Vegetation on and around large-scale buildings positively influences native tropical bird abundance and bird species richness

High population growth in the tropics is driving urbanisation, removing diverse natural ecosystems. This is causing native species to suffer while introduced synanthropes flourish. City planners are developing urban greenspace networks, in part trying to address this issue. Architects contribute to these greenspace networks by designing elevated and ground level green spaces on large-scale buildings. However, little evidence is available on whether building green spaces support native fauna. This is true for birds in tropical Singapore that support important ecosystem services and have existence value. Therefore, in this study, we conducted bird surveys and statistical analyses to determine, if and how vegetation on three building green space types (ground gardens, roof gardens and green walls) have a positive impact on native or introduced bird species. We found that elevated greenery (roof gardens and green walls) on large-scale buildings supported a higher richness of birds and abundance of urban native birds than control roofs and walls without vegetation. Ground gardens supported similar levels of native species as roof gardens but also a larger proportion of generalist synanthropes. However, we found no tropical forest habitat specialists across any space type. Therefore, we recommend roof gardens and ground gardens as a potential space for urban natives outside of a less competitive ground-level urban environment. Our study also found certain building design elements (height of elevated space, presence of specific plants) supported different species groups. Therefore, we suggest that these ecological requirements for different species groups are considered when designing a building’s green space.

     

Habitat- and matrix-related differences in species diversity and trait richness of vascular plants,Orthoptera and Lepidoptera in an urban landscape

Urban growth is considered to be a major driver of environmental change. Urbanisation can affect urban biodiversity in different ways. So far, most studies focused on the impact of urbanisation on single taxa in one habitat type. In this study, we used data of species inventories and GIS-based landscape elements to examine the effects of habitat size and landscape composition on the species diversity of three taxonomic groups (vascular plants, Orthoptera and Lepidoptera) in meadows and ruderal sites in the urban region of Basel, Switzerland. We also related the responses of three species traits (body size, dispersal ability and food specialisation) to habitat size in Orthoptera and Lepidoptera. We found that species of the different taxonomic groups differed in their response to habitat size and landscape composition both in meadows and ruderal sites depending on the traits examined. The species richness of Orthoptera and Lepidoptera was positively related to meadow size but not to the size of ruderal sites, while the opposite was true for plants. For Lepidoptera in ruderal sites, the percentage cover of ruderal area in the closer surroundings was a better predictor of species richness than habitat size per se. To sustain high levels of urban biodiversity, we recommend that urban planners develop adequate management strategies to satisfy the different requirements of various taxonomic groups and to increase the quality of green sites surrounding the target habitat.     

Species richness in urban parks and its drivers: A review of empirical evidence

There is growing recognition of urban areas as hosts for innovative ways to conserve and promote biodiversity. Parks, as one specific type of urban green space, constitute particularly important biodiversity hotspots in the cityscape. We reviewed empirical findings on the species richness in urban parks across all species groups that have been studied. The aim was to assess and discuss the overall species richness of urban parks, its community attributes and drivers. Search and subsequent selection process resulted in 62 papers from 25 different countries. For all examined species groups, the findings consistently show that parks are among the most species rich types of urban green spaces, but also that exotics constitute large shares, especially of plant species. Key ecological theories like the gradient approach and the island habitat ecological theory, and fundamental ecological relationships such as the species-area relationship are valid despite the manipulated ‘nature’ of parks and the surrounding urban matrix. Most studies surveyed large number of parks and applied ‘multi-scale’ approaches in tests of confounding variables, providing methodological strength. While matrix effects are consistently found to affect species richness negatively, the diversity of habitats and microhabitat heterogeneity contained in urban parks appears as the most decisive factor for the overall species richness. However, a constraint of research to date is the limitation of individual studies to one or a few species groups, rarely bridging between flora and fauna. Adopting ‘multi-species group’ approaches in future research is needed to further advance the understanding of the overall biodiversity of urban parks, and its drivers.     

