Beyond green space area: How landscape structure shapes urban bird communities in a megacity

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Lagunes-Díaz, Jonathan Morales-Contreras, and 1 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-6858525/v1 This work is licensed under a CC BY 4.0 License Status: Under Revision Version 1 posted 9 You are reading this latest preprint version Abstract Urbanization poses a significant threat to biodiversity, with urban green spaces providing crucial habitats for birds within cities. However, studies have often focused on site-scale variables when assessing urbanization effects on birds, neglecting the role of landscape structure on shaping bird communities. In this sense, the effects of landscape structure on birds requires further research in urban environments. We aimed to assess how landscape structure influences bird diversity in public green spaces across Mexico City. We surveyed birds in 20 public green spaces and estimated bird abundance, diversity and evenness for each study site. We estimated landscape composition and configuration metrics and analyzed their effects on bird responses at multiple spatial scales (100–900 m radius). We recorded 1,194 individual birds from 56 species during our surveys, where bird assemblages were dominated by few synanthropic bird species. Landscape composition was more important than landscape configuration for birds. Deforested landscapes promoted bird abundance, while lower forest and grass cover enhanced bird species richness and diversity. Evenness was most strongly influenced by urban landscapes with higher grass cover and lower proportions of pavement and built-up areas. Our findings highlight the importance of integrating landscape-scale ecological principles into urban planning to enhance avian diversity and promote sustainable urban ecosystems. Prioritizing landscapes with high forest (native and non-native) and grass cover, and minimizing and managing impervious surfaces, could improve the quality of urban green spaces for birds and the environmental services they provide. Urban bird assemblages public parks urban landscape pattern multi-scale approach Figures Figure 1 Figure 2 Figure 3 INTRODUCTION The replacement of native habitats within the urbanization process lead to biodiversity decline, species extinction (Sun et al. 2024 ) and biotic homogenization (Devictor et al. 2007 ; Croci et al. 2008 ). Despite the many negative impacts of urbanization on avian assemblages (reviewed in Chace and Walsh 2006 ), several studies have documented the importance of urban green spaces for retaining an important proportion of the world’s avifauna (Aronson et al. 2017 ; Ibáñez-Álamo et al. 2017 ; MacGregor-Fors et al. 2021 ), providing important habitat and refuge for birds within cities (Fernández-Juricic and Jokimäki 2001 ; Vasquez and Wood 2022 ). Overall, studies in urban environments have demonstrated that bird species richness decreases with urbanization while bird density may increase (e.g. Marzluff 2001 , Chace and Walsh 2006 , van Heezik et al. 2008 ). Multiple studies conducted at the site scale (i.e. measuring environmental variables within urban green space) reported higher bird abundance and richness with larger urban green space (Gavareski 1976 ; Evans et al. 2009 ; Dale 2018 ; Leveau et al. 2022 ; Thompson et al. 2022 ) conducting to the generalization that larger urban green spaces contain higher bird diversity. However, the majority of studies have centered on urban green space local predictors like size (Gavareski 1976 ; Dale 2018 ; Leveau et al. 2022 ; Thompson et al. 2022 ) or evaluating gradients of urbanization (Xie et al. 2019 ), neglecting the influence of landscape structure in shaping avian assemblages. At the landscape scale (i.e. measuring environmental variables beyond the limits of the urban green space), the amount of woody cover or tree cover is an important predictor for the presence of many bird species (Mörtberg 2001 ; Melles et al. 2003 ) and positively relates to bird species richness and abundance (Ikin et al. 2013 ; Callaghan et al. 2018 ; Chong et al. 2019 ; Villaseñor et al. 2021 ; Mao et al. 2023 ). Species richness and functional diversity are negatively affected by increasing sealed cover (Schütz and Schulze 2015 ) and construction land (Mao et al. 2023 ) in the landscape, whereas the connectivity of urban green spaces enhanced the abundance of different birds (Pellissier et al. 2012 ; Kang et al. 2015 ) and species richness (Oliver et al. 2011 ). Overall, although local-site variables are highly important for birds, their abundance and diversity in urban ecosystems also depend on patterns (e.g. landscape forest cover, impervious surface) and processes (e.g. dispersion) operating at landscape scales. Implementing urban planning strategies and policymaking that reduce or reverse the negative impacts of the urbanization process on biodiversity is imperative, and those strategies rely on the understanding of the effects of the urban landscape pattern on biodiversity. Here, we estimated bird abundance, species richness, diversity, and assemblage evenness in urban greens paces across Mexico City, and examined the influence of landscape structure on bird responses. Since biological responses to landscape structure can be scale-dependent, and, as we do not know a priori the best spatial scale of bird responses to landscape structure (the so-called ‘scale of effect’, Jackson and Fahrig 2015 ), we used a multi-scale analysis and measured landscape variables at different nested scales surrounding study sites. We predicted that landscape composition is more important than landscape configuration for birds. We also expected that species richness and diversity would be highly and positively associated to forest cover in the urban landscape, given the importance of such land cover for birds. To our knowledge, few studies at the landscape scale have used a multi-scale approach when evaluating the influence of landscape structure on birds within urban ecosystems (but see Huang et al. 2015 , Callaghan et al. 2018 ). METHODS Our study was carried out in Central Mexico in Mexico City (19°35'34.08" N, 99°21'53.64" W and 19°02'53.52" N, 98°56'25.08" W, 1494.3 km 2 ). Mean annual temperature is 16ºC and average altitude is 2240 masl and is surrounded by mountains (Carrera-Hernández and Gaskin 2007 ). The urbanized land cover occupies most of the territory, but remnants of native vegetation (coniferous forests, wetlands, pastures) are still present in the south of the city (PAOT 2018 ). This megacity (~ 21 million people, INEGI 2020 ) has considerable amounts of public green areas. By 2017, the city had 67.3 km 2 of urban green spaces (SEDEMA 2020 ). Study site selection To select study sites, we used a map of the study area and use the database of urban green spaces of Mexico City (SEDEMA 2020 ). Then we selected 20 public green spaces to carry out bird surveys (Fig. 1 ). We selected those urban green spaces of public access with no time restriction for public entry and composed mainly of forested vegetation (i.e. high cover of trees), as many public green spaces were almost treeless. Selected urban green spaces were separated at least 2 km from each other, this to avoid overlapping of influencing landscape metrics and to ensure spatial independence of samples. Northeast side of the study area was underrepresented because the smaller number of forested public green spaces (Fig. 1 ). Bird surveys We conducted point counts (Bibby et al. 2000 ) to sample birds in each of the selected urban green spaces. We recorded all birds seen or heard up to a 50 m radius during 15 min. The number of point counts at each urban green space was proportional to its size (1–2 point counts per site), and the points were separated at least 200 m (Bibby et al. 2000 ). All sites were visited once in the morning (20 min after dawn) only on working days (Monday-Thursday) from February to April 2024. Urban landscape composition and configuration We performed a supervised classification for characterizing landscape features at our study area: we used two images from the Planet remote sensing platform, from the “Tropical Normalized Analytic Monthly series basemaps” (PBC 2024), made available through the Norway’s International Climate and Forests Initiative (NICFI). We selected one image for August 2023, the image for the most recent rainy season, when plant phenology allows for better differentiation of its spectral signatures from other classes. We visually selected points for sampling the different land use and land cover classes on a Google Satellite image (Google 2024 ), using the Quickmapservices plugin for the QGIS software (QGIS Development Team 2024 ). The classes we used for the sampling were: asphalt, built-up, water, grass and forest; to avoid mixing in very different spectral signatures, thus reducing noise in the training phase, we subdivided built-up classes into concrete, waterproofed concrete and metal-roof. At least 15 samples were selected from each class. We then used the modules terra (Hijmans 2023 ) for raster processing, randomForest (Liaw and Wiener 2002 ) for classification with the random forest algorithm and caret (Kuhn 2008 ) for predictive model building and analysis, in the R statistical computing language (R Development Core Team 2022 ). The selected image had high accuracy, with a benchmarked median Kappa of 0.925. On the classified image, we used the module landscape metrics (Hesselbarth et al. 2019 ) and calculated forest (ha), pavement (ha), grass (ha) and built (ha) cover as variables of landscape composition and estimated the density of patches of forest patches and the mean distance between forest patches as variables of landscape configuration. It should be noted that the coverage called "forest" does not correspond to a native forest but to forest structure vegetation that may be composed of native and/or exotic tree species. We calculated each of the landscape metrics at nine concentric buffers (i.e. nine spatial scales, radius 100-900m, by 100m) from the geographic center of each study site to evaluate the spatial scale of effect (Jackson and Fahrig 2012 ). Bird abundance, diversity and assemblage structure For diversity estimates, we first checked the sample completeness of each forest site using the sample coverage index suggested by Chao and Jost ( 2012 ). The sample coverage index represents the percentage of the total number of individuals in an assemblage that belongs to the species found in the sample (Chao and Jost 2012 ). Since sample coverage varied among study sites, ranging from 0.68 to 0.95 (Table S1 ), we estimated species diversity by using both observed and expected data based on the rarefaction and extrapolation procedure suggested by Chao and Jost ( 2012 ) using the iNEXT package for R (Hsieh et al. 2022 ). Specifically, we estimated Hill numbers of order 0 and 1 as measures of bird diversity ( 0 D , 1 D , hereafter). 