Making it in the city: Public data reveal patterns and drivers of terrestrial mammal occurrence across urban Philippines

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Using 43,604 publicly available GBIF records, this study analyzed occurrence patterns and predictors for 205 Philippine terrestrial mammal species in urban areas, while accounting for sampling/knowledge biases and using GIS overlays with Sentinel-2 urban land-use maps; it classified “urban mammals” as species with at least one record intersecting urban boundaries. Sixty-six species (32%) were recorded in urban areas, with strong taxonomic/order differences and bats overrepresented, consistent with phylogenetically structured urban tolerance. Urban occurrence was more likely for species with larger geographic ranges, herbivorous diets, and associations with taller forest canopies, and species appearing in urban areas also had more publications and observations, indicating documentation bias. A major caveat noted in the abstract is that the presence of threatened/endemic/declining taxa may reflect persistence under habitat contraction rather than true adaptation to urban environments. The paper does not explicitly discuss endometriosis or adenomyosis; it was included in the corpus via a keyword match in the upstream search index.

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Making it in the city: Public data reveal patterns and drivers of terrestrial mammal occurrence across urban Philippines | 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 Making it in the city: Public data reveal patterns and drivers of terrestrial mammal occurrence across urban Philippines Angeline Mampang Alcala, Princess Giulian Vianney B. Salon, Angelo R. Agduma, and 6 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-9367707/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Urbanisation is rapidly reshaping mammal communities; however, its effects remain poorly understood in tropical archipelagos with high endemism, such as the Philippines. Using 43,604 publicly available GBIF records, we examined the patterns and predictors of terrestrial mammal occurrence in urban areas of the Philippines while accounting for sampling and knowledge biases. Of the 205 mammal species, 66 (32%) were recorded in urban areas, indicating that urban assemblages represent a substantial but taxonomically uneven subset of mammalian species in the Philippines. Urban occurrence differed significantly across orders and families, with bats being strongly overrepresented, indicating a phylogenetically structured urban tolerance. Species were more likely to occur in urban areas when they had larger geographic range sizes, herbivorous diets, and were associated with landscapes with taller forest canopies, highlighting the role of ecological generality and vegetation structure in urban persistence. Although urban assemblages were dominated by widespread and least threatened species, they also included threatened, endemic, and declining taxa, suggesting that some records reflect persistence under habitat contraction rather than true adaptation to urban environments. Our findings further showed that species occurring in urban areas tended to have more publications and observation records. Our findings indicate that urban mammal assemblages are shaped by species’ ecological tolerance, range characteristics, and conservation status, but are also filtered through spatial biases inherent in biodiversity observations and documentation. Conservation Biology Terrestrial Ecology Conservation Island ecosystems Tropics Urban Systems Wildlife Figures Figure 1 Figure 2 Figure 3 Figure 4 Introduction Urbanisation, characterised by the expansion of urban populations and built environments, represents a major global social and environmental shift (Alberti, 2023 ; Chen, 2007 ; Simkin et al., 2022 ). Currently, more than 50% of the global population resides in cities, with urban areas projected to absorb most of the population growth from 2009 to 2050, particularly in Asia, where countries such as the Philippines are experiencing rapid expansion (Johnson et al., 2021 ; Mojares, 2013 ). By 2050, over 70% of the global population is predicted to live in cities, further accelerating the urban sprawl. Urbanisation transforms landscapes by altering land cover and hydrological systems (Grimm et al., 2008 ), leading to habitat degradation, fragmentation, and isolation, which significantly impact biodiversity at multiple scales (Villaseñor et al., 2014 ). It is widely recognised as a significant threat to biodiversity worldwide (Bell and Donnelly, 2006 ; Garden et al., 2006 ). This is particularly concerning in Southeast Asia, including the Philippines, where economic growth is driven by tourism and globalisation, alongside rapid urban expansion, degrading ecosystem structure and function (Olfato-Parojinog and Dagamac, 2024 ). Therefore, understanding the ecological processes at the boundary between wilderness and urban areas is critical for effective biodiversity conservation (Villaseñor et al., 2014 ). Urbanisation significantly impacts terrestrial mammals, with wide-ranging consequences for the ecosystem services they provide. Mammals are a crucial focus of conservation efforts because their interactions with humans make them effective indicators of ecological change (Bowyer et al., 2019 ). Urbanisation poses multiple threats to wildlife, including land-use change, leading to habitat fragmentation, reduced food availability, and noise and light pollution, often resulting in declines in species richness, diversity, and abundance, particularly among larger and specialist species (Villaseñor et al., 2014 ). It reshapes the distribution of terrestrial mammals by altering habitat availability and quality. Studies have shown that urban expansion displaces larger mammals, with species richness and occurrence declining most markedly in hotter cities with sparse green spaces (Melgares, 2023 ). In addition to shifts in species composition, urbanisation drives distinct behavioural and physiological changes among mammals, influencing breeding strategies and movement patterns, which further complicate conservation efforts in urban landscapes (Hahs et al., 2023 ). Urban mammals have multiple biological characteristics that improve their chances of survival in urban habitats, including increased litter size, varied diets, behavioural adaptability, and changes in body size (Santini et al., 2019 ). Numerous species have adapted to survive in environments dominated by humans using resources such as human food and waste, enabling them to maintain stable populations despite habitat change (Santini et al., 2019 ). The capacity of these mammals to inhabit various environments, such as private gardens and public parks, demonstrates their ecological adaptability and ability to withstand urbanisation (Gallo et al., 2022 ). Additionally, urbanisation leads mammals to adapt by becoming more active at night to avoid humans, potentially leading to human-wildlife conflicts. Therefore, research on mammalian behaviour in cities is crucial for wildlife conservation (Ritzel and Gallo, 2020 ). The Philippines harbours exceptional terrestrial biodiversity, characterised by high levels of endemism across numerous taxonomic groups (Agduma et al., 2023 ; Posa et al., 2008 ). Despite their biological significance, terrestrial mammals face mounting extinction risks driven by habitat loss due to agricultural expansion, urbanisation, and consumption (K. Tanalgo et al., 2025 ; Tanalgo and Hughes, 2019 , 2018 ). Understanding how species are distributed across habitat types and how anthropogenic pressures shape these patterns remains critically understudied in the Philippines, particularly in urban environments (Dela Torre et al., 2025 ). Given the rapid pace of urban growth in the region, understanding which mammal species thrive in urban areas is crucial for developing effective conservation strategies and prioritising the protection of critical populations. However, studying terrestrial mammal occurrence in Philippine urban areas presents a fundamental challenge due to limited data availability from the lack of systematic field studies focused on these environments (Dela Torre et al., 2025 ; Olfato-Parojinog and Dagamac, 2024 ). However, recent advancements in biodiversity documentation have provided new opportunities to address this gap. The increasing digitalisation of biodiversity records and the growth of large, publicly available databases have made more occurrence data accessible and interoperable (Tanalgo et al., 2023 ). For example, international initiatives, such as the Global Biodiversity Information Facility (GBIF), standardise data storage and make research-ready occurrence records openly accessible, further expanding the data landscape available to researchers studying the persistence of Philippine terrestrial mammals in urban environments (Tanalgo, 2025a ). Here, using publicly available datasets from the GBIF, we aimed to characterise the diversity and distribution of terrestrial mammals across urban areas in the Philippines and identify the intrinsic and ecological drivers of their occurrence. We also aimed to understand the patterns of research efforts based on knowledge gap indicators and compare them according to taxonomic groups, conservation status, and levels of endemism. We hypothesised that urban occurrence among Philippine terrestrial mammals is non-random across taxonomic groups and ecological statuses. First, we predicted that common and least-threatened species would be more widespread in urban environments than range-restricted species because broadly distributed taxa are more likely to encounter, tolerate, and persist in human-modified landscapes. Second, we predicted that smaller-bodied species with wider geographic ranges and broader habitat breadths would be more likely to occur in urban areas, as these traits are often linked to greater ecological flexibility in disturbed environments. Third, we predicted that species occurring in urban areas would be more frequently studied and better represented in biodiversity databases than species absent from urban settings, reflecting a strong sampling bias towards accessible, human-dominated landscapes where research efforts and opportunistic observations are concentrated. Materials and Methods Data overview We compiled terrestrial mammal occurrence records for the Philippines from the Global Biodiversity Information Facility (GBIF) on 5 April 2025 (Global Biodiversity Information Facility, 2025 ). Our dataset included all available records from 1950 to 2025 for 205 terrestrial mammal species in the country. We first curated the georeferenced occurrence data by checking for anomalous or problematic records in QGIS and removing records that fell outside the Philippine geopolitical boundaries. We also cross-checked species names against the Catalogue of Life Checklist Bank to ensure taxonomic consistency (GBIF & Catalogue of Life, 2025).. In addition, we excluded domesticated species, particularly dogs ( Canis lupus familiaris ) and cats ( Felis catus ), from the dataset. The initial download comprised 63,959 occurrence records, which were reduced to 43,604 after cleaning and filtering the data. To assess mammal occurrence in urban areas, we integrated the cleaned occurrence dataset with urbanisation and human settlement layers in QGIS. We overlaid occurrence points onto high-resolution (10 × 10 m) urban land-use maps derived from Sentinel-2 data and used spatial sampling tools in QGIS to determine whether the species records intersected with the mapped urban boundaries (Karra et al., 2021 ). Based on this spatial overlap, we classified each species as occurring or not occurring in the urban areas. Because mammal occurrence patterns in the Philippines remain incompletely documented, we did not attempt to assign species to finer categories of urban adaptation, such as urban avoiders, adapters, or exploiters (e.g. Santini et al. ( 2019 )). Instead, we used a conservative operational definition and considered “urban