Impacts of urban sprawl on species richness of plants,butterflies, gastropods and birds: not only built-up area matters

Urban growth is a major factor of global environmental change and has important impacts on biodiversity, such as changes in species composition and biotic homogenization. Most previous studies have focused on effects of urban area as a general measure of urbanization, and on few or single taxa. Here, we analyzed the impacts of the different components of urban sprawl (i.e., scattered and widespread urban growth) on species richness of a variety of taxonomic groups covering mosses, vascular plants, gastropods, butterflies, and birds at the habitat and landscape scales. Besides urban area, we considered the average age, imperviousness, and dispersion degree of urban area, along with human population density, to disentangle the effects of the different components of urban sprawl on biodiversity. The study was carried out in the Swiss Plateau that has undergone substantial urban sprawl in recent decades.Vascular plants and birds showed the strongest responses to urban sprawl, especially at the landscape scale, with non-native and ruderal plants proliferating and common generalist birds increasing at the expense of specialist birds as urban sprawl grew. Overall, urban area had the greatest contribution on such impacts, but additional effects of urban dispersion (i.e., increase of non-native plants) and human population density (i.e., increases of ruderal plants and common generalist birds) were found. Our findings support the hypothesis that negative impacts of urban sprawl on biodiversity can be reduced by compacting urban growth while still avoiding the formation of very densely populated areas.     

What shapes plant and animal diversity on urban golf courses?

Recent concern over increasing loss of biodiversity has prompted considerable interest in the role of urban green spaces as reservoirs of local biodiversity. This study assessed the diversity of three indicator taxa - plants, ants and birds - on golf courses spanning a wide range of environmental variation in terms of climate, elevation, course age, size and connectivity to native woodland. Species richness and community composition was further compared between contrasting on-course habitat types that reflect different management intensities. We identified a set of taxon-specific environmental correlates indicating an intricate interplay of landscape- and local-scale variables that affect local species diversity. Our results show that floristic diversity is positively associated with the amount of rainfall, whereas ant and bird diversity are related to local-scale factors, particularly the number of trees and the size of water features on a site. The amount of on-course native habitat was a strong predictor of plant and ant diversity and was also associated with the number of unique species at the site level; this reinforces the value of remnant habitat patches as local biodiversity reservoirs that represent mini hot-spots in an otherwise species-poor urban landscape. Community composition for all three taxa differed markedly between non-playing and playing areas, with boundary and remnant habitats generally having more diverse, species-rich communities. Our results suggest that local floral and faunal biodiversity on urban golf courses can be enhanced by creating woody non-playing areas and, especially, by preserving, restoring or expanding remnant habitats.     

Effects of urbanization on species richness: A review of plants and animals

Many studies have described the effects of urbanization on species richness. These studies indicate that urbanization can increase or decrease species richness, depending on several variables. Some of these variables include: taxonomic group, spatial scale of analysis, and intensity of urbanization. Recent reviews of birds (the most-studied group) indicate that species richness decreases with increasing urbanization in most cases but produces no change or even increases richness in some studies. Here I expand beyond the bird studies by reviewing 105 studies on the effects of urbanization on the species richness of non-avian species: mammals, reptiles, amphibians, invertebrates and plants. For all groups, species richness tends to be reduced in areas with extreme urbanization (i.e., central urban core areas). However, the effects of moderate levels of urbanization (i.e., suburban areas) vary significantly among groups. Most of the plant studies (about 65%) indicate increasing species richness with moderate urbanization whereas only a minority of invertebrate studies (about 30%) and a very small minority of non-avian vertebrate studies (about 12%) show increasing species richness. Possible explanations for these results are discussed, including the importance of nonnative species importation, spatial heterogeneity, intermediate disturbance and scale as major factors influencing species richness.     

Distribution of neotropical migratory bird species across an urbanizing landscape

Urbanization leads to long-term modification of landscapes by habitat loss, fragmentation, and the creation of new habitats. Species’ distributions respond to these modifications of habitat availability, but the combination of parameters and scale at which habitat alteration most strongly influences species distributions is poorly understood. We evaluated responses of neotropical migratory birds, a group known to be sensitive to habitat modification, across a gradient of urbanization in the southeastern United States. Thirteen Breeding Bird Survey routes, each with 40 to 50 point counts, were used to quantify species richness across the gradient of urbanization extending from downtown areas of Columbus, GA to natural woodlands. Buffers of 100, 200, and 1000 m radii were constructed from remote images around each counting point to quantify land-use with the goal of evaluating land-use parameters and scales that best described spatial variation in migrant bird species richness. Within each buffer we quantified the proportion of each cover type and within the 1000 m buffers we included several configuration parameters. We used an information-theoretic approach to separate models whose predictor variables were land-use parameters. Because measures of landscape configuration were all correlated with urban cover, these variables were entered separately. In 2002, the best model was composed of large-scale urban cover (negative effect) and mid-scale mixed hardwoods (negative and positive effect) and transitional cover (negative and positive effect) as well as the interaction between the latter two terms (positive effect). In 2003, the best model was composed of weighted edge density (negative effect), mid-scale mixed hardwood cover (negative and positive effect) and large scale transitional cover (positive effect) and the interaction between mixed hardwoods and weighted edge density (positive effect). Our results indicate that large scale habitat attributes influence the local species richness of migrant birds more than smaller scales. These results also indicate that urbanization, through increased urban cover or increasing edge contrast, has strong negative effects on species richness. Our findings support the contention that the conservation value of small woodlots in urban settings may be minimal and suggest that conservation of migratory birds will be best achieved by giving higher priority to sites where urban cover is still low and by preserving large areas of “green space” in urbanizing landscapes. The negative influence of urban cover combined with relatively minor effects of non-urban habitats on distributions of neotropical migratory birds indicates that continued urbanization of landscapes is a serious concern for conservation efforts.     

