Terrestrial vertebrates in urban ecosystems: developing hypotheses for the Gwynns Falls Watershed in Baltimore,Maryland

We review the literature on terrestrial vertebrates in urban and urbanizing areas in terms of the new paradigm in ecology that focuses on the origin, spatial pattern, and current influences of habitat patches. Studies of terrestrial vertebrates have documented differences in species composition along an urban-rural gradient and have identified some of the key land use features associated with these differences. We use information from a database developed for the Gwynns Falls Watershed in Baltimore City and County, Maryland to demonstrate how a study of forest-dependent breeding birds could be used to identify mechanisms that account for differences in the bird communities of forest patches in urban areas, and to provide useful information to planners and managers involved in wildlife conservation and management.     

Associations between multiscale landscape characteristics and breeding bird abundance and diversity across urban-rural gradients in Northeastern Georgia, USA

Urbanization is rapidly changing the southeastern US landscape, particularly in Georgia within and around the Atlanta metropolitan area, the fastest-growing Southeastern population center. Landscape characteristics related to human disturbance, landscape composition, and landscape configuration vary across the urban-rural gradient and are easily and inexpensively evaluated using geographic information systems (GIS). Previous studies have suggested that avian communities and populations respond in scale-dependent ways to urbanization-related landscape changes through changes in abundance and diversity. We conducted a 2-year study (2007?C2008) of the response of breeding bird population abundance, community abundance, species richness, and relative diversity in young and mature single-family residential areas to 15 landscape characteristics measured at multiple spatial scales across urban-rural development intensity gradients near Athens, Georgia. We grouped birds by species, entire site assemblage, and by functional guild affiliation. Our models suggest strong relationships between the landscape and both community and population abundance of birds, as well as a somewhat weaker relationship with species richness. We detected only a weak relationship between breeding season relative diversity and landscape characteristics; we hypothesize a possible seasonal component to that relationship based on an earlier study. Our results suggest that widely available geospatial metrics of human disturbance and landscape pattern can be used to model breeding bird abundance and diversity across urban-rural gradients. This study sets the framework for a landscape scale understanding of the effects of housing developments and development intensity on breeding birds in northeastern Georgia and such a landscape-based approach to modeling species numbers represents a valuable tool in natural resources management.     

The pattern of distribution and diversity of avifauna over an urbanizing tropical landscape

An investigation was done to determine the occurrence and composition of avian fauna community in the urbanizing city of Nairobi, Kenya. We conducted bird counts in sample sites randomly distributed over the Nairobi landscape within a two-year period. The proportion of seven different land cover types derived from within a 500 m radius of classified satellite image described the habitat condition of each sample site. Multivariate analysis of the site and species data indicated that savannah vegetation, forest and agriculture land cover types were the main environmental gradients that differentiated the sample sites. Four clusters of sample sites occurred on the ordination plane according to canonical correspondence analysis (CCA). From 50 families of birds observed, species related to bush and scrub habitats occurred at a rate of about 31 %, followed by grassland species at 20 % and forest species at about 16 % rate. Out of CCA, five functional groups of birds were distinguishable. The proportion of sites occupied by birds per functional group and mean count of individual birds declined significantly, p?<?0.0001, as groups of birds changed from being urban-related to savannah and then to forest-related species. One cluster comprised birds marked as having conservation concern status and related mainly to savannah vegetation of scrub and grasslands. In this landscape context, features such as native savannah vegetation and woodland accompanied by a process of controlled land use, could greatly mitigate negative impacts of ecosystem degradation on the sensitive tropical urban bird biodiversity.     

Seasonal dynamics of bird communities in urban forests of a Mediterranean city (Montpellier,Southern France)

We evaluated the richness, composition and abundance of bird communities in three urban forests of Mediterranean France during winter and spring. The urban forests differed in size, composition, structure, age of vegetation, and location relative to the city centre. Estimated species richness across all three urban forest parks was 45 species. Twenty six (26) species were recorded in both winter and spring, whereas ten species were recorded only in spring, and six were recorded only in winter. Distribution, turnover, and type of bird foraging guild were related to characteristics of each urban forest and season. During spring migration, more species were recorded in sampling units (250?×?250 m) within the largest and most natural urban forest located in the outskirts of Montpellier, whereas during winter, more species were recorded in the most urbanized park (i.e., a botanical garden dominated by exotic vegetation), which was located in the city centre. Insectivorous birds were more abundant in winter, whereas seedeaters associated with wooded habitats were recorded more frequently in spring. Our results suggested that different kinds of urban forests are important modulators of urban bird communities and they are necessary to maintain the diversity of migratory and resident birds as well as increasing the environmental quality of urban areas.     

