Lean birds in the city: body size and condition of house sparrows along the urbanization gradient

1. Urbanized habitats differ from natural ones in several ecological features, including climate, food availability, strength of predation and competition. Although the effects of urbanization on avian community composition are well known, there is much less information about how individual birds are affected by these human-generated habitat differences. 2. In this study we investigated the relationships between the morphological characteristics and the degree of habitat urbanization in house sparrows, Passer domesticus (Linne 1758) . We collected data for more than 1000 non-breeding adult birds in Hungary between 1997 and 2006, from seven sites including farmlands, suburban areas and city centres. 3. We found that the body mass, tarsus length and body condition of free-living sparrows differed among the sites: birds in more urbanized habitats were consistently smaller and in worse condition than birds in more rural habitats. A composite measure of habitat urbanization (based on building density, road density and vegetation cover) explained over 75% of variance between sites in the studied traits, after we controlled for the effects of sex, year, season and time of capture. 4. The difference in body mass between rural and urban sparrows was significant when birds were kept in aviaries under identical conditions, with constant ad libitum food availability. It is therefore unlikely that the reduced body size and condition of urban sparrows are a consequence of reduced access to food for adults (e.g. due to strong competition), or their short-term responses to high food predictability (e.g. by strategic mass regulation). 5. We suggest that habitat differences in nestling development or adaptive divergence of sparrow populations due to distinct environmental conditions (such as differing predation pressure) may account for the differences along the urbanization gradient.     

Response to Predation Risk in Urban and Rural House Sparrows

Habitat urbanization may change the density of predators, and it is often assumed that such changes lead to altered predation risk for urban populations of their prey. Although it is difficult to study predation hazard directly, behavior responses of prey species may be informative in inferring such habitat differences. In this study, we compared the risk‐taking behavior of urban and rural house sparrows (Passer domesticus) after simulated attacks by two of their important predators (sparrowhawk Accipiter nisus and domestic cat Felis catus). The birds were startled by moving dummies of these predators and respective control objects, and their risk taking was estimated as their latency to feed after the startle. We found that sparrows responded more strongly (had longer post‐startle feeding latencies) to sparrowhawk attacks than to the control object, and their responses differed between the habitats. First, risk taking of urban birds strongly decreased with age (older birds had longer latencies than young birds), while there was no such age difference in rural birds. Second, young urban birds responded less strongly, while older urban birds responded more strongly to the sparrowhawk than the same age groups of rural birds, respectively. We did not succeed in evoking antipredatory response by simulated cat attacks, because birds responded similarly to the dummy and the control object. Our results support that predation risk, posed at least by avian predators, is different in urban and rural habitats of house sparrows. The increased wariness of older, hence presumably more experienced, urban birds implies that sparrows may be more exposed to predation in cities.     

Does urbanization select for weak competitors in house sparrows?

How urbanization affects animal populations is in the focus of current ecological research. Existing theory of this topic suggests that the cities' more constant food supplies and lower predation pressure lead to a high proportion of weak competitors in urban populations. To evaluate this hypothesis, we tested whether competitive performance differs between differently urbanized populations of house sparrows Passer domesticus. We previously showed that wild urban sparrows are smaller and leaner than rural conspecifics, and this difference persists for months under identical captive conditions. Here we compared several aspects of their competitiveness (fighting, scrambling and searching for food) in captive mixed flocks of urban and rural birds. We found that sparrows exhibited consistent individual differences in competitiveness, but these differences were not related either to the degree of urbanization of their original habitats or to their body mass. Moreover, the variance in competitive abilities also did not differ between birds from more and less urbanized habitats. Thus our results did not support the hypothesis that urbanization shifts population structure towards an over‐abundance of weak competitors in house sparrows. We discuss possible explanations why sparrow populations may not differ in competitiveness despite the smaller body mass of urban birds.     

北京市麻雀体质水平沿城市化梯度的变化

麻雀是城市和乡村均有分布的鸟类物种.快速的城市化正在使城市麻雀的栖息地和食物资源大量减少.以北京市为例,研究了麻雀的体质水平沿城市化梯度的变化,以期为我国城市化过程中的鸟类保护提供参考.研究结果表明城市高层楼房居民区、低层楼房居民区以及大学校园中麻雀的体质指数显著低于郊区环境,而城市公园和平房居民区麻雀的体质水平则与郊区环境的差异较小.随采样点城市化水平的增加,麻雀的体质水平呈下降趋势,高度城市化的环境导致麻雀的体质水平下降.在城市化过程中,增加城市公园的数量及居民区的植被覆盖量可为麻雀等鸟类提供必要的生活资源,从而实现城市化过程中的生物多样性保护.     