mammals” as species with at least one recorded occurrence within the urban boundaries of the Philippines. Knowledge gap indicators We accounted for research efforts across Philippine terrestrial mammals using Google Scholar hit counts and GBIF occurrence frequency records as indicators of knowledge levels and gaps for each terrestrial mammal species occurring in urban areas. The frequency of publications per species and observation records has been shown to serve as a reliable proxy for the research effort dedicated to a given species, and this approach has been applied in previous mammal studies in the Philippines (Tanalgo et al., 2025 ). For this analysis, we searched Google Scholar ( https://scholar.google.com.ph/ ) using species-specific queries to quantify how well each terrestrial mammal species in the Philippines has been studied. We expected species with more Google Scholar hits to reflect greater research efforts. We chose Google Scholar because it is open-access, requires no institutional subscription, and provides broader coverage of the published literature per species than Scopus and the Web of Science. We used the search syntax [“species X” + “Philippines”] following Tanalgo et al. ( 2025 ) to retrieve all publications containing the species name for each species in our analyses. The search results were filtered to include only records from the Philippines. We log 10 -transformed all records for the analysis. Explanatory variables We compiled species-level data for Philippine terrestrial mammals, including the IUCN Red List conservation status, country-level endemism, and life-history traits. We included body size, trophic level, diel activity, range size, and habitat breadth as intrinsic variables potentially influencing mammal occurrence in urban areas, extracted from Tanalgo et al. ( 2025 ), along with forest height (Potapov et al., 2021 ) and mean surface temperature as extrinsic variables representing landscape structure and climatic conditions. We expected that intrinsic traits would be strong predictors of mammal occurrence in urban areas. Large-bodied species were predicted to be less likely to occur in urban environments because they generally require larger home ranges and are more sensitive to habitat loss and human disturbances. In contrast, small-bodied mammals are expected to occur more frequently in urban areas because their size allows them to exploit fine-scale microhabitats, including tree holes, canopies, foliage, and built structures (Santini et al., 2019 ). Species with broader habitat breadths and wider geographic ranges are predicted to have higher urban occurrence, reflecting greater ecological flexibility (Tanalgo et al., 2025 ). We further expected omnivorous and nocturnal species to fare better in urban environments because they can exploit diverse food resources and reduce their direct encounters with humans. For extrinsic variables, we predicted that greater surrounding forest cover would increase mammal occurrence, even within urban landscapes, by providing refuge and supplementary habitat. In contrast, higher surface temperatures are expected to reduce the occurrence of forest-dependent and thermally sensitive species, many of which are endemic to the Philippines and occupy narrow ecological niches in the forest understory. Data analysis Prior to formal analysis, all datasets were checked for statistical assumptions. First, we performed a Chi-square test of association (χ 2 ) to determine whether there was a significant difference in the distribution of non-urban and urban mammals in terms of taxonomic groups, conservation status, and endemism. We modelled the probability of urban occurrence of mammal species using a generalised linear mixed model (GLMM) with a binomial error structure. The predictor variables included body mass, trophic level, diel activity, geographic range size, habitat breadth, forest canopy height and mean surface temperature. This framework allowed us to test whether variations in urban occurrence were associated with species traits, environmental conditions, and extent of species distribution. We further tested whether mammal species occurring in urban areas were disproportionately represented in the biodiversity knowledge, as reflected by publication outputs and occurrence records. To do so, we fitted a second set of GLMM with publication density and the number of GBIF records as separate response variables. The predictors included urban occurrence, extinction risk, and endemism. We expected species recorded from urban areas to show higher publication density and more GBIF records than species restricted to other habitat types, consistent with a bias towards taxa occupying more accessible, human-dominated landscapes. Continuous variables were z -standardised to facilitate comparisons across predictors. For both models, taxonomic family was included as a random effect to account for phylogenetic structure, and model selection was performed using the Akaike Information Criterion (AIC), with the best-supported model identified as the one with the lowest AIC value. All statistical analyses were conducted using the open-source Jamovi software (version 2.7.17; The jamovi project, 2026 ), and data visualisation was performed using GraphPad Prism (version 11; GraphPad Prism, 2022). Results Diversity and drivers of urban mammal occurrence We examined the occurrence of Philippine terrestrial mammal species in urban areas and the factors associated with their presence in these areas. Of the 205 terrestrial mammals included in our analysis, 32% were recorded in urban areas across the country. Among mammalian orders, Chiroptera comprised the largest proportion of urban mammals (62%, n = 41 species), followed by Rodentia (18.18%, n = 12 species). Urban occurrence differed significantly across both orders (χ² = 46.83, p < 0.001) and families (χ² = 66.52, p < 0.001) ( Fig. 1 ). Urban occurrence varied significantly according to conservation status (χ² = 12.97, p = 0.024). Most urban species (71%, n = 47 species) were classified as having a Least Concern. Notably, however, 15% of the species recorded in urban areas were threatened, including Vulnerable, Endangered and Critically Endangered taxa. A substantial proportion of urban mammals were endemic to the Philippines (48%, n = 32 species), with significant differences in endemicity between urban and non-urban groups (χ² = 10.97, p < 0.001). In terms of population trends, 41% (n = 27 species) of urban mammals showed stable trends, whereas 33% showed decreasing trends (χ² = 9.06, p = 0.028). We built four models to predict the occurrence of terrestrial mammals in urban areas in the Philippines using biological, geographical, and extrinsic traits while accounting for discovery bias. Among these, the full model provided the best fit to the data (AICc = 163.479). Binomial GLMMs showed that herbivorous species were more likely to occur in urban areas than carnivores and omnivores (β = 1.553, p = 0.013) and species with larger geographical ranges (β = 1.393, p = 0.008). Among the extrinsic variables, urban mammals were also associated with areas of higher forest canopy height (β = 1.974, p < 0.001) ( Fig. 2 ). Relationship of urban occurrence and knowledge gaps We tested whether mammal occurrence in urban areas contributed to taxonomic bias in Google Scholar hits and GBIF occurrence. Urban occurrence was positively associated with both Google Scholar hits (β = 0.460, p < 0.001) and GBIF occurrence records (β = 0.998, p < 0.001), with a significantly stronger effect on the latter than on the former. These results suggest that species occurring in urban areas are generally more detectable and better documented across their ranges, rather than simply being more frequently recorded within cities. The interaction between non-endemic status and urban occurrence was significant only for the GBIF records (β = 0.360, p = 0.048) (Fig. 3 a). Extinction risk was associated only with GBIF records (β = −0.583, p < 0.001), demonstrating that less threatened species tended to have more records than threatened taxa. Endemism was associated with Google Scholar hits, with non-endemic species tending to receive more citations than endemic species (β = 0.613, p < 0.001). Discussion Urbanisation is expected to intensify globally, yet its effects on biodiversity remain poorly understood across many taxonomic groups and regions, particularly in the Philippines. Here, we provide the first nationwide assessment of urban mammal occurrence in the country using publicly available records from the Global Biodiversity Information Facility (GBIF). By integrating biological, geographical, and extrinsic predictors, we established a baseline for understanding mammal distributions in urban landscapes and evaluated whether urban occurrence shapes the existing knowledge gaps in mammal occurrence data. Diversity of urban mammals Public observation data revealed that approximately one-third of terrestrial mammals in the Philippines are found in urban areas. We found significant differences in urban occurrence across both order and family levels, indicating that phylogenetic identity and the suite of life-history traits it encapsulates are significant determinants of urban occurrence than previously expected under random assembly. Our results are consistent with the growing evidence from urban mammal studies in other regions that phylogenetic conservatism in urban tolerance, mediated through traits such as body size, dietary breadth, and reproductive rate, produces non-random assemblage structures (Ritzel and Gallo, 2020 ; Santini et al., 2019 ). Bats were markedly overrepresented among Philippine urban mammals, comprising 62% of the species recorded. Our findings suggest that Philippine bats are highly tolerant of the effects of urbanisation compared with other mammalian groups (Jung and Threlfall, 2018 , 2016 ; Moretto et al., 2022 ). For example, many Philippine bats, particularly pteropodid fruit bats and various insectivorous species, readily exploit anthropogenic structures for roosting, with bridges, building eaves, and urban canopies serving as functional analogues to natural caves and old-growth tree hollows (Dela Torre et al., 2025 ; Tanalgo et al., 2025 ). Several interrelated traits explain the high affinity of bats for urbanisation. The high aerial mobility of many bats, especially cosmopolitan and smaller-bodied insectivorous bats, may substantially reduce commuting costs between urban matrices and adjacent natural habitats, allowing bats to exploit both simultaneously, a foraging flexibility that is unavailable to most ground-dwelling mammals (Avila-Flores et al., 2022 ; Jung and Kalko, 2010 ; Russo and Ancillotto, 2015 ). Some species of fruit bats forage in urban green areas over shorter distances and exploit food resources that are more readily available in the city (Meade et al., 2021 ). Egert-Berg et al. ( 2021 ) hypothesizes that fruit bats might be seeking a refuge from predation and benefit from warmer micro-climates and easier navigation due to abundant landmarks in cities. Meanwhile, Páez et al. ( 2018 ) proposed that habitat loss, which increases the time and energy costs required to search for a new foraging patch, and the availability of non-native, more constant food sources in urban areas may cause flying foxes to become residents in urban environments. Finally, light-induced insect aggregation in urban areas provides a reliable and predictable prey subsidy for common insectivorous species, potentially making urban environments more profitable than naturally lit forest habitats and increasing the likelihood of detecting more bat species in these settings (Avila-Flores and Fenton, 2005 ; Li and Wilkins, 2022 ; Schoeman, 2016 ). Nonetheless, the role of urban areas as alternative favourable foraging grounds for many bat species creates an ecological and evolutionary trap, which makes urban areas appear favourable but negatively affects bat populations and behaviour in the long term (Tanalgo et al., 2025 ). Although rodents tend to dominate urban areas in terms of abundance, only 