0 D (i.e. species richness) is not sensitive to species abundance, only presences are counted, whereas 1 D (the exponential of Shannon’s entropy index) weights species in proportion to their frequency (Chao et al. 2014 ), and it can be interpreted as the number of common or typical species in the assemblage (Jost 2006 ). We considered the abundance of each species to construct rank-abundance plots of each of the study sites (Fig. 2 ) and estimated the evenness factor proposed by Jost ( 2010 ). The evenness factor EF represents the proportion of dominant species in the assemblage, it ranges between 1 (a perfectly even assemblage), whereas values near to 0 would a highly dominated assemblage (Jost 2010 ). Estimation of the scale of landscape effect We performed Generalized Linear Models following a multiscale approach to first detect the so called “scale of effect” (Jackson and Fahrig 2012 ), this is the spatial scale that yields the strongest species-landscape relationship. We quantified the relationship between each landscape metric considered in this study (i.e. forest cover, grass cover, pavement cover, built cover, density of forest patches, mean distance between forest patches) and each response variable (i.e. abundance, 0 D , 1 D , EF ) at each spatial scale (nine scales, from 100-900m radius). We used Gaussian distribution error for continuous variables (i.e. 1 D , EF ) and count variables of abundance and 0 D were modeled with a Poisson distribution error (Crawley 2007 ). Then, we plotted the value of AICc as a dependent variable against landscape size to identify the scale that best yields the best-fitted associations between each response variable and each landscape predictor. We estimated a total of 30 scales of effect values (5 response variables x 6 landscape metrics; Table S2) selected by the lowest sample-corrected Akaike information criterion (AICc) value among sets of models (Table S3). Influence of landscape structure on bird abundance and diversity We conducted Principal Component Analysis (PCA) of the six-landscape variables measured at the identified scale of effect (Table S2) to reduce multidimensional landscape metrics into a reduced number of indices. Since the spatial scale of effect was different for each combination of response and predictor variables Table S3, we conducted one PCA for each response variable (Fig S1 -S5). In all cases, we retained the first three components since they accounted for ~ 85% of the total variance in each data set (Table S4-S5). To evaluate the relative effect of landscape variables (contained in each of the three retained Principal Components) on each response variable (abundance, 0 D , 1 D, EF ) we used an information-theoretic approach and multi-model inference (Burnham and Anderson 2002 ). For each response variable we performed 16 models, representing all combinations of the predictors (i.e. three PCA axes for each case) and the null model (intercept only). Then, we classified the models from the lowest (the best-supported model) to the highest (the least-supported model) considering their AICc value. After, we summed the Akaike weights of each PCA component appearing in the models, which is considered a proxy of the relative importance of each PCA component on each response variable (Burnham and Anderson 2002 ; Giam and Olden 2016 ). We found no collinearity among the multiple models (VIF < 2) after checking for the Variance Inflation Factor (VIF) for each predictor. We used a normal distribution for 1 D , EF , and Poisson distribution for abundance and 0 D data. All analyses were conducted in R (R Development Core Team 2022 ). RESULTS Bird assemblages in urban green spaces We recorded a total of 1,194 individuals from 56 species and 23 families (Table 1). Most of the identified bird species were resident breeders in Mexico City (26 species, 65.4%). A large proportion were migratory, non-breeding species (15 species, 27.2%), while the remaining identified birds (4 species, 7.2%) were alien species (Table 1). The three most abundant species were generalist, conspicuous urban residents: the Inca Dove (181 individuals), the introduced and now feral Rock Pigeon (168 individuals), and the opportunistic House Sparrow (156 individuals), each accounting for more than 150 individuals. All three dominated most bird assemblages (Table 1, Fig. 2). Next, bird assemblages were dominated by other abundant species that had expanded their ranges significantly and have invasive and acclimatization abilities: the House Finch (73 individuals), the White-winged Dove (67 individuals), and the Great-tailed Grackle (66 individuals). Additionally, we recorded high abundances of the common and widespread Yellow-rumped Warbler (63 individuals) and the Rufous-backed Robin (63 individuals) (Table 1, Fig. 2). Other bird species showed abundances ranging from 5 to 25 individuals (Table 1), including migratory species (e.g., American Robin, Cassin’s Vireo, Nashville Warbler, Wilson’s Warbler; Table 1), hummingbirds (Beryllina Hummingbird, Broad-billed Hummingbird), and non-native invasive species (Eurasian Collared-Dove, Monk Parakeet) among others (Table 1). Finally, species with low abundances included raptors (Accipitridae, Falconidae), migrants (Parulidae) and resident birds (Tyrannidae), among others (Table 1). We observed that half of bird assemblages (50%) within public green spaces exhibited intermediate evenness (EF = 0.5-0.7; Fig. 2), whereas 40% (8) or urban green spaces showed highly dominated assemblages (EF < 0.5; Fig. 2), and only 10% (2) of the studied sites exhibited more even assemblages (EF < 0.7; Fig. 2). Influence of landscape structure on bird assemblages Landscape composition had a greater influence on bird responses than landscape configuration, with forest patch density being the only configuration metric that affected bird assemblages. The overall abundance of birds was negatively influenced by a linear combination of lower pavement cover and higher grass cover (PC2, Fig. 3a). In contrast, a combination of lower forest cover and higher built cover, along with greater forest patch density (PC1, PC3; Fig. 3a) positively influenced bird abundance. Bird species richness and the number of common bird species ( 1 D ) were both negatively and mainly affected by highly urbanized landscapes with lower proportion of forest and grass cover (PC2; Fig. 3b,c), and higher forest patch density (PC3: Fig. 3b,c). A higher proportion of built and pavement cover negatively influenced bird species richness and 1 D , but this predictor was less important (PC1; Fig. 3b,c). Finally, the evenness of bird assemblages was mainly and positively influenced by landscapes with lower pavement cover and high grass cover (PC3; Fig 3d), whereas landscapes with high built cover, and lower forest cover distributed in many fragments decreased evenness of bird assemblages (PC1, PC2; Fig 3d). DISCUSSION Bird assemblages Our results indicated that bird assemblages inhabiting public green spaces in Mexico City are dominated by a few synanthropic bird species that thrive in urban environments. These findings are consistent with other studies conducted in temperate (Donnelly and Marzluff 2004 ; Villegas and Garitano-Zavala 2010 ; Ikin et al. 2013 ), Mediterranean (Kurucz et al. 2021 ), subtropical (van Rensburg et al. 2009 ) and tropical (Ortega-Álvarez and MacGregor-Fors 2009 ; MacGregor-Fors and Ortega-Álvarez 2011a ; Chamberlain et al. 2017 ) cities globally, where dominant bird species are generalists better suited to exploit urban environments. Our findings also align with studies that report the dominance of a few exotic species and granivores in urbanized areas (reviewed in Chace and Walsh 2006 ; Kurucz et al. 2021 ) and within our study area (Charre et al. 2013 ). Altogether, these findings underscore the homogenizing effects of urbanization on bird assemblages, which have been observed in other urban environments (e.g. Devictor et al. 2007 ; van Rensburg et al. 2009 ; Sidemo-Holm et al. 2022 ), where a small number of generalist species adapt and thrive over more specialized native birds. More than 400 bird species have been reported for the Mexico City Metropolitan Area (Ramírez-Albores et al. 2024 ), with more specialized bird species found in higher numbers towards the city’s borders (MacGregor-Fors and Ortega-Álvarez 2011a ) where large extentions of native vegetation still surround the more urbanized areas (Arriaga et al. 2000 ). Nevertheless, the dominance of synanthropic and generalist bird species in urban green spaces raises concerns on the loss of biodiversity and ecosystem function within urbanized landscapes. The higher number of bird species documented for our study area (~ 400 species; I don´t lilRamírez-Albores et al. 2024 ) includes all types of urban green spaces such as university campuses, sports centers, ecological reserves, and wetlands. Similarly, other studies that report higher species richness (e.g. Charre et al. 2013 , Ramírez-Albores et al. 2024 , Oropeza-Sánchez et al. 2025 ) have compiled data from various sources or over multiple seasons. Our study, however focuses specifically on public green spaces, which are different from other green spaces like arboreta (e.g. Charre et al. 2013 ; Oropeza-Sánchez et al. 2025 ), private gardens, or controlled-access urban green areas. This suggests that urban public urban green spaces represent a relatively small proportion of the total bird species that inhabit the city. Although bird assemblages in public green spaces are primarily dominated by a few synanthropic species, we also recorded a notable proportion of migratory species. Of the 55 species identified, 15 were migratory, highlighting the importance of urban landscapes in providing habitat for migratory birds. This finding suggests that, despite the dominance of generalist species, public green spaces offer valuable resources for birds, particularly those requiring biodiversity-friendly infrastructure, such as large trees and forested areas (Amaya-Espinel and Hostetler 2019 ). These results underscore the potential for targeted management and restoration efforts to enhance public green spaces that support greater diversity of bird species. Influence of landscape structure on bird assemblages Our results demonstrated that highly deforested landscapes, with a high proportion of impervious surface and built cover, are associated with increased bird abundance in the city. This aligns with previous studies that reported higher bird abundance in more urbanized areas (Donnelly and Marzluff 2004 ; Ortega-Álvarez and MacGregor-Fors 2009 ; MacGregor-Fors and Ortega-Álvarez 2011b ; Chamberlain et al. 2017 ; Kurucz et al. 2021 ). Particularly, our results agree with studies that analyzed land cover types as predictors, showing increased bird abundance with higher built cover (Lim and Sodhi 2004 ; Amaya-Espinel et al. 2019 ). Overall, our findings confirm that urbanized areas can support high bird abundance, even though these assemblages are simplified and dominated by a few species. Landscape structure had contrasting effects on bird abundance, richness, and evenness. Richness and evenness were lower in deforested landscapes with high proportions of built cover, whereas bird abundance was higher in these areas. These results are consistent with previous studies that show lower bird richness in areas with greater building presence (Chamberlain et al. 2007 ), building height (Leveau and Leveau 2016 ), built cover (Carbó-Ramírez and Zuria 2011 ), higher building densities, and more road coverage (Amaya-Espinel et al. 2019 ). Although built infrastructure can provide important resources for birds (Mainwaring 2015 ; Partridge and Clark 2018 ) it does not constitute a biodiversity-friendly land cover that promotes bird species richness and evenness within our study area. In contrast, landscapes with higher grass cover and less pavement were associated with greater bird species richness and evenness, confirming that grass and lawns within urban landscapes function as a source of resources for birds (Sánchez-Sotomayor et al. 2023 ) and provide a more bird-friendly environment than impervious surface like built and pavement cover. Bird species richness followed the expected pattern, decreasing in landscapes with lower proportions of vegetation (both forest and grass cover) and higher proportions of impervious surfaces, adding to the broader trend of reduced bird species richness with increased urbanization (Melles et al. 2003 ; Chamberlain et al. 2017 ). Our results emphasize the importance of forest cover in maintaining overall bird species richness within cities (Villegas and Garitano-Zavala 2010 ; Ikin et al. 2013 ; Callaghan et al. 2018 ) particularly for native birds (Villaseñor et al. 2021 ). Furthermore, we observed the same response pattern for the number of common species in the bird assemblages, reinforcing the critical role of forested land cover in sustaining high bird diversity in urban environments. Implications for conservation Our findings highlight the significant role that landscape composition and configuration play in shaping bird assemblages in urban environments, underscoring the need for urban planning strategies that prioritize the expansion and maintenance of vegetation and biodiversity-friendly infrastructure across landscapes. Specifically, our results suggested that urban conservation efforts should focus on protecting and