Diversity,abundance, and species composition of ants in urban green spaces

Urbanization threatens biodiversity, yet the number and scope of studies on urban arthropod biodiversity are relatively limited. We sampled ant communities in three urban habitats (forest remnants, community gardens, vacant lots) in Detroit and Toledo, USA, to compare species richness, abundance, and species composition. We measured 24 site characteristics to examine relationships between richness and composition and habitat patch size, vegetation, and urban features. Ant richness was higher in forests (26) than in gardens (14) and intermediate in vacant lots (20). Ant richness in gardens and vacant lots negatively correlated with abundance of an exotic ant species (Tetramorium caespitum); thus this ant may affect native ant richness in urban habitats. Ant composition differed with habitat type, and abundance was lowest in forests. Site characteristics varied with habitat type: forests were larger, had more woody plants, higher woody plant richness, more branches, and leaf litter whereas lots and gardens had more concrete and buildings. Vacant lots had taller herbaceous vegetation, and gardens had higher forb richness, density, and more bare ground. Differences in vegetation did not correlate with ant richness, but several vegetation factors (e.g. patch size, number and size of trees, leaf litter, and amount of concrete and buildings) correlated with differences in ant species composition. Additional factors relating to soil, nests, or microclimatic factors may also be important for urban ant communities. Implications for biodiversity conservation in urban ecosystems are discussed.     

Influence of habitat type and distance from source area on bird taxonomic and functional diversity in a Neotropical megacity

The Neotropical region has been subjected to massive urbanization, which poses high risks for some global biodiversity hotspots and losses of ecosystem functions and services. In this study, we investigate how distance from large patches of native forests (source areas) and vegetation (green)/and infrastructure (gray) characteristics affect bird species richness and functional diversity in São Paulo megacity, southeastern Brazil. We analyzed the effects of source areas and green/gray characteristics on species richness and functional diversity (richness, evenness, and divergence) indices. We detected 231 bird species, and our data confirmed our predictions: (1) bird species richness in urbanized habitats was found to be (~?50–85%) lower than in source habitats; (2) species richness and trait composition significantly decreased as the distance from the source area increased, while functional richness was not affected by this metric; and (3) shrub and herbaceous covers and maximum height of trees were positively correlated with species richness and unique functional traits regarding habitat, diet, foraging and nesting strata and dispersal ability of birds in the forest-urban matrix. The number of buildings was negatively correlated with bird species richness and functional richness. Maximum height of buildings caused dramatic declines in functional evenness. Functional divergence was notably lower in sites with high shrub cover. Our study stresses the complexity of vegetation embedded in large Neotropical urban settlements and the need to maintain large protected areas surrounding megacities to mitigate the impacts of urbanization on birds.

     

Drivers of biodiversity patterns in parks of a growing South American megacity

How urban habitats contribute to biodiversity conservation is a key challenge in a rapidly urbanising world. Urban parks can provide important habitats for native species, but previous studies are geographically biased; fast growing megacities, in particular in South America, are clearly understudied. To assess habitat functions and underlying drivers in parks of Santiago de Chile, we analysed the assemblages of wild growing plant species in two ubiquitous park habitat types (grasslands, wooded areas) in 15 parks (150 plots) along an urban-rural gradient. We first used linear contrasts to compare species richness, beta diversity and the proportion of introduced species. We then tested for the explanatory value of environmental variables operating at different spatial scales (plot, park, urban matrix). Unlike in most previous studies, biodiversity patterns were not related to the position of the parks on the urban-rural gradient. Introduced species, mostly from Europe, generally dominated both habitat types (>90 %). Socio-economic (population growth or density), but not spatial, variables were retained in most models. Maintenance intensity was most influential in predicting species assemblages, complemented by park age in wooded areas. A high proportion of European grassland species indicates a trend of homogenisation in park grassland at a cross-continental scale. We conclude that habitat functions of urban parks for native species that have been mainly demonstrated for Europe cannot be generalised to South American megacities. This highlights the need for innovative and locally appropriate conservation approaches (e.g., re-introduction of native species) to foster biodiversity functions in urban parks of South American megacities.     