Novelty Responses in a Bird Assemblage Inhabiting an Urban Area

Neophobia, or the hesitancy to approach a novel food item, object, or place, is an important factor influencing the foraging behavior of animals. Environmental factors (e.g. rapid anthropogenic changes, migration into new habitats) are associated with novelty in feeding ecology and may affect neophobic responses. Mechanisms that underlie the differential neophobic response may involve complex interactions with the environment: post-fledging experience in a greater diversity of habitats or in habitats that are more complex may contribute to reduced neophobia. In a previous study, it was observed that some urbanized species, in particular house sparrows ( Passer domesticus ) and shiny cowbirds ( Molothrus bonariensis ) show high levels of neophobia. This study was carried out in a suburban marsh of Cortaderia selloana , a relatively simple and predictable ecosystem as compared to urban areas. For this reason, in the present study, we explored novelty responses of bird species inhabiting an urban area, representing a complex environment. The results were compared to those obtained previously in the suburban marsh. We found unexpectedly high levels of neophobia in house sparrows, but shiny cowbirds showed a somewhat neophilic response. In the presence of novel objects, house sparrows tended to enter the feeders alone, while shiny cowbirds tended to forage in groups. We found no differences in latencies to forage or in visit duration between habitat types, but the proportion of individuals that visited the feeders when novel objects were present was lower in the urban area for house sparrows and eared doves ( Zenaida auriculata ). The results are discussed in the context of invasion success and feeding innovation in shiny cowbirds.     

Behavioral Syndromes Break Down in Urban Song Sparrow Populations

Animals in urban habitats face a number of unique stresses, including the necessity of dealing with high levels of human activity. Growing research suggests that: (1) inherent traits, as opposed to learned behavior, influence which species invade urban habitats, and (2) individuals exhibit behavioral syndromes that limit behavioral flexibility. As a result, perhaps only animals with inherently bold personalities successfully settle in areas of high human activity, and such animals may also exhibit correlated variation in other behavioral traits, such as territorial aggression. In this study, we examine boldness and aggression in several urban and rural populations of song sparrows (Melospiza melodia). We found that urban birds were both bolder toward humans and also showed higher levels of aggression. We found a correlation between boldness and aggression in all populations combined, but no correlation within urban populations. Our results agree with other recent studies of song sparrow behavior, suggesting that greater boldness and aggression are general features of urban song sparrow populations, and a lack of a correlation between boldness and aggression in urban habitats is a general phenomenon as well. Urban habitats may select for bold and aggressive birds, and yet the traits can vary independently. These results add to a small number of studies which find that behavioral syndromes break down in potentially high quality habitats.     

Individual and Sex Differences in Habituation and Neophobia in House Sparrows (Passer domesticus)

Heterogeneity in the environment favours foragers that are flexible (phenotypically plastic). However, consistent individual differences in behaviour (personality), such as in risk‐taking, might affect an individual's ability to find food, avoid predators and adjust to new conditions. It is now well known that personalities exist in many taxa, but much less is known about individual variation in plasticity. We measured the tendency to begin foraging across different levels of risk in house sparrows (Passer domesticus), using a behavioural reaction norm framework to simultaneously assess personality and plasticity. We asked whether individuals were consistently different across risk levels, and whether they differed in habituation and neophobia, both of which were treated as cases of plasticity. We found that males habituated more than females by beginning to feed sooner after repeated instances of a human disturbance in the presence of an initially unfamiliar object. Individuals of both sexes also exhibited consistent differences across trials, but did not differ in the rate of habituation beyond the difference between the sexes. When a novel object was placed in the foraging area, both sexes exhibited similar degrees of neophobia by delaying feeding. The magnitude of this change varied among birds, indicating individual differences in neophobia. Our results indicate that both personality and individual variation in plasticity exist but should be treated as independent phenomena. The presence of variation in plasticity implies that the raw material necessary for selection on neophobia exists, and that if heritable, plasticity in risk‐taking across contexts could evolve.     