18% of rodent species have been recorded in urban habitats in the Philippines. Globally, commensal rodents, particularly those of the genus Rattus , are among the most ubiquitous and ecologically dominant urban mammals (Feng and Himsworth, 2014 ). Interestingly, although R. rattus and R. novergicus are common urban commensals worldwide (Feng et al., 2025 ), they are poorly represented in urban GBIF records in the Philippines. This pattern is unlikely to indicate that species are rare in cities. Instead, it may reflect how biodiversity data are collected, with greater attention often given to species of ecological or conservation interest than to widespread pest species (Boakes et al., 2010 ; Bowler et al., 2022 ). We argue that in urban areas, R. rattus and R. norvegicus may also be overlooked in surveys or left unreported because their presence is often expected. Consequently, its low representation in the GBIF probably reflects underreporting rather than a low occurrence, highlighting an important limitation of public occurrence data for describing urban mammal communities. Moreover, rodent diversity in the Philippines is dominated by a rich radiation of endemic, forest-dependent species, many of which occupy narrow ecological niches in montane and old-growth forest habitats with no history of commensal association with humans (Heaney et al., 1998 ; Lawrence R. Heaney et al., 2016 ; Lawrence Richard Heaney et al., 2016 ). Our study shows that rodent species represented in Philippine urban assemblages are likely taxonomically and ecologically distinct, and the presence of native species in urban areas may reflect incidental habitat use or range compression into disturbed landscapes rather than functional synanthropy in urban areas. Our findings indicate that urban landscapes tend to favour generalist and synanthropic small mammals, whereas forest-dependent species decline, resulting in reduced community diversity, which explains the patterns observed in our study. Ecological status of urban mammals The predominance of least threatened species among urban mammals (71%) is expected, as they are often characterised by broad distributions, large population sizes, and generalist ecological requirements that are congruent with their tolerance of anthropogenic disturbance (Wells et al., 2014 ). The pattern observed in our study suggests that mammal records in urban areas are dominated by species that are less threatened and have broader geographical ranges. This supports our initial hypothesis that urbanisation tends to favour cosmopolitan species that are generally more adaptable to human-modified environments than other species. Interestingly, at least 15% of urban species are considered threatened according to the IUCN Red List, suggesting that urban areas may serve as both refuges and threats to terrestrial mammals in the Philippines. In tropical biodiversity hotspots subjected to rapid land-use change, urban and peri-urban habitats can serve as de facto refuge environments for species that can no longer maintain viable populations in their surrounding landscapes (Boakes et al., 2024 ; Maclagan et al., 2018 ; Tanalgo et al., 2025 ). In the Philippine archipelago, where forest cover loss has been among the most severe globally, the presence of threatened mammals in cities is likely a symptom of widespread habitat contraction rather than an indicator of urban ecological suitability. Distinguishing between these two processes, genuine urban adaptation and forced habitat use under landscape stress, is empirically important, but we acknowledge that this cannot be resolved from occurrence data alone and warrants targeted demographic and behavioural investigations. Furthermore, the significantly high proportion of Philippine endemics among urban mammals is particularly intriguing in this study. The Philippines harbours the highest density of endemic non-volant mammal species per unit area of any country globally (Heaney et al., 1998 ; Heaney et al., 2016 ), and some endemic representation in urban assemblages is therefore not unexpected. However, the statistical significance of this pattern and its co-occurrence with a substantial proportion of declining species further suggest a dynamic that is more consistent with habitat compression than with urban adaptation. Endemic species in the Philippines are disproportionately forest-dependent and have experienced severe habitat loss across their limited geographic ranges (Tanalgo et al., 2025 ); therefore, urban occurrence may represent the residual persistence of such species in structurally complex fragments of urban green spaces rather than the active colonisation of novel urban environments (Gallo et al., 2017 ; Maclagan et al., 2018 ). Under this interpretation, the apparently high endemic richness of Philippine urban areas should be viewed not as a conservation opportunity, but as a biodiversity risk indicator, a signal that endemic taxa are being pushed into increasingly marginal and structurally impoverished habitats. For example, a recent global analysis showed that approximately 33% of studies reported the presence of species worthy of conservation, such as rare, endemic, or endangered species, in a total of 365 species within urban refugia (Gentili et al., 2024 ). Our findings further highlight the need to avoid viewing urban ecosystems as biologically irrelevant, especially in tropical regions, where urban expansion often overlaps with areas of high endemicity. Drivers of urban mammal occurrence Our models identified herbivory, large geographic range size, and forest canopy height as significant positive predictors of urban mammal occurrence in the Philippines, with the full model providing the best fit. Contrary to our expectations, body size did not significantly affect the occurrence of urban areas in this study. The positive effects of herbivory can be ecologically interpreted in the context of the Philippine urban landscape, where ornamental fruiting trees in parks, roadsides, and private gardens constitute a persistent and predictable food resource accessible to frugivorous species, particularly to pteropodid bats (Meade et al., 2021 ). Unlike animal-based food resources, which are spatially variable and ephemeral in urban environments, fruiting trees represent temporally structured but spatially stable resources that can support regular and repeated exploitation (Lim et al., 2018 ; Páez et al., 2018 , 2018 ). Although we argue that the dominance of herbivorous bats and rodents among Philippine urban mammals likely drives this relationship, it should not be generalised to imply that herbivory is broadly advantageous in urban mammal assemblages, where diverse feeding guilds exhibit complex and context-dependent urban responses. The positive association between geographical range size and urban occurrence is consistent with macroecological predictions and aligns with the results of global and regional syntheses of mammal ecology. Wide-ranging species typically exhibit broad environmental tolerances, high dispersal capacities, and generalist resource use, which function as pre-adaptations to urban environments rather than as responses specifically selected for these conditions (Sol et al., 2013 ). Range size may also serve as a proxy for population size and persistence capacity, such that wide-ranging species are more likely to maintain viable subpopulations across heterogeneous landscapes, including urban patches (Santini et al., 2019 ). Moreover, the positive association between the persistence of mammals and the surrounding forest canopy in urban areas is ecologically plausible and practically important. We argue that rather than portraying Philippine urban mammals as species tolerant of simplified habitats, our results suggest that mammal richness in cities is linked to the vertical complexity of urban vegetation in degraded ecosystems (Camargo et al., 2018 ). In urban areas, tall tree canopies may provide temporary refuge as roosting substrates, thermal buffering, shelter from disturbance and predators, and access to flowering and fruiting resources across canopy layers, which can potentially recruit mammal species in urban areas (Antoniazzi et al., 2026 ; Tratalos et al., 2007 ). Our findings underscore the importance of retaining substantial tree cover in and around urbanised areas to support a wider range of species and preserve their ecological roles across transitional landscapes (Jovanović et al., 2025 ). Knowledge bias in urban species We found a strong positive relationship between urban mammal occurrence and both Google Scholar publication density and GBIF occurrence records, indicating that species recorded in urban areas also tend to be more widely represented across other habitat types in the country. Rather than reflecting urban occurrence alone, this pattern suggests that mammals capable of extending into urban landscapes are generally those with broader ecological distributions and wider realised habitat use. Such species are likely to possess traits linked to environmental tolerance and flexibility, allowing them to persist in human-dominated settings and across a wider range of nonurban environments. Consequently, their occurrence is more frequently documented in both the scientific literature and biodiversity databases. This creates an important bias in biodiversity knowledge, whereby species that include urban areas within their distribution appear better known overall, whereas species confined to less accessible or less modified habitats are comparatively underrepresented. Such an imbalance may distort conservation priorities by overstating the apparent knowledge base for habitat-flexible species and obscuring the status and data limitations of more restricted taxa. Most of the urban mammal species recorded in the Philippines were least threatened, as threatened and endemic species are more often associated with forested, less disturbed, or remote habitats, which are often difficult to sample because of access and security (Hilario-Husain et al., 2024 ; Tanalgo et al., 2025 ). We further contend that the lower number of urban records for threatened and endemic mammals may reflect both sampling biases of these species and ecological filtering by urbanisation, where only species with wider range sizes that can withstand urban conditions persist and are recorded. Overall, our findings indicate that urban mammal assemblages are shaped by a combination of species’ ecological tolerance, range characteristics, and conservation status, and are influenced by spatial biases inherent in biodiversity observations. Our findings have practical implications for the interpretation and weighting of biodiversity data for conservation planning in the Philippines and beyond. Distribution models, species richness maps, and extinction risk assessments calibrated primarily on existing occurrence data will implicitly over-represent urban-tolerant, non-endemic, and least-concern taxa at the expense of ecologically distinctive fauna that confer global conservation significance to the archipelago (Hughes et al., 2021 ). Correcting for this bias will require deliberate redistribution of survey effort toward historically undersampled regions and taxonomic groups, particularly bats, which dominate urban assemblages and likely generate the strongest urban sampling artefact in the Philippine mammal data. This will also require investment in data infrastructure that can accommodate taxonomic uncertainty and track occurrence data for recently described and poorly known endemic taxa (Tanalgo et al., 2023 ; Tanalgo, 2025a , 2025b ). The significant relationship between urban mammal occurrence and data abundance identified here should serve as an empirical prompt for such rebalancing rather than as a justification for concentrating future research in environments with the greatest existing knowledge. Synthesis Our results suggest that urbanisation in the Philippines is not merely restructuring mammal communities but also reshaping how biodiversity is detected, recorded, and interpreted. Urban environments appear