The impact of urbanisation on avian species: The inextricable link between people and birds

This study investigates the impact of urbanisation on birdlife in a major city. Line transects and point counts were used to survey birds in three habitat types: parkland, residential areas and business/industrial areas. Abundance, richness and diversity of assemblages were determined for all bird species and for those birds native to the area. Behaviours of birds, and of human residents in relation to birds, in these urban areas were documented, including all instances of avian aggression. Bird species, including a subset of native bird species, have greater abundance and richness in parklands. Overall species diversity is greatest in residential habitat types, but native diversity is greatest in parklands. Introduced species are most abundant in business/industrial habitat types. The most frequent aggressive encounters were initiated by noisy miners Manorina melanocephalas, one of the four most common species throughout all habitat types (other common species include the rainbow lorikeet Trichoglossus haematodus, rock dove Columba livia and common myna Sturnus tristis). Other behaviours involved birds utilising food and roost resources and were classified as being caused by active and passive human behaviours. These outcomes indicate that local changes to the environment can impact the bird species by providing different food and roost resources. Human residents and local governments have a range of tools to modify the diversity of urban areas. Further research is needed to determine alternative definitions of modification, such as defining it as open space, and investigating the health of the avian populations in urban areas.     

Impact of urban structure on avian diversity along the Truckee River, USA

Urban riparian habitats are potentially important resources for native birds in arid ecosystems. Most studies have assessed the value of urban riparian habitat in terms of vegetation and natural resources; however, the surrounding land use and infrastructure may determine the viability of urban habitat. We studied the impact of urban structure, the combination of land use, infrastructure and vegetation variables that work together to shape the urban environment, on avian riparian habitat in the Truckee Meadows, Nevada, USA. Land use and infrastructure explained avian species richness and abundance better than local vegetation alone, but community resemblance was more strongly correlated to vegetation. Avian species guilds responded differentially to surrounding land use, suggesting there may be a functional difference between land use types. The best models for bird diversity used urban structure (both land use and vegetation) to describe potential habitat. Urban structure describes urban habitat in ways that vegetation variables alone cannot. Studies that ignore land use and infrastructure and other socioeconomic variables are likely missing key functional differences within urban ecosystems, and may miss the potential for compatible development that encourages both biodiversity and urban growth.     

Relation between green spaces and bird community structure in an urban area in Southeast Brazil

Increased urbanization typically leads to an increase in abundance of a few species and a reduction in bird species richness. Understanding the structure of biotic communities in urban areas will allow us to propose management techniques and to decrease conflicts between wild species and human beings. The objective of this study was to describe the structure of the bird community in an urban ecosystem. The study was carried out in the city of Taubaté in southeastern Brazil. Point-counts were established in areas with different levels of tree density ranging from urban green spaces to predominantly built-up areas. We looked for a correlation between the richness/abundance of birds and the size of the area surveyed, the number of houses, the number of tree species and the number of individual trees. The results of multiple regression showed that bird richness had a direct relationship with vegetation complexity. The abundance and diversity of tree species were better predictors of bird species than the number of houses and size of the area surveyed. We discuss implications of this study for conservation and management of bird diversity in urban areas, such as the need to increase green areas containing a large diversity of native plant species.     

Small mammal assemblages in fragmented shrublands of urban areas of Central Chile

Mediterranean-type ecosystems are one of the most affected environments by habitat loss and fragmentation due to urban development, however only few studies have evaluated the effects of urbanization on the biodiversity of remnant fragments in these ecosystems. This study aims to evaluate the effects of urban development over small mammal assemblages inhabiting isolated forest fragments of an urban area of Chilean Mediterranean zone. We compared abundance and richness of small mammal assemblages of six remnant fragments within an urban matrix, and six fragments similar in area and habitat characteristics with those of urban area, but surrounded by a rural matrix. We found that small mammal assemblages differ considerably among fragments types (urban vs rural), with lack of endemic species from urban fragments and with high proportion of introduced rodents in urban fragments. Furthermore abundance of small mammals was higher in rural than in urban fragments. In urban areas small mammal abundance and richness were not correlated with any of the explanatory variables assessed (woody cover, flora heterogeneity, fragment area, perimeter/area ratio). However in rural fragments small mammal richness was negatively correlated with flora heterogeneity and the abundance of small mammals was positively correlated with perimeter/area ratio. These results reveal important differences within the effects of fragmentation over small mammal assemblages among the two types of fragments assessed. Our findings suggest that in forest fragments isolated by urbanization, larger areas with good quality habitats are not sufficient to maintain native small mammal population.     