Behavioural Syndromes in Urban and Rural Populations of Song Sparrows

Animals in urban habitats are often noticeably bold in the presence of humans. Such boldness may arise due to habituation, as urban animals learn, through repeated exposure, that passing humans do not represent a threat. However, there is growing research suggesting that: (1) inherent traits, as opposed to learned behaviour, influence which species invade urban habitats, and (2) individuals exhibit individual personality traits that limit behavioural flexibility, with the possible result that not all individuals would be able to demonstrate an appropriate level of boldness in urban environments. As a result, perhaps only birds with inherently bold personalities could successfully settle in an area of high human disturbance, and further, we might also expect to see the existence of behavioural syndromes, where boldness is correlated with variation in other behavioural traits such as aggression. In this study, we examined boldness and territorial aggression in urban and rural populations of song sparrows. We found that urban birds were bolder towards humans and that urban birds also showed higher levels of territorial aggression. We also found an overall correlation between boldness and territorial aggression, suggesting that urban boldness may be part of a behavioural syndrome. However, we see no correlation between boldness and aggression in the urban population, and thus, more work is needed to determine the mechanisms accounting for high levels of boldness and aggression urban song sparrows.     

Urbanization and its effects on personality traits: a result of microevolution or phenotypic plasticity?

Human‐altered environmental conditions affect many species at the global scale. An extreme form of anthropogenic alteration is the existence and rapid increase of urban areas. A key question, then, is how species cope with urbanization. It has been suggested that rural and urban conspecifics show differences in behaviour and personality. However, (i) a generalization of this phenomenon has never been made; and (ii) it is still unclear whether differences in personality traits between rural and urban conspecifics are the result of phenotypic plasticity or of intrinsic differences. In a literature review, we show that behavioural differences between rural and urban conspecifics are common and taxonomically widespread among animals, suggesting a significant ecological impact of urbanization on animal behaviour. In order to gain insight into the mechanisms leading to behavioural differences in urban individuals, we hand‐raised and kept European blackbirds (Turdus merula) from a rural and a nearby urban area under common‐garden conditions. Using these birds, we investigated individual variation in two behavioural responses to the presence of novel objects: approach to an object in a familiar area (here defined as neophilia), and avoidance of an object in a familiar foraging context (defined as neophobia). Neophilic and neophobic behaviours were mildly correlated and repeatable even across a time period of one year, indicating stable individual behavioural strategies. Blackbirds from the urban population were more neophobic and seasonally less neophilic than blackbirds from the nearby rural area. These intrinsic differences in personality traits are likely the result of microevolutionary changes, although we cannot fully exclude early developmental influences.     

Inter-individual variability in fear of humans and relative brain size of the species are related to contemporary urban invasion in birds

Background

Urbanization is the most prevailing cause of habitat transformation worldwide, differing from others by its intense levels of human activity. Despite its obvious impact on wildlife, it is still unclear why and how some species are able to adapt to urban settings. One possibility is that fear of humans and vehicles could preclude most species from invading cities. Species entering urban environments might be those that are more tolerant of human disturbance (i.e., tame species). Alternatively or in addition, urban invaders could be a fraction of variable species, with “tame” individuals invading urban habitats and other individuals remaining in rural areas.

Methodology

Using the contemporary urban invasion by birds in a recently established South American city, we tested both hypotheses by relating interspecific differences in invasiveness to their flight initiation distances (i.e., the distances at which birds flee from approaching cars, FID), as well as to their relative brain size (RBS), a correlate of measures of behavioral flexibility.

Principal Findings

Urban invasiveness was not significantly related to species'' average rural FIDs but positively related to their RBS and inter-individual variability in FID. Moreover, FIDs were consistently lower in urban than in rural conspecifics, and the FIDs of urban individuals were within the lower-range distribution of their rural conspecifics. RBS indirectly influenced urban invasion through its positive effect on inter-individual variability in FID.

Conclusions/Significance

Urban invaders do not appear to be individuals from apparently tame species, but rather tame individuals from species with a variable response regarding fear of people. Given the positive relationship between RBS and inter-individual variability in FID, our results suggest that behavioural flexibility should be regarded as a specific trait encompassing variability among individuals. Further research is needed to ascertain the neurophysiological mechanisms underlying the relationship between brain size and inter-individual variability in behavioural traits.     