to favour mobile, generalist, and disturbance-tolerant species, whereas many endemic, threatened, and forest-dependent mammals remain ecologically marginal and poorly represented in available datasets. This pattern implies that urban biodiversity data may capture not only biological responses to land-use change but also the uneven visibility of species across human-altered landscapes. Consequently, species that persist in cities are more likely to be observed, recorded, and studied, whereas those that are most vulnerable to urban expansion may be systematically overlooked. We also acknowledge the important limitations of our study, particularly in the classification of urban species and the spatial extent used for defining urban areas. These choices may affect which species are considered urban-associated and, in turn, the patterns inferred from the analyses. Future studies would benefit from more standardised and ecologically explicit definitions of urban habitats across the urban–rural gradient. A more balanced assessment of Philippine mammal biodiversity will also require targeted sampling beyond major urban centres, especially in undersurveyed habitats and among poorly known endemic lineages, to better resolve how intrinsic and extrinsic factors shape species persistence in urban landscapes. Our analysis was also limited to presence–absence observations and therefore, cannot fully capture the broader ecological responses of mammals to urbanisation. We call for more field-based studies and systematic observations aimed at understanding species persistence and responses to urban environments. Such efforts are essential for fully assessing the impacts of urbanisation on mammal species and for informing more effective urban planning and conservation strategies. Nonetheless, our current work further underscores the value of openly accessible biodiversity databases, particularly the Global Biodiversity Information Facility (GBIF), in revealing the broad patterns of species occurrence in urban landscapes (Sweet et al., 2022 ). By aggregating records across taxa, regions, and time, these datasets provide an important foundation for detecting how biodiversity is distributed in human-dominated environments and how species respond to ongoing environmental changes (Li et al., 2019 ; Sultana and Storch, 2021 ). Although these records are subject to biases in sampling and detectability, they offer an essential starting point for large-scale urban biodiversity research, particularly in regions where systematic field surveys are limited (Beck et al., 2013 ; Hughes et al., 2021 ). Thus, GBIF and similar platforms can serve as critical baseline resources for advancing urban ecology, guiding hypothesis development, identifying knowledge gaps and informing future field-based investigations. More broadly, biodiversity knowledge from urban areas in the Philippines remains far more limited than that from other habitat types (Olfato-Parojinog and Dagamac 2024 ), with mammals being particularly underrepresented. To help narrow these gaps, we outline several priorities for strengthening biodiversity research in urban landscapes in the Philippines ( Fig. 4 ) . Importantly, the occurrence of conservation-relevant species in urban areas should not be interpreted as evidence of successful adaptation to urban environments. Instead, it may reflect habitat constriction, whereby species are forced into increasingly fragmented and degraded refuges embedded within expanding urban matrixes. In this context, we argue that urban areas may function as temporary refuges for some mammals in the Philippines, but such persistence may mask deeper ecological instability. As the human population in the Philippines continues to grow, the conversion of previously intact ecosystems into urban landscapes is likely to intensify, generating novel ecosystems to which many mammal species must adapt to. We argue that urbanisation may also generate ecological and evolutionary traps, in which apparently suitable habitats expose mammals to chronic disturbances, novel risks, and reduced long-term fitness (Gentili et al., 2024 ; Russo et al., 2024 ; Tanalgo et al., 2025 ). Thus, the urban presence of mammals, particularly ecologically unique species, may represent not only resilience but also a more fragile and transient form of persistence under increasing anthropogenic pressures. Declarations Data Availability Statement Data used for this study is available in https://doi.org/10.15468/dl.6ju8sc Ethics Declaration Not applicable Conflict of Interest The authors declare no conflict of interest. References Agduma AR, Garcia FG, Cabasan MT, Pimentel J, Ele RJ, Rubio M, Murray S, Hilario-Husain BA, Cruz D, Abdullah KC, Balase S, Tanalgo SM, K.C (2023) Overview of priorities, threats, and challenges to biodiversity conservation in the southern Philippines. 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Edradan","email":"","orcid":"https://orcid.org/0009-0007-4396-9910","institution":"Ecology and Conservation Research Laboratory (Eco/Con Lab), Department of Biological Sciences, College of Science and Mathematics, University of Southern Mindanao, Kabacan 9407, Cotabato, Philippines","correspondingAuthor":false,"prefix":"","firstName":"Janine","middleName":"C.","lastName":"Edradan","suffix":""},{"id":620422214,"identity":"dff7bdc4-870a-47e5-8c8c-40d51c4e5604","order_by":6,"name":"Pearl Ansel S. Dela Serna","email":"","orcid":"https://orcid.org/0009-0005-9095-1073","institution":"Ecology and Conservation Research Laboratory (Eco/Con Lab), Department of Biological Sciences, College of Science and Mathematics, University of Southern Mindanao, Kabacan 9407, Cotabato, Philippines","correspondingAuthor":false,"prefix":"","firstName":"Pearl","middleName":"Ansel S. Dela","lastName":"Serna","suffix":""},{"id":620422215,"identity":"43372845-4d27-43a2-9a21-0a79a1d1c01d","order_by":7,"name":"Kier C. Dela Cruz","email":"","orcid":"https://orcid.org/0009-0003-6255-5642","institution":"Ecology and Conservation Research Laboratory (Eco/Con Lab), Department of Biological Sciences, College of Science and Mathematics, University of Southern Mindanao, Kabacan 9407, Cotabato, Philippines","correspondingAuthor":false,"prefix":"","firstName":"Kier","middleName":"C. Dela","lastName":"Cruz","suffix":""},{"id":620422216,"identity":"04a7bdf4-34bc-4454-8fb1-28212a122d12","order_by":8,"name":"Krizler C. Tanalgo","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA0UlEQVRIiWNgGAWjYDACHgaGAwwFDAz8IE5CATE6wFoMGBgkG0BaDIjUwgDSYnAAyiAI7HnOPjz4w8Am3/j86sQPDwwY5PnFDhCwhbfd4ICEQZrlthtvN0sAHWY4c3YCAS38bAwHDAwOG5jdOLsBpCXB4DYxWhIM/hsYzzi7+QdxWnjbGA4cMABaxN+7jUhbzhxjONhgkGwgcYN3m0WCgQRhv7D3pDF//FFhZ8Dff3bzzR8VNvL80gS0IIAEWKUEscpBgP8AKapHwSgYBaNgJAEAKU9AeuQbnUAAAAAASUVORK5CYII=","orcid":"https://orcid.org/0000-0003-4140-336X","institution":"Ecology and Conservation Research Laboratory (Eco/Con Lab), Department of Biological Sciences, College of Science and Mathematics, University of Southern Mindanao, Kabacan 9407, Cotabato, Philippines","correspondingAuthor":true,"prefix":"","firstName":"Krizler","middleName":"C.","lastName":"Tanalgo","suffix":""}],"badges":[],"createdAt":"2026-04-09 11:10:13","currentVersionCode":1,"declarations":{"humanSubjects":false,"vertebrateSubjects":false,"conflictsOfInterestStatement":false,"humanSubjectEthicalGuidelines":false,"humanSubjectConsent":false,"humanSubjectClinicalTrial":false,"humanSubjectCaseReport":false,"vertebrateSubjectEthicalGuidelines":false},"doi":"10.21203/rs.3.rs-9367707/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-9367707/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":108475429,"identity":"37d7c503-9aa7-4820-83c4-9c4a11e1d691","added_by":"auto","created_at":"2026-05-05 06:42:50","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":11188502,"visible":true,"origin":"","legend":"\u003cp\u003ePhylogram of terrestrial Philippine mammals recorded and observed in urban areas of the Philippines, classified according to order and risk of extinction.\u003c/p\u003e","description":"","filename":"Figure1.png","url":"https://assets-eu.researchsquare.com/files/rs-9367707/v1/1ea619c8b9acdcf5700f69af.png"},{"id":108475430,"identity":"3f70e938-7cb4-458a-934e-05fe960506f0","added_by":"auto","created_at":"2026-05-05 06:42:50","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":1448009,"visible":true,"origin":"","legend":"\u003cp\u003eForest plot showing the best Generalised Linear Mixed Model (GLMM) predicting the effects of biological, geographical, extrinsic, and discovery bias on urban occurrence and observation of terrestrial Philippine mammals. The exact \u003cem\u003ep\u003c/em\u003e-values are indicated. The dots indicate the estimates, and the whiskers indicate the 95% confidence interval.\u003c/p\u003e","description":"","filename":"Figure2.png","url":"https://assets-eu.researchsquare.com/files/rs-9367707/v1/c24ae335161df86e1cf0000c.png"},{"id":108804286,"identity":"c7f34d4a-977a-451c-b8b8-4217563ebb79","added_by":"auto","created_at":"2026-05-08 15:18:57","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":1049059,"visible":true,"origin":"","legend":"\u003cp\u003eForest plot showing the best Generalised Linear Mixed Model (GLMM) predicting the effects of urban occurrence, extinction risk, and endemism of terrestrial Philippine mammals on Google Scholar hits and GBIF occurrence records.\u003c/p\u003e","description":"","filename":"Figure3.png","url":"https://assets-eu.researchsquare.com/files/rs-9367707/v1/b840928cd24f5947c6e4d11b.png"},{"id":108475432,"identity":"a1ef80c7-9df7-4337-aded-8bb3cee9d548","added_by":"auto","created_at":"2026-05-05 06:42:50","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":12556059,"visible":true,"origin":"","legend":"\u003cp\u003eSummary of key recommendations for improving the biodiversity knowledge of urban mammals in the Philippines.\u003c/p\u003e","description":"","filename":"Figure4.png","url":"https://assets-eu.researchsquare.com/files/rs-9367707/v1/218696781131735b46c4e0f7.png"},{"id":108811559,"identity":"7e979616-931f-40c7-8dae-c15df3d37325","added_by":"auto","created_at":"2026-05-08 16:05:37","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":25800050,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-9367707/v1/0c53ae80-bb27-425c-9c47-f03c6cb8d167.pdf"}],"financialInterests":"The authors declare no competing interests.","formattedTitle":"\u003cp\u003e\u003cstrong\u003eMaking it in the city: Public data reveal patterns and drivers of terrestrial mammal occurrence across urban Philippines\u003c/strong\u003e\u003c/p\u003e","fulltext":[{"header":"Introduction","content":"\u003cp\u003eUrbanisation, characterised by the expansion of urban populations and built environments, represents a major global social and environmental shift (Alberti, \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Chen, \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Simkin et al., \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Currently, more than 50% of the global population resides in cities, with urban areas projected to absorb most of the population growth from 2009 to 2050, particularly in Asia, where countries such as the Philippines are experiencing rapid expansion (Johnson et al., \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Mojares, \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). By 2050, over 70% of the global population is predicted to live in cities, further accelerating the urban sprawl. Urbanisation transforms landscapes by altering land cover and hydrological systems (Grimm et al., \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2008\u003c/span\u003e), leading to habitat degradation, fragmentation, and isolation, which significantly impact biodiversity at multiple scales (Villase\u0026ntilde;or et al., \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). It is widely recognised as a significant threat to biodiversity worldwide (Bell and Donnelly, \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Garden et al., \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). This is particularly concerning in