Influence of human population size and the built environment on avian assemblages in urban green spaces

While studies of how habitat patch dynamics structures avian communities in urban environments has received some attention, there is considerably less known of how the built environment and human population size may influence the structuring of urban bird communities. We investigated bird populations in Pittsburgh, Pennsylvania (U.S.A.) through replicated point count surveys of breeding birds in 50 parks and cemeteries of varying sizes. We counted 4,435 individual birds in 441 counts. Of the 61 species detected, 27 were rare (detected at <8 points). Migratory species accounted for 46.1 % of all individuals, while 23.2 % of all individuals were of introduced species. Species richness increased significantly with green area size, as did the number of rare species. Species diversity decreased significantly with an increase in the proportion of individuals of introduced species; in particular, cavity nesters were less abundant when introduced species were present. Elements of the built urban environment including commercial development and transportation corridors were associated with significant reductions in park-wide species richness, mean number of species per point, and mean number of individual birds recorded per point. Human population size was positively related to increased numbers of individuals of introduced species, but a lower mean number of species per point. Ours is among the first to identify specific relationships between avian population characteristics and human population size, as few other studies have specifically incorporated human population size into a local, fine grain study design. Our data suggest that human population size is an important parameter that can be measured independently of characteristics of the built environment and the physical characteristics of the park itself as a correlate of avian diversity and abundance. Our study points to a variety of trade-offs needed to manage habitat for birds in urban settings.     

Urban park area and age determine the richness of native and exotic plants in parks of a Latin American city: Santiago as a case study

Plant diversity and composition in urban parks is determined by socio-economics, environmental, and ecological drivers. These drivers give rise to urban spaces with unique compositions of flora, consisting of planted and spontaneous species. The present study aimed to determine the contribution of native and exotic species in urban parks of a Latin American city, and to assess the related role of park area, age and socio-economic status. We also evaluated administration type (public or private) and the effect of environmental factors on plant richness. We hypothesized that the composition of park vegetation differs according to urban-rural gradient. To determine flora composition, two transects (100 m long by 1–3 m wide) per park were selected in 49 parks. In each of these, we identified all the vascular plants found (woody and herbs, planted and spontaneous), which were then classified as native or exotic. We conducted ANCOVAs in order to determine the effect of five independent variables and one factor, on native and exotic plant richness. Of 550 recorded species, 16.2% were native and 83.8% exotic. Number of plant species per park varied between 42 and 146. The parks are known urban habitats containing the highest planted and spontaneously occurring exotic diversity in the city of Santiago, contrasting with other types of habitats. Likewise, parks in Santiago are habitats with low levels of planted and spontaneously occurring native diversity, compared to the parks of Europe, America North and Asia. Ours results show that park area and age affected native plant richness, while exotic plant richness was determined only by park age. We rejected all other possible drivers. Finally, according to the low frequency of native species, we propose that Santiago’s parks could be gradually reoriented towards ex-situ conservation of native plants.     

Patterns of human disturbance and response by small mammals and birds in chaparral near urban development

We report on the extent of disturbance (including habitat alteration and road and trail proliferation) in chaparral near urban development and analyze the effects of disturbance on small mammal and resident bird species. Disturbance patterns were evaluated in a 6700 ha study area in southern California: effects on mammals and birds were investigated by analyzing relationships between vegetation structure and animal species richness and abundance. Disturbance was prevalent throughout the study area and included extensive human-altered habitat (from past human activities such as vegetation clearing, human-caused fires, refuse dumping, and vegetation trampling) and 157 km of roads and trails. A nonsignificant trend was found between human-altered habitat and proximity to development, but human-altered habitat was significantly associated with roadway proximity. Trails were also more frequent near urban development and roads. Small mammals responded strongly to disturbance-related vegetation changes, while birds showed little or no response. Mammals endemic to chaparral vegetation were less diverse and abundant in disturbed sites, whereas disturbance-associated species increased in abundance. Close proximity of urban development to natural areas resulted in alteration of natural habitat and proliferation of roads and trails. Variation in life history traits between birds and mammals may affect response to disturbance and influence persistence if disturbance continues. Conservation efforts must recognize the potential for habitat damage and associated declines in native animal species caused by disturbance near urbanization and implement strategies to reduce these threats.     