restoring forested areas and grasslands to enhance bird diversity. The positive responses of birds to forest cover in our study indicated a valuable opportunity to improve urban landscapes for avian populations. Notably, the forest cover in our study did not necessarily consist of native vegetation but included a mix of native and non-native trees, suggesting that even urban green spaces with non-native plant species can be improved by introducing a more complex forest structure to better support bird diversity. In addition, policies aimed at reducing impervious surfaces and promoting green infrastructure—such as green roofs, permeable pavements, and expanded green corridors—could significantly enhance habitat quality for birds in urban areas. The benefits of such measures extend beyond birds, contributing to the broader ecological health of urban environments. Given the homogenizing effects of urbanization on bird assemblages, it is essential to integrate ecological principles into urban planning. Enhancing connectivity between urban green spaces and surrounding natural landscapes, reducing impervious surface areas, and increasing vegetation cover could create more sustainable and resilient environments for both migratory and resident bird species. Such conservation efforts would not only benefit avian populations but also foster biodiversity and ecological resilience in the face of ongoing urbanization. Public green spaces within megacities, therefore, hold substantial potential for conserving bird assemblages and promoting a more sustainable, biodiverse urban environment. Declarations ACKNOWLEDGMENTS MMR received a postdoctoral scholarship from SECIHTI (Secretaría de Ciencia, Humanidades, Tecnología e Innovación) and support from Idea Wild. We are thankful to Oscar G. Brito for field assistance. FUNDING SECIHTI supported this work through a postdoctoral grant awarded to MMR. COMPETING INTERESTS The authors have no relevant financial or non-financial interests to disclose. AUTHOR CONTRIBUTIONS Marisela Martínez-Ruiz developed the study conception and design. Material preparation and data collection were performed by Marisela Martínez-Ruiz, Jonathan Morales-Contreras and Elio G. Lagunes-Díaz. Analyses were performed by Marisela Martínez-Ruiz. The first draft of the manuscript was written by Marisela Martínez-Ruiz and all authors commented on previous versions of the manuscript. All authors read and approved the final manuscript. DATA AVAILABILITY The datasets generated and analyzed during the current study are available from the corresponding author on reasonable request References Amaya-Espinel JD, Hostetler M, Henríquez C, Bonacic C (2019) The influence of building density on Neotropical bird communities found in small urban parks. 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Mexico City Jackson HB, Fahrig L (2015) Are ecologists conducting research at the optimal scale? Glob Ecol Biogeogr 24:52–63. https://doi.org/10.1111/geb.12233 Jackson HB, Fahrig L (2012) What size is a biologically relevant landscape? Landsc Ecol 27:929–941. https://doi.org/10.1007/s10980-012-9757-9 Jost L (2006) Entropy and diversity. Oikos 113:363–375. https://doi.org/10.1111/j.2006.0030-1299.14714.x Jost L (2010) The relation between evenness and diversity. Diversity 2:207–232. https://doi.org/10.3390/d2020207 Kang W, Minor ES, Park CR, Lee D (2015) Effects of habitat structure, human disturbance, and habitat connectivity on urban forest bird communities. Urban Ecosyst 18:857–870. https://doi.org/10.1007/s11252-014-0433-5 Kuhn M (2008) Building predictive models in r using the caret package. J Stat Softw 28:1–26. https://doi.org/10.18637/jss.v028.i05 Kurucz K, Purger JJ, Batáry P (2021) Urbanization shapes bird communities and nest survival, but not their food quantity. Glob Ecol Conserv 26:e01475. https://doi.org/https://doi.org/10.1016/j.gecco.2021.e01475 Leveau LM, Bocelli ML, Quesada-Acuña SG et al (2022) Bird diversity-environment relationships in urban parks and cemeteries of the Neotropics during breeding and non-breeding seasons. PeerJ 10:1–19. https://doi.org/10.7717/peerj.14496 Leveau LM, Leveau CM (2016) Does urbanization affect the seasonal dynamics of bird communities in urban parks? Urban Ecosyst 19:631–647. https://doi.org/10.1007/s11252-016-0525-5 Liaw A, Wiener M (2002) Classification and regression by randomForest. R News 2:18–22 Lim HC, Sodhi NS (2004) Responses of avian guilds to urbanisation in a tropical city. Landsc Urban Plan 66:199–215. https://doi.org/10.1016/S0169-2046(03)00111-7 MacGregor-Fors I, Escobar-Ibáñez JF, Schondube JE et al (2021) The urban contrast: A nationwide assessment of avian diversity in Mexican cities. Sci Total Environ 753:141915. https://doi.org/https://doi.org/10.1016/j.scitotenv.2020.141915 MacGregor-Fors I, Ortega-Álvarez R (2011a) Fading from the forest: Bird community shifts related to urban park site-specific and landscape traits. Urban Urban Green 10:239–246. https://doi.org/https://doi.org/10.1016/j.ufug.2011.03.004 MacGregor-Fors I, Ortega-Álvarez R (2011b) Fading from the forest: Bird community shifts related to urban park site-specific and landscape traits. Urban Urban Green 10:239–246. https://doi.org/10.1016/j.ufug.2011.03.004 Mainwaring MC (2015) The use of man-made structures as nesting sites by birds: A review of the costs and benefits. J Nat Conserv 25:17–22. https://doi.org/https://doi.org/10.1016/j.jnc.2015.02.007 Mao Q, Sun J, Deng Y et al (2023) Assessing Effects of Multi-Scale Landscape Pattern and Habitats Attributes on Taxonomic and Functional Diversity of Urban River Birds. Diversity 15(4):486. https://doi.org/10.3390/d15040486 Marzluff JM (2001) Worldwide urbanization and its effects on birds. In: Marzluff JM, Bowman R, Donnelly R (eds) Avian Ecology and Conservation in an Urbanizing World. Springer US, Boston, MA, pp 19–47 Melles S, Glenn S, Martin K (2003) Urban Bird Diversity and Landscape Complexity: Species-environment associations along a multiscale habitat gradient. Conserv Ecol 7(1):5. URL: http://www.consecol.org/vol7/iss1/art5/ Mörtberg UM (2001) Resident bird species in urban forest remnants; landscape and habitat perspectives. Landsc Ecol 16:193–203. https://doi.org/10.1023/A:1011190902041 Oliver A, Hong-Wa C, Devonshire J et al (2011) Aviafauna richness enhanced in large, isolated urban parks. Landsc Urban Plan 102:215–225. https://doi.org/10.1016/j.landurbplan.2011.04.007 Oropeza-Sánchez MT, Solano-Zavaleta I, Cuandón-Hernández WL et al (2025) Urban green spaces with high connectivity and complex vegetation promote occupancy and richness of birds in a tropical megacity. Urban Ecosyst 28:1–18. https://doi.org/10.1007/s11252-024-01612-3 Ortega-Álvarez R, MacGregor-Fors I (2009) Living in the big city: Effects of urban land-use on bird community structure, diversity, and composition. Landsc Urban Plan 90:189–195. https://doi.org/10.1016/j.landurbplan.2008.11.003 PAOT (2018) Las áreas verdes de la Ciudad de México, una visión integral. Centro PAOT, Mexico City Partridge DR, Clark JA (2018) Urban green roofs provide habitat for migrating and breeding birds and their arthropod prey. PLoS ONE 13:e0202298. https://doi.org/10.1371/journal.pone.0202298 PBC PL (2024) Planet application program interface: In space for life on earth Pellissier V, Cohen M, Boulay A, Clergeau P (2012) Birds are also sensitive to landscape composition and configuration within the city centre. Landsc Urban Plan 104:181–188. https://doi.org/https://doi.org/10.1016/j.landurbplan.2011.10.011 QGIS Development Team (2024) QGIS Geographic Information System. http://qgis.org R Development Core Team (2022) R: a language and environment for statistical computing Ramírez-Albores JE, Sánchez-González LA, Pérez-Suárez M et al (2024) Greenspaces as shelters for the conservation of bird diversity in a big city. Urban Ecosyst 27:2047–2059. https://doi.org/10.1007/s11252-024-01573-7 Sánchez-Sotomayor D, Martín-Higuera A, Gil-Delgado JA et al (2023) Artificial grass in parks as a potential new threat for urban bird communities. Bird Conserv Int 33:e16. https://doi.org/ Schütz C, Schulze CH (2015) Functional diversity of urban bird communities: effects of landscape composition, green space area and vegetation cover. Ecol Evol 5:5230–5239. https://doi.org/https://doi.org/10.1002/ece3.1778 SEDEMA (2020) Inventario de Áreas Verdes Ciudad de México. Secretaría del Medio Ambiente de la Ciudad de México, Mexico City Sidemo-Holm W, Ekroos J, Reina García S et al (2022) Urbanization causes biotic homogenization of woodland bird communities at multiple spatial scales. Glob Chang Biol 28:6152–6164. https://doi.org/https://doi.org/10.1111/gcb.16350 Sun B, Lu Y, Yang Y et al (2024) Urbanization affects spatial variation and species similarity of bird diversity distribution. Sci Adv 8:eade3061. https://doi.org/10.1126/sciadv.ade3061 Thompson R, Tamayo M, Sigurðsson S (2022) Urban bird diversity: does abundance and richness vary unexpectedly with green space attributes? J Urban Ecol 8:juac017. https://doi.org/10.1093/jue/juac017 van Heezik Y, Smyth A, Mathieu R (2008) Diversity of native and exotic birds across an urban gradient in a New Zealand city. Landsc Urban Plan 87:223–232. https://doi.org/https://doi.org/10.1016/j.landurbplan.2008.06.004 van Rensburg BJ, Peacock DS, Robertson MP (2009) Biotic homogenization and alien bird species along an urban gradient in South Africa. Landsc Urban Plan 92:233–241. https://doi.org/https://doi.org/10.1016/j.landurbplan.2009.05.002 Vasquez AV, Wood EM (2022) Urban parks are a refuge for birds in park-poor areas. Front Ecol Evol 10 Villaseñor NR, Escobar MAH, Hernández HJ (2021) Can aggregated patterns of urban woody vegetation cover promote greater species diversity, richness and abundance of native birds? Urban Urban Green 61:127102. https://doi.org/https://doi.org/10.1016/j.ufug.2021.127102 Villegas M, Garitano-Zavala Á (2010) Bird community responses to different urban conditions in La Paz, Bolivia. Urban Ecosyst 13:375–391. https://doi.org/10.1007/s11252-010-0126-7 Xie S, Xu YS W, et al (2019) The effect of habitat changes along the urbanization gradient for breeding birds: an example from the Xiong’an New Area. PeerJ 7:e7961. https://doi.org/https://doi.org/10.7717/peerj.7961 Table Table 1. List of bird species registered during 15 min point counts in 20 public parks of Mexico City from February to April 2024. Ab = Abundance; Habitat use: G = Generalist species, F = forest species; F-G = Species associated to forested habitats but able to exploit different land covers; Status: A = Alien, B = Breeding, NB = non-breeding. Habitat use for non-breeding birds refers to habitat identified for the species during the non-breeding season. Habitat use and status of birds within Mexico City was retrieved from Birds of the World (www.birdsoftheworld.org). Family Species Common name Ab Habitat use Status Accipitridae Accipiter striatus Sharp-shinned Hawk 2 F-G B Parabuteo unicinctus Harris's Hawk 7 G B Aegithalidae Psaltriparus minimus Bushtit 5 G B Cardinalidae Pheucticus melanocephalus Black-headed Grosbeak 3 G NB Piranga ludoviciana Western Tanager 5 G NB Piranga rubra Summer Tanager 8 G A Columbidae Columba livia Rock Pigeon 168 G B Columbina inca Inca Dove 181 G A Streptopelia decaocto Eurasian Collared-Dove 21 G B Zenaida asiatica White-winged Dove 67 G B Zenaida macroura Mourning Dove 1 G B Emberizidae Melospiza melodia Song Sparrow 5 G B Falconidae Falco sparverius American kestrel 2 G B Fringilidae Haemorhous mexicanus House Finch 73 G B Spinus psaltria Lesser Goldfinch 8 G B Icteridae Icterus abeillei Black-backed Oriole 5 G NB Icterus bullockii Bullock's Oriole 9 G NB Icterus cuculatus Hooded Oriole 3 G B Molothrus aeneus Bronzed Cowbird 3 G B Quiscalus mexicanus Great-tailed Grackle 66 G B Mimidae Toxostoma curvirostre Curved-billed Thrasher 10 G B Passerellidae Melozone fusca Canyon Towhee 17 G A Passeridae Passer domesticus House Sparrow 156 F-G B Picidae Dryobates scalaris Ladder-backed Woodpecker 6 F-G B Melanerpes formicivorus Acorn Woodpecker 1 F-G NB Sphyracus varius Yellow-bellied Sapsucker 2 F-G NB Parulidae Cardellina pusilla Wilson's Warbler 18 G NB Leiothlypis ruficapilla Nashville Warbler 25 F-G NB