Can a forest specialist live in a city? Behavioural plasticity fosters the presence of a protected carnivore in urban areas

Can a forest specialist live in a city? Behavioural plasticity fosters the presence of a protected carnivore in urban areas | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Can a forest specialist live in a city? Behavioural plasticity fosters the presence of a protected carnivore in urban areas Leonardo Ancillotto, Andrea Amici, Dorian Grelli, Francesca Vercillo, and 1 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-9043132/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted 4 You are reading this latest preprint version Abstract Urbanisation is reshaping mammalian communities worldwide, yet forest specialists are generally considered poor urban adapters. The pine marten Martes martes , for long regarded as an indicator of intact woodlands, has recently shown signs of expansion into human-modified landscapes. Here, we combined citizen-science records and camera-trapping data to reconstruct the distribution of the pine marten in urban areas of Italy. Moreover, we used camera traps to investigate activity patterns, and spatiotemporal niche overlaps of the pine marten along an urban–natural gradient in Italy. Verified reports from 2000–2025 confirm the species presence in multiple Italian cities, including several regional capitals. In the Insugherata Urban Park in Rome, camera traps recorded 192 independent detections in one year, including 46 pine martens, 79 red foxes Vulpes vulpes , 33 Eurasian red squirrels Sciurus vulgaris , and 34 humans. Pine martens exhibited a predominantly crepuscular activity, with significant peaks at dawn and dusk, differing from the more nocturnal red fox. Temporal overlap between martens and foxes was low in the urban park (Δ₁ = 0.38), but overlap with red squirrels (Δ₁ = 0.84) and humans (Δ₁ = 0.77) was high, suggesting flexible diel adjustments to urban pressures. Compared to a natural reference site, pine martens in the city displayed reduced synchrony with foxes but increased overlap with both prey and human activity, indicating behavioural adaptation to anthropogenic environments. Our findings suggested a growing capacity of M. martes to persist in urban green spaces by modulating temporal activity and resource use. The increase of urban records of pine marten highlights the need to reconsider its ecological classification and integrate cities into future conservation strategies for this carnivore. Martes martes urban ecology temporal niche camera trapping behavioural plasticity Italy Figures Figure 1 Figure 2 Figure 3 Figure 4 Introduction Urban ecosystems are increasingly recognized as dynamic areas which may host a surprising diversity of wildlife (Werner 2011 ; Gallo et al. 2017 ). Urban ecology define urban areas are densely populated regions with extensive infrastructure and services, typically forming cities or large towns (Gottero et al. 2023 ). Conversely, periurban areas are transitional zones located on the outskirts of cities where urban and rural characteristics coexist and land use is rapidly changing (Gottero et al. 2023 ). Rural areas are regions with low population density, limited infrastructure, and predominantly agricultural or natural landscapes (Gottero et al. 2023 ). Wild mammals in urban areas face a range of threats primarily linked to habitat loss, fragmentation, and human activity (Soulsbury and White 2015 ; Santini et al. 2019 ). Urban expansion reduces and isolates natural habitats, making it difficult for mammals to find food, mates, and shelter (Dickman 1987 ; Villaseñor et al. 2014 ; Monteiro and Lira 2020 ). Roads and traffic cause high mortality through vehicle collisions, while light, noise, and chemical pollution disrupt behaviour, communication, and health (Birnie-Gauvin et al. 2016 ; Ritzel and Gallo 2020 ; Viviano et al. 2023 ). Domestic pets and invasive species increase predation and competition pressures (Mori et al. 2019 ; Miller et al. 2022 ; Sogliani et al. 2023 ). Additionally, direct conflict with humans (e.g., predations upon poultry or livestock), often harmful for wildlife, poses further risks (Soulsbury and White 2015 ). Combined, these pressures often lead to declines in population size, genetic diversity, and overall ecosystem stability. In particular, in Europe, several mammal species have successfully started to thrive in urban environments, exploiting anthropogenic resources and novel ecological opportunities (Santini et al. 2019 ). Urban mammals are typically ecological generalists, characterized by plasticity in space use, broad dietary spectra, and behavioural flexibility (Santini et al. 2019 ; Mori et al. 2025a ). Among them, native species including carnivores (e.g., red fox Vulpes vulpes and stone marten Martes foina ), bats (e.g., the Kuhl’s pipistrelle Pipistrellus kuhlii ) and small mammals (e.g., the Western hedgehog Erinaceus europaeus and the Eurasian red squirrel Sciurus vulgaris ) have become well-established urban dwellers (Russo and Ancillotto 2015 ; Ancillotto et al. 2016 , 2025 ; Bateman and Fleming 2012 ; Capon et al. 2021 ; Turner et al. 2022 ; Cox et al. 2025 ; Dondina et al., 2025 ). Additionally, a number of non-native species, such as the Eastern grey squirrel Sciurus carolinensis or the coypu Myocastor coypus , have thrived in European cities, benefiting from low predation pressure, human-made structures (e.g., abandoned buildings used as denning-sites) and frequent human tolerance or even support (Mori et al. 2025c ; Santicchia et al. 2024 ; Viviano et al. 2024 , 2025 ). Conversely, species tightly associated with specific or fragile habitats, such as wetlands or old-growth forests, tend to be poorly suitable to survive in urbanized ecosystems, and may undergo local extinctions as a result (Ancillotto et al. 2024 , 2025 ). Indeed, during the last decade, a growing number of studies have documented formerly forest-associated mammals expanding into peri-urban and urban areas. Examples include the European badger Meles meles and the crested porcupine Hystrix cristata , which are increasingly observed using urban shrublands as denning sites and foraging in parks and suburban gardens (Davison et al. 2008 ; Lovari et al. 2017 ; Geiger et al. 2018 ; Mori et al. 2022a ). Within the Mustelidae family, small Mustela species (namely, M. nivalis , M. putorius ) are rarely recorded both in natural environments and, for most, in cities (Zabala et al. 2005 ; Łopucki et al. 2019 ), whereas the genus Martes is represented almost exclusively by M. foina (Duduś et al. 2014 ; Capon et al. 2021 ; Cox et al. 2025 ; Mori et al. 2025a ). This species is highly synanthropic, omnivorous, and behaviourally plastic, exploiting urban and rural structures e.g., attics, archaeological sites, and roof spaces as diurnal resting places (Herr et al. 2009 ; Tóth et al. 2009 ; Balestrieri et al. 2019 ). The strictly nocturnal habits and tolerance to human presence make M. foina one of the most successful carnivores in European cities (Mori et al. 2025a ). In contrast, the pine marten M. martes has long been regarded as a forest specialist and is considered as an indicator species of well-preserved forest ecosystems (Clevenger 1993 ; Lombardini et al. 2015 ; Manzo et al. 2018 ). Balestrieri et al. ( 2019 ) suggested that distance to watercourses, and the cover of deciduous forest and shrubland, are the main factors shaping pine marten occurrence across its range, whereas the stone marten is more closely associated with higher human population density and extent of built-up areas. Consequently, although pine martens may locally select areas with intermediate levels of human modifications (Vergara et al. 2016 ), increasing human density and urbanization of lowland and hilly landscapes have been predicted to represent major limiting factors to the pine marten. Furthermore, it has been suggested that a competition for resources may occur between stone and pine martens (Wereszczuk and Zalewski 2015 ; Gazzola and Balestrieri 2020 ). Moreover, the more diurnal activity patterns of the pine marten compared to the stone marten may increase the risk of encounters with humans and domestic carnivores (Zielinski et al. 1983 ; Torretta et al. 2016 , 2017 ; Mori et al. 2022b ). In recent times, evidence from Italy and other parts of Europe suggests that this view may be changing, with pine martens increasingly occurring in rural, thus fragmented, landscapes (Pereboom et al. 2008 ; Balestrieri et al. 2010 ; Vergara et al. 2016 ; Manzo et al. 2018 , 2024 ). However, pine martens tend to avoid highly anthropized environments, possibly due to aversion to humans and domestic carnivores such as dogs and cats (Mori et al. 2022b ). In rural and natural landscapes, the species is known to compete with the red fox, and cases of intraguild predation have also been documented, potentially representing an additional constraint in urban habitats (Storch et al. 1990 ; Lindström et al. 1995 ), where more opportunistic carnivores may reach very high population densities. Against this background, the aim of the present study was to collect available records of pine martens in urban environments in Italy, and to describe their activity rhythms in urban environment and to relate them to those of the main potential prey (i.e., the Eurasian red squirrel Sciurus vulgaris ) and competitors (i.e., humans and the red fox). Moreover, we compared these patterns with those recorded in natural protected areas far from urban disturbance (Mori et al. 2025b ). We hypothesized that in urban settings, pine martens may exhibit temporal trade-offs to minimize overlap with humans and domestic carnivores, as well as to maintain a high overlap with potential prey species, maintaining partial diurnality to avoid competition or predation by red foxes, with which they share both space and dietary niches. Material and Methods Do pine martens occur in cities? We reviewed all publicly available records of pine marten occurring in urban environments within Italy up to December 2025. Our investigation focused on regional capital cities, which were selected as representative urban centres with consistent administrative boundaries and accessible information. Pine marten records from all the other provincial capital cities were also included. All reports of pine martens within boundaries of these cities were collected and verified, based on photographic or otherwise certified evidence. Records were considered as valid only when provided with a spatial precision < 100 meters, and were assigned to urban habitats whenever coordinates were within the administrative boundaries of the considered cities i.e., not encompassing the entire province and yet also including peripheral areas and urban parks. Type of records included: (i) carcasses and injured individuals (i.e., museum specimens and Wildlife Rescue Center data), (ii) photographs gathered from multiple complementary resources (i.e., social networks and citizen science platforms) and (iii) scientific literature. Publicly available photographs were systematically searched across social networks (i.e., Facebook, Instagram, Flickr, X/Twitter), considering only posts that were visible without restrictions or user login. In parallel, all georeferenced and photo-documented observations of the pine marten in Italy were downloaded from the citizen science platforms iNaturalist ( www.inaturalist.org . Accessed on 06.10.2025) and Ornitho.it ( www.ornitho.it . Accessed on 06.10.2025). To complement citizen-generated data, a comprehensive literature search was conducted using Zoological Records, Scopus, and Google Scholar. The searches included both Italian and English language publications and employed all relevant keyword combinations of “Mart*”, “ Martes martes ”, “urban”, “city”, and “Italy”, connected by Boolean operators. Example search strings included (“ Martes martes ” OR Martes OR marten OR martor*) AND (urban OR city OR “urban area”) AND (Italy OR Italia). Equivalent combinations in Italian were also used. Only records providing explicit urban locations or referred to observations within cities were retained. Duplicates between sources were identified and removed after cross-checking location, date, and image content. For each city we screened, we recorded whether reports of M. martes were available, the year of each observation, the status of the animal (alive or dead), and the source of the information (e.g., citizen science platform, social media, or peer-reviewed publication). The concurrent presence of the stone marten in the same city was also recorded by using the same methods specified for the pine marten, to enable comparative analyses between the two congeners. All photographic records were visually inspected by at least two researchers to verify species identifications, based on diagnostic morphological traits (following Raichev 2018 ). In order to assess whether pine martens occur in cities with specific characteristics, we downloaded city size (in squared km; ISTAT) and total amount of tree cover (ISPRA). We then built a generalized linear model (GLM) with a binomial error structure, using pine marten occurrence as binary response variable, city size and tree cover as explaining variables; additionally, to account for potential competition, we also included a binary covariate indicating the occurrence, within the same city, of the congeneric stone marten. We then selected the set of variables best explaining the observed patterns of occurrence by performing a stepwise backward model selection procedure according to AIC values (Burnham and Anderson 2002 ), using the stepAIC function in the MASS package (Ripley et al. 2013 ) for R version 4.2.2. (R Core Team 2021 ) i.e., considering the best model(s) the one minimizing AIC values and those featuring delta-AIC values < 2 from the best model. Do urban martens change their behaviour? In order to assess pine marten behavioural patterns in terms of activity rhythms, we set a camera trapping sampling in urban and natural sites. The first site was an urban Nature Reserve (Riserva Naturale dell’Insugherata, hereafter RNI) in Rome, one of the largest protected green areas within the municipality of Rome, Italy, covering approximately 730 hectares (Todini and Crosti 2020 ). Located in the northwestern sector of the city, the park features a diverse mosaic of Mediterranean ecosystems, including oak woodlands, cork oak Quercus suber L. forests, shrublands, and open grasslands. Its topography is characterized by hills and deep valleys, with elevations ranging from 50 to 150 meters above sea level. The Insugherata urban park supports high biodiversity, providing habitat for numerous plant and animal species (Todini and Crosti 2020 ), yet stretching inside the city, its boundaries being at < 2 km from Rome’s city centre. Additionally, the park contains remnants of ancient Roman roads and agricultural terraces, reflecting a long history of human interaction with the local landscape (Todini and Crosti 2020 ). The climate is typically Mediterranean, with an annual mean temperature of about 16°C ± 1°C (range: 3–37°C) in 2020–2025, featuring warm summers and mild winters, and an average annual precipitation of approximately 850 mm ± 50 mm, with rainfall mostly concentrated in autumn and winter. Seasonal temperature variations are moderate, reflecting typical conditions of the city and its surrounding hilly areas (Ancillotto et al. 2024 ). Three 32-MP camera traps (© Coolife, China) were deployed within the RNI at 30–80 cm above ground level and operated continuously (24 h/day). Each device was programmed to capture a 60-s video sequence upon detection of an animal, a setting chosen to reduce motion blur and thereby improve taxonomic resolution (O’Connell et al. 2011 ). Field sampling was conducted from September 2023 to August 2024, yielding 954 trap-nights and covering the whole reserve, the only one in Rome metropolitan area hosting pine martens (Ancillotto et al. 2024 ). Cameras were installed at three fixed, georeferenced points, secured to trees or rocks using ropes and chains, and positioned along the nearest wildlife trail to randomly generated sampling points created in QGIS (QGIS Development Team 2019 ). To reduce the likelihood of pseudoreplication, stations were spaced a minimum of 1.2 km apart (O’Connell et al. 2011 ). We compared the activity rhythms of pine martens, their prey, and competitor species in urban areas with those of the same species in a natural area at the same latitude in Italy (Monte Rufeno Nature Reserve, approx. 