, Brashares JS. The influence of human disturbance on wildlife nocturnality. Science. 2018;360(6394):1232–5. pmid:29903973
- 112. Nickel BA, Suraci JP, Allen ML, Wilmers CC. Human presence and human footprint have non-equivalent effects on wildlife spatiotemporal habitat use. Biol Conserv. 2020;241:108383.
- 113. Fagre AC, Cohen LE, Eskew EA, Farrell M, Glennon E, Joseph MB, et al. Assessing the risk of human-to-wildlife pathogen transmission for conservation and public health. Ecol Lett. 2022;25(6):1534–49. pmid:35318793
- 114. Jägerbrand AK, Alatalo JM. Effects of human trampling on abundance and diversity of vascular plants, bryophytes and lichens in alpine heath vegetation, Northern Sweden. Springerplus. 2015;4:95. pmid:25774335
- 115. Gosnell H, Abrams J. Amenity migration: diverse conceptualizations of drivers, socioeconomic dimensions, and emerging challenges. GeoJournal. 2009;76(4):303–22.
- 116. Aydoğan B, Güney S. Mobilizing capitals, forging terroir: Lifestyle migration and envisioning a “better rural landscape”. J Rural Stud. 2025;119:103714.
- 117. Li C, Fang Y, Wang Y, Xu Y, Zong Z, Yang Y. How can tourism help to revitalize the countryside? Content analysis based on the case of tourism enabling rural revitalization. Environ Dev Sustain. 2024;26:20333–54.
- 118. Freiberg JA, Callegaro RM. Agricultural environments contribute to bird biodiversity and birdwatching in rural landscapes. Agroforest Syst. 2025;99:102.
- 119. Schwoerer T, Dawson NG. Small sight-Big might: Economic impact of bird tourism shows opportunities for rural communities and biodiversity conservation. PLoS One. 2022;17(7):e0268594. pmid:35793337
- 120. Larson CL, Reed SE, Merenlender AM, Crooks KR. Effects of recreation on animals revealed as widespread through a global systematic review. PLoS One. 2016;11(12):e0167259. pmid:27930730
- 121. DeRose‐Wilson AL, Hunt KL, Monk JD, Catlin DH, Karpanty SM, Fraser JD. Piping plover chick survival negatively correlated with beach recreation. J Wildl Manag. 2018;82(8):1608–16.
- 122. Hentati-Sundberg J, Berglund P-A, Hejdström A, Olsson O. COVID-19 lockdown reveals tourists as seabird guardians. Biol Conserv. 2021;254:108950. pmid:35719894
- 123. Suh J. Environmental characteristics of urban-rural farming migrants in the Republic of Korea and their significance for rural sustainability. Local Environ. 2019;24(8):663–77.
- 124. Vizuete B, Oteros-Rozas E, García-Llorente M. Role of the neo-rural phenomenon and the new peasantry in agroecological transitions: a literature review. Agric Hum Values. 2024;41(3):1277–97.
- 125. Zollet S, Qu M. Creative newcomers and local food culture: revitalization, sustainability, and place-making between rural and urban. Food, Cult Soc. 2024;:1–19.
- 126. Holloway L. ‘Hell on earth and paradise all at the same time’: the production of smallholding space in the British countryside. Area. 2000;32(3):307–15.
- 127. Sutherland WJ. Restoring a sustainable countryside. Trends Ecol Evol. 2002;17(3):148–50.
- 128. Beaumelle L, Tison L, Eisenhauer N, Hines J, Malladi S, Pelosi C, et al. Pesticide effects on soil fauna communities – a meta‐analysis. J Appl Ecol. 2023;60(7):1239–53.
- 129. Tuck SL, Winqvist C, Mota F, Ahnström J, Turnbull LA, Bengtsson J. Land-use intensity and the effects of organic farming on biodiversity: a hierarchical meta-analysis. J Appl Ecol. 2014;51(3):746–55. pmid:25653457
- 130. Schneider MK, Lüscher G, Jeanneret P, Arndorfer M, Ammari Y, Bailey D, et al. Gains to species diversity in organically farmed fields are not propagated at the farm level. Nat Commun. 2014;5:4151. pmid:24958283
- 131. Batáry P, Gallé R, Korányi D, Lakatos T, Deák B, Gallé-Szpisjak N. Biodiversity and human well-being trade-offs and synergies in villages. Nat Sustain. 2025;:1–11.
- 132. González-Leonardo M, Rowe F, Fresolone-Caparrós A. Rural revival? The rise in internal migration to rural areas during the COVID-19 pandemic. Who moved and Where?. J Rural Stud. 2022;96:332–42. pmid:36439404
- 133. Karsten L. Counterurbanisation: Why settled families move out of the city again. J Hous Built Environ. 2020;35:429–42.
- 134. Gkartzios M. ‘Leaving Athens’: narratives of counterurbanisation in times of crisis. J Rural Stud. 2013;32:158–67.
- 135. Scott M, Gkartzios M, Halfacree K. Introducing climate-related counterurbanisation: individual adaptation or societal maladaptation?. Habitat Int. 2024;143:102970.
- 136. Dinis A. Tourism, immigrants and lifestyle entrepreneurship: the (in)coming of people as a key factor for sustainability of low-density territories – a case study in Portugal. Tourism, Hospitality & Event Management. Springer International Publishing. 2021. p. 149–82.
- 137. Ramani A, Alcedo J, Bloom N. How working from home reshapes cities. Proc Natl Acad Sci U S A. 2024;121(45):e2408930121. pmid:39471226
- 138. Steinführer A, Osterhage F, Tippel C, Kreis J, Moldovan A. Urban–rural migration in Germany: a decision in favour of ‘the rural’ or against ‘the urban’?. J Rural Stud. 2024;111:103431.
- 139. Goodwin CED, Bütikofer L, Hatfield JH, Evans PM, Bullock JM, Storkey J. Multi-tier archetypes to characterise British landscapes, farmland and farming practices. Environ Res Lett. 2022;17:095002.
- 140. Václavík T, Lautenbach S, Kuemmerle T, Seppelt R. Mapping global land system archetypes. Global Environ Change. 2013;23(6):1637–47.
- 141. Citron DT, Iyer S, Reiner RC, Smith DL. Activity space maps: a novel human mobility data set for quantifying time spent at risk. Epidemiology. 2021.
- 142. Thorve S, Baek YY, Swarup S, Mortveit H, Marathe A, Vullikanti A, et al. High resolution synthetic residential energy use profiles for the United States. Sci Data. 2023;10(1):76. pmid:36746951
- 143. Brown CF, Brumby SP, Guzder-Williams B, Birch T, Hyde SB, Mazzariello J, et al. Dynamic world, near real-time global 10 m land use land cover mapping. Sci Data. 2022;9(1).
- 144. Oostwegel LJN, Schorlemmer D, Guéguen P. From footprints to functions: a comprehensive global and semantic building footprint dataset. Sci Data. 2025;12(1):1699. pmid:41145547
- 145. Brown CF, Kazmierski MR, Pasquarella VJ, Rucklidge WJ, Samsikova M, Zhang C. AlphaEarth Foundations: an embedding field model for accurate and efficient global mapping from sparse label data. arXiv. 2025.