Leiothlypis virginiae Virginia's Warbler 1 G NB Mniotilta varia Black-and-white Warbler 6 F B Myioborus pictus Painted redstart 1 G NB Setophaga coronata Yellow-rumped Warbler 63 F-G NB Setophaga nigrescens Black-throated Gray Warbler 1 F NB Setophaga occidentalis Hermit Warbler 1 G B Setophaga petechia Yellow Warbler 2 F-G NB Setophaga townsendi Townsend's Warbler 9 G B Polioptilidae Polioptila caerulea Blue-gray Gnatcatcher 12 G A Psittacidae Myiopsitta monachus Monk Parakeet 10 F B Ptiliogonatidae Ptiliogonys cinereus Gray Silky-flycatcher 1 G B Thraupidae Diglossa baritula Cinnamon-bellied Flowerpiercer 1 W-G B Threskiornithidae Plegadis chihi White-faced Ibis 1 F-G B Trochilidae Basilinna leucotis White-eared Hummingbird 3 F-G B Cynanthus latirostris Broad-billed Hummingbird 5 F B Lampornis clemenciae Blue-throated Mountain-gem 2 F-G B Saucerottia beryllina Berylline Hummingbird 26 G B Trogloditidae Thryomanes bewickii Bewick's Wren 23 G B Troglodytes aedon Northern House Wren 7 F-G B Turdidae Turdus migratorius American Robin 33 F-G B Turdus rufopalliatus Rufous-backed Robin 63 F B Tyrannidae Contopus pertinax Greater Pewee 3 G NB Myiarchus cinerascens Ash-throated Flycatcher 2 G B Pyrocephalus rubinus Vermilion Flycatcher 8 G B Tyrannus melancholicus Tropical Kingbird 1 F-G B Tyrannus vociferans Cassin's Kingbird 5 – – Empidonax sp. – 1 G NB Vireonidae Vireo cassinni Vireo cassinii 26 G B Additional Declarations No competing interests reported. Supplementary Files SupplementaryMaterial200525.docx Cite Share Download PDF Status: Under Revision Version 1 posted Editorial decision: Revision requested 16 Sep, 2025 Reviews received at journal 10 Sep, 2025 Reviews received at journal 29 Aug, 2025 Reviewers agreed at journal 14 Jul, 2025 Reviewers agreed at journal 14 Jul, 2025 Reviewers invited by journal 14 Jul, 2025 Editor assigned by journal 12 Jun, 2025 Submission checks completed at journal 12 Jun, 2025 First submitted to journal 09 Jun, 2025 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-6858525","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":485387592,"identity":"3820acfe-b45e-4f09-9489-5c014f3a3357","order_by":0,"name":"Marisela Martínez-Ruiz","email":"","orcid":"","institution":"National Autonomous University of Mexico","correspondingAuthor":false,"prefix":"","firstName":"Marisela","middleName":"","lastName":"Martínez-Ruiz","suffix":""},{"id":485387593,"identity":"8ae33d1a-8648-47b6-b912-286df8a1cdd3","order_by":1,"name":"Elio G. Lagunes-Díaz","email":"","orcid":"","institution":"Instituto de Ecología, A.C. (INECOL)","correspondingAuthor":false,"prefix":"","firstName":"Elio","middleName":"G.","lastName":"Lagunes-Díaz","suffix":""},{"id":485387594,"identity":"d08e9174-e490-4394-90e4-da2a65eb7d45","order_by":2,"name":"Jonathan Morales-Contreras","email":"","orcid":"","institution":"El Colegio de la Frontera Sur","correspondingAuthor":false,"prefix":"","firstName":"Jonathan","middleName":"","lastName":"Morales-Contreras","suffix":""},{"id":485387595,"identity":"d699eacf-5d9c-4827-84ba-0b82078507ac","order_by":3,"name":"María del Coro Arizmendi","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAAjElEQVRIiWNgGAWjYLCCDyTrYJxBshZmHpKU87f3Pnxs23ZHXreBO02CKC0SZ44bG+e2PTPcdoB3swFRWgwk0tikc9sOMwK1bHxAvBbLtsP2QC0bDhCvhbHtcCLxtkicOcZs2HPucPK2w8T6hb+9jfHBj7LDttuO924jLsQQgJlE9aNgFIyCUTAK8AAAN8AsreQMw5cAAAAASUVORK5CYII=","orcid":"","institution":"National Autonomous University of Mexico","correspondingAuthor":true,"prefix":"","firstName":"María","middleName":"del Coro","lastName":"Arizmendi","suffix":""}],"badges":[],"createdAt":"2025-06-10 03:23:16","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-6858525/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-6858525/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":86949015,"identity":"32c26973-1a94-479a-a383-bd180d3127ff","added_by":"auto","created_at":"2025-07-17 13:42:31","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":464531,"visible":true,"origin":"","legend":"\u003cp\u003eLocation of 20 urban public parks in Mexico City. Size of orange circles indicates the largest spatial scale (900 m radius) considered in this study. We also show one of these urban public parks in detail, including the nine concentric spatial scales used to extract landscape structure variables\u003c/p\u003e","description":"","filename":"1.png","url":"https://assets-eu.researchsquare.com/files/rs-6858525/v1/0459fd5fffd9179ee8c22c1e.png"},{"id":86949013,"identity":"e536789f-b713-49a6-b45d-19d8059fa597","added_by":"auto","created_at":"2025-07-17 13:42:31","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":185152,"visible":true,"origin":"","legend":"\u003cp\u003eRank abundance plots of birds recorded in 20 urban parks in Mexico City. The four species dominating each assemblage are indicated: \u003cem\u003eCin Columbina inca, Cli Columba livia, Zas Zenaida asiatica, Mfu Melozone fusca, Pdo Passer domesticus, Sco Setophaga coronata, Qme Quiscalus mexicanus, Hme Haemorhus mexicanus, Iab Icterus abeillei, Tmi Turdus migratorius, Tru Turdus rufopalliatus, Sbe Saucerottia beryllina, Vca Vireo cassinni, Lru Leiothlypis ruficapilla, Sde Streptopelia decaocto, Tvo Tyrannus vociferans. \u003c/em\u003eNumbers in blue indicate the evenness factor (EF = \u003csup\u003e2\u003c/sup\u003eD/\u003csup\u003e0\u003c/sup\u003eD)\u003c/p\u003e","description":"","filename":"2.png","url":"https://assets-eu.researchsquare.com/files/rs-6858525/v1/d28e59f6f8c94f9f80daefdf.png"},{"id":86949012,"identity":"17c645fc-a3e2-4104-9f07-97691be6d6dd","added_by":"auto","created_at":"2025-07-17 13:42:31","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":166555,"visible":true,"origin":"","legend":"\u003cp\u003eSee image above for figure legend.\u003c/p\u003e","description":"","filename":"3.png","url":"https://assets-eu.researchsquare.com/files/rs-6858525/v1/f5701bff0bece52da24e7c39.png"},{"id":86949965,"identity":"fcdc162c-e7d2-4930-b338-b0290c931f58","added_by":"auto","created_at":"2025-07-17 13:58:32","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1554803,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6858525/v1/e329ad0c-653a-4241-b64c-86f3fb591678.pdf"},{"id":86949029,"identity":"31f788da-8b2a-49ee-b063-00567ec17f09","added_by":"auto","created_at":"2025-07-17 13:42:32","extension":"docx","order_by":0,"title":"","display":"","copyAsset":false,"role":"supplement","size":32338038,"visible":true,"origin":"","legend":"","description":"","filename":"SupplementaryMaterial200525.docx","url":"https://assets-eu.researchsquare.com/files/rs-6858525/v1/b55dfce46e66e9625a06959c.docx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Beyond green space area: How landscape structure shapes urban bird communities in a megacity","fulltext":[{"header":"INTRODUCTION","content":"\u003cp\u003eThe replacement of native habitats within the urbanization process lead to biodiversity decline, species extinction (Sun et al. \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) and biotic homogenization (Devictor et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Croci et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). Despite the many negative impacts of urbanization on avian assemblages (reviewed in Chace and Walsh \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2006\u003c/span\u003e), several studies have documented the importance of urban green spaces for retaining an important proportion of the world\u0026rsquo;s avifauna (Aronson et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Ib\u0026aacute;\u0026ntilde;ez-\u0026Aacute;lamo et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; MacGregor-Fors et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), providing important habitat and refuge for birds within cities (Fern\u0026aacute;ndez-Juricic and Jokim\u0026auml;ki \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Vasquez and Wood \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eOverall, studies in urban environments have demonstrated that bird species richness decreases with urbanization while bird density may increase (e.g. Marzluff \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2001\u003c/span\u003e, Chace and Walsh \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2006\u003c/span\u003e, van Heezik et al. \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). Multiple studies conducted at the site scale (i.e. measuring environmental variables within urban green space) reported higher bird abundance and richness with larger urban green space (Gavareski \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e1976\u003c/span\u003e; Evans et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Dale \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Leveau et al. \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Thompson et al. \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2022\u003c/span\u003e) conducting to the generalization that larger urban green spaces contain higher bird diversity. However, the majority of studies have centered on urban green space local predictors like size (Gavareski \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e1976\u003c/span\u003e; Dale \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Leveau et al. \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Thompson et al. \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2022\u003c/span\u003e) or evaluating gradients of urbanization (Xie et al. \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2019\u003c/span\u003e), neglecting the influence of landscape structure in shaping avian assemblages.\u003c/p\u003e\u003cp\u003eAt the landscape scale (i.e. measuring environmental variables beyond the limits of the urban green space), the amount of woody cover or tree cover is an important predictor for the presence of many bird species (M\u0026ouml;rtberg \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Melles et al. \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2003\u003c/span\u003e) and positively relates to bird species richness and abundance (Ikin et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Callaghan et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Chong et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Villase\u0026ntilde;or et al. \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Mao et al. \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Species richness and functional diversity are negatively affected by increasing sealed cover (Sch\u0026uuml;tz and Schulze \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2015\u003c/span\u003e) and construction land (Mao et al. \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) in the landscape, whereas the connectivity of urban green spaces enhanced the abundance of different birds (Pellissier et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Kang et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2015\u003c/span\u003e) and species richness (Oliver et al. \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). Overall, although local-site variables are highly important for birds, their abundance and diversity in urban ecosystems also depend on patterns (e.g. landscape forest cover, impervious surface) and processes (e.g. dispersion) operating at landscape scales.\u003c/p\u003e\u003cp\u003eImplementing urban planning strategies and policymaking that reduce or reverse the negative impacts of the urbanization process on biodiversity is imperative, and those strategies rely on the understanding of the effects of the urban landscape pattern on biodiversity. Here, we estimated bird abundance, species richness, diversity, and assemblage evenness in urban greens paces across Mexico City, and examined the influence of landscape structure on bird responses. Since biological responses to landscape structure can be scale-dependent, and, as we do not know a priori the best spatial scale of bird responses to landscape structure (the so-called \u0026lsquo;scale of effect\u0026rsquo;, Jackson and Fahrig \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2015\u003c/span\u003e), we used a multi-scale analysis and measured landscape variables at different nested scales surrounding study sites. We predicted that landscape composition is more important than landscape configuration for birds. We also expected that species richness and diversity would be highly and positively associated to forest cover in the urban landscape, given the importance of such land cover for birds. To our knowledge, few studies at the landscape scale have used a multi-scale approach when evaluating the influence of landscape structure on birds within urban ecosystems (but see Huang et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2015\u003c/span\u003e, Callaghan et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2018\u003c/span\u003e).