3,000 ha; 275–774 m a.s.l.; Mori et al. 2025b ), during the same study period. This rural area is characterized by a mosaic of deciduous oak woodlands, scrublands, pastures, and riparian habitats, where 30 fixed camera traps were deployed across the full altitudinal range, spaced at least 800 m apart and active 24 h/day, recording 60-second videos per trigger (see Mori et al. 2025b ). Cameras yielded over 9,000 camera-days of sampling effort. Camera-traps were checked once a month to retrieve recordings and replace dead batteries. For each video, metadata were entered into a structured dataset including: species identity, date, time, diel phase (diurnal/nocturnal), station ID, lighting condition (day/night), and meteorological season (spring, summer, autumn, winter). Videos for which species identification was not possible (8.8% of all wildlife records) were excluded from analyses. Camera trapping was conducted under all necessary authorizations and in full compliance with national and international regulations, including the Italian Legislative Decree 196/2003 and the European Regulation 2016/679 on privacy. Informative panels were installed at monitoring sites to notify the public about the presence of the cameras and the objectives of the study. Each panel included a QR code providing access to project documentation and ensuring that any recordings containing humans were deleted within 72 hours from all storage devices. Direct contact with the project manager was also provided in each panel to address any concern. All camera-trap records were compiled into a dataset containing information on species identity, date, time, and camera-trap station. Species identification was independently performed by three of the authors using a double-blind protocol. To reduce pseudoreplication, consecutive detections of the same species at the same camera-trap station occurring within a 30-minute interval were merged into a single event (Monterroso et al. 2014 ). The dataset retained the original dates and times, i.e. solar hour recorded on each video file to minimize temporal bias. Nighttime was defined as the period from one hour after sunset to one hour before sunrise (Carnevali et al. 2016 ), while dawn and dusk were delineated according to astronomical circadian twilight. These time periods were calculated using the NightDay package (Hughes-Brandl 2018 ) for R version 4.1.2 (R Core Team 2018). This framework distinguishes canonical diel niches based on relative activity across daily periods and identifies both bimodal and trimodal activity distributions (Gerber et al. 2024 ). Activity rhythms of pine martens were analyzed using the overlap package (Meredith & Ridout 2014 ). Temporal overlap (Δ) was quantified with three key counterparts in urban areas: the main competitor (red fox: Baltrunaite 2001 ; Lanszki et al. 2007 ; Takahashi et al. 2025 ), a representative prey (red squirrel: Twining et al. 2020 ), and humans. We also compared these overlap values with those calculated for the same species in the Monte Rufeno Nature Reserve (Mori et al. 2025b ). Distinguishing between pine and stone martens based on external morphological characteristics may be challenging, particularly in nocturnal videos from camera traps. However, following Manzo et al. ( 2012 ) and Raichev ( 2018 ), we identified several key diagnostic traits including the colour of the nose leather (dark in the pine marten, pale in the stone marten) and overall coat appearance. Pine martens generally show a darker, more uniformly brown pelage, whereas stone martens tend to display paler, greyer tones. Differences in body shape were also considered: pine martens typically present a slenderer profile, with proportionally longer tail and limbs and a more pointed muzzle, while stone martens appear stockier, with a relatively shorter tail and broader head. When an individual showed mixed / poorly identifiable traits (in 12% records), the record was conservatively assigned to Martes sp. and excluded from further analyses. The throat patch provided an additional distinguishing feature. In pine martens, it is usually yellow to orange, smaller, irregularly shaped, and often discontinuous; in stone martens, it is bright white, larger, more regularly outlined, and frequently extends toward the forelimbs. When individuals exhibited intermediate or conflicting traits, or when there was disagreement among reviewers, records were conservatively classified as Martes sp. These records were excluded from temporal activity analyses because of the documented ecological and behavioural differences between the two species (Wereszczuk and Zalewski, 2015 ; Torretta et al., 2016 , 2017 ; Balestrieri et al., 2019 ; Gazzola and Balestrieri, 2020 ). The Mardia–Watson–Wheeler (MWW) test was performed using the circular package in R (Lund et al. 2017 ) to assess whether the overlaps in the natural area differed significantly from those in the urban area (Massara et al. 2018 ). The Δ₁ estimator was always used, as at least one species in each pairwise comparison had fewer than 75 independent detections (Monterroso et al. 2014 ). Overlap values were interpreted as low ( 75%), or very high (> 90%) (Monterroso et al. 2014 ). Confidence intervals (95% CIs) for Δ were obtained from 999 bootstrap replicates with percentile-based limits (Monterroso et al. 2014 ). Deviations from a uniform (random) activity distribution over the 24-hour cycle were tested using the Hermans–Rasson (HR) test (Landler et al. 2019 ). Spatial niche overlap between species pairs was evaluated with the Pianka index (Pianka 1974 ; Mori et al. 2020 ), which ranges from 0 (no overlap) to 1 (complete overlap). The Pianka index was computed through the formula: O species1/ species2 = (Σ P i species1 × P i species2 ) / √(Σ P i species1 ² × Σ P i species2 ²), where P i species1 and P i species2 are the proportion of records of species 1 and species 2, respectively. The spatial overlap was considered as ‘high’ when O > 0.75 and as ‘very high’ when O > 0.90; conversely it was considered as intermediate with 0.50 < O < 0.75, and low when O < 0.50 (Monterroso et al., 2014 ). Given the low seasonal number of detections, we computed spatiotemporal overlaps for a total year. Results Do pine marten occur in cities? We retrieved records of martens from 26 Italian cities across the entire national territory, including both mainland and the two largest islands (Sardinia and Sicily: Fig. 1 , Table S1 in Supplementary Material 1). Namely, 6 cities featured records of both pine and stone marten, and 4 pine marten only. Amongst the latter, 3 out of 4 are out of the known range of stone marten (i.e., from islands). Thus, the pine marten occurred only in 10 out of 26 sampled cities, for a total of 84 records (Supplementary Material 1). The model selected by our AIC-based procedure, and thus best explaining the observed patterns of occurrence of the pine marten in our set of Italian cities, were the ones featuring city altitude and percent cover of urban wooded areas within the city boundaries, exerting a negative and positive effect upon the probability of occurrence of the pine marten in a given city, respectively (Table 1 ; Fig. 2 ). Namely, pine marten occurrence exceeded 50% in cities featuring > 40% of tree covers within their municipality, and dropped below 30% at altitude values above 600 m a.s.l. Table 1 Model selection on the probability of occurrence of Martes martes in urban areas of Italy (N = 26) as explained by city altitude, percent wooded cover within the city boundaries, city size, and co-occurrence of M. foina . Model rank Altitude Percent wooded cover Presence of Martes foina City size AIC 1 NA 0.025 ± 0.016 * NA NA 32.79 2 -0.002 ± 0.001 0.025 ± 0.016 * NA NA 33.75 3 -0.002 ± 0.001 0.025 ± 0.016 * -1.793 ± 1.086 NA 36.37 Do urban martens change their behaviour in urban areas? We recorded a total of 192 independent detections of the target species in RNI: 46 pine martens, 79 red foxes, 33 red squirrels and 34 humans. These data were compared to those collected by Mori et al. ( 2025b ) in the Monte Rufeno Reserve: 195 independent records of pine marten, 813 of red fox, 85 of red squirrels and 424 humans. No stone marten was detected in the study areas. In the urban area, the pine marten activity showed significant peaks at dawn and dusk, red fox activity peaked in the late afternoon hours, red squirrel activity in the early morning and afternoon, and human activity in mid-morning and mid-afternoon (Hermans–Rasson tests; r = 62.48–81.33, all p < 0.001). Temporal overlap between urban and natural areas was low to moderate for all species, except for the red fox, which showed a high degree of overlap in activity peaks between urban and natural areas (Fig. 3 ). We found significant differences in the temporal overlaps of pine martens with red squirrels, red foxes, and humans between urban and natural areas (Mann–Whitney–Wilcoxon test, W = 143.29–181.03, all p < 0.001). Specifically, in urban environments, pine martens exhibited increased temporal overlap with red squirrels and with humans, whereas their temporal overlap with red foxes remarkably declined (Fig. 4 ). Spatial overlaps (Pianka index, O) between pine martens and red squirrels and between pine martens and humans were high in both urban (respectively 0.84 and 0.77) and natural (respectively 0.75 and 0.78) areas. Conversely, spatial overlap between pine martens and red foxes was low in both urban (O = 0.38) and natural (O = 0.27) areas. Discussion In this work, we provided the first evidence of pine martens occurring in urban environments across Italy, likely reflecting a rise in urban detections, which might in turn indicate an ongoing process of urban colonisation by a species historically regarded as a forest specialist, rather than an actual increase in population size (but see Robinson et al. 2017 ). Museum records also indicate the long-standing presence of the stone marten in urban areas since at least the 19th century, whereas confirmed records of the pine marten date only to recent decades and remain relatively sporadic (Ancillotto et al. 2024 , 2025 ). The compilation of verified urban records across multiple Italian cities, combined with detailed camera-trap data from Rome in comparison to neighboring natural areas, may suggest that the pine marten is exhibiting both spatial and temporal behavioural plasticity, possibly allowing it to persist in human-dominated landscapes (Manzo et al. 2025 ). This result challenges long-standing assumptions about the species’ strict dependence on mature forests (Clevenger 1993 ; Lombardini et al. 2015 ) and is in line with recent findings of range expansion into agricultural and peri-urban areas (Balestrieri et al. 2010 ; Grelli et al. 2014 ; Manzo et al. 2018 , 2024 ), as also confirmed by increasing road mortality events (Supplementary Material 1). Indeed, considering that many urban records of the European pine marten are associated with road kills (see Supplementary Material 1), this pattern may suggest that remnant woodland patches within cities function as ecological traps for the species (see Ancillotto et al. 2025 ). In other words, these forest remnants may attract individuals attempting to colonize urban areas by following existing habitat connectivity. Once inside the urban matrix, the pine marten may be less well adapted than the stone marten to cope with anthropogenic pressures. Consequently, although environmental corridors may facilitate urban entry, survival prospects within the city appear comparatively low, potentially leading to increased mortality rather than successful establishment. However, the pine marten has undergone a remarkable range expansion in Italy over the last two decades, now occupying lowland areas where it was once considered absent (Balestrieri et al. 2010 , 2015 ; Manzo et al. 2018 ). This expansion has been partly revealed through citizen-science platforms, wildlife rescue center records, and increasing public awareness of wildlife conservation (Grelli et al. 2014 ; Manzo et al. 2025 ). Reports of pine martens in peri-urban areas, where the species was previously unrecorded, are becoming increasingly frequent, whereas earlier records from similar contexts referred exclusively to stone martens. This pattern may indicate an ongoing process of urban colonization, or, in general, an increase in use of human-modified habitats (Grelli et al. 2014 ). The presence of pine martens within several regional and provincial capital cities, verified through citizen science, Wildlife Rescue Centers and social media data, reflects a broader ecological trend in which formerly habitat-specialist mammals are colonising novel environments, possibly as a response to landscape modification and climatic changes (Balestrieri et al. 2010 ; Santini et al. 2019 ; Ancillotto et al. 2024 , 2025 ; Manzo et al. 2024 ). The RNI in Rome, although embedded within a densely populated area, still hosts structural elements, e.g. continuous woodland patches and abundant tree cover (Todini and Crosti 2020 ), which provide all habitat and resource requirements by the species (Manzo et al. 2025 ). These findings suggest that green corridors and urban parks can function as “ecological stepping stones” facilitating movement and settlement of forest mammals in metropolitan matrices (Monteiro and Lira 2020 ). Furthermore, the presence of pine marten in capital cities could be a response not only to landscape modifications and climate change, but also to behavioural changes both in martens and humans, driving to a higher reciprocal tolerance (Mori et al. 2025c ). The importance of profitable patches of natural habitats for pine martens within anthropogenic landscapes e.g., natural woodlands embedded in urban parks, was also revealed by our modeling exercise, highlighting that cities with low values of wooded cover are less likely to host the species. Additionally, the negative effect of altitude upon the probability of presence of the pine marten is actually inconsistent with previous work from the alpine region, and Bulgaria, where the species seems to favour higher elevations (Doykin et al. 2017 ; Fonda et al. 2021 ). Nonetheless, this suggests that the pine marten is actually not a specialist of higher-altitude forests, yet that this latter habitat type is associated to areas where the species occurred after it past decline (Sainsbury et al. 2019 ), since low-altitude forests were those historically more impacted (Widmer et al. 2025 ). The camera-trapping results also provide a more species-based perspective on whether and how pine martens may fine-tune their temporal behaviour to the peculiar challenges posed by urban environments, as already shown in more natural contexts similarly affected by human disturbance (Mirante et al. 2024 ). Although three camera traps may seem limited, they effectively cover the study area (RNI), the only site in Rome where the pine marten has been regularly recorded (Ancillotto et al. 2024 ). Therefore, we are confident that increasing the number of camera traps would not substantially alter our results. The comparison of activity rhythms between urban and natural parks in fact revealed some changes in temporal activity patterns by pine martens in response to the urban context. While in natural areas the species is predominantly crepuscular with occasional diurnal activity (Zielinski et al. 1983 ; Torretta et al. 2016 ), in Rome, it exhibited bimodal peaks at dawn and dusk with moderate overlap with human activity. This partial diurnality may represent a trade-off between avoiding nocturnal competitors (particularly the red fox, as also evident by our results) and exploiting diurnal prey, e.g., red squirrels and birds. However, given that both squirrels and birds may be more vulnerable at nights while resting in nests or roosts (particularly for a species with remarkable arboreal skills as the pine marten), competition avoidance appears the most convincing hypothesis. The reduction in temporal overlap with red foxes observed in urban conditions supports this interpretation and may reflect active temporal niche partitioning to reduce intraguild predation risk or competition for resources (Lindström et al. 1995 ). Avoidance of foxes in urban contexts may also be due to the high densities of red foxes in anthropogenic environments in comparison to more natural ones, a factor that may induce behavioural adjustments in pine martens, as those we observed (Padovani et al. 2021 ). Such plasticity has already been reported in other mesocarnivores able to thrive in anthropogenic environments, where shifts in activity rhythms are crucial for coexistence and access to resources (Massara et al. 2018 ; Mori et al. 2025a ). Pine martens in the urban park showed increased temporal and spatial overlap with humans relative to those in natural habitats. While this might initially suggest habituation to human presence, it could also be a by-product of shared use of the same temporal windows associated with reduced fox activity and higher prey availability. Similar behavioural tolerance to anthropogenic disturbance has been observed in stone martens, a close congener widely recognized as synanthropic (Herr et al. 2009 ; Capon et al. 2021 ). The absence of the stone marten in our study area might have facilitated the pine marten colonisation of this part of Rome, reducing direct competition and allowing niche expansion. However, also the opposite scenario is possible, i.e. the local disappearance of the stone marten is due to the local arrival of the pine marten. Future studies should test whether pine martens can persist in areas where stone martens are well established, or whether interspecific exclusion mechanisms operate between the two congeners (Wereszczuk and Zalewski 2015 ; Gazzola and Balestrieri 2020 ). The high spatial overlap between pine martens and red squirrels in both urban and natural settings further emphasizes their potential trophic and ecological link. Urban parks hosting stable red squirrel populations likely represent valuable foraging grounds for pine martens, as squirrels constitute a potential prey item (Twining et al. 2020 ). However, the increased proximity to humans and domestic animals may on one side limit competition with red foxes and on the other side expose them to elevated risks of road mortality, disturbance, or disease transmission, echoing challenges faced by other urban carnivores (Soulsbury and White 2015 ; Ritzel and Gallo 2020 ; Mori et al. 2022b ). Whether pine martens can sustain long-term viable populations in urban settings will depend on their ability to mitigate these pressures and maintain access to suitable foraging and breeding sites. For instance, in future studies, throat patterns could potentially be used to identify individual martens (Raichev 2018 ). However, our camera-trap images often captured animals in lateral view, making even species-level identification challenging. Nevertheless, the ability to recognise whether the same pine marten individuals persist in a given area would provide valuable insights into their degree of site fidelity, adaptation to urban environments, and coexistence dynamics with the red fox. From a conservation perspective, the colonisation of urban parks by pine martens may have positive ecological implications (Caryl et al. 2012 ). As an opportunistic predator, the species could contribute to controlling rodent populations (including invasive rats Rattus rattus and R. norvegicus , and grey squirrels Sciurus carolinensis : McNicol et al. 2020 ), and regulating other urban mesofauna, potentially restoring some trophic balance within urban ecosystems (Caryl et al. 2012 ). Moreover, its presence in cities may enhance public awareness of biodiversity conservation, triggering citizen engagement through platforms such as iNaturalist or Ornitho.it, which proved invaluable for assessing its distribution. Nonetheless, increased visibility of carnivores in human environments can also lead to conflicts, particularly if perceived as threats to pets or property (Soulsbury and White 2015 ). Clear communication and coexistence strategies will therefore be essential to ensure tolerance and minimize negative interactions. The ongoing expansion of the pine marten into lowland and urbanised landscapes may also have implications for monitoring and management policies, especially since the species is listed under the Annex IV of the Habitats Directive, thus being considered as conservation-relevant and in need of monitoring for assessing its conservation status across the EU. Traditional conservation frameworks which consider the pine marten exclusively as a forest indicator species may need revision to reflect its emerging ecological plasticity (Vergara et al. 2016 ). Urban green spaces, when managed with connectivity and vegetation diversity in mind, could serve as refugia supporting the persistence of native mammals such as rodents (Dondina et al. 2025 ) and, in turn, carnivores (Ancillotto et al. 2024 ). In summary, our findings highlight that the pine marten, once considered an emblematic species of pristine forest ecosystems, is exhibiting unexpected behavioural and ecological flexibility, allowing it to exploit urban habitats without fully altering its natural activity patterns. This adaptive response summarises the dynamic interactions shaping mammalian assemblages in cities, where human presence, resource distribution, and interspecific competition drive rapid ecological change (Mori et al. 2025a ). Continued monitoring will be essential to determine whether these early colonisation patterns lead to stable, reproducing urban populations or represent transient incursions facilitated by peri-urban connectivity. Declarations Author Contribution Statement LA and EM conceived this work, conducted the analyses and wrote the first draft; AA, DG and FV collected data and participated in writing up all drafts. All authors approved the MS before the submission. Conflict of interest the authors have no conflict of interest to declare. Ethics approval Not applicable. Funding No funding was received for this research. Acknowledgements LA and EM were funded by the National Recovery and Resilience Plan (NRRP), Mission 4 Component 2 Investment 1.4 - Call for tender No. 3138 of 16 December 2021, rectified by Decree n.3175 of 18 December 2021 of Italian Ministry of University and Research funded by the European Union – NextGenerationEU; Project code CN_00000033, Concession Decree No. 1034 of 17 June 2022 adopted by the Italian Ministry of University and Research, CUP B83C22002930006, Project title “National Biodiversity Future Center - NBFC”. Isabella Vitali and Cristina Mascalchi (CNR IRET Sesto Fiorentino) also provided valuable technical, administrative and management support. Camera-trapping in RNI was implemented as part of the Activity 5.2 of the Spoke 5 of the PNRR National Biodiversity Future Center project (see Mori et al., 2025a; Viviano et al., 2025; see Acknowledgements). The “U.O. Manutenzione e Valorizzazione del Verde Urbano” office of the Municipality of Roma provided us with permits to set camera traps within urban parks. Data availability statement Data showed in this paper are uploaded as Supplementary Material 1. References Ancillotto L, Tomassini A, Russo D (2016) The fancy city life: Kuhl’s pipistrelle, Pipistrellus kuhlii , benefits from urbanisation. Wildl Res 42(7):598–606. https://doi.org/10.1071/WR15003 Ancillotto L, Amori G, Capizzi D, Cignini B, Zapparoli M, Mori E (2024) No city for wetland species: habitat associations affect mammal persistence in urban areas. Proc R Soc B 291:20240079. https://doi.org/10.1098/rspb.2024.0079 Ancillotto L, Guerri G, Agnelli P, Bonora L, Maggioni M, Morabito M, Mori E (2025) Past present: extinction debt of forest mammals from urban areas. Biol Conserv 306:111143. https://doi.org/10.1016/j.biocon.2025.111143 Balestrieri A, Remonti L, Ruiz-González A, Gómez-Moliner BJ, Vergara M, Prigioni C (2010) Range expansion of the pine marten ( Martes martes ) in an agricultural landscape matrix (NW Italy). Mammal Biol 75:412–419. https://doi.org/10.1016/j.mambio.2009.05.003 Balestrieri A, Remonti L, Ruiz-Gonzalez R, Zenato M, Gazzola A, Vergara M, Dettori EE, Saino N, Capelli E, Gomez-Moliner BJ, Guidali F, Prigioni C (2015) Distribution and habitat use by pine marten Martes martes in a riparian corridor crossing intensively cultivated lowlands. Ecol Res 30:153–162. https://doi.org/10.1007/s11284-014-1220-8 Balestrieri A, Mori E, Menchetti M, Ruiz-González A, Milanesi P (2019) Far from the madding crowd: Tolerance toward human disturbance shapes distribution and connectivity patterns of closely related Martes spp. Popul Ecol 61:289–299. https://doi.org/10.1002/1438-390X.12001 Balestrieri R, Vento R, Viviano A, Mori E, Gili C, Monti F (2023) Razorbills Alca torda in Italian seas: a massive irruption of historical relevance and role of social network monitoring. Animals 13(4):656. https://doi.org/10.3390/ani13040656 Baltrunaite L (2001) Feeding habits, food niche overlap of red fox ( Vulpes vulpes L.) and pine marten ( Martes martes L.) in hilly moraine highland. Lith Ecologija 2:27–31 Bateman PW, Fleming PA (2012) Big city life: carnivores in urban environments. J Zool 287:1–23. https://doi.org/10.1111/j.1469-7998.2011.00887.x Birnie-Gauvin K, Peiman KS, Gallagher AJ, De Bruijn R, Cooke SJ (2016) Sublethal consequences of urban life for wild vertebrates. Environ Rev 24:416–425. https://doi.org/10.1139/er-2016-0029 Burnham KP, Anderson DR (2002) Model selection and multimodel inference: a practical information-theoretic approach, 2nd edn. Springer Editions, Berlin Heidelberg Capon M, Lysaniuk B, Godard V, Clauzel C, Simon L (2021) Characterizing the landscape compositions of urban wildlife encounters: the case of the stone marten ( Martes foina ), the red fox ( Vulpes vulpes ) and the hedgehog ( Erinaceus europaeus ) in the Greater Paris area. Urb Ecosyst 24:885–903. https://doi.org/10.1007/s11252-020-01071-6 Carnevali L, Lovari S, Monaco A, Mori E (2016) Nocturnal activity of a diurnal species, the northern chamois, in a predator-free Alpine area. Behav Proc 126:101–107. https://doi.org/10.1016/j.beproc.2016.03.013 Caryl FM, Quine CP, Park KJ (2012) Martens in the matrix: the importance of nonforested habitats for forest carnivores in fragmented landscapes. J Mammal 93(2):464–474. https://doi.org/10.1644/11-MAMM-A-149.1 Clevenger AP (1993) The European pine marten Martes martes in the Balearic Islands, Spain. Mammal Rev 23:65–72. https://doi.org/10.1111/j.1365-2907.1993.tb00417.x Cox K, Gouwy J, Mergeay J, Neyrinck S, Van Den Berge K (2025) Rapid Colonisation of Synanthropic Stone Martens in a Highly Urbanised Region: Insights From Temporal and Spatial Analysis. Ecol Evol 15:e71392. https://doi.org/10.1111/j.1365-2907.1993.tb00417.x Davison J, Huck M, Delahay RJ, Roper TJ (2008) Urban badger setts: characteristics, patterns of use and management implications. J Zool 275(2):190–200. https://doi.org/10.1111/j.1469-7998.2008.00424.x Dickman CR (1987) Habitat fragmentation and vertebrate species richness in an urban environment. J Appl Ecol 24:337–351. https://doi.org/10.2307/2403879 Dondina O, Tirozzi P, Viviano A, Mori E, Orioli V, Tommasi N, Tanzi A, Bazzoli A, Caprio E, Patetta C, Pastore MC, Bani L, Ancillotto L (2025) Spatial and habitat determinants of small-mammal biodiversity in urban green areas: Lessons for nature-based solutions. Urb Urb Green 104:128641 Doykin N, Popova E, Paraskova M, Zlatanov V, Zlatanova D (2017) Pine marten ( Martes martes L.): distribution, habitat preference and activity in Vitosha Nature Park, Bulgaria. Proceedings of Seminar Of Ecology–2016 with International Participation 2016: 26–33 Duduś L, Zalewski A, Kozioł O, Jakubiec Z, Król N (2014) Habitat selection by two predators in an urban area: the stone marten and red fox in Wrocław (SW Poland). Mamm Biol 79(1):71–76. https://doi.org/10.1016/j.mambio.2013.08.001 Fonda F, Chiatante G, Meriggi A, Mustoni A, Armanini M, Mosini A, Spada A, Lombardini M, Righetti D, Granata M, Capelli E, Pontarini R, Roux Poignant G, Balestrieri A (2021) Spatial distribution of the pine marten ( Martes martes ) and stone marten ( Martes foina ) in the Italian Alps. Mammal Biol 101:345–356. https://doi.org/10.1007/s42991-020-00098-8 Gallo T, Fidino M, Lehrer EW, Magle SB (2017) Mammal diversity and metacommunity dynamics in urban green spaces: implications for urban wildlife conservation. Ecol Appl 27(8):2330–2341. https://doi.org/10.1002/eap.1611 Gazzola A, Balestrieri A (2020) Nutritional ecology provides insights into competitive interactions between closely related Martes species. Mamm Rev 50:82–90. https://doi.org/10.1111/mam.12177 Geiger M, Taucher AL, Gloor S, Hegglin D, Bontadina F (2018) In the footsteps of city foxes: evidence for a rise of urban badger populations in Switzerland. Hystrix 29(2):236–238. https://doi.org/10.4404/hystrix-00069-2018 Gerber BD, Devarajan K, Farris ZJ, Fidino M (2024) A model-based hypothesis framework to define and estimate the diel niche via the Diel.Niche R package. J Anim Ecol 93:132–146. https://doi.org/10.1111/1365-2656.14035 Gottero E, Larcher F, Cassatella C (2023) Defining and regulating peri-urban areas through a landscape planning approach: The case study of Turin Metropolitan Area (Italy). Land 12:217. https://doi.org/10.3390/land12010217 Grelli D, Paoloni D, Vercillo F, Ragni B (2014) Un racconto dalla città: la presenza della martora a Perugia (Italia centrale). Hystrix 25 suppl.: 108 Herr J, Schley L, Roper TJ (2009) Socio-spatial organization of urban stone martens. J Zool 277:54–62. https://10.1111/j.1469-7998.2008.00510.x Hughes-Brandl M (2018) Package ‘NightDay’. Night and day boundary plot function. https://cran.r-project.org/web/packages/NightDay/NightDay.pdf . Accessed on 07.10.2025 Landler L, Ruxton GD, Malkemper EP (2019) The Hermans-Rasson test as a powerful alternative to the Rayleigh test for circular statistics in biology. BMC Ecol 19:1–8. https://doi.org/10.1186/s12898-019-0246-8 Lanszki J, Zalewski A, Horváth G (2007) Comparison of red fox Vulpes vulpes and pine marten Martes martes food habits in a deciduous forest in Hungary. Wildl Biol 13:258–271. https://doi.org/10.2981/0909-6396 (2007)13[258:CORFVV]2.0.CO;2 Lindström ER, Brainerd SM, Helldin JO, Overskaug K (1995) Pine marten-red fox interactions: a case of intraguild predation? Ann Zool Fennici 32:123–130 Lombardini M, Cinerari CE, Murru M, Vidus Rosin A, Mazzoleni L, Meriggi A (2015) Habitat requirements of Eurasian pine marten Martes martes in a Mediterranean environment. Mamm Res 60(2):97–105. https://doi.org/10.1007/s13364-014-0211-z Łopucki R, Klich D, Kitowski I (2019) Are small carnivores urban avoiders or adapters: Can they be used as indicators of well-planned green areas? Ecol Indic 101:1026–1031. https://doi.org/10.1016/j.ecolind.2019.02.016 Lovari S, Corsini MT, Guazzini B, Romeo G, Mori E (2017) Suburban ecology of the crested porcupine in a heavily poached area: a global approach. Eur J Wildl Res 63:10. https://doi.org/10.1007/s10344-016-1075-0 Lund U, Agostinelli C, Arai H, Gagliardi A, Portugues EG, Giunchi D, Irisson JO, Pocernich M, Rotolo F (2017) Package circular. http://mirrors.ucr.ac.cr/CRAN/web/packages/circular/circular.pdf . Accessed on 03.10.2025 Massara RL, Paschoal AMO, Bailey LL, Doherty PF, Barreto MF, Chiarello AG (2018) Effect of humans and pumas on the temporal activity of ocelots in protected areas of Atlantic Forest. Mamm Biol 92:86–93. https://doi.org/10.1016/j.mambi o.2018.04.009 Manzo E, Bartolommei P, Rowcliffe JM, Cozzolino R (2012) Estimation of population density of European pine marten in central Italy using camera trapping. Acta Theriol 57:165–172. https://doi.org/10.4404/hystrix-23.2-7099 Manzo E, Bartolommei P, Giuliani A, Gentile G, Dessì-Fulgheri F, Cozzolino R (2018) Habitat selection of European pine marten in Central Italy: from a tree dependent to a generalist species. Mamm Res 63:357–367. https://doi.org/10.1007/s13364-018-0374-0 Manzo E, Iannarilli F, Dell'Agnello F, Bartolommei P, Bonacchi A, Gasperini S, Cozzolino R (2024) Assessing the co-occurrence of European pine marten ( Martes martes ) with humans and domestic cats on a Mediterranean island. Ecol Evol 14(12):e70651. https://doi.org/10.1002/ece3.70651 Manzo E, Vercillo F, Fedi C (2025) Martora Martes martes . In: Loy A, Bon M, Di Febbraro M, Baisero D, Amori G (eds) Atlas of mammals in Italy. Edizioni Belvedere, Latina, Italy, pp 234–237 McNicol CM, Bavin D, Bearhop S, Ferryman M, Gill R, Goodwin CE, MacPherson J, Silk MJ, McDonald RA (2020) Translocated native pine martens Martes martes alter short-term space use by invasive non‐native grey squirrels Sciurus carolinensis . J Appl Ecol 57:903–913 Meredith M, Ridout M (2014) Overview of the overlap package. http://cran.cs.wwu.edu/web/packages/overlap/vignettes/overlap.pdf Accessed on 07.10.2025 Miller KF, Wilson DJ, Hartley S, Innes JG, Fitzgerald NB, Miller P, van Heezik Y (2022) Invasive urban mammalian predators: distribution and multi-scale habitat selection. Biology 11(10):1527. https://doi.org/10.3390/biology11101527 Mirante D, Ancillotto L, Zampetti A, Coiro G, Pisa G, Santocchi C, Giuliani M, Santini L (2024) Fine-tuning coexistence: Wildlife's short-term responses to dynamic human disturbance patterns. Glob Ecol Conserv 54:e03053. https://doi.org/10.1016/j.gecco.2024.e03053 Monteiro MCM, Lira PK (2020) Metropolitan mammals: understanding the threats inside an urban protected area. Oecol Australis 24:661–675. https://doi.org/10.4257/oeco.2020.2403.10 Monterroso P, Alves PC, Ferreras P (2014) Plasticity in circadian activity patterns of mesocarnivores in southwestern Europe: Implications for species coexistence. Behav Ecol Sociobiol 68:1403–1417. https://doi.org/10.1007/s00265-014-1748-1 Mori E, Menchetti M, Camporesi A, Cavigioli L, Tabarelli de Fatis K, Girardello M (2019) License to kill? Domestic cats affect a wide range of native fauna in a highly biodiverse Mediterranean country. Front Ecol Evol 7:477. https://doi.org/10.3389/fevo.2019.00477 Mori E, Bagnato S, Serroni P, Sangiuliano A, Rotondaro F, Marchianò V, Cascini V, Poerio L, Ferretti F (2020) Spatiotemporal mechanisms of coexistence in an European mammal community in a protected area of southern Italy. J Zool 310:232–245. https://doi.org/10.1111/jzo.12743 Mori E, Molteni R, Ancillotto L, Ficetola GF, Falaschi M (2022a) Spatial ecology of crested porcupine in a metropolitan landscape. Urb Ecosyst 25(6):1797–1803. https://doi.org/10.1007/s11252-022-01264-1 Mori E, Fedele E, Greco I, Giampaoli Rustichelli M, Massolo A, Miniati S, Puppo F, Santini G, Zaccaroni M (2022b) Spatiotemporal activity of the pine marten Martes martes : Insights from an island population. Ecol Res 37:102–114. https://doi.org/10.1111/1440-1703.12269 Mori E, Lazzeri L, Maggioni M, Viviano A, Guerri G, Morabito M, Martini S, Dondina O, Sogliani D, Scarfò M, Ancillotto L (2025a) Camera-traps and the city: spatiotemporal adaptations of wildlife to urban environments. Ecol Sol Evid 6:e70115. https://doi.org/10.1002/2688-8319.70115 Mori E, Palombi A, Blasi MC, Viviano A, Piazzai M, Colonnelli L, Lazzeri L, Spilinga C, Montioni F, Bombara G, Sogliani D, Ancillotto L (2025b) Spatiotemporal partitioning between sympatric hares and their predators. Eur J Wildl Res 71:127. https://doi.org/10.1007/s10344-025-01979-4 Mori E, Marchi V, Dondina O, Viviano A, Di Bari P, Balestrieri R, Corradetti M, Ancillotto L (2025c) Citizen eyes on elusive wildlife: assessing public appreciation for urban wild mammals. Ambio https://doi.org/10.1007/s13280-025-02315-5 O’Connell AF, Nichols JD, Karanth KU (2011) Camera traps in animal ecology: methods and analyses, camera traps in animal ecology: methods and analyses. Springer Science & Business Media, Tokyo, Japan Padovani R, Shi Z, Harris S (2021) Are British urban foxes ( Vulpes vulpes ) bold? The importance of understanding human–wildlife interactions in urban areas. Ecol Evol 11:835–851. https://doi.org/10.1002/ece3.7087 Pereboom V, Mergey M, Villerette N, Helder R, Gerard JF, Lode T (2008) Movement patterns, habitat selection, and corridor use of a typical woodland-dweller species, the European pine marten ( Martes martes ), in fragmented landscape. Can J Zool 86(9):983–991. https://doi.org/10.1139/Z08-076 Pianka ER (1974) Niche overlap and diffuse competition. Proc Nat Ac Sci USA 71:2141–2145. https://doi.org/10.1073/pnas.71.5.2141 QGIS Development Team (2019) QGIS Geographic Information System. Open Source Geospatial Foundation Project. http://qgis.osgeo.org . Accessed on 06.10.2025 R Core Team (2021) R: A language and environment for statistical computing. R Foundation for Statistical Computing. https://www.R-project.org . Accessed on 07.10.2025 Raichev E (2018) Determination of stone marten ( Martes foina ) and pine marten ( Martes martes ) in natural habitats using camera traps. Agricult Sci Technol 10:160–163 Ripley B, Venables B, Bates DM, Hornik K, Gebhardt A, Firth D, Ripley MB (2013) Package ‘mass’. Cran R 538:822 Ritzel K, Gallo T (2020) Behavior change in urban mammals: a systematic review. Front Ecol Evol 8:576665. https://doi.org/10.3389/fevo.2020.576665 Robinson OJ, Ruiz-Gutierrez V, Fink D (2017) Correcting for bias in distribution modelling for rare species using citizen science data. Divers Distrib 24:460–472. https://doi.org/10.1111/ddi.12698 Russo D, Ancillotto L (2015) Sensitivity of bats to urbanization: a review. Mamm Biol 80(3):205–212. https://doi.org/10.3389/fevo.2020.576665 Sainsbury KA, Shore RF, Schofield H, Croose E, Campbell RD, Mcdonald RA (2019) Recent history, current status, conservation and management of native mammalian carnivore species in Great Britain. Mamm Rev 49:171–188. https://doi.org/10.1111/mam.12150 Santicchia F, Tranquillo C, Wauters LA, Palme R, Panzeri M, Preatoni D, Bisi F, Martinoli A (2024) Physiological stress response to urbanisation differs between native and invasive squirrel species. Sci Tot Environ 992:171336. https://doi.org/10.1016/j.scitotenv.2024.171336 Santini L, González-Suárez M, Russo D, Gonzalez‐Voyer A, von Hardenberg A, Ancillotto L (2019) One strategy does not fit all: determinants of urban adaptation in mammals. Ecol Lett 22:365–376. https://doi.org/10.1111/ele.13199 Silvertown J (2009) A new dawn for citizen science. Trends Ecol Evol 24:467–471. https://doi.org/10.1016/j.tree.2009.03.017 Sogliani D, Mori E, Lovari S, Lazzeri L, Longoni A, Tabarelli De Fatis K, Sabatini P, Di Nicola MR, Russo D (2023) Citizen science and diet analysis shed light on dog-wildlife interactions in Italy. Biodiv Conserv 32(13):4461–4479. https://doi.org/10.1007/s10531-023-02707-7 Soulsbury CD, White PC (2015) Human–wildlife interactions in urban areas: a review of conflicts, benefits and opportunities. Wildl Res 42:541–553. https://doi.org/10.1071/WR14229 Storch I, Lindström E, de Jounge J (1990) Diet and habitat selection of the pine marten in relation to competition with the red fox. Acta Theriol 35:311–320 Takahashi Y, Yagisawa R, Tsuji Y (2025) Niche separation of sympatric carnivores, red foxes ( Vulpes vulpes ) and Japanese martens ( Martes melampus ) in Northern Japan. Mamm Stud 51:1. https://doi.org/10.3106/ms2024-0075 Todini A, Crosti R (2020) Il cinghiale ( Sus scrofa ) come determinante di cambiamenti di vegetazione in una foresta urbana mediterranea: impatto sulla biodiversità di un’area protetta. Forest@ 17:71–77. https://doi.org/10.3832/efor3284-017 Torretta E, Serafini M, Puopolo F, Schenone L (2016) Spatial and temporal adjustments allowing the coexistence among carnivores in Liguria (NW Italy). Acta Ethol 19:123–132. https://doi.org/10.1007/s10211-015-0231-y Torretta E, Mosini A, Piana M, Tirozzi P, Serafini M, Puopolo F, Saino N, Balestrieri A (2017) Time partitioning in mesocarnivore communities from different habitats of NW Italy: insights into martens’ competitive abilities. Behaviour 154:241–266. https://doi.org/10.1163/1568539X-00003420 Tóth M, Bárány A, Kis R (2009) An evaluation of stone marten ( Martes foina ) records in the city of Budapest, Hungary. Acta Zool Ac Scient Hung 55(2):199–209 Turner J, Freeman R, Carbone C (2022) Using citizen science to understand and map habitat suitability for a synurbic mammal in an urban landscape: The hedgehog Erinaceus europaeus . Mamm Rev 52:291–303. https://doi.org/10.1111/mam.12278 Twining JP, Montgomery WI, Tosh DG (2020) The dynamics of pine marten predation on red and grey squirrels. Mammal Biol 100:285–293. https://doi.org/10.1007/s42991-020-00031-z Vergara M, Cushman SA, Urra F, Ruiz-González A (2016) Shaken but not stirred: multiscale habitat suitability modeling of sympatric marten species ( Martes martes and Martes foina ) in the northern Iberian Peninsula. Landsc Ecol 31:1241–1260. https://doi.org/10.1007/s10980-015-0307-0 Villaseñor NR, Driscoll DA, Escobar MA, Gibbons P, Lindenmayer DB (2014) Urbanization impacts on mammals across urban-forest edges and a predictive model of edge effects. PLoS ONE 9(5):e97036. https://doi.org/10.1371/journal.pone.0097036 Viviano A, D’Amico M, Mori E (2023) Aliens on the road: surveying wildlife roadkill to assess the risk of biological invasion. Biology 12:850. https://doi.org/10.3390/biology12060850 Viviano A, De Meo I, Mori E, Sergiacomi C, Paletto A (2024) Public perception and acceptance of coypu Myocastor coypus removal in urban areas: influences of age and education. Sci Nat 111(5):42. https://doi.org/10.1007/s00114-024-01928-2 Viviano A, Ancillotto L, Dondina O, Burchielli A, Miccolis D, Mori E (2025) What a camera trap survey can reveal about the behaviour of an invasive species: Insights from coypus Myocastor coypus in an urban park of central Italy. Appl Anim Behav Sci 284:106534. https://doi.org/10.1016/j.applanim.2025.106534 Wereszczuk A, Zalewski A (2015) Spatial niche segregation of sympatric stone marten and pine marten–avoidance of competition or selection