Southeast Asia, including the Philippines, where economic growth is driven by tourism and globalisation, alongside rapid urban expansion, degrading ecosystem structure and function (Olfato-Parojinog and Dagamac, \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Therefore, understanding the ecological processes at the boundary between wilderness and urban areas is critical for effective biodiversity conservation (Villase\u0026ntilde;or et al., \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2014\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eUrbanisation significantly impacts terrestrial mammals, with wide-ranging consequences for the ecosystem services they provide. Mammals are a crucial focus of conservation efforts because their interactions with humans make them effective indicators of ecological change (Bowyer et al., \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Urbanisation poses multiple threats to wildlife, including land-use change, leading to habitat fragmentation, reduced food availability, and noise and light pollution, often resulting in declines in species richness, diversity, and abundance, particularly among larger and specialist species (Villase\u0026ntilde;or et al., \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). It reshapes the distribution of terrestrial mammals by altering habitat availability and quality. Studies have shown that urban expansion displaces larger mammals, with species richness and occurrence declining most markedly in hotter cities with sparse green spaces (Melgares, \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). In addition to shifts in species composition, urbanisation drives distinct behavioural and physiological changes among mammals, influencing breeding strategies and movement patterns, which further complicate conservation efforts in urban landscapes (Hahs et al., \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eUrban mammals have multiple biological characteristics that improve their chances of survival in urban habitats, including increased litter size, varied diets, behavioural adaptability, and changes in body size (Santini et al., \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Numerous species have adapted to survive in environments dominated by humans using resources such as human food and waste, enabling them to maintain stable populations despite habitat change (Santini et al., \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). The capacity of these mammals to inhabit various environments, such as private gardens and public parks, demonstrates their ecological adaptability and ability to withstand urbanisation (Gallo et al., \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Additionally, urbanisation leads mammals to adapt by becoming more active at night to avoid humans, potentially leading to human-wildlife conflicts. Therefore, research on mammalian behaviour in cities is crucial for wildlife conservation (Ritzel and Gallo, \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe Philippines harbours exceptional terrestrial biodiversity, characterised by high levels of endemism across numerous taxonomic groups (Agduma et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Posa et al., \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). Despite their biological significance, terrestrial mammals face mounting extinction risks driven by habitat loss due to agricultural expansion, urbanisation, and consumption (K. Tanalgo et al., \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2025\u003c/span\u003e; Tanalgo and Hughes, \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2019\u003c/span\u003e, \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Understanding how species are distributed across habitat types and how anthropogenic pressures shape these patterns remains critically understudied in the Philippines, particularly in urban environments (Dela Torre et al., \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Given the rapid pace of urban growth in the region, understanding which mammal species thrive in urban areas is crucial for developing effective conservation strategies and prioritising the protection of critical populations. However, studying terrestrial mammal occurrence in Philippine urban areas presents a fundamental challenge due to limited data availability from the lack of systematic field studies focused on these environments (Dela Torre et al., \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2025\u003c/span\u003e; Olfato-Parojinog and Dagamac, \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). However, recent advancements in biodiversity documentation have provided new opportunities to address this gap. The increasing digitalisation of biodiversity records and the growth of large, publicly available databases have made more occurrence data accessible and interoperable (Tanalgo et al., \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). For example, international initiatives, such as the Global Biodiversity Information Facility (GBIF), standardise data storage and make research-ready occurrence records openly accessible, further expanding the data landscape available to researchers studying the persistence of Philippine terrestrial mammals in urban environments (Tanalgo, \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2025a\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eHere, using publicly available datasets from the GBIF, we aimed to characterise the diversity and distribution of terrestrial mammals across urban areas in the Philippines and identify the intrinsic and ecological drivers of their occurrence. We also aimed to understand the patterns of research efforts based on knowledge gap indicators and compare them according to taxonomic groups, conservation status, and levels of endemism.\u003c/p\u003e \u003cp\u003eWe hypothesised that urban occurrence among Philippine terrestrial mammals is non-random across taxonomic groups and ecological statuses. First, we predicted that common and least-threatened species would be more widespread in urban environments than range-restricted species because broadly distributed taxa are more likely to encounter, tolerate, and persist in human-modified landscapes. Second, we predicted that smaller-bodied species with wider geographic ranges and broader habitat breadths would be more likely to occur in urban areas, as these traits are often linked to greater ecological flexibility in disturbed environments. Third, we predicted that species occurring in urban areas would be more frequently studied and better represented in biodiversity databases than species absent from urban settings, reflecting a strong sampling bias towards accessible, human-dominated landscapes where research efforts and opportunistic observations are concentrated.\u003c/p\u003e"},{"header":"Materials and Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eData overview\u003c/h2\u003e \u003cp\u003eWe compiled terrestrial mammal occurrence records for the Philippines from the Global Biodiversity Information Facility (GBIF) on 5 April 2025 (Global Biodiversity Information Facility, \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Our dataset included all available records from 1950 to 2025 for 205 terrestrial mammal species in the country. We first curated the georeferenced occurrence data by checking for anomalous or problematic records in QGIS and removing records that fell outside the Philippine geopolitical boundaries. We also cross-checked species names against the Catalogue of Life Checklist Bank to ensure taxonomic consistency (GBIF \u0026amp; Catalogue of Life, 2025).. In addition, we excluded domesticated species, particularly dogs (\u003cem\u003eCanis lupus familiaris\u003c/em\u003e) and cats (\u003cem\u003eFelis catus\u003c/em\u003e), from the dataset. The initial download comprised 63,959 occurrence records, which were reduced to 43,604 after cleaning and filtering the data.\u003c/p\u003e \u003cp\u003eTo assess mammal occurrence in urban areas, we integrated the cleaned occurrence dataset with urbanisation and human settlement layers in QGIS. We overlaid occurrence points onto high-resolution (10 \u0026times; 10 m) urban land-use maps derived from Sentinel-2 data and used spatial sampling tools in QGIS to determine whether the species records intersected with the mapped urban boundaries (Karra et al., \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Based on this spatial overlap, we classified each species as occurring or not occurring in the urban areas. Because mammal occurrence patterns in the Philippines remain incompletely documented, we did not attempt to assign species to finer categories of urban adaptation, such as urban avoiders, adapters, or exploiters (e.g. Santini et al. (\u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2019\u003c/span\u003e)). Instead, we used a conservative operational definition and considered \u0026ldquo;urban mammals\u0026rdquo; as species with at least one recorded occurrence within the urban boundaries of the Philippines.\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eKnowledge gap indicators\u003c/h3\u003e\n\u003cp\u003eWe accounted for research efforts across Philippine terrestrial mammals using Google Scholar hit counts and GBIF occurrence frequency records as indicators of knowledge levels and gaps for each terrestrial mammal species occurring in urban areas. The frequency of publications per species and observation records has been shown to serve as a reliable proxy for the research effort dedicated to a given species, and this approach has been applied in previous mammal studies in the Philippines (Tanalgo et al., \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). For this analysis, we searched Google Scholar (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://scholar.google.com.ph/\u003c/span\u003e\u003cspan address=\"https://scholar.google.com.ph/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) using species-specific queries to quantify how well each terrestrial mammal species in the Philippines has been studied. We expected species with more Google Scholar hits to reflect greater research efforts. We chose Google Scholar because it is open-access, requires no institutional subscription, and provides broader coverage of the published literature per species than Scopus and the Web of Science. We used the search syntax [\u0026ldquo;species X\u0026rdquo; + \u0026ldquo;Philippines\u0026rdquo;] following Tanalgo et al. (\u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2025\u003c/span\u003e) to retrieve all publications containing the species name for each species in our analyses. The search results were filtered to include only records from the Philippines. We log\u003csub\u003e10\u003c/sub\u003e-transformed all records for the analysis.\u003c/p\u003e\n\u003ch3\u003eExplanatory variables\u003c/h3\u003e\n\u003cp\u003eWe compiled species-level data for Philippine terrestrial mammals, including the IUCN Red List conservation status, country-level endemism, and life-history traits. We included body size, trophic level, diel activity, range size, and habitat breadth as intrinsic variables potentially influencing mammal occurrence in urban areas, extracted from Tanalgo et al. (\u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2025\u003c/span\u003e), along with forest height (Potapov et al., \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) and mean surface temperature as extrinsic variables representing landscape structure and climatic conditions.\u003c/p\u003e \u003cp\u003eWe expected that intrinsic traits would be strong predictors of mammal occurrence in urban areas. Large-bodied species were predicted to be less likely to occur in urban environments because they generally require larger home ranges and are more sensitive to habitat loss and human disturbances. In contrast, small-bodied mammals are expected to occur more frequently in urban areas because their size allows them to exploit fine-scale microhabitats, including tree holes, canopies, foliage, and built structures (Santini et al., \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Species with broader habitat breadths and wider geographic ranges are predicted to have higher urban occurrence, reflecting greater ecological flexibility (Tanalgo et al., \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). We further expected omnivorous and nocturnal species to fare better in urban environments because they can exploit diverse food resources and reduce their direct encounters with humans.