\u003c/p\u003e"},{"header":"METHODS","content":"\u003cp\u003eOur study was carried out in Central Mexico in Mexico City (19\u0026deg;35'34.08\" N, 99\u0026deg;21'53.64\" W and 19\u0026deg;02'53.52\" N, 98\u0026deg;56'25.08\" W, 1494.3 km\u003csup\u003e2\u003c/sup\u003e). Mean annual temperature is 16\u0026ordm;C and average altitude is 2240 masl and is surrounded by mountains (Carrera-Hern\u0026aacute;ndez and Gaskin \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). The urbanized land cover occupies most of the territory, but remnants of native vegetation (coniferous forests, wetlands, pastures) are still present in the south of the city (PAOT \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). This megacity (~\u0026thinsp;21\u0026nbsp;million people, INEGI \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) has considerable amounts of public green areas. By 2017, the city had 67.3 km\u003csup\u003e2\u003c/sup\u003e of urban green spaces (SEDEMA \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e\u003ch2\u003eStudy site selection\u003c/h2\u003e\u003cp\u003eTo select study sites, we used a map of the study area and use the database of urban green spaces of Mexico City (SEDEMA \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Then we selected 20 public green spaces to carry out bird surveys (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). We selected those urban green spaces of public access with no time restriction for public entry and composed mainly of forested vegetation (i.e. high cover of trees), as many public green spaces were almost treeless. Selected urban green spaces were separated at least 2 km from each other, this to avoid overlapping of influencing landscape metrics and to ensure spatial independence of samples. Northeast side of the study area was underrepresented because the smaller number of forested public green spaces (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003c/div\u003e\n\u003ch3\u003eBird surveys\u003c/h3\u003e\n\u003cp\u003eWe conducted point counts (Bibby et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2000\u003c/span\u003e) to sample birds in each of the selected urban green spaces. We recorded all birds seen or heard up to a 50 m radius during 15 min. The number of point counts at each urban green space was proportional to its size (1\u0026ndash;2 point counts per site), and the points were separated at least 200 m (Bibby et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2000\u003c/span\u003e). All sites were visited once in the morning (20 min after dawn) only on working days (Monday-Thursday) from February to April 2024.\u003c/p\u003e\n\u003ch3\u003eUrban landscape composition and configuration\u003c/h3\u003e\n\u003cp\u003eWe performed a supervised classification for characterizing landscape features at our study area: we used two images from the Planet remote sensing platform, from the \u0026ldquo;Tropical Normalized Analytic Monthly series basemaps\u0026rdquo; (PBC 2024), made available through the Norway\u0026rsquo;s International Climate and Forests Initiative (NICFI). We selected one image for August 2023, the image for the most recent rainy season, when plant phenology allows for better differentiation of its spectral signatures from other classes. We visually selected points for sampling the different land use and land cover classes on a Google Satellite image (Google \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), using the \u003cem\u003eQuickmapservices\u003c/em\u003e plugin for the QGIS software (QGIS Development Team \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). The classes we used for the sampling were: asphalt, built-up, water, grass and forest; to avoid mixing in very different spectral signatures, thus reducing noise in the training phase, we subdivided built-up classes into concrete, waterproofed concrete and metal-roof. At least 15 samples were selected from each class. We then used the modules \u003cem\u003eterra\u003c/em\u003e (Hijmans \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) for raster processing, \u003cem\u003erandomForest\u003c/em\u003e (Liaw and Wiener \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2002\u003c/span\u003e) for classification with the random forest algorithm and caret (Kuhn \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2008\u003c/span\u003e) for predictive model building and analysis, in the R statistical computing language (R Development Core Team \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). The selected image had high accuracy, with a benchmarked median Kappa of 0.925.\u003c/p\u003e\u003cp\u003eOn the classified image, we used the module landscape metrics (Hesselbarth et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) and calculated forest (ha), pavement (ha), grass (ha) and built (ha) cover as variables of landscape composition and estimated the density of patches of forest patches and the mean distance between forest patches as variables of landscape configuration. It should be noted that the coverage called \"forest\" does not correspond to a native forest but to forest structure vegetation that may be composed of native and/or exotic tree species. We calculated each of the landscape metrics at nine concentric buffers (i.e. nine spatial scales, radius 100-900m, by 100m) from the geographic center of each study site to evaluate the spatial scale of effect (Jackson and Fahrig \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2012\u003c/span\u003e).\u003c/p\u003e\n\u003ch3\u003eBird abundance, diversity and assemblage structure\u003c/h3\u003e\n\u003cp\u003eFor diversity estimates, we first checked the sample completeness of each forest site using the sample coverage index suggested by Chao and Jost (\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). The sample coverage index represents the percentage of the total number of individuals in an assemblage that belongs to the species found in the sample (Chao and Jost \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). Since sample coverage varied among study sites, ranging from 0.68 to 0.95 (Table \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e), we estimated species diversity by using both observed and expected data based on the rarefaction and extrapolation procedure suggested by Chao and Jost (\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2012\u003c/span\u003e) using the iNEXT package for R (Hsieh et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Specifically, we estimated Hill numbers of order 0 and 1 as measures of bird diversity (\u003csup\u003e0\u003c/sup\u003e\u003cem\u003eD\u003c/em\u003e, \u003csup\u003e1\u003c/sup\u003e\u003cem\u003eD\u003c/em\u003e, hereafter). \u003csup\u003e0\u003c/sup\u003e\u003cem\u003eD\u003c/em\u003e (i.e. species richness) is not sensitive to species abundance, only presences are counted, whereas \u003csup\u003e1\u003c/sup\u003e\u003cem\u003eD\u003c/em\u003e (the exponential of Shannon\u0026rsquo;s entropy index) weights species in proportion to their frequency (Chao et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2014\u003c/span\u003e), and it can be interpreted as the number of common or typical species in the assemblage (Jost \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2006\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eWe considered the abundance of each species to construct rank-abundance plots of each of the study sites (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e) and estimated the evenness factor proposed by Jost (\u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). The evenness factor \u003cem\u003eEF\u003c/em\u003e represents the proportion of dominant species in the assemblage, it ranges between 1 (a perfectly even assemblage), whereas values near to 0 would a highly dominated assemblage (Jost \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2010\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\n\u003ch3\u003eEstimation of the scale of landscape effect\u003c/h3\u003e\n\u003cp\u003eWe performed Generalized Linear Models following a multiscale approach to first detect the so called \u0026ldquo;scale of effect\u0026rdquo; (Jackson and Fahrig \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2012\u003c/span\u003e), this is the spatial scale that yields the strongest species-landscape relationship. We quantified the relationship between each landscape metric considered in this study (i.e. forest cover, grass cover, pavement cover, built cover, density of forest patches, mean distance between forest patches) and each response variable (i.e. abundance, \u003csup\u003e\u003cem\u003e0\u003c/em\u003e\u003c/sup\u003e\u003cem\u003eD\u003c/em\u003e, \u003csup\u003e\u003cem\u003e1\u003c/em\u003e\u003c/sup\u003e\u003cem\u003eD\u003c/em\u003e, \u003cem\u003eEF\u003c/em\u003e) at each spatial scale (nine scales, from 100-900m radius). We used Gaussian distribution error for continuous variables (i.e. \u003csup\u003e\u003cem\u003e1\u003c/em\u003e\u003c/sup\u003e\u003cem\u003eD\u003c/em\u003e, \u003cem\u003eEF\u003c/em\u003e) and count variables of abundance and \u003csup\u003e\u003cem\u003e0\u003c/em\u003e\u003c/sup\u003e\u003cem\u003eD\u003c/em\u003e were modeled with a Poisson distribution error (Crawley \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). Then, we plotted the value of AICc as a dependent variable against landscape size to identify the scale that best yields the best-fitted associations between each response variable and each landscape predictor. We estimated a total of 30 scales of effect values (5 response variables x 6 landscape metrics; Table S2) selected by the lowest sample-corrected Akaike information criterion (AICc) value among sets of models (Table S3).\u003c/p\u003e\u003cdiv id=\"Sec8\" class=\"Section2\"\u003e\u003ch2\u003eInfluence of landscape structure on bird abundance and diversity\u003c/h2\u003e\u003cp\u003eWe conducted Principal Component Analysis (PCA) of the six-landscape variables measured at the identified scale of effect (Table S2) to reduce multidimensional landscape metrics into a reduced number of indices. Since the spatial scale of effect was different for each combination of response and predictor variables Table S3, we conducted one PCA for each response variable (Fig \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e-S5). In all cases, we retained the first three components since they accounted for ~\u0026thinsp;85% of the total variance in each data set (Table S4-S5).