of optimal habitat? PLoS ONE 10:e0139852. https://doi.org/10.1016/10.1371/journal.pone.0139852 Werner P (2011) The ecology of urban areas and their functions for species diversity. Landsc Ecol Engin 7(2):231–240. https://doi.org/10.1007/s11355-011-0153-4 Widmer S, Riedel S, Babbi M, Herzog F, Wohlgemuth T, Kessler M, Dengler J (2025) One century of change: stronger diversity decline in lowland than in mountain grasslands in central Europe. Glob Change Biol 31:e70529. https://doi.org/10.1111/gcb.70529 Zabala J, Zuberogoitia I, Martínez-Climent JA (2005) Site and landscape features ruling the habitat use and occupancy of the polecat ( Mustela putorius ) in a low density area: a multiscale approach. Eur J Wildl Res 51(3):157–162. https://doi.org/10.1007/s10344-005-0094-z Zielinski WJ, Spencer WD, Barrett RH (1983) Relationship between food habits and activity patterns of pine martens. J Mammal 64(3):387–396. https://doi.org/10.2307/1380351 Supplementary Files SUPPLEMENTARYMATERIAL1.docx Cite Share Download PDF Status: Under Review Version 1 posted Reviewers agreed at journal 24 Mar, 2026 Reviewers invited by journal 19 Mar, 2026 Editor assigned by journal 06 Mar, 2026 First submitted to journal 05 Mar, 2026 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-9043132","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":608817648,"identity":"6ca685d3-6403-4733-97cc-837f182b94e2","order_by":0,"name":"Leonardo Ancillotto","email":"","orcid":"","institution":"Istituto di Ricerca sugli Ecosistemi Terrestri Consiglio Nazionale delle Ricerche","correspondingAuthor":false,"prefix":"","firstName":"Leonardo","middleName":"","lastName":"Ancillotto","suffix":""},{"id":608817649,"identity":"914ac3a3-3e1a-4cdd-a5d0-09cdf34a52f9","order_by":1,"name":"Andrea Amici","email":"","orcid":"","institution":"Terna SpA","correspondingAuthor":false,"prefix":"","firstName":"Andrea","middleName":"","lastName":"Amici","suffix":""},{"id":608817650,"identity":"7e71dc56-8520-48b1-bac6-0d72eaf98b4b","order_by":2,"name":"Dorian Grelli","email":"","orcid":"","institution":"Umbria Region: Regione Umbria","correspondingAuthor":false,"prefix":"","firstName":"Dorian","middleName":"","lastName":"Grelli","suffix":""},{"id":608817651,"identity":"38d4b1b2-a396-45c6-bd30-5108a0578405","order_by":3,"name":"Francesca Vercillo","email":"","orcid":"","institution":"Regione Umbria","correspondingAuthor":false,"prefix":"","firstName":"Francesca","middleName":"","lastName":"Vercillo","suffix":""},{"id":608817652,"identity":"6305cccb-d0a5-4602-89eb-0512b05283ee","order_by":4,"name":"Emiliano Mori","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA9ElEQVRIiWNgGAWjYFACxsYDCVDmgY9/bCAiBLQ0wLUcnNmQBhEhZA9cATNvw2FUEWyAX+xww4EHNXfkdNt7Hx6cueO83dp2oAhDjU00Li2SsxOBDjv2zNjszHGDAx/P3E7edgYownAsLbcBhxaD2yAtbIcTt91IYzg4g+12stkBoAhjw2GcWuzBWv5BtBzmYTuXbHb+IX4tBtJALYltUC28bQfszG4QsEUCZEti32GgX44BHXYmOcHsxkOQ73D7hX92+sOHP74dljM73sb84UOFnb3Z+fSHDz7U2ODUggESwSoTiFUOAvakKB4Fo2AUjIKRAQCtS3ScJVYLMwAAAABJRU5ErkJggg==","orcid":"https://orcid.org/0000-0001-8108-7950","institution":"IRET CNR: Istituto di Ricerca sugli Ecosistemi Terrestri Consiglio Nazionale delle Ricerche","correspondingAuthor":true,"prefix":"","firstName":"Emiliano","middleName":"","lastName":"Mori","suffix":""}],"badges":[],"createdAt":"2026-03-05 17:47:28","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-9043132/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-9043132/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":105234193,"identity":"eb629572-3577-4ffa-86d5-d6fa784daf49","added_by":"auto","created_at":"2026-03-23 19:22:49","extension":"jpg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":94739,"visible":true,"origin":"","legend":"\u003cp\u003eNumber of pine marten records in Italian cities (N = 10 out of the 26 sampled), and range of observation years. Green diamonds highlight the occurrence of the stone marten in the same city (Table S1 in Supplementary Material 1).\u003c/p\u003e","description":"","filename":"Figure1.jpg","url":"https://assets-eu.researchsquare.com/files/rs-9043132/v1/1ed85215e3abc5a02a2e0f5f.jpg"},{"id":105234190,"identity":"415253ca-f223-4d47-b4fd-acebeb4e15c3","added_by":"auto","created_at":"2026-03-23 19:22:48","extension":"jpg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":626691,"visible":true,"origin":"","legend":"\u003cp\u003eProbability of pine marten occurrence as a function of percent of urban tree cover (left panel) and altitude (right panel, in metres above sea level) in 26 Italian cities. The line is the modelled trend and grey areas represent 95% confidence intervals.\u003c/p\u003e","description":"","filename":"Figure2.jpg","url":"https://assets-eu.researchsquare.com/files/rs-9043132/v1/1301eaf6ff71f876fb1da8af.jpg"},{"id":105234191,"identity":"a9a13248-3f76-4086-b1ca-7e7a9a3fe3fe","added_by":"auto","created_at":"2026-03-23 19:22:48","extension":"jpg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":1007500,"visible":true,"origin":"","legend":"\u003cp\u003eAnnual overlap of activity rhythms between target species (from left to right: pine marten, red fox, red squirrel and human) in urban (Insugherata urban park) and natural areas (Monte Rufeno Nature Reserve), represented by the shaded areas. Coefficients of overlap Δ₁ and relevant 95% confidence intervals (95%CIs) are expressed above each plot. Species silhouettes from \u003ca href=\"http://www.phylopic.org/\"\u003ewww\u003c/a\u003e\u003ca href=\"http://www.phylopic.org/\"\u003e.\u003c/a\u003e\u003ca href=\"http://www.phylopic.org/\"\u003ephylopic\u003c/a\u003e\u003ca href=\"http://www.phylopic.org/\"\u003e.\u003c/a\u003e\u003ca href=\"http://www.phylopic.org/\"\u003eorg\u003c/a\u003e.\u003c/p\u003e","description":"","filename":"Figure3.jpg","url":"https://assets-eu.researchsquare.com/files/rs-9043132/v1/3ce2fdd722242affa4e7276e.jpg"},{"id":105234192,"identity":"d6a6d7de-155d-464b-a524-0b366884e246","added_by":"auto","created_at":"2026-03-23 19:22:48","extension":"jpg","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":2992350,"visible":true,"origin":"","legend":"\u003cp\u003eAnnual overlap (shaded areas) of activity rhythms between pine martens and red squirrels, pine martens and red foxes, and pine martens and humans in urban (RNI, left) and natural (Monte Rufeno Nature Reserve, right) areas. Values of bootstrap tests are shown. Species silhouettes from \u003ca href=\"http://www.phylopic.org/\"\u003ewww.phylopic.org\u003c/a\u003e.\u003c/p\u003e","description":"","filename":"Figure4.jpg","url":"https://assets-eu.researchsquare.com/files/rs-9043132/v1/00e8cc7b80011c67ba6b12d7.jpg"},{"id":105564222,"identity":"18284aab-7aca-41ca-9626-1b041b2b3bab","added_by":"auto","created_at":"2026-03-27 12:49:03","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":5414763,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-9043132/v1/46f21019-03fa-4c22-a3b1-b5f253ddda3f.pdf"},{"id":105234194,"identity":"dda37a7c-dcbe-4c14-9397-1df99395ca6c","added_by":"auto","created_at":"2026-03-23 19:22:49","extension":"docx","order_by":8,"title":"","display":"","copyAsset":false,"role":"supplement","size":28783,"visible":true,"origin":"","legend":"","description":"","filename":"SUPPLEMENTARYMATERIAL1.docx","url":"https://assets-eu.researchsquare.com/files/rs-9043132/v1/7c6d441e17927afdb44b26d3.docx"}],"financialInterests":"","formattedTitle":"Can a forest specialist live in a city? Behavioural plasticity fosters the presence of a protected carnivore in urban areas","fulltext":[{"header":"Introduction","content":"\u003cp\u003eUrban ecosystems are increasingly recognized as dynamic areas which may host a surprising diversity of wildlife (Werner \u003cspan citationid=\"CR87\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Gallo et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Urban ecology define urban areas are densely populated regions with extensive infrastructure and services, typically forming cities or large towns (Gottero et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Conversely, periurban areas are transitional zones located on the outskirts of cities where urban and rural characteristics coexist and land use is rapidly changing (Gottero et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Rural areas are regions with low population density, limited infrastructure, and predominantly agricultural or natural landscapes (Gottero et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eWild mammals in urban areas face a range of threats primarily linked to habitat loss, fragmentation, and human activity (Soulsbury and White \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Santini et al. \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Urban expansion reduces and isolates natural habitats, making it difficult for mammals to find food, mates, and shelter (Dickman \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e1987\u003c/span\u003e; Villase\u0026ntilde;or et al. \u003cspan citationid=\"CR82\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Monteiro and Lira \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Roads and traffic cause high mortality through vehicle collisions, while light, noise, and chemical pollution disrupt behaviour, communication, and health (Birnie-Gauvin et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Ritzel and Gallo \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Viviano et al. \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Domestic pets and invasive species increase predation and competition pressures (Mori et al. \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Miller et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Sogliani et al. \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Additionally, direct conflict with humans (e.g., predations upon poultry or livestock), often harmful for wildlife, poses further risks (Soulsbury and White \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Combined, these pressures often lead to declines in population size, genetic diversity, and overall ecosystem stability. In particular, in Europe, several mammal species have successfully started to thrive in urban environments, exploiting anthropogenic resources and novel ecological opportunities (Santini et al. \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Urban mammals are typically ecological generalists, characterized by plasticity in space use, broad dietary spectra, and behavioural flexibility (Santini et al. \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Mori et al. \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2025a\u003c/span\u003e). Among them, native species including carnivores (e.g., red fox \u003cem\u003eVulpes vulpes\u003c/em\u003e and stone marten \u003cem\u003eMartes foina\u003c/em\u003e), bats (e.g., the Kuhl\u0026rsquo;s pipistrelle \u003cem\u003ePipistrellus kuhlii\u003c/em\u003e) and small mammals (e.g., the Western hedgehog \u003cem\u003eErinaceus europaeus\u003c/em\u003e and the Eurasian red squirrel \u003cem\u003eSciurus vulgaris\u003c/em\u003e) have become well-established urban dwellers (Russo and Ancillotto \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Ancillotto et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2016\u003c/span\u003e, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2025\u003c/span\u003e; Bateman and Fleming \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Capon et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Turner et al. \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Cox et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2025\u003c/span\u003e; Dondina et al., \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Additionally, a number of non-native species, such as the Eastern grey squirrel \u003cem\u003eSciurus carolinensis\u003c/em\u003e or the coypu \u003cem\u003eMyocastor coypus\u003c/em\u003e, have thrived in European cities, benefiting from low predation pressure, human-made structures (e.g., abandoned buildings used as denning-sites) and frequent human tolerance or even support (Mori et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2025c\u003c/span\u003e; Santicchia et al. \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Viviano et al. \u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e2024\u003c/span\u003e, \u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e2025\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eConversely, species tightly associated with specific or fragile habitats, such as wetlands or old-growth forests, tend to be poorly suitable to survive in urbanized ecosystems, and may undergo local extinctions as a result (Ancillotto et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2024\u003c/span\u003e, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Indeed, during the last decade, a growing number of studies have documented formerly forest-associated mammals expanding into peri-urban and urban areas. Examples include the European badger \u003cem\u003eMeles meles\u003c/em\u003e and the crested porcupine \u003cem\u003eHystrix cristata\u003c/em\u003e, which are increasingly observed using urban shrublands as denning sites and foraging in parks and suburban gardens (Davison et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Lovari et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Geiger et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Mori et al. \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2022a\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eWithin the Mustelidae family, small \u003cem\u003eMustela\u003c/em\u003e species (namely, \u003cem\u003eM. nivalis\u003c/em\u003e, \u003cem\u003eM. putorius\u003c/em\u003e) are rarely recorded both in natural environments and, for most, in cities (Zabala et al. \u003cspan citationid=\"CR89\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Łopucki et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2019\u003c/span\u003e), whereas the genus \u003cem\u003eMartes\u003c/em\u003e is represented almost exclusively by \u003cem\u003eM. foina\u003c/em\u003e (Duduś et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Capon et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Cox et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2025\u003c/span\u003e; Mori et al. \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2025a\u003c/span\u003e). This species is highly synanthropic, omnivorous, and behaviourally plastic, exploiting urban and rural structures e.g., attics, archaeological sites, and roof spaces as diurnal resting places (Herr et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; T\u0026oacute;th et al. \u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Balestrieri et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). The strictly nocturnal habits and tolerance to human presence make \u003cem\u003eM. foina\u003c/em\u003e one of the most successful carnivores in European cities (Mori et al. \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2025a\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIn contrast, the pine marten \u003cem\u003eM. martes\u003c/em\u003e has long been regarded as a forest specialist and is considered as an indicator species of well-preserved forest ecosystems (Clevenger \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e1993\u003c/span\u003e; Lombardini et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Manzo et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Balestrieri et al. (\u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) suggested that distance to watercourses, and the cover of deciduous forest and shrubland, are the main factors shaping pine marten occurrence across its range, whereas the stone marten is more closely associated with higher human population density and extent of built-up areas. Consequently, although pine martens may locally select areas with intermediate levels of human modifications (Vergara et al. \u003cspan citationid=\"CR81\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), increasing human density and urbanization of lowland and hilly landscapes have been predicted to represent major limiting factors to the pine marten. Furthermore, it has been suggested that a competition for resources may occur between stone and pine martens (Wereszczuk and Zalewski \u003cspan citationid=\"CR86\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Gazzola and Balestrieri \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Moreover, the more diurnal activity patterns of the pine marten compared to the stone marten may increase the risk of encounters with humans and domestic carnivores (Zielinski et al. \u003cspan citationid=\"CR90\" class=\"CitationRef\"\u003e1983\u003c/span\u003e; Torretta et al. \u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e2016\u003c/span\u003e, \u003cspan citationid=\"CR77\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Mori et al. \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2022b\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIn recent times, evidence from Italy and other parts of Europe suggests that this view may be changing, with pine martens increasingly occurring in rural, thus fragmented, landscapes (Pereboom et al. \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Balestrieri et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Vergara et al. \u003cspan citationid=\"CR81\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Manzo et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2018\u003c/span\u003e, \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). However, pine martens tend to avoid highly anthropized environments, possibly due to aversion to humans and domestic carnivores such as dogs and cats (Mori et al. \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2022b\u003c/span\u003e). In rural and natural landscapes, the species is known to compete with the red fox, and cases of intraguild predation have also been documented, potentially representing an additional constraint in urban habitats (Storch et al. \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Lindstr\u0026ouml;m et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e1995\u003c/span\u003e), where more opportunistic carnivores may reach very high population densities.\u003c/p\u003e \u003cp\u003eAgainst this background, the aim of the present study was to collect available records of pine martens in urban environments in Italy, and to describe their activity rhythms in urban environment and to relate them to those of the main potential prey (i.e., the Eurasian red squirrel \u003cem\u003eSciurus vulgaris\u003c/em\u003e) and competitors (i.e., humans and the red fox). Moreover, we compared these patterns with those recorded in natural protected areas far from urban disturbance (Mori et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2025b\u003c/span\u003e). We hypothesized that in urban settings, pine martens may exhibit temporal trade-offs to minimize overlap with humans and domestic carnivores, as well as to maintain a high overlap with potential prey species, maintaining partial diurnality to avoid competition or predation by red foxes, with which they share both space and dietary niches.\u003c/p\u003e"},{"header":"Material and Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eDo pine martens occur in cities?