\u003c/p\u003e \u003cp\u003eFor extrinsic variables, we predicted that greater surrounding forest cover would increase mammal occurrence, even within urban landscapes, by providing refuge and supplementary habitat. In contrast, higher surface temperatures are expected to reduce the occurrence of forest-dependent and thermally sensitive species, many of which are endemic to the Philippines and occupy narrow ecological niches in the forest understory.\u003c/p\u003e \u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003eData analysis\u003c/h2\u003e \u003cp\u003ePrior to formal analysis, all datasets were checked for statistical assumptions. First, we performed a Chi-square test of association (χ\u003csup\u003e2\u003c/sup\u003e) to determine whether there was a significant difference in the distribution of non-urban and urban mammals in terms of taxonomic groups, conservation status, and endemism. We modelled the probability of urban occurrence of mammal species using a generalised linear mixed model (GLMM) with a binomial error structure. The predictor variables included body mass, trophic level, diel activity, geographic range size, habitat breadth, forest canopy height and mean surface temperature. This framework allowed us to test whether variations in urban occurrence were associated with species traits, environmental conditions, and extent of species distribution.\u003c/p\u003e \u003cp\u003eWe further tested whether mammal species occurring in urban areas were disproportionately represented in the biodiversity knowledge, as reflected by publication outputs and occurrence records. To do so, we fitted a second set of GLMM with publication density and the number of GBIF records as separate response variables. The predictors included urban occurrence, extinction risk, and endemism. We expected species recorded from urban areas to show higher publication density and more GBIF records than species restricted to other habitat types, consistent with a bias towards taxa occupying more accessible, human-dominated landscapes. Continuous variables were \u003cem\u003ez\u003c/em\u003e-standardised to facilitate comparisons across predictors. For both models, taxonomic family was included as a random effect to account for phylogenetic structure, and model selection was performed using the Akaike Information Criterion (AIC), with the best-supported model identified as the one with the lowest AIC value.\u003c/p\u003e \u003cp\u003eAll statistical analyses were conducted using the open-source Jamovi software (version 2.7.17; The jamovi project, \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2026\u003c/span\u003e), and data visualisation was performed using GraphPad Prism (version 11; GraphPad Prism, 2022).\u003c/p\u003e \u003c/div\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003eDiversity and drivers of urban mammal occurrence\u003c/h2\u003e \u003cp\u003eWe examined the occurrence of Philippine terrestrial mammal species in urban areas and the factors associated with their presence in these areas. Of the 205 terrestrial mammals included in our analysis, 32% were recorded in urban areas across the country. Among mammalian orders, Chiroptera comprised the largest proportion of urban mammals (62%, n\u0026thinsp;=\u0026thinsp;41 species), followed by Rodentia (18.18%, n\u0026thinsp;=\u0026thinsp;12 species). Urban occurrence differed significantly across both orders (χ\u0026sup2; = 46.83, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001) and families (χ\u0026sup2; = 66.52, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001) \u003cb\u003e(\u003c/b\u003eFig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e\u003cb\u003e).\u003c/b\u003e\u003c/p\u003e \u003cp\u003eUrban occurrence varied significantly according to conservation status (χ\u0026sup2; = 12.97, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.024). Most urban species (71%, n\u0026thinsp;=\u0026thinsp;47 species) were classified as having a Least Concern. Notably, however, 15% of the species recorded in urban areas were threatened, including Vulnerable, Endangered and Critically Endangered taxa. A substantial proportion of urban mammals were endemic to the Philippines (48%, n\u0026thinsp;=\u0026thinsp;32 species), with significant differences in endemicity between urban and non-urban groups (χ\u0026sup2; = 10.97, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001). In terms of population trends, 41% (n\u0026thinsp;=\u0026thinsp;27 species) of urban mammals showed stable trends, whereas 33% showed decreasing trends (χ\u0026sup2; = 9.06, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.028).\u003c/p\u003e \u003cp\u003eWe built four models to predict the occurrence of terrestrial mammals in urban areas in the Philippines using biological, geographical, and extrinsic traits while accounting for discovery bias. Among these, the full model provided the best fit to the data (AICc\u0026thinsp;=\u0026thinsp;163.479). Binomial GLMMs showed that herbivorous species were more likely to occur in urban areas than carnivores and omnivores (β\u0026thinsp;=\u0026thinsp;1.553, p\u0026thinsp;=\u0026thinsp;0.013) and species with larger geographical ranges (β\u0026thinsp;=\u0026thinsp;1.393, p\u0026thinsp;=\u0026thinsp;0.008). Among the extrinsic variables, urban mammals were also associated with areas of higher forest canopy height (β\u0026thinsp;=\u0026thinsp;1.974, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001) \u003cb\u003e(\u003c/b\u003eFig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e\u003cb\u003e).\u003c/b\u003e\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eRelationship of urban occurrence and knowledge gaps\u003c/h3\u003e\n\u003cp\u003eWe tested whether mammal occurrence in urban areas contributed to taxonomic bias in Google Scholar hits and GBIF occurrence. Urban occurrence was positively associated with both Google Scholar hits (β\u0026thinsp;=\u0026thinsp;0.460, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001) and GBIF occurrence records (β\u0026thinsp;=\u0026thinsp;0.998, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), with a significantly stronger effect on the latter than on the former. These results suggest that species occurring in urban areas are generally more detectable and better documented across their ranges, rather than simply being more frequently recorded within cities. The interaction between non-endemic status and urban occurrence was significant only for the GBIF records (β\u0026thinsp;=\u0026thinsp;0.360, p\u0026thinsp;=\u0026thinsp;0.048) (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ea).\u003c/p\u003e \u003cp\u003eExtinction risk was associated only with GBIF records (β = \u0026minus;0.583, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), demonstrating that less threatened species tended to have more records than threatened taxa. Endemism was associated with Google Scholar hits, with non-endemic species tending to receive more citations than endemic species (β\u0026thinsp;=\u0026thinsp;0.613, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eUrbanisation is expected to intensify globally, yet its effects on biodiversity remain poorly understood across many taxonomic groups and regions, particularly in the Philippines. Here, we provide the first nationwide assessment of urban mammal occurrence in the country using publicly available records from the Global Biodiversity Information Facility (GBIF). By integrating biological, geographical, and extrinsic predictors, we established a baseline for understanding mammal distributions in urban landscapes and evaluated whether urban occurrence shapes the existing knowledge gaps in mammal occurrence data.\u003c/p\u003e \u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003eDiversity of urban mammals\u003c/h2\u003e \u003cp\u003ePublic observation data revealed that approximately one-third of terrestrial mammals in the Philippines are found in urban areas. We found significant differences in urban occurrence across both order and family levels, indicating that phylogenetic identity and the suite of life-history traits it encapsulates are significant determinants of urban occurrence than previously expected under random assembly. Our results are consistent with the growing evidence from urban mammal studies in other regions that phylogenetic conservatism in urban tolerance, mediated through traits such as body size, dietary breadth, and reproductive rate, produces non-random assemblage structures (Ritzel and Gallo, \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Santini et al., \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eBats were markedly overrepresented among Philippine urban mammals, comprising 62% of the species recorded. Our findings suggest that Philippine bats are highly tolerant of the effects of urbanisation compared with other mammalian groups (Jung and Threlfall, \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2018\u003c/span\u003e, \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Moretto et al., \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). For example, many Philippine bats, particularly pteropodid fruit bats and various insectivorous species, readily exploit anthropogenic structures for roosting, with bridges, building eaves, and urban canopies serving as functional analogues to natural caves and old-growth tree hollows (Dela Torre et al., \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2025\u003c/span\u003e; Tanalgo et al., \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Several interrelated traits explain the high affinity of bats for urbanisation. The high aerial mobility of many bats, especially cosmopolitan and smaller-bodied insectivorous bats, may substantially reduce commuting costs between urban matrices and adjacent natural habitats, allowing bats to exploit both simultaneously, a foraging flexibility that is unavailable to most ground-dwelling mammals (Avila-Flores et al., \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Jung and Kalko, \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Russo and Ancillotto, \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Some species of fruit bats forage in urban green areas over shorter distances and exploit food resources that are more readily available in the city (Meade et al., \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Egert-Berg et al. (\u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) hypothesizes that fruit bats might be seeking a refuge from predation and benefit from warmer micro-climates and easier navigation due to abundant landmarks in cities. Meanwhile, P\u0026aacute;ez et al. (\u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) proposed that habitat loss, which increases the time and energy costs required to search for a new foraging patch, and the availability of non-native, more constant food sources in urban areas may cause flying foxes to become residents in urban environments. Finally, light-induced insect aggregation in urban areas provides a reliable and predictable prey subsidy for common insectivorous species, potentially making urban environments more profitable than naturally lit forest habitats and increasing the likelihood of detecting more bat species in these settings (Avila-Flores and Fenton, \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Li and Wilkins, \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Schoeman, \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Nonetheless, the role of urban areas as alternative favourable foraging grounds for many bat species creates an ecological and evolutionary trap, which makes urban areas appear favourable but negatively affects bat populations and behaviour in the long term (Tanalgo et al., \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2025\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eAlthough rodents tend to dominate urban areas in terms of abundance, only 18% of rodent species have been recorded in urban habitats in the Philippines. Globally, commensal rodents, particularly those of the genus \u003cem\u003eRattus\u003c/em\u003e, are among the most ubiquitous and ecologically dominant urban mammals (Feng and Himsworth, \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Interestingly, although \u003cem\u003eR. rattus\u003c/em\u003e and \u003cem\u003eR. novergicus\u003c/em\u003e are common urban commensals worldwide (Feng et al., \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2025\u003c/span\u003e), they are poorly represented in urban GBIF records in the Philippines. This pattern is unlikely to indicate that species are rare in cities. Instead, it may reflect how biodiversity data are collected, with greater attention often given to species of ecological or conservation interest than to widespread pest species (Boakes et al., \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Bowler et al., \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). We argue that in urban areas, \u003cem\u003eR. rattus\u003c/em\u003e and \u003cem\u003eR. norvegicus\u003c/em\u003e may also be overlooked in surveys or left unreported because their presence is often expected. Consequently, its low representation in the GBIF probably reflects underreporting rather than a low occurrence, highlighting an important limitation of public occurrence data for describing urban mammal communities.