Composition and conservation value of bird assemblages of urban ‘habitat islands’: Do pedestrian traffic and landscape variables exert an influence?

Urban forest fragments can potentially be ‘habitat islands’ for native birds, but pedestrian traffic could influence the composition and conservation value of their bird assemblages by reducing the representation of disturbance-sensitive species. Winter bird assemblages and pedestrian traffic rates were documented in twelve forested parks in suburban Melbourne, Australia. Native birds comprised 87% of sightings, but two exotic species were common. Bird species' distribution among the parks was significantly nested. However, park rank orders for bird assemblage nestedness and mean pedestrian traffic rate were not correlated and only one species' distribution was nested with respect to this variable. The observed correlation between park area and bird assemblage nestedness may have been partly due to passive sampling. Degree of isolation of parks influenced nestedness less than park area. Forested parks provided habitat for species that occur at intermediate to low densities in other suburban habitats and for ecological specialists otherwise rare in suburbia. Larger parks contained more species than, and many of the native species found in, smaller parks. However, smaller parks each had 25% of the native bird species inhabiting the suite of fragments and thus had both aesthetic value and some potential to enhance city-dwellers' awareness about biodiversity conservation.     

Forest bird communities across a gradient of urban development

This study examined native bird communities in forest patches across a gradient of urbanization. We used field data and multivariate statistical techniques to examine the effects of landscape context, roads, traffic noise, and vegetation characteristics on bird community composition in the North Carolina Piedmont (U.S.A.). Landscape-level variables, particularly those related to urbanization, were most important in structuring forest bird communities. Specifically, we found that road density and amount of urban land cover were the best predictors of species composition. We found that urban and rural bird communities were quite distinct from each other. Rural communities had more long-distance migrants and forest interior species but species richness did not differ between the communities. Our results suggest some specific guidelines to target bird species of interest both inside and outside of urban areas. For example, if increasing numbers of migratory species is of primary concern, then conservation areas should be located outside of urban boundaries or in areas with low road density. However, if maximizing species richness is the focus, location of the conservation area may not be as important if the conservation area is surrounded by at least 50 m of forest habitat in all directions.     

The causal response of avian communities to suburban development: a quasi-experimental,longitudinal study

Correlations between urbanization and biodiversity are well known, but the causes driving such associations are lacking. We used a long-term, quasi-experimental approach to study the responses of avian communities to suburban and exurban development around Seattle, WA, USA. We measured indices of bird abundance, reproduction, and survival for 12 years at many locations, including 5 forest ‘reserves,’ 10 existing ‘developments,’ and 11 ‘changing’ sites where ongoing development converted forests to single-family residential neighborhoods. In the first few years of clearing, building, and occupation of new neighborhoods by humans avian communities shifted from those typical of second-growth forest to those more characteristic of developments. During this time avian diversity increased and numerical dominance by abundant birds declined. Species that adapted and exploited development reproduced more successfully there than did forest-dependent species that avoided development. Adults of species that thrived in developments attained equal annual survival across reserved to developed landscapes, while species that avoided neighborhoods tended to survive poorly outside of reserves. The humans living in our study areas frequently fed birds and provided nest boxes. These actions were positively correlated with increases in secondary cavity nesting and seed eating birds. Humans also maintained outdoor cats and 11 % of humans both fed birds and let their cats outside. These actions were negatively correlated with the abundance of birds regularly using feeders. We suggest that a key management goal in urban ecosystems is the maintenance of avian diversity because a diverse avifauna engages a diversity of humans.     

Relative importance of habitat quantity, structure, and spatial pattern to birds in urbanizing environments

Urbanization reduces the quantity of native vegetation and alters its local structure and regional spatial pattern. These changes cause local extirpations of bird species associated with native vegetation and increases in the abundance and number of bird species associated with human activity. We used 54–1 km² landscapes in the Seattle, Washington, USA metropolitan area to determine (1) the relative importance of habitat quantity, structure, and pattern to bird diversity and abundance and (2) whether housing developments can be managed to mitigate the negative impacts of urbanization on forest bird diversity. In general, bird species richness was high and many native forest species were retained where urban landcover comprised less than 52% of the landscape, tree density (especially that of evergreens) remained at least 9.8 trees/ha in developments, and forest was at least 64% aggregated across the landscape. These results suggest that the quantity, structure, and pattern of forested habitat affected breeding bird diversity in urbanizing landscapes. However, habitat pattern appeared less influential than other habitat attributes when results from all community- and population-level analyses were considered. Conservation of native birds in reserves can be supplemented by managing the amount, composition, structural complexity, and—to a lesser extent—arrangement of vegetation in neighborhoods.     