\u003c/p\u003e\u003cp\u003eTo evaluate the relative effect of landscape variables (contained in each of the three retained Principal Components) on each response variable (abundance, \u003csup\u003e\u003cem\u003e0\u003c/em\u003e\u003c/sup\u003e\u003cem\u003eD\u003c/em\u003e, \u003csup\u003e\u003cem\u003e1\u003c/em\u003e\u003c/sup\u003e\u003cem\u003eD, EF\u003c/em\u003e) we used an information-theoretic approach and multi-model inference (Burnham and Anderson \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2002\u003c/span\u003e). For each response variable we performed 16 models, representing all combinations of the predictors (i.e. three PCA axes for each case) and the null model (intercept only). Then, we classified the models from the lowest (the best-supported model) to the highest (the least-supported model) considering their AICc value. After, we summed the Akaike weights of each PCA component appearing in the models, which is considered a proxy of the relative importance of each PCA component on each response variable (Burnham and Anderson \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Giam and Olden \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). We found no collinearity among the multiple models (VIF\u0026thinsp;\u0026lt;\u0026thinsp;2) after checking for the Variance Inflation Factor (VIF) for each predictor. We used a normal distribution for \u003csup\u003e\u003cem\u003e1\u003c/em\u003e\u003c/sup\u003e\u003cem\u003eD\u003c/em\u003e, \u003cem\u003eEF\u003c/em\u003e, and Poisson distribution for abundance and \u003csup\u003e\u003cem\u003e0\u003c/em\u003e\u003c/sup\u003e\u003cem\u003eD\u003c/em\u003e data. All analyses were conducted in R (R Development Core Team \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e\u003c/div\u003e"},{"header":"RESULTS","content":"\u003cp\u003e\u003cem\u003eBird assemblages in urban green spaces\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eWe recorded a total of 1,194 individuals from 56 species and 23 families (Table 1). Most of the identified bird species were resident breeders in Mexico City (26 species, 65.4%). A large proportion were migratory, non-breeding species (15 species, 27.2%), while the remaining identified birds (4 species, 7.2%) were alien species (Table 1).\u003c/p\u003e\n\u003cp\u003eThe three most abundant species were generalist, conspicuous urban residents: the Inca Dove (181 individuals), the introduced and now feral Rock Pigeon (168 individuals), and the opportunistic House Sparrow (156 individuals), each accounting for more than 150 individuals. All three dominated most bird assemblages (Table 1, Fig. 2).\u003c/p\u003e\n\u003cp\u003eNext, bird assemblages were dominated by other abundant species that had expanded their ranges significantly and have invasive and acclimatization abilities: the House Finch (73 individuals), the White-winged Dove (67 individuals), and the Great-tailed Grackle (66 individuals). Additionally, we recorded high abundances of the common and widespread Yellow-rumped Warbler (63 individuals) and the Rufous-backed Robin (63 individuals) (Table 1, Fig. 2).\u003c/p\u003e\n\u003cp\u003eOther bird species showed abundances ranging from 5 to 25 individuals (Table 1), including migratory species (e.g., American Robin, Cassin\u0026rsquo;s Vireo,\u0026nbsp;Nashville Warbler, Wilson\u0026rsquo;s Warbler; Table 1), hummingbirds (Beryllina Hummingbird,\u0026nbsp;Broad-billed Hummingbird), and non-native invasive species (Eurasian Collared-Dove,\u0026nbsp;Monk Parakeet) among others (Table 1). Finally, species with low abundances included raptors (Accipitridae, Falconidae), migrants (Parulidae) and resident birds (Tyrannidae), among others (Table 1).\u003c/p\u003e\n\u003cp\u003eWe observed that half of bird assemblages (50%) within public green spaces exhibited intermediate evenness (EF = 0.5-0.7; Fig. 2), whereas 40% (8) or urban green spaces showed highly dominated assemblages (EF \u0026lt; 0.5; Fig. 2), and only 10% (2) of the studied sites exhibited more even assemblages (EF \u0026lt; 0.7; Fig. 2).\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eInfluence of landscape structure on bird assemblages\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eLandscape composition had a greater influence on bird responses than landscape configuration, with forest patch density being the only configuration metric that affected bird assemblages.\u003c/p\u003e\n\u003cp\u003eThe overall abundance of birds was negatively influenced by a linear combination of lower pavement cover and higher grass cover (PC2, Fig. 3a). In contrast, a combination of lower forest cover and higher built cover, along with greater forest patch density (PC1, PC3; Fig. 3a) positively influenced bird abundance.\u003c/p\u003e\n\u003cp\u003eBird species richness and the number of common bird species (\u003csup\u003e1\u003c/sup\u003e\u003cem\u003eD\u003c/em\u003e) were both negatively and mainly affected by highly urbanized landscapes with lower proportion of forest and grass cover (PC2; Fig. 3b,c), and higher forest patch density (PC3: Fig. 3b,c). \u0026nbsp;A higher proportion of built and pavement cover negatively influenced bird species richness and \u003csup\u003e1\u003c/sup\u003e\u003cem\u003eD\u003c/em\u003e, but this predictor was less important (PC1; Fig. 3b,c).\u003c/p\u003e\n\u003cp\u003eFinally, the evenness of bird assemblages was mainly and positively influenced by landscapes with lower pavement cover and high grass cover (PC3; Fig 3d), whereas landscapes with high built cover, and lower forest cover distributed in many fragments decreased evenness of bird assemblages (PC1, PC2; Fig 3d).\u003c/p\u003e"},{"header":"DISCUSSION","content":"\u003cdiv id=\"Sec13\" class=\"Section2\"\u003e\u003ch2\u003eBird assemblages\u003c/h2\u003e\u003cp\u003eOur results indicated that bird assemblages inhabiting public green spaces in Mexico City are dominated by a few synanthropic bird species that thrive in urban environments. These findings are consistent with other studies conducted in temperate (Donnelly and Marzluff \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Villegas and Garitano-Zavala \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Ikin et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2013\u003c/span\u003e), Mediterranean (Kurucz et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), subtropical (van Rensburg et al. \u003cspan citationid=\"CR70\" class=\"CitationRef\"\u003e2009\u003c/span\u003e) and tropical (Ortega-\u0026Aacute;lvarez and MacGregor-Fors \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; MacGregor-Fors and Ortega-\u0026Aacute;lvarez \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2011a\u003c/span\u003e; Chamberlain et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2017\u003c/span\u003e) cities globally, where dominant bird species are generalists better suited to exploit urban environments. Our findings also align with studies that report the dominance of a few exotic species and granivores in urbanized areas (reviewed in Chace and Walsh \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Kurucz et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) and within our study area (Charre et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Altogether, these findings underscore the homogenizing effects of urbanization on bird assemblages, which have been observed in other urban environments (e.g. Devictor et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; van Rensburg et al. \u003cspan citationid=\"CR70\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Sidemo-Holm et al. \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), where a small number of generalist species adapt and thrive over more specialized native birds. More than 400 bird species have been reported for the Mexico City Metropolitan Area (Ram\u0026iacute;rez-Albores et al. \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), with more specialized bird species found in higher numbers towards the city\u0026rsquo;s borders (MacGregor-Fors and Ortega-\u0026Aacute;lvarez \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2011a\u003c/span\u003e) where large extentions of native vegetation still surround the more urbanized areas (Arriaga et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2000\u003c/span\u003e). Nevertheless, the dominance of synanthropic and generalist bird species in urban green spaces raises concerns on the loss of biodiversity and ecosystem function within urbanized landscapes.\u003c/p\u003e\u003cp\u003eThe higher number of bird species documented for our study area (~\u0026thinsp;400 species; I don\u0026acute;t lilRam\u0026iacute;rez-Albores et al. \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) includes all types of urban green spaces such as university campuses, sports centers, ecological reserves, and wetlands. Similarly, other studies that report higher species richness (e.g. Charre et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2013\u003c/span\u003e, Ram\u0026iacute;rez-Albores et al. \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2024\u003c/span\u003e, Oropeza-S\u0026aacute;nchez et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2025\u003c/span\u003e) have compiled data from various sources or over multiple seasons. Our study, however focuses specifically on public green spaces, which are different from other green spaces like arboreta (e.g. Charre et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Oropeza-S\u0026aacute;nchez et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2025\u003c/span\u003e), private gardens, or controlled-access urban green areas. This suggests that urban public urban green spaces represent a relatively small proportion of the total bird species that inhabit the city.\u003c/p\u003e\u003cp\u003eAlthough bird assemblages in public green spaces are primarily dominated by a few synanthropic species, we also recorded a notable proportion of migratory species. Of the 55 species identified, 15 were migratory, highlighting the importance of urban landscapes in providing habitat for migratory birds. This finding suggests that, despite the dominance of generalist species, public green spaces offer valuable resources for birds, particularly those requiring biodiversity-friendly infrastructure, such as large trees and forested areas (Amaya-Espinel and Hostetler \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). These results underscore the potential for targeted management and restoration efforts to enhance public green spaces that support greater diversity of bird species.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec14\" class=\"Section2\"\u003e\u003ch2\u003eInfluence of landscape structure on bird assemblages\u003c/h2\u003e\u003cp\u003eOur results demonstrated that highly deforested landscapes, with a high proportion of impervious surface and built cover, are associated with increased bird abundance in the city. This aligns with previous studies that reported higher bird abundance in more urbanized areas (Donnelly and Marzluff \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Ortega-\u0026Aacute;lvarez and MacGregor-Fors \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; MacGregor-Fors and Ortega-\u0026Aacute;lvarez \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2011b\u003c/span\u003e; Chamberlain et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Kurucz et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Particularly, our results agree with studies that analyzed land cover types as predictors, showing increased bird abundance with higher built cover (Lim and Sodhi \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Amaya-Espinel et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Overall, our findings confirm that urbanized areas can support high bird abundance, even though these assemblages are simplified and dominated by a few species.\u003c/p\u003e\u003cp\u003eLandscape structure had contrasting effects on bird abundance, richness, and evenness. Richness and evenness were lower in deforested landscapes with high proportions of built cover, whereas bird abundance was higher in these areas. These results are consistent with previous studies that show lower bird richness in areas with greater building presence (Chamberlain et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2007\u003c/span\u003e), building height (Leveau and Leveau \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), built cover (Carb\u0026oacute;-Ram\u0026iacute;rez and Zuria \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2011\u003c/span\u003e), higher building densities, and more road coverage (Amaya-Espinel et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Although built infrastructure can provide important resources for birds (Mainwaring \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Partridge and Clark \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) it does not constitute a biodiversity-friendly land cover that promotes bird species richness and evenness within our study area. In contrast, landscapes with higher grass cover and less pavement were associated with greater bird species richness and evenness, confirming that grass and lawns within urban landscapes function as a source of resources for birds (S\u0026aacute;nchez-Sotomayor et al. \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) and provide a more bird-friendly environment than impervious surface like built and pavement cover.