\u003c/p\u003e \u003cp\u003eMoreover, rodent diversity in the Philippines is dominated by a rich radiation of endemic, forest-dependent species, many of which occupy narrow ecological niches in montane and old-growth forest habitats with no history of commensal association with humans (Heaney et al., \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Lawrence R. Heaney et al., \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Lawrence Richard Heaney et al., \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Our study shows that rodent species represented in Philippine urban assemblages are likely taxonomically and ecologically distinct, and the presence of native species in urban areas may reflect incidental habitat use or range compression into disturbed landscapes rather than functional synanthropy in urban areas.\u003c/p\u003e \u003cp\u003eOur findings indicate that urban landscapes tend to favour generalist and synanthropic small mammals, whereas forest-dependent species decline, resulting in reduced community diversity, which explains the patterns observed in our study.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec12\" class=\"Section2\"\u003e \u003ch2\u003eEcological status of urban mammals\u003c/h2\u003e \u003cp\u003eThe predominance of least threatened species among urban mammals (71%) is expected, as they are often characterised by broad distributions, large population sizes, and generalist ecological requirements that are congruent with their tolerance of anthropogenic disturbance (Wells et al., \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). The pattern observed in our study suggests that mammal records in urban areas are dominated by species that are less threatened and have broader geographical ranges. This supports our initial hypothesis that urbanisation tends to favour cosmopolitan species that are generally more adaptable to human-modified environments than other species.\u003c/p\u003e \u003cp\u003eInterestingly, at least 15% of urban species are considered threatened according to the IUCN Red List, suggesting that urban areas may serve as both refuges and threats to terrestrial mammals in the Philippines. In tropical biodiversity hotspots subjected to rapid land-use change, urban and peri-urban habitats can serve as \u003cem\u003ede facto\u003c/em\u003e refuge environments for species that can no longer maintain viable populations in their surrounding landscapes (Boakes et al., \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Maclagan et al., \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Tanalgo et al., \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). In the Philippine archipelago, where forest cover loss has been among the most severe globally, the presence of threatened mammals in cities is likely a symptom of widespread habitat contraction rather than an indicator of urban ecological suitability. Distinguishing between these two processes, genuine urban adaptation and forced habitat use under landscape stress, is empirically important, but we acknowledge that this cannot be resolved from occurrence data alone and warrants targeted demographic and behavioural investigations.\u003c/p\u003e \u003cp\u003eFurthermore, the significantly high proportion of Philippine endemics among urban mammals is particularly intriguing in this study. The Philippines harbours the highest density of endemic non-volant mammal species per unit area of any country globally (Heaney et al., \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Heaney et al., \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), and some endemic representation in urban assemblages is therefore not unexpected. However, the statistical significance of this pattern and its co-occurrence with a substantial proportion of declining species further suggest a dynamic that is more consistent with habitat compression than with urban adaptation. Endemic species in the Philippines are disproportionately forest-dependent and have experienced severe habitat loss across their limited geographic ranges (Tanalgo et al., \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2025\u003c/span\u003e); therefore, urban occurrence may represent the residual persistence of such species in structurally complex fragments of urban green spaces rather than the active colonisation of novel urban environments (Gallo et al., \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Maclagan et al., \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Under this interpretation, the apparently high endemic richness of Philippine urban areas should be viewed not as a conservation opportunity, but as a biodiversity risk indicator, a signal that endemic taxa are being pushed into increasingly marginal and structurally impoverished habitats. For example, a recent global analysis showed that approximately 33% of studies reported the presence of species worthy of conservation, such as rare, endemic, or endangered species, in a total of 365 species within urban refugia (Gentili et al., \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Our findings further highlight the need to avoid viewing urban ecosystems as biologically irrelevant, especially in tropical regions, where urban expansion often overlaps with areas of high endemicity.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec13\" class=\"Section2\"\u003e \u003ch2\u003eDrivers of urban mammal occurrence\u003c/h2\u003e \u003cp\u003eOur models identified herbivory, large geographic range size, and forest canopy height as significant positive predictors of urban mammal occurrence in the Philippines, with the full model providing the best fit. Contrary to our expectations, body size did not significantly affect the occurrence of urban areas in this study. The positive effects of herbivory can be ecologically interpreted in the context of the Philippine urban landscape, where ornamental fruiting trees in parks, roadsides, and private gardens constitute a persistent and predictable food resource accessible to frugivorous species, particularly to pteropodid bats (Meade et al., \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Unlike animal-based food resources, which are spatially variable and ephemeral in urban environments, fruiting trees represent temporally structured but spatially stable resources that can support regular and repeated exploitation (Lim et al., \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; P\u0026aacute;ez et al., \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2018\u003c/span\u003e, \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Although we argue that the dominance of herbivorous bats and rodents among Philippine urban mammals likely drives this relationship, it should not be generalised to imply that herbivory is broadly advantageous in urban mammal assemblages, where diverse feeding guilds exhibit complex and context-dependent urban responses.\u003c/p\u003e \u003cp\u003eThe positive association between geographical range size and urban occurrence is consistent with macroecological predictions and aligns with the results of global and regional syntheses of mammal ecology. Wide-ranging species typically exhibit broad environmental tolerances, high dispersal capacities, and generalist resource use, which function as pre-adaptations to urban environments rather than as responses specifically selected for these conditions (Sol et al., \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Range size may also serve as a proxy for population size and persistence capacity, such that wide-ranging species are more likely to maintain viable subpopulations across heterogeneous landscapes, including urban patches (Santini et al., \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eMoreover, the positive association between the persistence of mammals and the surrounding forest canopy in urban areas is ecologically plausible and practically important. We argue that rather than portraying Philippine urban mammals as species tolerant of simplified habitats, our results suggest that mammal richness in cities is linked to the vertical complexity of urban vegetation in degraded ecosystems (Camargo et al., \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). In urban areas, tall tree canopies may provide temporary refuge as roosting substrates, thermal buffering, shelter from disturbance and predators, and access to flowering and fruiting resources across canopy layers, which can potentially recruit mammal species in urban areas (Antoniazzi et al., \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2026\u003c/span\u003e; Tratalos et al., \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). Our findings underscore the importance of retaining substantial tree cover in and around urbanised areas to support a wider range of species and preserve their ecological roles across transitional landscapes (Jovanović et al., \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2025\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec14\" class=\"Section2\"\u003e \u003ch2\u003eKnowledge bias in urban species\u003c/h2\u003e \u003cp\u003eWe found a strong positive relationship between urban mammal occurrence and both Google Scholar publication density and GBIF occurrence records, indicating that species recorded in urban areas also tend to be more widely represented across other habitat types in the country. Rather than reflecting urban occurrence alone, this pattern suggests that mammals capable of extending into urban landscapes are generally those with broader ecological distributions and wider realised habitat use. Such species are likely to possess traits linked to environmental tolerance and flexibility, allowing them to persist in human-dominated settings and across a wider range of nonurban environments. Consequently, their occurrence is more frequently documented in both the scientific literature and biodiversity databases. This creates an important bias in biodiversity knowledge, whereby species that include urban areas within their distribution appear better known overall, whereas species confined to less accessible or less modified habitats are comparatively underrepresented. Such an imbalance may distort conservation priorities by overstating the apparent knowledge base for habitat-flexible species and obscuring the status and data limitations of more restricted taxa.