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.     

Effects of urbanization on breeding birds in European towns: Impacts of species traits

Urbanization acts as a filter on bird species behavioral traits so that only few species can tolerate urban constraints. We analyzed how behavioral traits (nesting, feeding, and migratory habits) of breeding bird species affect their frequency of occurrence in the urban centers of 38 European towns. We used binary logistic regression analysis to predict the bird species traits belonging to each trait group. A total of 108 species (21% of the European breeding bird species) were found to breed in the European town centers. According to our broad-scale analyses the bird species most frequently breeding in town centers nest in buildings and/or buildings have diverse diets, in trees (40%) and are resident omnivores, or relied on seeds or fruits as their sources of food. However, almost all bird species also fed on arthropods (92%) during the breeding season. Only a few urban bird species bred on the ground. Four out of the studied 108 species were non-native and five species were predators. Our broad-scale results from Europe indicate that bird species with different behavioral traits can respond differently to urbanization. Bird species that nest in cavities/buildings have diverse diets, that benefit a resident way-of-life, may have an advantage in living and settling in European town centers. Our results from Europe may provide insights related to the development of bird assemblages in the urban core areas of the New World.     

Value of an urban habitat for the native Mediterranean avifauna

Mediterranean landscapes resulted from complex land uses that produced a mosaic of extensive crops, grasslands, scrublands and scattered woodlands. During the twentieth century the decrease in traditional agriculture triggered a decrease in open habitats and an increase in forests. In the meantime urban centres grew dramatically. Both spread of forest and urban areas have been suspected to participate in the decline of typical Mediterranean bird species and, in general, to cause faunal loss. However, modern cities offer a variety of landscape types and, in the Mediterranean, their value for native bird species has been little assessed. We compared the bird communities from an urban landscape, including built up and natural or semi natural units within the limits of the city of Montpellier, to the bird communities from non-urban habitats (cropland, grassland and woodland) located nearby but away from direct urban influence. Fifty four percent of the bird species recorded in the non-urban habitats also occurred in the urban landscape. On average, estimated species richness in the urban landscape was similar to values obtained for non-urban habitats. Within the urban landscape species richness was lowest in the dense historical centre and highest in the residential areas. The residential areas and urban woods were suitable habitats for most generalist species but also for several more specialized species recorded in the non-urban habitats. Some species actually reached their highest observation frequency in the urban landscape. Urban landscape was least favourable to the same farmland and open-habitat specialists that have been negatively affected by agricultural abandonment in the region. Finally, five of the species common in the urban landscape had an unfavorable conservation status in 2004 in the EU. This study emphasizes that Mediterranean urban areas have the potential to host a diverse native bird community. Finding ways to improve their carrying capacity for the local avifauna might be a worthwhile objective for animal conservation.     

Bird responses to housing development in intensively managed agricultural landscapes

Housing a growing human population is a global issue and designing environmentally friendly developments requires identifying the species likely to be negatively impacted and finding mitigation solutions. Existing studies that consider fragmentation of natural habitats have limited application in countries such as Britain where a prime target for development is agricultural land where decades of intensive management have already diminished biodiversity. Here I used citizen science data on the abundance of 146 breeding and wintering birds to develop models linking abundance to human population density and habitat features. I used these as a proxy for the urbanisation process, finding that impacts of urbanisation were species-specific and context dependent. Low-density developments benefited a high proportion of birds, with wetland birds benefitting most and farmland birds least, but as human densities increased further, up to 75% of species were negatively impacted. Almost half of species currently occurring at 14 flagship residential development sites were predicted to decline based on projected human population density increases, with a third predicted to increase. Presence of wetlands, canopy cover and patches of trees all benefited certain species but efforts to identify more detailed habitat associations were hampered by collinearity among variables. I conclude that even in heavily degraded agricultural landscapes, a high proportion of species will be negatively impacted by residential development and that some will require spared land to persist in the wider landscape. As no single habitat benefited the entire bird community, urban planners wishing to design bird-friendly developments will need to make difficult decisions over which aspects of the bird community to prioritise.