Urbanization,nestling growth and reproductive success in a moderately declining house sparrow population

Ecological conditions are likely to change with increasing urbanization, influencing the demography and size of animal populations. Although one of the most tightly linked species to humans, the house sparrow has been suffering a significant decline worldwide, especially in European cities. Several factors have been proposed to explain this conspicuous loss of urban sparrows, but studies evaluating these factors are usually restricted to Britain where the decline was very drastic, and it is unclear whether similar or different processes are affecting urban populations of the species elsewhere. In this study we investigated the reproductive success of urban and rural sparrows in a central European country, Hungary where our census data indicate a moderate decline during the last decade. We found that rural pairs produced more and larger fledglings than suburban pairs, and the difference remained consistent in two years with very contrasting meteorological conditions during breeding. This difference is likely explained by habitat differences in nestling diet, because we found that 1) rural parents provided large prey items more often than suburban parents, 2) birds from differently urbanized habitats produced fledglings of similar number and size in captivity under identical rearing conditions with ample food for nestlings, and 3) in a cross‐fostering experiment, nestlings tended to grow larger in rural than in suburban nests irrespective of their hatching environment. These results agree with those found in a recent British study, indicating that poor nestling development and survival due to inadequate diet may be widespread phenomena in urbanized habitats.     

Personality and urbanization: behavioural traits and <Emphasis Type="Italic">DRD4</Emphasis> SNP830 polymorphisms in great tits in Barcelona city

Most examples of adaptation to the urban environment relate to plasticity processes rather than to natural selection. Personality, however, defined as consistent individual differences in behaviour related to exploration, caution, and neophobia, is a good behavioural candidate character to study natural selection in relation to the urban habitat due to its heritable variation. The aim of this paper was to analyse variation in personality by comparing urban and forest great tits Parus major using standard tests of exploratory behaviour and boldness. We studied personality in 130 wild great tits captured in Barcelona city and nearby forests and found that urban birds were more explorative and bolder towards a novel object than forest birds. Genotype frequencies of the DRD4 SNP830 polymorphism, a gene region often associated with personality variation, varied significantly between forest and urban birds. Behavioural scores, however, were not correlated with this polymorphism in our population. Exploration scores correlated to boldness for forest birds but not for urban birds. Our findings suggest that the novel selection pressures of the urban environment favour the decoupling of behavioural traits that commonly form behavioural syndromes in the wild.     

Urbanization and the temporal patterns of social networks and group foraging behaviors

Urbanization causes dramatic and rapid changes to natural environments, which can lead the animals inhabiting these habitats to adjust their behavioral responses. For social animals, urbanized environments may alter group social dynamics through modification of the external environment (e.g., resource distribution). This might lead to changes in how individuals associate or engage in group behaviors, which could alter the stability and characteristics of social groups. However, the potential impacts of urban habitat use, and of habitat characteristics in general, on the nature and stability of social associations remain poorly understood. Here, we quantify social networks and dynamics of group foraging behaviors of black‐capped chickadees (N = 82, Poecile atricapillus), at four urban and four rural sites weekly throughout the nonbreeding season using feeders with radio frequency identification of individual birds. Because anthropogenic food sources in urban habitats (e.g., bird feeders) provide abundant and reliable resources, we predicted that social foraging associations may be of less value in urban groups, and thus would be less consistent than in rural groups. Additionally, decreased variability of food resources in urban habitats could lead to more predictable foraging patterns (group size, foraging duration, and the distribution of foraging events) in contrast to rural habitats. Networks were found to be highly consistent through time in both urban and rural habitats. No significant difference was found in the temporal clumping of foraging events between habitats. However, as predicted, the repeatability of the clumping of foraging events in time was significantly higher in urban than rural habitats. Our results suggest that individuals living in urban areas have more consistent foraging behaviors throughout the nonbreeding season, whereas rural individuals adjust their tactics due to less predictable foraging conditions. This first examination of habitat‐related differences in the characteristics and consistency of social networks along an urbanization gradient suggests that anthropic habitat use results in subtle modifications in social foraging patterns. Future studies should examine potential implications of these differences for variation in predation risk, energy intake, and information flow.     