\u003c/p\u003e \u003cp\u003eMost of the urban mammal species recorded in the Philippines were least threatened, as threatened and endemic species are more often associated with forested, less disturbed, or remote habitats, which are often difficult to sample because of access and security (Hilario-Husain et al., \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Tanalgo et al., \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). We further contend that the lower number of urban records for threatened and endemic mammals may reflect both sampling biases of these species and ecological filtering by urbanisation, where only species with wider range sizes that can withstand urban conditions persist and are recorded. Overall, our findings indicate that urban mammal assemblages are shaped by a combination of species\u0026rsquo; ecological tolerance, range characteristics, and conservation status, and are influenced by spatial biases inherent in biodiversity observations.\u003c/p\u003e \u003cp\u003eOur findings have practical implications for the interpretation and weighting of biodiversity data for conservation planning in the Philippines and beyond. Distribution models, species richness maps, and extinction risk assessments calibrated primarily on existing occurrence data will implicitly over-represent urban-tolerant, non-endemic, and least-concern taxa at the expense of ecologically distinctive fauna that confer global conservation significance to the archipelago (Hughes et al., \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Correcting for this bias will require deliberate redistribution of survey effort toward historically undersampled regions and taxonomic groups, particularly bats, which dominate urban assemblages and likely generate the strongest urban sampling artefact in the Philippine mammal data. This will also require investment in data infrastructure that can accommodate taxonomic uncertainty and track occurrence data for recently described and poorly known endemic taxa (Tanalgo et al., \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Tanalgo, \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2025a\u003c/span\u003e, \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2025b\u003c/span\u003e). The significant relationship between urban mammal occurrence and data abundance identified here should serve as an empirical prompt for such rebalancing rather than as a justification for concentrating future research in environments with the greatest existing knowledge.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec15\" class=\"Section2\"\u003e \u003ch2\u003eSynthesis\u003c/h2\u003e \u003cp\u003eOur results suggest that urbanisation in the Philippines is not merely restructuring mammal communities but also reshaping how biodiversity is detected, recorded, and interpreted. Urban environments appear to favour mobile, generalist, and disturbance-tolerant species, whereas many endemic, threatened, and forest-dependent mammals remain ecologically marginal and poorly represented in available datasets. This pattern implies that urban biodiversity data may capture not only biological responses to land-use change but also the uneven visibility of species across human-altered landscapes. Consequently, species that persist in cities are more likely to be observed, recorded, and studied, whereas those that are most vulnerable to urban expansion may be systematically overlooked.\u003c/p\u003e \u003cp\u003eWe also acknowledge the important limitations of our study, particularly in the classification of urban species and the spatial extent used for defining urban areas. These choices may affect which species are considered urban-associated and, in turn, the patterns inferred from the analyses. Future studies would benefit from more standardised and ecologically explicit definitions of urban habitats across the urban\u0026ndash;rural gradient. A more balanced assessment of Philippine mammal biodiversity will also require targeted sampling beyond major urban centres, especially in undersurveyed habitats and among poorly known endemic lineages, to better resolve how intrinsic and extrinsic factors shape species persistence in urban landscapes. Our analysis was also limited to presence\u0026ndash;absence observations and therefore, cannot fully capture the broader ecological responses of mammals to urbanisation. We call for more field-based studies and systematic observations aimed at understanding species persistence and responses to urban environments. Such efforts are essential for fully assessing the impacts of urbanisation on mammal species and for informing more effective urban planning and conservation strategies.\u003c/p\u003e \u003cp\u003eNonetheless, our current work further underscores the value of openly accessible biodiversity databases, particularly the Global Biodiversity Information Facility (GBIF), in revealing the broad patterns of species occurrence in urban landscapes (Sweet et al., \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). By aggregating records across taxa, regions, and time, these datasets provide an important foundation for detecting how biodiversity is distributed in human-dominated environments and how species respond to ongoing environmental changes (Li et al., \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Sultana and Storch, \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Although these records are subject to biases in sampling and detectability, they offer an essential starting point for large-scale urban biodiversity research, particularly in regions where systematic field surveys are limited (Beck et al., \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Hughes et al., \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Thus, GBIF and similar platforms can serve as critical baseline resources for advancing urban ecology, guiding hypothesis development, identifying knowledge gaps and informing future field-based investigations. More broadly, biodiversity knowledge from urban areas in the Philippines remains far more limited than that from other habitat types (Olfato-Parojinog and Dagamac \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), with mammals being particularly underrepresented. To help narrow these gaps, we outline several priorities for strengthening biodiversity research in urban landscapes in the Philippines \u003cb\u003e(\u003c/b\u003eFig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e\u003cb\u003e)\u003c/b\u003e.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eImportantly, the occurrence of conservation-relevant species in urban areas should not be interpreted as evidence of successful adaptation to urban environments. Instead, it may reflect habitat constriction, whereby species are forced into increasingly fragmented and degraded refuges embedded within expanding urban matrixes. In this context, we argue that urban areas may function as temporary refuges for some mammals in the Philippines, but such persistence may mask deeper ecological instability. As the human population in the Philippines continues to grow, the conversion of previously intact ecosystems into urban landscapes is likely to intensify, generating novel ecosystems to which many mammal species must adapt to. We argue that urbanisation may also generate ecological and evolutionary traps, in which apparently suitable habitats expose mammals to chronic disturbances, novel risks, and reduced long-term fitness (Gentili et al., \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Russo et al., \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Tanalgo et al., \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Thus, the urban presence of mammals, particularly ecologically unique species, may represent not only resilience but also a more fragile and transient form of persistence under increasing anthropogenic pressures.\u003c/p\u003e \u003c/div\u003e "},{"header":"Declarations","content":"\u003cdiv id=\"Sec16\" class=\"Section2\"\u003e \u003ch2\u003eData Availability Statement\u003c/h2\u003e \u003cp\u003eData used for this study is available in \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.15468/dl.6ju8sc\u003c/span\u003e\u003cspan address=\"10.15468/dl.6ju8sc\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec17\" class=\"Section2\"\u003e \u003ch2\u003eEthics Declaration\u003c/h2\u003e \u003cp\u003eNot applicable\u003c/p\u003e \u003c/div\u003e\u003cp\u003e \u003ch2\u003eConflict of Interest\u003c/h2\u003e \u003cp\u003eThe authors declare no conflict of interest.\u003c/p\u003e \u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAgduma AR, Garcia FG, Cabasan MT, Pimentel J, Ele RJ, Rubio M, Murray S, Hilario-Husain BA, Cruz D, Abdullah KC, Balase S, Tanalgo SM, K.C (2023) Overview of priorities, threats, and challenges to biodiversity conservation in the southern Philippines. 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Biodivers Conserv 23:2289\u0026ndash;2303. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s10531-014-0723-5\u003c/span\u003e\u003cspan address=\"10.1007/s10531-014-0723-5\" 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":true,"hideJournal":true,"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":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Conservation, Island ecosystems, Tropics, Urban Systems, Wildlife","lastPublishedDoi":"10.21203/rs.3.rs-9367707/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-9367707/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eUrbanisation is rapidly reshaping mammal communities; however, its effects remain poorly understood in tropical archipelagos with high endemism, such as the Philippines. Using 43,604 publicly available GBIF records, we examined the patterns and predictors of terrestrial mammal occurrence in urban areas of the Philippines while accounting for sampling and knowledge biases. Of the 205 mammal species, 66 (32%) were recorded in urban areas, indicating that urban assemblages represent a substantial but taxonomically uneven subset of mammalian species in the Philippines. Urban occurrence differed significantly across orders and families, with bats being strongly overrepresented, indicating a phylogenetically structured urban tolerance. Species were more likely to occur in urban areas when they had larger geographic range sizes, herbivorous diets, and were associated with landscapes with taller forest canopies, highlighting the role of ecological generality and vegetation structure in urban persistence. Although urban assemblages were dominated by widespread and least threatened species, they also included threatened, endemic, and declining taxa, suggesting that some records reflect persistence under habitat contraction rather than true adaptation to urban environments. Our findings further showed that species occurring in urban areas tended to have more publications and observation records. Our findings indicate that urban mammal assemblages are shaped by species\u0026rsquo; ecological tolerance, range characteristics, and conservation status, but are also filtered through spatial biases inherent in biodiversity observations and documentation.\u003c/p\u003e","manuscriptTitle":"Making it in the city: Public data reveal patterns and drivers of terrestrial mammal occurrence across urban Philippines","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-05-05 06:42:41","doi":"10.21203/rs.3.rs-9367707/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"c475674d-79b7-4caa-ae6f-7170fed0821a","owner":[],"postedDate":"May 5th, 2026","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[{"id":66029580,"name":"Conservation Biology"},{"id":66029581,"name":"Terrestrial Ecology"}],"tags":[],"updatedAt":"2026-05-05T06:42:41+00:00","versionOfRecord":[],"versionCreatedAt":"2026-05-05 06:42:41","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-9367707","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-9367707","identity":"rs-9367707","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}

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