\u003c/h2\u003e \u003cp\u003e We reviewed all publicly available records of pine marten occurring in urban environments within Italy up to December 2025. Our investigation focused on regional capital cities, which were selected as representative urban centres with consistent administrative boundaries and accessible information. Pine marten records from all the other provincial capital cities were also included. All reports of pine martens within boundaries of these cities were collected and verified, based on photographic or otherwise certified evidence. Records were considered as valid only when provided with a spatial precision\u0026thinsp;\u0026lt;\u0026thinsp;100 meters, and were assigned to urban habitats whenever coordinates were within the administrative boundaries of the considered cities i.e., not encompassing the entire province and yet also including peripheral areas and urban parks. Type of records included: (i) carcasses and injured individuals (i.e., museum specimens and Wildlife Rescue Center data), (ii) photographs gathered from multiple complementary resources (i.e., social networks and citizen science platforms) and (iii) scientific literature. Publicly available photographs were systematically searched across social networks (i.e., Facebook, Instagram, Flickr, X/Twitter), considering only posts that were visible without restrictions or user login. In parallel, all georeferenced and photo-documented observations of the pine marten in Italy were downloaded from the citizen science platforms iNaturalist (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e\u003ca href=\"http://www.inaturalist.org\" target=\"_blank\"\u003ewww.inaturalist.org\u003c/a\u003e\u003c/span\u003e\u003cspan address=\"http://www.inaturalist.org\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e. Accessed on 06.10.2025) and Ornitho.it (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e\u003ca href=\"http://www.inaturalist.org\" target=\"_blank\"\u003ewww.ornitho.it\u003c/a\u003e\u003c/span\u003e\u003cspan address=\"http://www.ornitho.it\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e. Accessed on 06.10.2025). To complement citizen-generated data, a comprehensive literature search was conducted using Zoological Records, Scopus, and Google Scholar. The searches included both Italian and English language publications and employed all relevant keyword combinations of \u0026ldquo;Mart*\u0026rdquo;, \u0026ldquo;\u003cem\u003eMartes martes\u003c/em\u003e\u0026rdquo;, \u0026ldquo;urban\u0026rdquo;, \u0026ldquo;city\u0026rdquo;, and \u0026ldquo;Italy\u0026rdquo;, connected by Boolean operators. Example search strings included (\u0026ldquo;\u003cem\u003eMartes martes\u003c/em\u003e\u0026rdquo; OR \u003cem\u003eMartes\u003c/em\u003e OR marten OR martor*) AND (urban OR city OR \u0026ldquo;urban area\u0026rdquo;) AND (Italy OR Italia). Equivalent combinations in Italian were also used. Only records providing explicit urban locations or referred to observations within cities were retained. Duplicates between sources were identified and removed after cross-checking location, date, and image content. For each city we screened, we recorded whether reports of \u003cem\u003eM. martes\u003c/em\u003e were available, the year of each observation, the status of the animal (alive or dead), and the source of the information (e.g., citizen science platform, social media, or peer-reviewed publication). The concurrent presence of the stone marten in the same city was also recorded by using the same methods specified for the pine marten, to enable comparative analyses between the two congeners. All photographic records were visually inspected by at least two researchers to verify species identifications, based on diagnostic morphological traits (following Raichev \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2018\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIn order to assess whether pine martens occur in cities with specific characteristics, we downloaded city size (in squared km; ISTAT) and total amount of tree cover (ISPRA). We then built a generalized linear model (GLM) with a binomial error structure, using pine marten occurrence as binary response variable, city size and tree cover as explaining variables; additionally, to account for potential competition, we also included a binary covariate indicating the occurrence, within the same city, of the congeneric stone marten. We then selected the set of variables best explaining the observed patterns of occurrence by performing a stepwise backward model selection procedure according to AIC values (Burnham and Anderson \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2002\u003c/span\u003e), using the \u003cem\u003estepAIC\u003c/em\u003e function in the MASS package (Ripley et al. \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2013\u003c/span\u003e) for R version 4.2.2. (R Core Team \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) i.e., considering the best model(s) the one minimizing AIC values and those featuring delta-AIC values\u0026thinsp;\u0026lt;\u0026thinsp;2 from the best model.\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eDo urban martens change their behaviour?\u003c/h3\u003e\n\u003cp\u003eIn order to assess pine marten behavioural patterns in terms of activity rhythms, we set a camera trapping sampling in urban and natural sites. The first site was an urban Nature Reserve (Riserva Naturale dell\u0026rsquo;Insugherata, hereafter RNI) in Rome, one of the largest protected green areas within the municipality of Rome, Italy, covering approximately 730 hectares (Todini and Crosti \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Located in the northwestern sector of the city, the park features a diverse mosaic of Mediterranean ecosystems, including oak woodlands, cork oak \u003cem\u003eQuercus suber\u003c/em\u003e L. forests, shrublands, and open grasslands. Its topography is characterized by hills and deep valleys, with elevations ranging from 50 to 150 meters above sea level. The Insugherata urban park supports high biodiversity, providing habitat for numerous plant and animal species (Todini and Crosti \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), yet stretching inside the city, its boundaries being at \u0026lt;\u0026thinsp;2 km from Rome\u0026rsquo;s city centre. Additionally, the park contains remnants of ancient Roman roads and agricultural terraces, reflecting a long history of human interaction with the local landscape (Todini and Crosti \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). The climate is typically Mediterranean, with an annual mean temperature of about 16\u0026deg;C\u0026thinsp;\u0026plusmn;\u0026thinsp;1\u0026deg;C (range: 3\u0026ndash;37\u0026deg;C) in 2020\u0026ndash;2025, featuring warm summers and mild winters, and an average annual precipitation of approximately 850 mm\u0026thinsp;\u0026plusmn;\u0026thinsp;50 mm, with rainfall mostly concentrated in autumn and winter. Seasonal temperature variations are moderate, reflecting typical conditions of the city and its surrounding hilly areas (Ancillotto et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Three 32-MP camera traps (\u0026copy; Coolife, China) were deployed within the RNI at 30\u0026ndash;80 cm above ground level and operated continuously (24 h/day). Each device was programmed to capture a 60-s video sequence upon detection of an animal, a setting chosen to reduce motion blur and thereby improve taxonomic resolution (O\u0026rsquo;Connell et al. \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). Field sampling was conducted from September 2023 to August 2024, yielding 954 trap-nights and covering the whole reserve, the only one in Rome metropolitan area hosting pine martens (Ancillotto et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Cameras were installed at three fixed, georeferenced points, secured to trees or rocks using ropes and chains, and positioned along the nearest wildlife trail to randomly generated sampling points created in QGIS (QGIS Development Team \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). To reduce the likelihood of pseudoreplication, stations were spaced a minimum of 1.2 km apart (O\u0026rsquo;Connell et al. \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). We compared the activity rhythms of pine martens, their prey, and competitor species in urban areas with those of the same species in a natural area at the same latitude in Italy (Monte Rufeno Nature Reserve, approx. 3,000 ha; 275\u0026ndash;774 m a.s.l.; Mori et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2025b\u003c/span\u003e), during the same study period. This rural area is characterized by a mosaic of deciduous oak woodlands, scrublands, pastures, and riparian habitats, where 30 fixed camera traps were deployed across the full altitudinal range, spaced at least 800 m apart and active 24 h/day, recording 60-second videos per trigger (see Mori et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2025b\u003c/span\u003e). Cameras yielded over 9,000 camera-days of sampling effort.\u003c/p\u003e \u003cp\u003eCamera-traps were checked once a month to retrieve recordings and replace dead batteries. For each video, metadata were entered into a structured dataset including: species identity, date, time, diel phase (diurnal/nocturnal), station ID, lighting condition (day/night), and meteorological season (spring, summer, autumn, winter). Videos for which species identification was not possible (8.8% of all wildlife records) were excluded from analyses. Camera trapping was conducted under all necessary authorizations and in full compliance with national and international regulations, including the Italian Legislative Decree 196/2003 and the European Regulation 2016/679 on privacy. Informative panels were installed at monitoring sites to notify the public about the presence of the cameras and the objectives of the study. Each panel included a QR code providing access to project documentation and ensuring that any recordings containing humans were deleted within 72 hours from all storage devices. Direct contact with the project manager was also provided in each panel to address any concern.\u003c/p\u003e \u003cp\u003eAll camera-trap records were compiled into a dataset containing information on species identity, date, time, and camera-trap station. Species identification was independently performed by three of the authors using a double-blind protocol. To reduce pseudoreplication, consecutive detections of the same species at the same camera-trap station occurring within a 30-minute interval were merged into a single event (Monterroso et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). The dataset retained the original dates and times, i.e. solar hour recorded on each video file to minimize temporal bias. Nighttime was defined as the period from one hour after sunset to one hour before sunrise (Carnevali et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), while dawn and dusk were delineated according to astronomical circadian twilight. These time periods were calculated using the \u003cem\u003eNightDay\u003c/em\u003e package (Hughes-Brandl \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) for R version 4.1.2 (R Core Team 2018). This framework distinguishes canonical diel niches based on relative activity across daily periods and identifies both bimodal and trimodal activity distributions (Gerber et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Activity rhythms of pine martens were analyzed using the overlap package (Meredith \u0026amp; Ridout \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Temporal overlap (Δ) was quantified with three key counterparts in urban areas: the main competitor (red fox: Baltrunaite \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Lanszki et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Takahashi et al. \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2025\u003c/span\u003e), a representative prey (red squirrel: Twining et al. \u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), and humans. We also compared these overlap values with those calculated for the same species in the Monte Rufeno Nature Reserve (Mori et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2025b\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eDistinguishing between pine and stone martens based on external morphological characteristics may be challenging, particularly in nocturnal videos from camera traps. However, following Manzo et al. (\u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2012\u003c/span\u003e) and Raichev (\u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2018\u003c/span\u003e), we identified several key diagnostic traits including the colour of the nose leather (dark in the pine marten, pale in the stone marten) and overall coat appearance. Pine martens generally show a darker, more uniformly brown pelage, whereas stone martens tend to display paler, greyer tones. Differences in body shape were also considered: pine martens typically present a slenderer profile, with proportionally longer tail and limbs and a more pointed muzzle, while stone martens appear stockier, with a relatively shorter tail and broader head. When an individual showed mixed / poorly identifiable traits (in 12% records), the record was conservatively assigned to \u003cem\u003eMartes\u003c/em\u003e sp. and excluded from further analyses.\u003c/p\u003e \u003cp\u003eThe throat patch provided an additional distinguishing feature. In pine martens, it is usually yellow to orange, smaller, irregularly shaped, and often discontinuous; in stone martens, it is bright white, larger, more regularly outlined, and frequently extends toward the forelimbs. When individuals exhibited intermediate or conflicting traits, or when there was disagreement among reviewers, records were conservatively classified as \u003cem\u003eMartes\u003c/em\u003e sp. These records were excluded from temporal activity analyses because of the documented ecological and behavioural differences between the two species (Wereszczuk and Zalewski, \u003cspan citationid=\"CR86\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Torretta et al., \u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e2016\u003c/span\u003e, \u003cspan citationid=\"CR77\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Balestrieri et al., \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Gazzola and Balestrieri, \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe Mardia\u0026ndash;Watson\u0026ndash;Wheeler (MWW) test was performed using the \u003cem\u003ecircular\u003c/em\u003e package in R (Lund et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2017\u003c/span\u003e) to assess whether the overlaps in the natural area differed significantly from those in the urban area (Massara et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). The Δ₁ estimator was always used, as at least one species in each pairwise comparison had fewer than 75 independent detections (Monterroso et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Overlap values were interpreted as low (\u0026lt;\u0026thinsp;50%), moderate (50\u0026ndash;75%), high (\u0026gt;\u0026thinsp;75%), or very high (\u0026gt;\u0026thinsp;90%) (Monterroso et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Confidence intervals (95% CIs) for Δ were obtained from 999 bootstrap replicates with percentile-based limits (Monterroso et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Deviations from a uniform (random) activity distribution over the 24-hour cycle were tested using the Hermans\u0026ndash;Rasson (HR) test (Landler et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Spatial niche overlap between species pairs was evaluated with the Pianka index (Pianka \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e1974\u003c/span\u003e; Mori et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), which ranges from 0 (no overlap) to 1 (complete overlap). The Pianka index was computed through the formula: O\u003csub\u003especies1/ species2\u003c/sub\u003e = (Σ P\u003csub\u003ei species1\u003c/sub\u003e\u0026times; P\u003csub\u003ei species2\u003c/sub\u003e) / \u0026radic;(Σ P\u003csub\u003ei species1\u003c/sub\u003e\u0026sup2; \u0026times; Σ P\u003csub\u003ei species2\u003c/sub\u003e \u0026sup2;), where P\u003csub\u003ei species1\u003c/sub\u003e and P\u003csub\u003ei species2\u003c/sub\u003e are the proportion of records of species 1 and species 2, respectively. The spatial overlap was considered as \u0026lsquo;high\u0026rsquo; when O\u0026thinsp;\u0026gt;\u0026thinsp;0.75 and as \u0026lsquo;very high\u0026rsquo; when O\u0026thinsp;\u0026gt;\u0026thinsp;0.90; conversely it was considered as intermediate with 0.50\u0026thinsp;\u0026lt;\u0026thinsp;O\u0026thinsp;\u0026lt;\u0026thinsp;0.75, and low when O\u0026thinsp;\u0026lt;\u0026thinsp;0.50 (Monterroso et al., \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Given the low seasonal number of detections, we computed spatiotemporal overlaps for a total year.\u003c/p\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003eDo pine marten occur in cities?\u003c/h2\u003e \u003cp\u003eWe retrieved records of martens from 26 Italian cities across the entire national territory, including both mainland and the two largest islands (Sardinia and Sicily: Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e, Table \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e in Supplementary Material 1). Namely, 6 cities featured records of both pine and stone marten, and 4 pine marten only. Amongst the latter, 3 out of 4 are out of the known range of stone marten (i.e., from islands). Thus, the pine marten occurred only in 10 out of 26 sampled cities, for a total of 84 records (Supplementary Material 1).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eThe model selected by our AIC-based procedure, and thus best explaining the observed patterns of occurrence of the pine marten in our set of Italian cities, were the ones featuring city altitude and percent cover of urban wooded areas within the city boundaries, exerting a negative and positive effect upon the probability of occurrence of the pine marten in a given city, respectively (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e; Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Namely, pine marten occurrence exceeded 50% in cities featuring\u0026thinsp;\u0026gt;\u0026thinsp;40% of tree covers within their municipality, and dropped below 30% at altitude values above 600 m a.s.l.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eModel selection on the probability of occurrence of \u003cem\u003eMartes martes\u003c/em\u003e in urban areas of Italy (N\u0026thinsp;=\u0026thinsp;26) as explained by city altitude, percent wooded cover within the city boundaries, city size, and co-occurrence of \u003cem\u003eM. foina\u003c/em\u003e.