Hunt or hide: How insularity and urbanization affect foraging decisions in lizards

Foraging decisions should reflect a balance between costs and benefits of alternative strategies. Predation risk and resource availability in the environment may be crucial in deciding how cautious individuals should behave during foraging. These costs and benefits will vary in time and context, meaning that animals should be able to adjust their foraging behaviour to new or altered environments. Studying how animals do this is essential to understand their survival in these environments. In this study, we investigated the effect of both insularity and urbanization on risk‐taking and neophobia during foraging in the Dalmatian wall lizard (Podarcis melisellensis). Small islets tend to have both a lower number of predators and less resources. Therefore, islet populations were expected to show more risk‐taking behaviour and less neophobia in a foraging context. Previous studies on behaviour of urban lizards have yielded inconsistent results, but due to a lack of both predators and arthropod prey in urban habitats, we expected urban lizards to also take more risks and behave less neophobic. We sampled several inhabited and uninhabited locations on Vis (Croatia) and surrounding islets. Risk‐taking behaviour was tested by measuring the latency of lizards to feed in the presence of a predator model, and neophobia by measuring the latency to feed in the presence of a novel object. We found that islet lizards do indeed take more risks and were less vigilant, but not less neophobic. Urban and rural lizards did not differ in any of these behaviours, which is in sharp contrast with previous work on mammals and birds. The behavioural differences between islet and island lizards were novel, but not unexpected findings and are in line with the theory of “island tameness”. The effect of urbanization on the behaviour of animals seems to be more complex and might vary among taxa.     

城市化对城市麻雀栖息地利用的影响:以北京市为例

本文以北京市为例研究了城市化水平不同的8个区域中麻雀(Passermontanus)的数量分布现状以及影响麻雀分布的栖息地因子。结果表明,越冬期和繁殖期的麻雀数量均与城市化程度呈显著的负相关关系;城市化程度高的城市中心商业区、高层居民区和城市主干道中的麻雀数量均很少;城市化程度较低的城乡结合区、公园、城市的平房区及古建筑区域中麻雀数量均较多;高校校园和低层楼房居民区,虽然城市化程度相对较高,但由于植被较丰富,麻雀数量也较多。平房面积、针叶树数量、阔叶树数量、空调数量、高层楼房的面积、硬化地面的面积、人流量及车流量是影响麻雀栖息地利用的重要因素。其中平房面积、针叶树、阔叶树、空调的数量增加,可为麻雀提供栖息条件而有利于麻雀的利用;硬化地面的面积、高层楼房的面积、人流量及车流量的增加,由于减少了杂草等麻雀的食物来源并增加了干扰,不利于麻雀的利用。结果表明,虽然麻雀是一个适应人类生活环境的物种,但在快速的城市化变迁中,它已表现出对高度城市化环境的不适应。在城市的规划与建设中,应考虑到为以麻雀为代表的城市鸟类提供生存必要的植被和繁殖场所,构建人鸟和谐共存的生态城市。     

Does urbanization affect selective pressures and life‐history strategies in the common blackbird (Turdus merula L.)?

Urbanization, one of the most extreme land‐use alterations, is currently spreading, and the number of species confronting these changes is increasing. However, contradictory results of previous studies impede a clear interpretation of which selective pressure (nest predation or food limitation) is more important in urban habitats compared with natural situations, and whether birds can confront them by adjusting their life‐history strategies. We investigated life‐history syndromes of three common blackbird (Turdus merula) populations differing in their human influence (urban, rural, and woodland). We analysed daily nest predation and nestling starvation rates to assess the relative importance of these selection pressures in each habitat. Simultaneously, several life‐history traits were investigated to determine if T. merula seem adapted to their main source of selection. Food limitation was more important in the city, whereas nest predation was the most important selective force in the forest. The rural habitat was characterized by an intermediate influence of these two factors. Life‐history syndromes, as the covariation of a suite of traits, confirmed these results because T. merula seem well adapted to the main cause of selection in each habitat. Our results are consistent with urbanization imposing new challenges on birds, and that they adaptively respond to them. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 759–766.     