     

Avian diversity in a suburban park system: current conditions and strategies for dealing with anticipated change

A growing trend towards increased urbanization emphasizes the role of suburban parks in wildlife conservation. Spatial planning aimed at maintaining biological diversity and functionality must consider how changes at landscape and more local scales will influence the biotic structure of urban areas. From May 2006 to July 2010, bird surveys were conducted in three metropolitan parks in Cleveland, Ohio, USA. Surveys were conducted with the goal of examining the effect of vegetation structure and adjacent land cover on the distribution and species richness of breeding birds within this park system. A total of 65 species were recorded throughout the study area. Avian species richness was linked to several habitat metrics, measured at both the local and landscape scale. Generally, species richness was highest at locations characterized by moderate forest cover. The proportion of canopy cover at survey sites related negatively to species richness and the density of understory vegetation showed a positive relationship with species diversity. Despite the influence of these three metrics, sensitivity analysis indicates that the density of understory vegetation is the most significant correlate to avian diversity within this suburban park system. Management actions aimed at providing habitat for the greatest diversity of breeding songbirds within the study area should allow for moderate canopy cover while retaining or improving the structural complexity of understory vegetatation.     

Impacts of traffic noise on anuran and bird communities

Anurans and birds rely on sound for a number of social behaviors. Species that use roadside habitats are exposed to traffic noise that can mask important social signals and directly affect the community diversity and composition. We evaluate the impact of traffic noise on anuran and bird species richness, species occurrence, and composition in Puerto Rico, where there is a high density of highways and cars that generate high levels of noise pollution. We compared paired forest sites near (100 m, n = 20, dB > 60) and far (>300 m, n = 20, dB < 60) from highways, with similar vegetation structure, but different levels of noise. We found that the anuran community was not affected by traffic noise. In contrast, bird species richness and occurrence were significantly lower in sites near the highway, and bird species composition also varied significantly. Bird species with low-frequency songs were only detected in sites far from highways. The differences in the ecology and communication behavior between anurans and birds could explain these results. Anurans mainly call at night, when traffic activity was low. In contrast, bird singing activity occurs during the day and overlaps with the high levels of traffic noise. In addition, in natural habitats, Puerto Rican anurans occur at high densities and form noisy choruses (>80 dB), which may allow them to tolerate high levels of anthropogenic noise.     

Forest cover is important across multiple scales for bird communities in vacant lots

Urban vacant lots can vary considerably in their vegetation structure, from dense, shrubby habitats to wooded remnant fragments that may provide habitat for a variety of birds. By identifying which features promote diverse bird communities, we can determine at which scale management practices should focus and the necessary habitat structure and composition features. We surveyed 150 vacant lots throughout Baltimore, Maryland for their bird communities, lot vegetation, and landscape-level forest cover. An ordination of the bird community indicated a response to a gradient of canopy cover and canopy height at the vacant lot. We also found that forest cover within 100 m of the vacant lot was the most important predictor of abundance for five bird species of interest. Species richness was spatially autocorrelated among sites, indicating that bird communities may also be driven by species’ dispersal and environmental gradients across the city. Overall, bird communities are responding to habitat features across multiple scales, from the vacant lot vegetation, to landscape-level forest cover, to city-wide dynamics. Thus, we recommend management practices to focus on increasing city-wide forest cover in order to increase species richness, yet with awareness regarding where the lot occurs within the city.     

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.

     

Ecosystem processes along an urban-to-rural gradient

In order to understand the effect of urban development on the functioning of forest ecosystems, during the past decade we have been studying red oak stands located on similar soil along an urban-rural gradient running from New York City ro rural Litchfield County, Connecticut. This paper summarizes the results of this work. Field measurements, controlled laboratory experiments, and reciprocal transplants documented soil pollution, soil hydrophobicity, litter decomposition rates, total soil carbon, potential nitrogen mineralization, nitrification, fungal biomass, and earthworm populations in forests along the 140 × 20 km study transect. The results revealed a complex urban-rural environmental gradient. The urban forests exhibit unique ecosystem structure and function in relation to the suburban and rural forest stands these are likely linked to stresses of the urban environment such as air pollution, which has also resulted in elevated levels of heavy metals in the soil, the positive effects of the heat island phenomenon, and the presence of earthworms. The data suggest a working model to guide mechanistic work on the ecology of forests along urban-to-rural gradients, and for comparison of different metropolitan areas.     