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"6\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\"\u0026plusmn;\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eModel rank\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAltitude\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003ePercent wooded cover\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003ePresence of \u003cem\u003eMartes foina\u003c/em\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eCity size\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eAIC\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e0.025\u0026thinsp;\u0026plusmn;\u0026thinsp;0.016 *\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eNA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eNA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e32.79\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-0.002\u0026thinsp;\u0026plusmn;\u0026thinsp;0.001\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e0.025\u0026thinsp;\u0026plusmn;\u0026thinsp;0.016 *\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eNA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eNA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e33.75\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-0.002\u0026thinsp;\u0026plusmn;\u0026thinsp;0.001\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e0.025\u0026thinsp;\u0026plusmn;\u0026thinsp;0.016 *\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-1.793\u0026thinsp;\u0026plusmn;\u0026thinsp;1.086\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eNA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e36.37\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eDo urban martens change their behaviour in urban areas?\u003c/h3\u003e\n\u003cp\u003eWe recorded a total of 192 independent detections of the target species in RNI: 46 pine martens, 79 red foxes, 33 red squirrels and 34 humans. These data were compared to those collected by Mori et al. (\u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2025b\u003c/span\u003e) in the Monte Rufeno Reserve: 195 independent records of pine marten, 813 of red fox, 85 of red squirrels and 424 humans. No stone marten was detected in the study areas. In the urban area, the pine marten activity showed significant peaks at dawn and dusk, red fox activity peaked in the late afternoon hours, red squirrel activity in the early morning and afternoon, and human activity in mid-morning and mid-afternoon (Hermans\u0026ndash;Rasson tests; r\u0026thinsp;=\u0026thinsp;62.48\u0026ndash;81.33, all p\u0026thinsp;\u0026lt;\u0026thinsp;0.001). Temporal overlap between urban and natural areas was low to moderate for all species, except for the red fox, which showed a high degree of overlap in activity peaks between urban and natural areas (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). We found significant differences in the temporal overlaps of pine martens with red squirrels, red foxes, and humans between urban and natural areas (Mann\u0026ndash;Whitney\u0026ndash;Wilcoxon test, W\u0026thinsp;=\u0026thinsp;143.29\u0026ndash;181.03, all p\u0026thinsp;\u0026lt;\u0026thinsp;0.001). Specifically, in urban environments, pine martens exhibited increased temporal overlap with red squirrels and with humans, whereas their temporal overlap with red foxes remarkably declined (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eSpatial overlaps (Pianka index, O) between pine martens and red squirrels and between pine martens and humans were high in both urban (respectively 0.84 and 0.77) and natural (respectively 0.75 and 0.78) areas. Conversely, spatial overlap between pine martens and red foxes was low in both urban (O\u0026thinsp;=\u0026thinsp;0.38) and natural (O\u0026thinsp;=\u0026thinsp;0.27) areas.\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eIn this work, we provided the first evidence of pine martens occurring in urban environments across Italy, likely reflecting a rise in urban detections, which might in turn indicate an ongoing process of urban colonisation by a species historically regarded as a forest specialist, rather than an actual increase in population size (but see Robinson et al. \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Museum records also indicate the long-standing presence of the stone marten in urban areas since at least the 19th century, whereas confirmed records of the pine marten date only to recent decades and remain relatively sporadic (Ancillotto et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2024\u003c/span\u003e, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). The compilation of verified urban records across multiple Italian cities, combined with detailed camera-trap data from Rome in comparison to neighboring natural areas, may suggest that the pine marten is exhibiting both spatial and temporal behavioural plasticity, possibly allowing it to persist in human-dominated landscapes (Manzo et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). This result challenges long-standing assumptions about the species\u0026rsquo; strict dependence on mature forests (Clevenger \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e1993\u003c/span\u003e; Lombardini et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2015\u003c/span\u003e) and is in line with recent findings of range expansion into agricultural and peri-urban areas (Balestrieri et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Grelli et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Manzo et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2018\u003c/span\u003e, \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), as also confirmed by increasing road mortality events (Supplementary Material 1).\u003c/p\u003e \u003cp\u003eIndeed, considering that many urban records of the European pine marten are associated with road kills (see Supplementary Material 1), this pattern may suggest that remnant woodland patches within cities function as ecological traps for the species (see Ancillotto et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). In other words, these forest remnants may attract individuals attempting to colonize urban areas by following existing habitat connectivity. Once inside the urban matrix, the pine marten may be less well adapted than the stone marten to cope with anthropogenic pressures. Consequently, although environmental corridors may facilitate urban entry, survival prospects within the city appear comparatively low, potentially leading to increased mortality rather than successful establishment. However, the pine marten has undergone a remarkable range expansion in Italy over the last two decades, now occupying lowland areas where it was once considered absent (Balestrieri et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2010\u003c/span\u003e, \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Manzo et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). This expansion has been partly revealed through citizen-science platforms, wildlife rescue center records, and increasing public awareness of wildlife conservation (Grelli et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Manzo et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Reports of pine martens in peri-urban areas, where the species was previously unrecorded, are becoming increasingly frequent, whereas earlier records from similar contexts referred exclusively to stone martens. This pattern may indicate an ongoing process of urban colonization, or, in general, an increase in use of human-modified habitats (Grelli et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2014\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe presence of pine martens within several regional and provincial capital cities, verified through citizen science, Wildlife Rescue Centers and social media data, reflects a broader ecological trend in which formerly habitat-specialist mammals are colonising novel environments, possibly as a response to landscape modification and climatic changes (Balestrieri et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Santini et al. \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Ancillotto et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2024\u003c/span\u003e, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2025\u003c/span\u003e; Manzo et al. \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). The RNI in Rome, although embedded within a densely populated area, still hosts structural elements, e.g. continuous woodland patches and abundant tree cover (Todini and Crosti \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), which provide all habitat and resource requirements by the species (Manzo et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). These findings suggest that green corridors and urban parks can function as \u0026ldquo;ecological stepping stones\u0026rdquo; facilitating movement and settlement of forest mammals in metropolitan matrices (Monteiro and Lira \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Furthermore, the presence of pine marten in capital cities could be a response not only to landscape modifications and climate change, but also to behavioural changes both in martens and humans, driving to a higher reciprocal tolerance (Mori et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2025c\u003c/span\u003e). The importance of profitable patches of natural habitats for pine martens within anthropogenic landscapes e.g., natural woodlands embedded in urban parks, was also revealed by our modeling exercise, highlighting that cities with low values of wooded cover are less likely to host the species. Additionally, the negative effect of altitude upon the probability of presence of the pine marten is actually inconsistent with previous work from the alpine region, and Bulgaria, where the species seems to favour higher elevations (Doykin et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Fonda et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Nonetheless, this suggests that the pine marten is actually not a specialist of higher-altitude forests, yet that this latter habitat type is associated to areas where the species occurred after it past decline (Sainsbury et al. \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2019\u003c/span\u003e), since low-altitude forests were those historically more impacted (Widmer et al. \u003cspan citationid=\"CR88\" class=\"CitationRef\"\u003e2025\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe camera-trapping results also provide a more species-based perspective on whether and how pine martens may fine-tune their temporal behaviour to the peculiar challenges posed by urban environments, as already shown in more natural contexts similarly affected by human disturbance (Mirante et al. \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Although three camera traps may seem limited, they effectively cover the study area (RNI), the only site in Rome where the pine marten has been regularly recorded (Ancillotto et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Therefore, we are confident that increasing the number of camera traps would not substantially alter our results.\u003c/p\u003e \u003cp\u003eThe comparison of activity rhythms between urban and natural parks in fact revealed some changes in temporal activity patterns by pine martens in response to the urban context. While in natural areas the species is predominantly crepuscular with occasional diurnal activity (Zielinski et al. \u003cspan citationid=\"CR90\" class=\"CitationRef\"\u003e1983\u003c/span\u003e; Torretta et al. \u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), in Rome, it exhibited bimodal peaks at dawn and dusk with moderate overlap with human activity. This partial diurnality may represent a trade-off between avoiding nocturnal competitors (particularly the red fox, as also evident by our results) and exploiting diurnal prey, e.g., red squirrels and birds. However, given that both squirrels and birds may be more vulnerable at nights while resting in nests or roosts (particularly for a species with remarkable arboreal skills as the pine marten), competition avoidance appears the most convincing hypothesis. The reduction in temporal overlap with red foxes observed in urban conditions supports this interpretation and may reflect active temporal niche partitioning to reduce intraguild predation risk or competition for resources (Lindstr\u0026ouml;m et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e1995\u003c/span\u003e). Avoidance of foxes in urban contexts may also be due to the high densities of red foxes in anthropogenic environments in comparison to more natural ones, a factor that may induce behavioural adjustments in pine martens, as those we observed (Padovani et al. \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Such plasticity has already been reported in other mesocarnivores able to thrive in anthropogenic environments, where shifts in activity rhythms are crucial for coexistence and access to resources (Massara et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Mori et al. \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2025a\u003c/span\u003e).\u003c/p\u003e \u003cp\u003ePine martens in the urban park showed increased temporal and spatial overlap with humans relative to those in natural habitats. While this might initially suggest habituation to human presence, it could also be a by-product of shared use of the same temporal windows associated with reduced fox activity and higher prey availability. Similar behavioural tolerance to anthropogenic disturbance has been observed in stone martens, a close congener widely recognized as synanthropic (Herr et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Capon et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). The absence of the stone marten in our study area might have facilitated the pine marten colonisation of this part of Rome, reducing direct competition and allowing niche expansion. However, also the opposite scenario is possible, i.e. the local disappearance of the stone marten is due to the local arrival of the pine marten. Future studies should test whether pine martens can persist in areas where stone martens are well established, or whether interspecific exclusion mechanisms operate between the two congeners (Wereszczuk and Zalewski \u003cspan citationid=\"CR86\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Gazzola and Balestrieri \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe high spatial overlap between pine martens and red squirrels in both urban and natural settings further emphasizes their potential trophic and ecological link. Urban parks hosting stable red squirrel populations likely represent valuable foraging grounds for pine martens, as squirrels constitute a potential prey item (Twining et al. \u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). However, the increased proximity to humans and domestic animals may on one side limit competition with red foxes and on the other side expose them to elevated risks of road mortality, disturbance, or disease transmission, echoing challenges faced by other urban carnivores (Soulsbury and White \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Ritzel and Gallo \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Mori et al. \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2022b\u003c/span\u003e). Whether pine martens can sustain long-term viable populations in urban settings will depend on their ability to mitigate these pressures and maintain access to suitable foraging and breeding sites. For instance, in future studies, throat patterns could potentially be used to identify individual martens (Raichev \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). However, our camera-trap images often captured animals in lateral view, making even species-level identification challenging. Nevertheless, the ability to recognise whether the same pine marten individuals persist in a given area would provide valuable insights into their degree of site fidelity, adaptation to urban environments, and coexistence dynamics with the red fox.\u003c/p\u003e \u003cp\u003eFrom a conservation perspective, the colonisation of urban parks by pine martens may have positive ecological implications (Caryl et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). As an opportunistic predator, the species could contribute to controlling rodent populations (including invasive rats \u003cem\u003eRattus rattus\u003c/em\u003e and \u003cem\u003eR. norvegicus\u003c/em\u003e, and grey squirrels \u003cem\u003eSciurus carolinensis\u003c/em\u003e: McNicol et al. \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), and regulating other urban mesofauna, potentially restoring some trophic balance within urban ecosystems (Caryl et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). Moreover, its presence in cities may enhance public awareness of biodiversity conservation, triggering citizen engagement through platforms such as iNaturalist or Ornitho.it, which proved invaluable for assessing its distribution. Nonetheless, increased visibility of carnivores in human environments can also lead to conflicts, particularly if perceived as threats to pets or property (Soulsbury and White \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Clear communication and coexistence strategies will therefore be essential to ensure tolerance and minimize negative interactions.\u003c/p\u003e \u003cp\u003eThe ongoing expansion of the pine marten into lowland and urbanised landscapes may also have implications for monitoring and management policies, especially since the species is listed under the Annex IV of the Habitats Directive, thus being considered as conservation-relevant and in need of monitoring for assessing its conservation status across the EU. Traditional conservation frameworks which consider the pine marten exclusively as a forest indicator species may need revision to reflect its emerging ecological plasticity (Vergara et al. \u003cspan citationid=\"CR81\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Urban green spaces, when managed with connectivity and vegetation diversity in mind, could serve as refugia supporting the persistence of native mammals such as rodents (Dondina et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2025\u003c/span\u003e) and, in turn, carnivores (Ancillotto et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). In summary, our findings highlight that the pine marten, once considered an emblematic species of pristine forest ecosystems, is exhibiting unexpected behavioural and ecological flexibility, allowing it to exploit urban habitats without fully altering its natural activity patterns. This adaptive response summarises the dynamic interactions shaping mammalian assemblages in cities, where human presence, resource distribution, and interspecific competition drive rapid ecological change (Mori et al. \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2025a\u003c/span\u003e). Continued monitoring will be essential to determine whether these early colonisation patterns lead to stable, reproducing urban populations or represent transient incursions facilitated by peri-urban connectivity.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e \u003ch2\u003eAuthor Contribution Statement\u003c/h2\u003e \u003cp\u003eLA and EM conceived this work, conducted the analyses and wrote the first draft; AA, DG and FV collected data and participated in writing up all drafts. All authors approved the MS before the submission.\u003c/p\u003e \u003ch2\u003eConflict of interest\u003c/h2\u003e \u003cp\u003ethe authors have no conflict of interest to declare.\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eEthics approval\u003c/strong\u003e \u003cp\u003eNot applicable.\u003c/p\u003e \u003c/p\u003e\u003ch2\u003eFunding\u003c/h2\u003e \u003cp\u003eNo funding was received for this research.