High urban population density of birds reflects their timing of urbanization

Living organisms generally occur at the highest population density in the most suitable habitat. Therefore, invasion of and adaptation to novel habitats imply a gradual increase in population density, from that at or below what was found in the ancestral habitat to a density that may reach higher levels in the novel habitat following adaptation to that habitat. We tested this prediction of invasion biology by analyzing data on population density of breeding birds in their ancestral rural habitats and in matched nearby urban habitats that have been colonized recently across a continental latitudinal gradient. We estimated population density in the two types of habitats using extensive point census bird counts, and we obtained information on the year of urbanization when population density in urban habitats reached levels higher than that of the ancestral rural habitat from published records and estimates by experienced ornithologists. Both the difference in population density between urban and rural habitats and the year of urbanization were significantly repeatable when analyzing multiple populations of the same species across Europe. Population density was on average 30 % higher in urban than in rural habitats, although density reached as much as 100-fold higher in urban habitats in some species. Invasive urban bird species that colonized urban environments over a long period achieved the largest increases in population density compared to their ancestral rural habitats. This was independent of whether species were anciently or recently urbanized, providing a unique cross-validation of timing of urban invasions. These results suggest that successful invasion of urban habitats was associated with gradual adaptation to these habitats as shown by a significant increase in population density in urban habitats over time.     

Behavioral plasticity and consistency in the naked-footed mouse (<Emphasis Type="Italic">Peromyscus nudipes</Emphasis>) with habitat disturbance

Very few studies have evaluated whether habitat disturbance affects behavioral consistency and plasticity. We measured shyness–boldness and exploration–avoidance for the first time in naked-footed mice (Peromyscus nudipes) in two adjacent habitats with differing level of disturbance in Costa Rica. Each habitat was measured in different, consecutive years. With these data, we explored the possibility of habitat differences in individual- and population-level behavioral plasticity, magnitude of behavior, and consistency. Mice in both habitats behaved consistently across time with high repeatability (i.e., showed personality). The strength of consistency of shyness–boldness and exploration–avoidance over time was higher in the disturbed habitat, and individuals in the undisturbed habitat were bolder and less exploratory. Behaviors were correlated with each other in both habitats, indicating that behavioral syndromes do not always break down in disturbed areas. Mice changed the magnitude of their response the second time they were tested, indicating population-level plasticity. There was also greater individual plasticity among mice in the undisturbed habitat. Our study suggests the possibility that habitat disturbance can affect behavioral plasticity and personality and that shyer/more exploratory individuals might colonize and/or persist in disturbed habitats. These results are preliminary and exploratory because we did not control for temporal differences between habitats.     

Linking fearfulness and coping styles in fish

Consistent individual differences in cognitive appraisal and emotional reactivity, including fearfulness, are important personality traits in humans, non-human mammals, and birds. Comparative studies on teleost fishes support the existence of coping styles and behavioral syndromes also in poikilothermic animals. The functionalist approach to emotions hold that emotions have evolved to ensure appropriate behavioral responses to dangerous or rewarding stimuli. Little information is however available on how evolutionary widespread these putative links between personality and the expression of emotional or affective states such as fear are. Here we disclose that individual variation in coping style predicts fear responses in Nile tilapia Oreochromis niloticus, using the principle of avoidance learning. Fish previously screened for coping style were given the possibility to escape a signalled aversive stimulus. Fearful individuals showed a range of typically reactive traits such as slow recovery of feed intake in a novel environment, neophobia, and high post-stress cortisol levels. Hence, emotional reactivity and appraisal would appear to be an essential component of animal personality in species distributed throughout the vertebrate subphylum.     

Linking optimal foraging behavior to bird community structure in an urban-desert landscape: field experiments with artificial food patches

Urban bird communities exhibit high population densities and low species diversity, yet mechanisms behind these patterns remain largely untested. We present results from experimental studies of behavioral mechanisms underlying these patterns and provide a test of foraging theory applied to urban bird communities. We measured foraging decisions at artificial food patches to assess how urban habitats differ from wildlands in predation risk, missed-opportunity cost, competition, and metabolic cost. By manipulating seed trays, we compared leftover seed (giving-up density) in urban and desert habitats in Arizona. Deserts exhibited higher predation risk than urban habitats. Only desert birds quit patches earlier when increasing the missed-opportunity cost. House finches and house sparrows coexist by trading off travel cost against foraging efficiency. In exclusion experiments, urban doves were more efficient foragers than passerines. Providing water decreased digestive costs only in the desert. At the population level, reduced predation and higher resource abundance drive the increased densities in cities. At the community level, the decline in diversity may involve exclusion of native species by highly efficient urban specialists. Competitive interactions play significant roles in structuring urban bird communities. Our results indicate the importance and potential of mechanistic approaches for future urban bird community studies.