Does urbanization affect the seasonal dynamics of bird communities in urban parks?

The environmental factors affecting the spatial dynamics of bird communities in urban parks are well understood, but much less attention has been paid to the seasonal dynamics of bird communities. Since migrant and resident human commensal birds might have contrasting responses to environmental factors of urban parks, we expected different seasonal dynamics among parks. On the other hand, because bird species can have different habitat relationships throughout the year, we also expected different responses of bird richness to environmental variables between breeding and non-breeding seasons. Bird surveys were conducted in 14 small urban parks (1–4 Ha) of Mar del Plata city (Argentina) for one full annual cycle. Bird richness changed between seasons, but bird abundance remained constant. Bird community composition did not vary between seasons, but urban parks near the urban center, with the highest pedestrian traffic and isolation to other green areas had the least seasonal change of composition. During the breeding season, bird richness was negatively affected by the percentage cover of high buildings surrounding the immediate limits of parks, whereas during the non-breeding season bird richness was not related with any environmental variable. Bird composition variation among parks was affected by the distance to the urban center during both seasons. Results showed that urbanization promotes a seasonal homogenization of bird communities in urban parks, probably by affecting the presence of migrant species and promoting the temporal stability of human commensal species.     

Floral biodiversity of allotment gardens and its contribution to urban green infrastructure

Allotment gardens are a key feature of the urban landscape, providing numerous benefits beyond those of food production. These include ecosystem services such as local climate modification, pollination, and providing a pleasant location for socialising. The biodiversity of flora underpins many of these ecosystem services. The aims of this study are to assess the spontaneous vascular flora of urban allotment gardens and derive estimates of the species richness and diversity. In doing so, the specific position of allotment gardens within urban green infrastructure will be revealed and the role of allotment users in shaping local biodiversity determined. 358 species of spontaneous flora were recorded in 11 representative allotment garden estates (total area 150 ha) in Poznań, Poland. Fourteen biodiversity features were taken into account. The investigated areas exhibited high plant species richness, high diversity, low synanthropisation level, and contained valuable geobotanical elements, including plants of key importance for the European Community. For these reasons, allotment gardens should be considered as biodiversity hotspots for native species within urban green infrastructure. Based on an analysis of the spatial and functional organization of 110 individual plots, four types of usage, and corresponding plant diversity have been distinguished. The composition of the flora depends mostly on spatial and functional organization of allotment gardens and, to a much lesser extent, on the habitat conditions, which on individual plots varied very little. The high species richness and diversity in allotment gardens has important consequences for urban planning.     

Effects of area size,heterogeneity, isolation,and disturbances on urban park avifauna in a highly populated tropical city

We assessed bird diversity and nesting activity in 54 urban parks in the core of a highly populated East-Asian tropical city to investigate the effects of area, isolation, habitat heterogeneity, and human disturbances on avifauna. Tree density and heterogeneity were lower while isolation was greater in small than in large or medium-sized parks. Yet, park features were often interrelated, with isolation variables negatively correlated to each other, heterogeneity positively correlated to area, and habitat variables mostly positively correlated with one another but variously to disturbances; whereas pedestrian densities wee positively correlated with green area proportions (ISO3) but negatively with the distances to large green areas (ISO2). Park size played a primary role by positively affecting species richness and numbers of nesting species, thus contributing to higher total and nesting species heterogeneity, and determining the occurrence of a greatest number of species. Pedestrian density, canopy cover, and ground heterogeneity played secondary roles, whereas isolation appeared less influential at the community level. Omnivores, granivores, and aerial insectivores appeared more abundant but were dominated by several common species. Nesting was restricted to even fewer species. While species varied in the occupancy responses to these factors, tree heterogeneity positively affected the nesting of Zosterops japonicus and cavity-nesting birds, and predator density negatively affected those of Lonchura punctulata and Streptopelia doves. Improved urban land use planning and park management incorporating these effects should be implemented to enhance breeding and avoid compositional homogenization for long-term sustainability of urban avian diversity.