\u003c/p\u003e\u003cp\u003eBird species richness followed the expected pattern, decreasing in landscapes with lower proportions of vegetation (both forest and grass cover) and higher proportions of impervious surfaces, adding to the broader trend of reduced bird species richness with increased urbanization (Melles et al. \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Chamberlain et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Our results emphasize the importance of forest cover in maintaining overall bird species richness within cities (Villegas and Garitano-Zavala \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Ikin et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Callaghan et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) particularly for native birds (Villase\u0026ntilde;or et al. \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Furthermore, we observed the same response pattern for the number of common species in the bird assemblages, reinforcing the critical role of forested land cover in sustaining high bird diversity in urban environments.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec15\" class=\"Section2\"\u003e\u003ch2\u003eImplications for conservation\u003c/h2\u003e\u003cp\u003eOur findings highlight the significant role that landscape composition and configuration play in shaping bird assemblages in urban environments, underscoring the need for urban planning strategies that prioritize the expansion and maintenance of vegetation and biodiversity-friendly infrastructure across landscapes. Specifically, our results suggested that urban conservation efforts should focus on protecting and restoring forested areas and grasslands to enhance bird diversity. The positive responses of birds to forest cover in our study indicated a valuable opportunity to improve urban landscapes for avian populations. Notably, the forest cover in our study did not necessarily consist of native vegetation but included a mix of native and non-native trees, suggesting that even urban green spaces with non-native plant species can be improved by introducing a more complex forest structure to better support bird diversity.\u003c/p\u003e\u003cp\u003eIn addition, policies aimed at reducing impervious surfaces and promoting green infrastructure\u0026mdash;such as green roofs, permeable pavements, and expanded green corridors\u0026mdash;could significantly enhance habitat quality for birds in urban areas. The benefits of such measures extend beyond birds, contributing to the broader ecological health of urban environments.\u003c/p\u003e\u003cp\u003eGiven the homogenizing effects of urbanization on bird assemblages, it is essential to integrate ecological principles into urban planning. Enhancing connectivity between urban green spaces and surrounding natural landscapes, reducing impervious surface areas, and increasing vegetation cover could create more sustainable and resilient environments for both migratory and resident bird species. Such conservation efforts would not only benefit avian populations but also foster biodiversity and ecological resilience in the face of ongoing urbanization. Public green spaces within megacities, therefore, hold substantial potential for conserving bird assemblages and promoting a more sustainable, biodiverse urban environment.\u003c/p\u003e\u003c/div\u003e"},{"header":"Declarations","content":"\u003cp\u003eACKNOWLEDGMENTS\u003c/p\u003e\n\u003cp\u003eMMR received a postdoctoral scholarship from SECIHTI (Secretar\u0026iacute;a de Ciencia, Humanidades, Tecnolog\u0026iacute;a e Innovaci\u0026oacute;n) and support from Idea Wild. We are thankful to Oscar G. Brito for field assistance.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eFUNDING\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eSECIHTI supported this work through a postdoctoral grant awarded to MMR.\u003c/p\u003e\n\u003cp\u003eCOMPETING INTERESTS\u003c/p\u003e\n\u003cp\u003eThe authors have no relevant financial or non-financial interests to disclose.\u003c/p\u003e\n\u003cp\u003eAUTHOR CONTRIBUTIONS\u003c/p\u003e\n\u003cp\u003eMarisela Mart\u0026iacute;nez-Ruiz developed the study conception and design. Material preparation and data collection were performed by Marisela Mart\u0026iacute;nez-Ruiz, Jonathan Morales-Contreras and Elio G. Lagunes-D\u0026iacute;az. Analyses were performed by Marisela Mart\u0026iacute;nez-Ruiz. The first draft of the manuscript was written by Marisela Mart\u0026iacute;nez-Ruiz and all authors commented on previous versions of the manuscript. All authors read and approved the final manuscript.\u003c/p\u003e\n\u003cp\u003eDATA AVAILABILITY\u003c/p\u003e\n\u003cp\u003eThe datasets generated and analyzed during the current study are available from the corresponding author on reasonable request\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAmaya-Espinel JD, Hostetler M, Henr\u0026iacute;quez C, Bonacic C (2019) The influence of building density on Neotropical bird communities found in small urban parks. 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Urban Ecosyst 13:375\u0026ndash;391. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s11252-010-0126-7\u003c/span\u003e\u003cspan address=\"10.1007/s11252-010-0126-7\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eXie S, Xu YS W, et al (2019) The effect of habitat changes along the urbanization gradient for breeding birds: an example from the Xiong\u0026rsquo;an New Area. PeerJ 7:e7961. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/https://doi.org/10.7717/peerj.7961\u003c/span\u003e\u003cspan address=\"10.7717/peerj.7961\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"},{"header":"Table","content":"\u003cp\u003eTable 1. List of bird species registered during 15 min point counts in 20 public parks of Mexico City from February to April 2024. Ab = Abundance; Habitat use: G = Generalist species, F = forest species; F-G = Species associated to forested habitats but able to exploit different land covers; Status: A = Alien, B = Breeding, NB = non-breeding. Habitat use for non-breeding birds refers to habitat identified for the species during the non-breeding season. Habitat use and status of birds within Mexico City was retrieved from Birds of the World (www.birdsoftheworld.org).\u003c/p\u003e\n\u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eFamily\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eSpecies\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eCommon name\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eAb\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eHabitat \u0026nbsp; \u0026nbsp; \u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp;use\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eStatus\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eAccipitridae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eAccipiter striatus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eSharp-shinned Hawk\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eF-G\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eParabuteo unicinctus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eHarris\u0026apos;s Hawk\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eAegithalidae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003ePsaltriparus minimus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eBushtit\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eCardinalidae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003ePheucticus melanocephalus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eBlack-headed Grosbeak\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eNB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003ePiranga ludoviciana\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eWestern Tanager\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eNB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003ePiranga rubra\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eSummer Tanager\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eA\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eColumbidae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eColumba livia\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eRock Pigeon\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e168\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eColumbina inca\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eInca Dove\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e181\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eA\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eStreptopelia decaocto\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eEurasian Collared-Dove\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e21\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eZenaida asiatica\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eWhite-winged Dove\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e67\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eZenaida macroura\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eMourning Dove\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eEmberizidae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eMelospiza melodia\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eSong Sparrow\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eFalconidae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eFalco sparverius\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eAmerican kestrel\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eFringilidae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eHaemorhous mexicanus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eHouse Finch\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e73\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eSpinus psaltria\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eLesser Goldfinch\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eIcteridae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eIcterus abeillei\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eBlack-backed Oriole\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eNB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eIcterus bullockii\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eBullock\u0026apos;s Oriole\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eNB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eIcterus cuculatus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eHooded Oriole\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eMolothrus aeneus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eBronzed Cowbird\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eQuiscalus mexicanus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eGreat-tailed Grackle\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e66\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eMimidae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eToxostoma curvirostre\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eCurved-billed Thrasher\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e10\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003ePasserellidae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eMelozone fusca\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eCanyon Towhee\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e17\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eA\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003ePasseridae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003ePasser domesticus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eHouse Sparrow\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e156\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eF-G\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003ePicidae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eDryobates scalaris\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eLadder-backed Woodpecker\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eF-G\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eMelanerpes