Conservation-priority grassland bird response to urban landcover and habitat fragmentation

As urbanization in the landscape increases, some urban centers are setting aside habitat for wildlife. This habitat may be particularly valuable to declining or conservation-priority species. One group in particular need of conservation actions that may benefit from habitat located in urban areas is grassland birds. Declines of grassland bird species have been particularly severe in the Midwestern U.S., where most grassland cover has been lost, fragmented, and surrounded by unsuitable habitat. Conservation efforts have focused on protecting large grasslands surrounded by minimal amounts of trees and development. Although urban development is considered hostile to grassland birds, this assumption has received little attention. In heavily fragmented landscapes where habitat is limited, urban grasslands may be of significant value to grassland birds. We examined grassland bird response to development and additional landscape and habitat variables in the greater Chicago metropolitan area. In 2012 and 2013, we surveyed bird communities in grassland patches along a gradient of urbanization and patch sizes. Density of Savannah Sparrows (Passerculus sandwichensis) increased with amount of development, while density of Sedge Wrens (Cistothorus platensis) decreased. Development did not appreciably impact Bobolinks (Dolichonyx oryzivorus), Dickcissels (Spiza americana), Eastern Meadowlarks (Sturnella magna), Grasshopper Sparrows (Ammodramus savannarum), or Henslow’s Sparrows (Ammodramus henslowii). Patch size had a positive effect on species densities. These results indicate that for many conservation-priority grassland birds, urban landcover surrounding grasslands generally has neutral rather than negative effects on habitat use. Therefore, grasslands in developed landscapes may provide valuable contributions to regional conservation efforts.     

Spontaneous plant colonization and bird visits of tropical extensive green roof

Green roofs can contribute to enrichment and conservation of urban ecology. An experimental green roof was established in humid-tropical Hong Kong to monitor over two years its spontaneous colonization by plants and bird visits. Some 94 voluntary vascular plant species from 26 families and 76 genera were established, with propagules brought mainly by birds and wind and secondarily inherited from soil seed bank. Plant species composition changed dynamically during the study period. They fall into three groups, namely dominant ruderal (herbaceous and sub-shrub) as surrogate of early-stage local grassland ecosystem succession, arboreal (trees and shrubs), and hygrophilous herb. Progressive increase in vegetation cover was accompanied by changes in species diversity and evenness. In addition, 16 bird species from 8 families and 14 genera were recorded. Ten species were residents and the six migrant species were winter visitors. Their food preference was mainly omnivore and insectivore. Winter and the second year encountered higher species richness, diversity, and evenness. Most vegetation parameters correlated positively with avian community indexes, signifying provision of sustenance by green-roof ecosystem to birds. Vegetation coverage correlated negatively with avian abundance, due to shunning by the abundant ground-foraging Tree Sparrow. Both local common ruderal plant species and common urban bird species can successfully establish and reproduce on the extensive green roof, confirming potentials for urban ecology and biodiversity enhancement and conservation even in densely-developed urban areas. The successful nurturing of naturalistic green roof offers new opportunities for green roof design that deviates from the predominant cultivated-horticultural approach.     

Avian haemosporidian parasites in an urban forest and their relationship to bird size and abundance

Urbanization has been identified as a threat to biodiversity due to landscape modifications. Studies of parasite ecology in urbanized areas lagged behind those made on macro organisms. Here we studied infection prevalence of haemosporidian parasites in an avian community of an urban forest from Germany, and its relationship with bird abundance and body mass. We used PCR to amplify a fragment of the mtDNA cyt b gene to determine the infection status of birds, and bird point counts to determine bird relative abundances. The avifauna was dominated by two small sized insectivore passerines (Parus major, Cyanistes caeruleus), representing ~40 % of the total bird records. The highest haemosporidian prevalence was recorded for Turdus philomelos (100 %) and for Fringilla coelebs (75 %). Bird abundance and body mass were positively associated with infection status for two haemosporidian genera: Plasmodium and Leucocytozoon. Infection rate was lower in juveniles compared to adult birds. We recorded a total of 7 Plasmodium, 26 Haemoproteus, and 10 Leucocytozoon lineages. Avian malaria (P. relictum) was detected infecting 5 individuals of P. major, the most abundant species in the community. These results, together with those of previous studies at the same site, suggest that potentially any of the genetic haemosporidian lineages detected in this urban forest can be transmitted across native and pet bird species, and to species of conservation concern housed at aviaries.