\u003c/p\u003e\u003ch2\u003eAcknowledgements\u003c/h2\u003e \u003cp\u003eLA and EM were funded by the National Recovery and Resilience Plan (NRRP), Mission 4 Component 2 Investment 1.4 - Call for tender No. 3138 of 16 December 2021, rectified by Decree n.3175 of 18 December 2021 of Italian Ministry of University and Research funded by the European Union \u0026ndash; NextGenerationEU; Project code CN_00000033, Concession Decree No. 1034 of 17 June 2022 adopted by the Italian Ministry of University and Research, CUP B83C22002930006, Project title \u0026ldquo;National Biodiversity Future Center - NBFC\u0026rdquo;. Isabella Vitali and Cristina Mascalchi (CNR IRET Sesto Fiorentino) also provided valuable technical, administrative and management support. Camera-trapping in RNI was implemented as part of the Activity 5.2 of the Spoke 5 of the PNRR National Biodiversity Future Center project (see Mori et al., 2025a; Viviano et al., 2025; see Acknowledgements). The \u0026ldquo;U.O. Manutenzione e Valorizzazione del Verde Urbano\u0026rdquo; office of the Municipality of Roma provided us with permits to set camera traps within urban parks.\u003c/p\u003e\u003ch2\u003eData availability statement\u003c/h2\u003e \u003cp\u003eData showed in this paper are uploaded as Supplementary Material 1.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAncillotto L, Tomassini A, Russo D (2016) The fancy city life: Kuhl\u0026rsquo;s pipistrelle, \u003cem\u003ePipistrellus kuhlii\u003c/em\u003e, benefits from urbanisation. Wildl Res 42(7):598\u0026ndash;606. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1071/WR15003\u003c/span\u003e\u003cspan address=\"10.1071/WR15003\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eAncillotto L, Amori G, Capizzi D, Cignini B, Zapparoli M, Mori E (2024) No city for wetland species: habitat associations affect mammal persistence in urban areas. Proc R Soc B 291:20240079. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1098/rspb.2024.0079\u003c/span\u003e\u003cspan address=\"10.1098/rspb.2024.0079\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eAncillotto L, Guerri G, Agnelli P, Bonora L, Maggioni M, Morabito M, Mori E (2025) Past present: extinction debt of forest mammals from urban areas. Biol Conserv 306:111143. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.biocon.2025.111143\u003c/span\u003e\u003cspan address=\"10.1016/j.biocon.2025.111143\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBalestrieri A, Remonti L, Ruiz-Gonz\u0026aacute;lez A, G\u0026oacute;mez-Moliner BJ, Vergara M, Prigioni C (2010) Range expansion of the pine marten (\u003cem\u003eMartes martes\u003c/em\u003e) in an agricultural landscape matrix (NW Italy). Mammal Biol 75:412\u0026ndash;419. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.mambio.2009.05.003\u003c/span\u003e\u003cspan address=\"10.1016/j.mambio.2009.05.003\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBalestrieri A, Remonti L, Ruiz-Gonzalez R, Zenato M, Gazzola A, Vergara M, Dettori EE, Saino N, Capelli E, Gomez-Moliner BJ, Guidali F, Prigioni C (2015) Distribution and habitat use by pine marten \u003cem\u003eMartes martes\u003c/em\u003e in a riparian corridor crossing intensively cultivated lowlands. Ecol Res 30:153\u0026ndash;162. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s11284-014-1220-8\u003c/span\u003e\u003cspan address=\"10.1007/s11284-014-1220-8\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBalestrieri A, Mori E, Menchetti M, Ruiz-Gonz\u0026aacute;lez A, Milanesi P (2019) Far from the madding crowd: Tolerance toward human disturbance shapes distribution and connectivity patterns of closely related \u003cem\u003eMartes\u003c/em\u003e spp. Popul Ecol 61:289\u0026ndash;299. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1002/1438-390X.12001\u003c/span\u003e\u003cspan address=\"10.1002/1438-390X.12001\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBalestrieri R, Vento R, Viviano A, Mori E, Gili C, Monti F (2023) Razorbills \u003cem\u003eAlca torda\u003c/em\u003e in Italian seas: a massive irruption of historical relevance and role of social network monitoring. Animals 13(4):656. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3390/ani13040656\u003c/span\u003e\u003cspan address=\"10.3390/ani13040656\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBaltrunaite L (2001) Feeding habits, food niche overlap of red fox (\u003cem\u003eVulpes vulpes\u003c/em\u003e L.) and pine marten (\u003cem\u003eMartes martes\u003c/em\u003e L.) in hilly moraine highland. Lith Ecologija 2:27\u0026ndash;31\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBateman PW, Fleming PA (2012) Big city life: carnivores in urban environments. J Zool 287:1\u0026ndash;23. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/j.1469-7998.2011.00887.x\u003c/span\u003e\u003cspan address=\"10.1111/j.1469-7998.2011.00887.x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBirnie-Gauvin K, Peiman KS, Gallagher AJ, De Bruijn R, Cooke SJ (2016) Sublethal consequences of urban life for wild vertebrates. Environ Rev 24:416\u0026ndash;425. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1139/er-2016-0029\u003c/span\u003e\u003cspan address=\"10.1139/er-2016-0029\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBurnham KP, Anderson DR (2002) Model selection and multimodel inference: a practical information-theoretic approach, 2nd edn. Springer Editions, Berlin Heidelberg\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eCapon M, Lysaniuk B, Godard V, Clauzel C, Simon L (2021) Characterizing the landscape compositions of urban wildlife encounters: the case of the stone marten (\u003cem\u003eMartes foina\u003c/em\u003e), the red fox (\u003cem\u003eVulpes vulpes\u003c/em\u003e) and the hedgehog (\u003cem\u003eErinaceus europaeus\u003c/em\u003e) in the Greater Paris area. Urb Ecosyst 24:885\u0026ndash;903. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s11252-020-01071-6\u003c/span\u003e\u003cspan address=\"10.1007/s11252-020-01071-6\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eCarnevali L, Lovari S, Monaco A, Mori E (2016) Nocturnal activity of a diurnal species, the northern chamois, in a predator-free Alpine area. Behav Proc 126:101\u0026ndash;107. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.beproc.2016.03.013\u003c/span\u003e\u003cspan address=\"10.1016/j.beproc.2016.03.013\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eCaryl FM, Quine CP, Park KJ (2012) Martens in the matrix: the importance of nonforested habitats for forest carnivores in fragmented landscapes. J Mammal 93(2):464\u0026ndash;474. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1644/11-MAMM-A-149.1\u003c/span\u003e\u003cspan address=\"10.1644/11-MAMM-A-149.1\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eClevenger AP (1993) The European pine marten \u003cem\u003eMartes martes\u003c/em\u003e in the Balearic Islands, Spain. Mammal Rev 23:65\u0026ndash;72. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/j.1365-2907.1993.tb00417.x\u003c/span\u003e\u003cspan address=\"10.1111/j.1365-2907.1993.tb00417.x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eCox K, Gouwy J, Mergeay J, Neyrinck S, Van Den Berge K (2025) Rapid Colonisation of Synanthropic Stone Martens in a Highly Urbanised Region: Insights From Temporal and Spatial Analysis. Ecol Evol 15:e71392. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/j.1365-2907.1993.tb00417.x\u003c/span\u003e\u003cspan address=\"10.1111/j.1365-2907.1993.tb00417.x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eDavison J, Huck M, Delahay RJ, Roper TJ (2008) Urban badger setts: characteristics, patterns of use and management implications. J Zool 275(2):190\u0026ndash;200. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/j.1469-7998.2008.00424.x\u003c/span\u003e\u003cspan address=\"10.1111/j.1469-7998.2008.00424.x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eDickman CR (1987) Habitat fragmentation and vertebrate species richness in an urban environment. J Appl Ecol 24:337\u0026ndash;351. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.2307/2403879\u003c/span\u003e\u003cspan address=\"10.2307/2403879\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eDondina O, Tirozzi P, Viviano A, Mori E, Orioli V, Tommasi N, Tanzi A, Bazzoli A, Caprio E, Patetta C, Pastore MC, Bani L, Ancillotto L (2025) Spatial and habitat determinants of small-mammal biodiversity in urban green areas: Lessons for nature-based solutions. Urb Urb Green 104:128641\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eDoykin N, Popova E, Paraskova M, Zlatanov V, Zlatanova D (2017) Pine marten (\u003cem\u003eMartes martes\u003c/em\u003e L.): distribution, habitat preference and activity in Vitosha Nature Park, Bulgaria. Proceedings of Seminar Of Ecology\u0026ndash;2016 with International Participation 2016: 26\u0026ndash;33\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eDuduś L, Zalewski A, Kozioł O, Jakubiec Z, Kr\u0026oacute;l N (2014) Habitat selection by two predators in an urban area: the stone marten and red fox in Wrocław (SW Poland). Mamm Biol 79(1):71\u0026ndash;76. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.mambio.2013.08.001\u003c/span\u003e\u003cspan address=\"10.1016/j.mambio.2013.08.001\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eFonda F, Chiatante G, Meriggi A, Mustoni A, Armanini M, Mosini A, Spada A, Lombardini M, Righetti D, Granata M, Capelli E, Pontarini R, Roux Poignant G, Balestrieri A (2021) Spatial distribution of the pine marten (\u003cem\u003eMartes martes\u003c/em\u003e) and stone marten (\u003cem\u003eMartes foina\u003c/em\u003e) in the Italian Alps. Mammal Biol 101:345\u0026ndash;356. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s42991-020-00098-8\u003c/span\u003e\u003cspan address=\"10.1007/s42991-020-00098-8\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eGallo T, Fidino M, Lehrer EW, Magle SB (2017) Mammal diversity and metacommunity dynamics in urban green spaces: implications for urban wildlife conservation. Ecol Appl 27(8):2330\u0026ndash;2341. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1002/eap.1611\u003c/span\u003e\u003cspan address=\"10.1002/eap.1611\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eGazzola A, Balestrieri A (2020) Nutritional ecology provides insights into competitive interactions between closely related \u003cem\u003eMartes\u003c/em\u003e species. Mamm Rev 50:82\u0026ndash;90. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/mam.12177\u003c/span\u003e\u003cspan address=\"10.1111/mam.12177\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eGeiger M, Taucher AL, Gloor S, Hegglin D, Bontadina F (2018) In the footsteps of city foxes: evidence for a rise of urban badger populations in Switzerland. Hystrix 29(2):236\u0026ndash;238. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.4404/hystrix-00069-2018\u003c/span\u003e\u003cspan address=\"10.4404/hystrix-00069-2018\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eGerber BD, Devarajan K, Farris ZJ, Fidino M (2024) A model-based hypothesis framework to define and estimate the diel niche via the Diel.Niche R package. J Anim Ecol 93:132\u0026ndash;146. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/1365-2656.14035\u003c/span\u003e\u003cspan address=\"10.1111/1365-2656.14035\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eGottero E, Larcher F, Cassatella C (2023) Defining and regulating peri-urban areas through a landscape planning approach: The case study of Turin Metropolitan Area (Italy). Land 12:217. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3390/land12010217\u003c/span\u003e\u003cspan address=\"10.3390/land12010217\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eGrelli D, Paoloni D, Vercillo F, Ragni B (2014) Un racconto dalla citt\u0026agrave;: la presenza della martora a Perugia (Italia centrale). Hystrix 25 suppl.: 108\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eHerr J, Schley L, Roper TJ (2009) Socio-spatial organization of urban stone martens. J Zool 277:54\u0026ndash;62. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://10.1111/j.1469-7998.2008.00510.x\u003c/span\u003e\u003cspan address=\"https://10.1111/j.1469-7998.2008.00510.x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eHughes-Brandl M (2018) Package \u0026lsquo;NightDay\u0026rsquo;. Night and day boundary plot function. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://cran.r-project.org/web/packages/NightDay/NightDay.pdf\u003c/span\u003e\u003cspan address=\"https://cran.r-project.org/web/packages/NightDay/NightDay.pdf\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e. Accessed on 07.10.2025\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLandler L, Ruxton GD, Malkemper EP (2019) The Hermans-Rasson test as a powerful alternative to the Rayleigh test for circular statistics in biology. BMC Ecol 19:1\u0026ndash;8. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1186/s12898-019-0246-8\u003c/span\u003e\u003cspan address=\"10.1186/s12898-019-0246-8\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLanszki J, Zalewski A, Horv\u0026aacute;th G (2007) Comparison of red fox \u003cem\u003eVulpes vulpes\u003c/em\u003e and pine marten \u003cem\u003eMartes martes\u003c/em\u003e food habits in a deciduous forest in Hungary. Wildl Biol 13:258\u0026ndash;271. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.2981/0909-6396\u003c/span\u003e\u003cspan address=\"10.2981/0909-6396\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e(2007)13[258:CORFVV]2.0.CO;2\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLindstr\u0026ouml;m ER, Brainerd SM, Helldin JO, Overskaug K (1995) Pine marten-red fox interactions: a case of intraguild predation? Ann Zool Fennici 32:123\u0026ndash;130\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLombardini M, Cinerari CE, Murru M, Vidus Rosin A, Mazzoleni L, Meriggi A (2015) Habitat requirements of Eurasian pine marten \u003cem\u003eMartes martes\u003c/em\u003e in a Mediterranean environment. Mamm Res 60(2):97\u0026ndash;105. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s13364-014-0211-z\u003c/span\u003e\u003cspan address=\"10.1007/s13364-014-0211-z\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eŁopucki R, Klich D, Kitowski I (2019) Are small carnivores urban avoiders or adapters: Can they be used as indicators of well-planned green areas? Ecol Indic 101:1026\u0026ndash;1031. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.ecolind.2019.02.016\u003c/span\u003e\u003cspan address=\"10.1016/j.ecolind.2019.02.016\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLovari S, Corsini MT, Guazzini B, Romeo G, Mori E (2017) Suburban ecology of the crested porcupine in a heavily poached area: a global approach. Eur J Wildl Res 63:10. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s10344-016-1075-0\u003c/span\u003e\u003cspan address=\"10.1007/s10344-016-1075-0\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLund U, Agostinelli C, Arai H, Gagliardi A, Portugues EG, Giunchi D, Irisson JO, Pocernich M, Rotolo F (2017) Package circular. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://mirrors.ucr.ac.cr/CRAN/web/packages/circular/circular.pdf\u003c/span\u003e\u003cspan address=\"http://mirrors.ucr.ac.cr/CRAN/web/packages/circular/circular.pdf\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e. Accessed on 03.10.2025\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMassara RL, Paschoal AMO, Bailey LL, Doherty PF, Barreto MF, Chiarello AG (2018) Effect of humans and pumas on the temporal activity of ocelots in protected areas of Atlantic Forest. Mamm Biol 92:86\u0026ndash;93. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.mambi o.2018.04.009\u003c/span\u003e\u003cspan address=\"10.1016/j.mambi o.2018.04.009\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eManzo E, Bartolommei P, Rowcliffe JM, Cozzolino R (2012) Estimation of population density of European pine marten in central Italy using camera trapping. Acta Theriol 57:165\u0026ndash;172. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.4404/hystrix-23.2-7099\u003c/span\u003e\u003cspan address=\"10.4404/hystrix-23.2-7099\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eManzo E, Bartolommei P, Giuliani A, Gentile G, Dess\u0026igrave;-Fulgheri F, Cozzolino R (2018) Habitat selection of European pine marten in Central Italy: from a tree dependent to a generalist species. Mamm Res 63:357\u0026ndash;367. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s13364-018-0374-0\u003c/span\u003e\u003cspan address=\"10.1007/s13364-018-0374-0\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eManzo E, Iannarilli F, Dell'Agnello F, Bartolommei P, Bonacchi A, Gasperini S, Cozzolino R (2024) Assessing the co-occurrence of European pine marten (\u003cem\u003eMartes martes\u003c/em\u003e) with humans and domestic cats on a Mediterranean island. Ecol Evol 14(12):e70651. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1002/ece3.70651\u003c/span\u003e\u003cspan address=\"10.1002/ece3.70651\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eManzo E, Vercillo F, Fedi C (2025) Martora \u003cem\u003eMartes martes\u003c/em\u003e. In: Loy A, Bon M, Di Febbraro M, Baisero D, Amori G (eds) Atlas of mammals in Italy. Edizioni Belvedere, Latina, Italy, pp 234\u0026ndash;237\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMcNicol CM, Bavin D, Bearhop S, Ferryman M, Gill R, Goodwin CE, MacPherson J, Silk MJ, McDonald RA (2020) Translocated native pine martens \u003cem\u003eMartes martes\u003c/em\u003e alter short-term space use by invasive non‐native grey squirrels \u003cem\u003eSciurus carolinensis\u003c/em\u003e. J Appl Ecol 57:903\u0026ndash;913\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMeredith M, Ridout M (2014) Overview of the overlap package. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://cran.cs.wwu.edu/web/packages/overlap/vignettes/overlap.pdf\u003c/span\u003e\u003cspan address=\"http://cran.cs.wwu.edu/web/packages/overlap/vignettes/overlap.pdf\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e Accessed on 07.10.2025\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMiller KF, Wilson DJ, Hartley S, Innes JG, Fitzgerald NB, Miller P, van Heezik Y (2022) Invasive urban mammalian predators: distribution and multi-scale habitat selection. Biology 11(10):1527. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3390/biology11101527\u003c/span\u003e\u003cspan address=\"10.3390/biology11101527\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMirante D, Ancillotto L, Zampetti A, Coiro G, Pisa G, Santocchi C, Giuliani M, Santini L (2024) Fine-tuning coexistence: Wildlife's short-term responses to dynamic human disturbance patterns. Glob Ecol Conserv 54:e03053. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.gecco.2024.e03053\u003c/span\u003e\u003cspan address=\"10.1016/j.gecco.2024.e03053\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMonteiro MCM, Lira PK (2020) Metropolitan mammals: understanding the threats inside an urban protected area. Oecol Australis 24:661\u0026ndash;675. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.4257/oeco.2020.2403.10\u003c/span\u003e\u003cspan address=\"10.4257/oeco.2020.2403.10\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMonterroso P, Alves PC, Ferreras P (2014) Plasticity in circadian activity patterns of mesocarnivores in southwestern Europe: Implications for species coexistence. Behav Ecol Sociobiol 68:1403\u0026ndash;1417. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s00265-014-1748-1\u003c/span\u003e\u003cspan address=\"10.1007/s00265-014-1748-1\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMori E, Menchetti M, Camporesi A, Cavigioli L, Tabarelli de Fatis K, Girardello M (2019) License to kill? Domestic cats affect a wide range of native fauna in a highly biodiverse Mediterranean country. Front Ecol Evol 7:477. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3389/fevo.2019.00477\u003c/span\u003e\u003cspan address=\"10.3389/fevo.2019.00477\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMori E, Bagnato S, Serroni P, Sangiuliano A, Rotondaro F, Marchian\u0026ograve; V, Cascini V, Poerio L, Ferretti F (2020) Spatiotemporal mechanisms of coexistence in an European mammal community in a protected area of southern Italy. J Zool 310:232\u0026ndash;245. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/jzo.12743\u003c/span\u003e\u003cspan address=\"10.1111/jzo.12743\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMori E, Molteni R, Ancillotto L, Ficetola GF, Falaschi M (2022a) Spatial ecology of crested porcupine in a metropolitan landscape. Urb Ecosyst 25(6):1797\u0026ndash;1803. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s11252-022-01264-1\u003c/span\u003e\u003cspan address=\"10.1007/s11252-022-01264-1\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMori E, Fedele E, Greco I, Giampaoli Rustichelli M, Massolo A, Miniati S, Puppo F, Santini G, Zaccaroni M (2022b) Spatiotemporal activity of the pine marten \u003cem\u003eMartes martes\u003c/em\u003e: Insights from an island population. Ecol Res 37:102\u0026ndash;114. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/1440-1703.12269\u003c/span\u003e\u003cspan address=\"10.1111/1440-1703.12269\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMori E, Lazzeri L, Maggioni M, Viviano A, Guerri G, Morabito M, Martini S, Dondina O, Sogliani D, Scarf\u0026ograve; M, Ancillotto L (2025a) Camera-traps and the city: spatiotemporal adaptations of wildlife to urban environments. Ecol Sol Evid 6:e70115. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1002/2688-8319.70115\u003c/span\u003e\u003cspan address=\"10.1002/2688-8319.70115\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMori E, Palombi A, Blasi MC, Viviano A, Piazzai M, Colonnelli L, Lazzeri L, Spilinga C, Montioni F, Bombara G, Sogliani D, Ancillotto L (2025b) Spatiotemporal partitioning between sympatric hares and their predators. Eur J Wildl Res 71:127. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s10344-025-01979-4\u003c/span\u003e\u003cspan address=\"10.1007/s10344-025-01979-4\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMori E, Marchi V, Dondina O, Viviano A, Di Bari P, Balestrieri R, Corradetti M, Ancillotto L (2025c) Citizen eyes on elusive wildlife: assessing public appreciation for urban wild mammals. Ambio \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s13280-025-02315-5\u003c/span\u003e\u003cspan address=\"10.1007/s13280-025-02315-5\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eO\u0026rsquo;Connell AF, Nichols JD, Karanth KU (2011) Camera traps in animal ecology: methods and analyses, camera traps in animal ecology: methods and analyses. Springer Science \u0026amp; Business Media, Tokyo, Japan\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ePadovani R, Shi Z, Harris S (2021) Are British urban foxes (\u003cem\u003eVulpes vulpes\u003c/em\u003e) bold? The importance of understanding human\u0026ndash;wildlife interactions in urban areas. Ecol Evol 11:835\u0026ndash;851. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1002/ece3.7087\u003c/span\u003e\u003cspan address=\"10.1002/ece3.7087\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ePereboom V, Mergey M, Villerette N, Helder R, Gerard JF, Lode T (2008) Movement patterns, habitat selection, and corridor use of a typical woodland-dweller species, the European pine marten (\u003cem\u003eMartes martes\u003c/em\u003e), in fragmented landscape. Can J Zool 86(9):983\u0026ndash;991. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1139/Z08-076\u003c/span\u003e\u003cspan address=\"10.1139/Z08-076\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ePianka ER (1974) Niche overlap and diffuse competition. Proc Nat Ac Sci USA 71:2141\u0026ndash;2145. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1073/pnas.71.5.2141\u003c/span\u003e\u003cspan address=\"10.1073/pnas.71.5.2141\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eQGIS Development Team (2019) QGIS Geographic Information System. Open Source Geospatial Foundation Project. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://qgis.osgeo.org\u003c/span\u003e\u003cspan address=\"http://qgis.osgeo.org\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e. Accessed on 06.10.2025\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eR Core Team (2021) R: A language and environment for statistical computing. R Foundation for Statistical Computing. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://www.R-project.org\u003c/span\u003e\u003cspan address=\"https://www.R-project.org\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e. Accessed on 07.10.2025\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eRaichev E (2018) Determination of stone marten (\u003cem\u003eMartes foina\u003c/em\u003e) and pine marten (\u003cem\u003eMartes martes\u003c/em\u003e) in natural habitats using camera traps. Agricult Sci Technol 10:160\u0026ndash;163\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eRipley B, Venables B, Bates DM, Hornik K, Gebhardt A, Firth D, Ripley MB (2013) Package \u0026lsquo;mass\u0026rsquo;. Cran R 538:822\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eRitzel K, Gallo T (2020) Behavior change in urban mammals: a systematic review. Front Ecol Evol 8:576665. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3389/fevo.2020.576665\u003c/span\u003e\u003cspan address=\"10.3389/fevo.2020.576665\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eRobinson OJ, Ruiz-Gutierrez V, Fink D (2017) Correcting for bias in distribution modelling for rare species using citizen science data. Divers Distrib 24:460\u0026ndash;472. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/ddi.12698\u003c/span\u003e\u003cspan address=\"10.1111/ddi.12698\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eRusso D, Ancillotto L (2015) Sensitivity of bats to urbanization: a review. Mamm Biol 80(3):205\u0026ndash;212. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3389/fevo.2020.576665\u003c/span\u003e\u003cspan address=\"10.3389/fevo.2020.576665\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSainsbury KA, Shore RF, Schofield H, Croose E, Campbell RD, Mcdonald RA (2019) Recent history, current status, conservation and management of native mammalian carnivore species in Great Britain. Mamm Rev 49:171\u0026ndash;188. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/mam.12150\u003c/span\u003e\u003cspan address=\"10.1111/mam.12150\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSanticchia F, Tranquillo C, Wauters LA, Palme R, Panzeri M, Preatoni D, Bisi F, Martinoli A (2024) Physiological stress response to urbanisation differs between native and invasive squirrel species. Sci Tot Environ 992:171336. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.scitotenv.2024.171336\u003c/span\u003e\u003cspan address=\"10.1016/j.scitotenv.2024.171336\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSantini L, Gonz\u0026aacute;lez-Su\u0026aacute;rez M, Russo D, Gonzalez‐Voyer A, von Hardenberg A, Ancillotto L (2019) One strategy does not fit all: determinants of urban adaptation in mammals. Ecol Lett 22:365\u0026ndash;376. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/ele.13199\u003c/span\u003e\u003cspan address=\"10.1111/ele.13199\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSilvertown J (2009) A new dawn for citizen science. Trends Ecol Evol 24:467\u0026ndash;471. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.tree.2009.03.017\u003c/span\u003e\u003cspan address=\"10.1016/j.tree.2009.03.017\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSogliani D, Mori E, Lovari S, Lazzeri L, Longoni A, Tabarelli De Fatis K, Sabatini P, Di Nicola MR, Russo D (2023) Citizen science and diet analysis shed light on dog-wildlife interactions in Italy. Biodiv Conserv 32(13):4461\u0026ndash;4479. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s10531-023-02707-7\u003c/span\u003e\u003cspan address=\"10.1007/s10531-023-02707-7\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSoulsbury CD, White PC (2015) Human\u0026ndash;wildlife interactions in urban areas: a review of conflicts, benefits and opportunities. Wildl Res 42:541\u0026ndash;553. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1071/WR14229\u003c/span\u003e\u003cspan address=\"10.1071/WR14229\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eStorch I, Lindstr\u0026ouml;m E, de Jounge J (1990) Diet and habitat selection of the pine marten in relation to competition with the red fox. Acta Theriol 35:311\u0026ndash;320\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eTakahashi Y, Yagisawa R, Tsuji Y (2025) Niche separation of sympatric carnivores, red foxes (\u003cem\u003eVulpes vulpes\u003c/em\u003e) and Japanese martens (\u003cem\u003eMartes melampus\u003c/em\u003e) in Northern Japan. Mamm Stud 51:1. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3106/ms2024-0075\u003c/span\u003e\u003cspan address=\"10.3106/ms2024-0075\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eTodini A, Crosti R (2020) Il cinghiale (\u003cem\u003eSus scrofa\u003c/em\u003e) come determinante di cambiamenti di vegetazione in una foresta urbana mediterranea: impatto sulla biodiversit\u0026agrave; di un\u0026rsquo;area protetta. Forest@ 17:71\u0026ndash;77. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3832/efor3284-017\u003c/span\u003e\u003cspan address=\"10.3832/efor3284-017\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eTorretta E, Serafini M, Puopolo F, Schenone L (2016) Spatial and temporal adjustments allowing the coexistence among carnivores in Liguria (NW Italy). Acta Ethol 19:123\u0026ndash;132. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s10211-015-0231-y\u003c/span\u003e\u003cspan address=\"10.1007/s10211-015-0231-y\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eTorretta E, Mosini A, Piana M, Tirozzi P, Serafini M, Puopolo F, Saino N, Balestrieri A (2017) Time partitioning in mesocarnivore communities from different habitats of NW Italy: insights into martens\u0026rsquo; competitive abilities. Behaviour 154:241\u0026ndash;266. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1163/1568539X-00003420\u003c/span\u003e\u003cspan address=\"10.1163/1568539X-00003420\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eT\u0026oacute;th M, B\u0026aacute;r\u0026aacute;ny A, Kis R (2009) An evaluation of stone marten (\u003cem\u003eMartes foina\u003c/em\u003e) records in the city of Budapest, Hungary. Acta Zool Ac Scient Hung 55(2):199\u0026ndash;209\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eTurner J, Freeman R, Carbone C (2022) Using citizen science to understand and map habitat suitability for a synurbic mammal in an urban landscape: The hedgehog \u003cem\u003eErinaceus europaeus\u003c/em\u003e. Mamm Rev 52:291\u0026ndash;303. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/mam.12278\u003c/span\u003e\u003cspan address=\"10.1111/mam.12278\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eTwining JP, Montgomery WI, Tosh DG (2020) The dynamics of pine marten predation on red and grey squirrels. Mammal Biol 100:285\u0026ndash;293. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s42991-020-00031-z\u003c/span\u003e\u003cspan address=\"10.1007/s42991-020-00031-z\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eVergara M, Cushman SA, Urra F, Ruiz-Gonz\u0026aacute;lez A (2016) Shaken but not stirred: multiscale habitat suitability modeling of sympatric marten species (\u003cem\u003eMartes martes\u003c/em\u003e and \u003cem\u003eMartes foina\u003c/em\u003e) in the northern Iberian Peninsula. Landsc Ecol 31:1241\u0026ndash;1260. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s10980-015-0307-0\u003c/span\u003e\u003cspan address=\"10.1007/s10980-015-0307-0\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eVillase\u0026ntilde;or NR, Driscoll DA, Escobar MA, Gibbons P, Lindenmayer DB (2014) Urbanization impacts on mammals across urban-forest edges and a predictive model of edge effects. PLoS ONE 9(5):e97036. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1371/journal.pone.0097036\u003c/span\u003e\u003cspan address=\"10.1371/journal.pone.0097036\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eViviano A, D\u0026rsquo;Amico M, Mori E (2023) Aliens on the road: surveying wildlife roadkill to assess the risk of biological invasion. Biology 12:850. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3390/biology12060850\u003c/span\u003e\u003cspan address=\"10.3390/biology12060850\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eViviano A, De Meo I, Mori E, Sergiacomi C, Paletto A (2024) Public perception and acceptance of coypu \u003cem\u003eMyocastor coypus\u003c/em\u003e removal in urban areas: influences of age and education. Sci Nat 111(5):42. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s00114-024-01928-2\u003c/span\u003e\u003cspan address=\"10.1007/s00114-024-01928-2\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eViviano A, Ancillotto L, Dondina O, Burchielli A, Miccolis D, Mori E (2025) What a camera trap survey can reveal about the behaviour of an invasive species: Insights from coypus \u003cem\u003eMyocastor coypus\u003c/em\u003e in an urban park of central Italy. Appl Anim Behav Sci 284:106534. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.applanim.2025.106534\u003c/span\u003e\u003cspan address=\"10.1016/j.applanim.2025.106534\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eWereszczuk A, Zalewski A (2015) Spatial niche segregation of sympatric stone marten and pine marten\u0026ndash;avoidance of competition or selection of optimal habitat? PLoS ONE 10:e0139852. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/10.1371/journal.pone.0139852\u003c/span\u003e\u003cspan address=\"10.1016/10.1371/journal.pone.0139852\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eWerner P (2011) The ecology of urban areas and their functions for species diversity. Landsc Ecol Engin 7(2):231\u0026ndash;240. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s11355-011-0153-4\u003c/span\u003e\u003cspan address=\"10.1007/s11355-011-0153-4\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eWidmer S, Riedel S, Babbi M, Herzog F, Wohlgemuth T, Kessler M, Dengler J (2025) One century of change: stronger diversity decline in lowland than in mountain grasslands in central Europe. Glob Change Biol 31:e70529. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/gcb.70529\u003c/span\u003e\u003cspan address=\"10.1111/gcb.70529\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eZabala J, Zuberogoitia I, Mart\u0026iacute;nez-Climent JA (2005) Site and landscape features ruling the habitat use and occupancy of the polecat (\u003cem\u003eMustela putorius\u003c/em\u003e) in a low density area: a multiscale approach. Eur J Wildl Res 51(3):157\u0026ndash;162. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s10344-005-0094-z\u003c/span\u003e\u003cspan address=\"10.1007/s10344-005-0094-z\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eZielinski WJ, Spencer WD, Barrett RH (1983) Relationship between food habits and activity patterns of pine martens. J Mammal 64(3):387\u0026ndash;396. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.2307/1380351\u003c/span\u003e\u003cspan address=\"10.2307/1380351\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"mammalian-biology","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"mamb","sideBox":"Learn more about [Mammalian Biology](https://link.springer.com/journal/42991)","snPcode":"42991","submissionUrl":"https://www.editorialmanager.com/mamb/default2.aspx","title":"Mammalian Biology","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Martes martes, urban ecology, temporal niche, camera trapping, behavioural plasticity, Italy","lastPublishedDoi":"10.21203/rs.3.rs-9043132/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-9043132/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eUrbanisation is reshaping mammalian communities worldwide, yet forest specialists are generally considered poor urban adapters. The pine marten \u003cem\u003eMartes martes\u003c/em\u003e, for long regarded as an indicator of intact woodlands, has recently shown signs of expansion into human-modified landscapes. Here, we combined citizen-science records and camera-trapping data to reconstruct the distribution of the pine marten in urban areas of Italy. Moreover, we used camera traps to investigate activity patterns, and spatiotemporal niche overlaps of the pine marten along an urban\u0026ndash;natural gradient in Italy. Verified reports from 2000\u0026ndash;2025 confirm the species presence in multiple Italian cities, including several regional capitals. In the Insugherata Urban Park in Rome, camera traps recorded 192 independent detections in one year, including 46 pine martens, 79 red foxes \u003cem\u003eVulpes vulpes\u003c/em\u003e, 33 Eurasian red squirrels \u003cem\u003eSciurus vulgaris\u003c/em\u003e, and 34 humans. Pine martens exhibited a predominantly crepuscular activity, with significant peaks at dawn and dusk, differing from the more nocturnal red fox. Temporal overlap between martens and foxes was low in the urban park (Δ₁ = 0.38), but overlap with red squirrels (Δ₁ = 0.84) and humans (Δ₁ = 0.77) was high, suggesting flexible diel adjustments to urban pressures. Compared to a natural reference site, pine martens in the city displayed reduced synchrony with foxes but increased overlap with both prey and human activity, indicating behavioural adaptation to anthropogenic environments. Our findings suggested a growing capacity of \u003cem\u003eM. martes\u003c/em\u003e to persist in urban green spaces by modulating temporal activity and resource use. The increase of urban records of pine marten highlights the need to reconsider its ecological classification and integrate cities into future conservation strategies for this carnivore.\u003c/p\u003e","manuscriptTitle":"Can a forest specialist live in a city? Behavioural plasticity fosters the presence of a protected carnivore in urban areas","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-03-23 19:22:44","doi":"10.21203/rs.3.rs-9043132/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"reviewerAgreed","content":"","date":"2026-03-24T15:04:39+00:00","index":0,"fulltext":""},{"type":"reviewersInvited","content":"","date":"2026-03-19T10:40:15+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2026-03-06T07:09:10+00:00","index":"","fulltext":""},{"type":"submitted","content":"Mammalian Biology","date":"2026-03-05T12:06:12+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"mammalian-biology","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"mamb","sideBox":"Learn more about [Mammalian Biology](https://link.springer.com/journal/42991)","snPcode":"42991","submissionUrl":"https://www.editorialmanager.com/mamb/default2.aspx","title":"Mammalian Biology","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"87941423-086f-4ece-87f3-2e0a6a5f3726","owner":[],"postedDate":"March 23rd, 2026","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2026-03-23T19:22:44+00:00","versionOfRecord":[],"versionCreatedAt":"2026-03-23 19:22:44","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-9043132","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-9043132","identity":"rs-9043132","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}