formicivorus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eAcorn Woodpecker\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eF-G\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eNB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eSphyracus varius\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eYellow-bellied Sapsucker\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eF-G\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eNB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eParulidae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eCardellina pusilla\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eWilson\u0026apos;s Warbler\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e18\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eNB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eLeiothlypis ruficapilla\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eNashville Warbler\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e25\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eF-G\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eNB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eLeiothlypis virginiae\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eVirginia\u0026apos;s Warbler\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eNB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eMniotilta varia\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eBlack-and-white Warbler\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eF\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eMyioborus pictus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003ePainted redstart\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eNB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eSetophaga coronata\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eYellow-rumped Warbler\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e63\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003eF-G\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eNB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eSetophaga nigrescens\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eBlack-throated Gray Warbler\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003eF\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eNB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eSetophaga occidentalis\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eHermit Warbler\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eSetophaga petechia\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eYellow Warbler\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eF-G\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eNB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eSetophaga townsendi\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eTownsend\u0026apos;s Warbler\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003ePolioptilidae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003ePolioptila caerulea\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eBlue-gray Gnatcatcher\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e12\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eA\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003ePsittacidae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eMyiopsitta monachus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eMonk Parakeet\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e10\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eF\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003ePtiliogonatidae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003ePtiliogonys cinereus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eGray Silky-flycatcher\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eThraupidae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eDiglossa baritula\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eCinnamon-bellied Flowerpiercer\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eW-G\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eThreskiornithidae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003ePlegadis chihi\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eWhite-faced Ibis\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eF-G\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eTrochilidae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eBasilinna leucotis\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eWhite-eared Hummingbird\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eF-G\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eCynanthus latirostris\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eBroad-billed Hummingbird\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eF\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eLampornis clemenciae\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eBlue-throated Mountain-gem\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eF-G\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eSaucerottia beryllina\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eBerylline Hummingbird\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e26\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eTrogloditidae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eThryomanes bewickii\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eBewick\u0026apos;s Wren\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e23\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eTroglodytes aedon\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eNorthern House Wren\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eF-G\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eTurdidae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eTurdus migratorius\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eAmerican Robin\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e33\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eF-G\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eTurdus rufopalliatus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eRufous-backed Robin\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e63\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eF\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eTyrannidae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eContopus pertinax\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eGreater Pewee\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eNB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eMyiarchus cinerascens\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eAsh-throated Flycatcher\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003ePyrocephalus rubinus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eVermilion Flycatcher\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\"\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eTyrannus melancholicus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eTropical Kingbird\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eF-G\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eTyrannus vociferans\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eCassin\u0026apos;s Kingbird\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e\u0026ndash;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e\u0026ndash;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eEmpidonax sp.\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003e\u0026ndash;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eNB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003eVireonidae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eVireo cassinni\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eVireo cassinii\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\"\u003e\n \u003cp\u003e26\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd\u003e\n \u003cp\u003eB\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"urban-ecosystems","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"ueco","sideBox":"Learn more about [Urban Ecosystems](https://www.springer.com/journal/11252)","snPcode":"11252","submissionUrl":"https://submission.nature.com/new-submission/11252/3","title":"Urban Ecosystems","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Urban bird assemblages, public parks, urban landscape pattern, multi-scale approach","lastPublishedDoi":"10.21203/rs.3.rs-6858525/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-6858525/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eUrbanization poses a significant threat to biodiversity, with urban green spaces providing crucial habitats for birds within cities. However, studies have often focused on site-scale variables when assessing urbanization effects on birds, neglecting the role of landscape structure on shaping bird communities. In this sense, the effects of landscape structure on birds requires further research in urban environments.\u003cem\u003e \u003c/em\u003eWe aimed to\u003cem\u003e \u003c/em\u003eassess how landscape structure influences bird diversity in public green spaces across Mexico City. We surveyed birds in 20 public green spaces and estimated bird abundance, diversity and evenness for each study site. We estimated landscape composition and configuration metrics and analyzed their effects on bird responses at multiple spatial scales (100–900 m radius). We recorded 1,194 individual birds from 56 species during our surveys, where bird assemblages were dominated by few synanthropic bird species. Landscape composition was more important than landscape configuration for birds. Deforested landscapes promoted bird abundance, while lower forest and grass cover enhanced bird species richness and diversity. Evenness was most strongly influenced by urban landscapes with higher grass cover and lower proportions of pavement and built-up areas. Our findings highlight the importance of integrating landscape-scale ecological principles into urban planning to enhance avian diversity and promote sustainable urban ecosystems. Prioritizing landscapes with high forest (native and non-native) and grass cover, and minimizing and managing impervious surfaces, could improve the quality of urban green spaces for birds and the environmental services they provide.\u003c/p\u003e","manuscriptTitle":"Beyond green space area: How landscape structure shapes urban bird communities in a megacity","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-07-17 13:42:27","doi":"10.21203/rs.3.rs-6858525/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2025-09-16T09:15:33+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-09-10T08:16:49+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-08-29T04:38:27+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"157970886676031705424566164533443747131","date":"2025-07-14T21:34:53+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"100252924376604368241555538018944864863","date":"2025-07-14T09:53:28+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-07-14T09:38:57+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-06-13T01:02:02+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2025-06-12T22:33:25+00:00","index":"","fulltext":""},{"type":"submitted","content":"Urban Ecosystems","date":"2025-06-10T03:13:44+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"urban-ecosystems","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"ueco","sideBox":"Learn more about [Urban Ecosystems](https://www.springer.com/journal/11252)","snPcode":"11252","submissionUrl":"https://submission.nature.com/new-submission/11252/3","title":"Urban Ecosystems","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"9e640b12-99b1-4376-a804-3ea9675690ca","owner":[],"postedDate":"July 17th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"in-revision","subjectAreas":[],"tags":[],"updatedAt":"2026-03-31T07:41:44+00:00","versionOfRecord":[],"versionCreatedAt":"2025-07-17 13:42:27","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-6858525","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-6858525","identity":"rs-6858525","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}