Urban Colonization Research Articles

Genetics of urban colonization: neutral and adaptive variation in coyotes (Canis latrans) inhabiting the New York metropolitan area

Genetics of urban colonization: neutral and adaptive variation in coyotes (<i>Canis latrans</i>) inhabiting the New York metropolitan area

Offspring defense by an urban raptor responds to human subsidies and ritual animal-feeding practices.

There is a growing interest in the behavioural and life history mechanisms that allow animal species to cope with rapidly expanding urban habitats, which impose frequent proximity to humans. A particular case of behavioral bottleneck (i.e. conflicting interests) faced by animals in urban environments is how they will modulate the defence of their offspring against the potential danger represented by humans, an aspect that has received scarce research attention. We examined the nest defense against humans by a dense breeding population of a raptor, the Black Kite Milvus migrans, within the megacity of Delhi (India). Here, kites live on a diet dominated by human waste and meat offered through religiously motivated bird feeding practices. Nest defense levels increased with the number of offspring, and with the progression of the breeding season. Defense also intensified close to ritual-feeding areas and with increasing human waste in the streets, suggesting synergistic effects of food availability, parental investment, personality-boldness and habituation to humans, with consequent attenuation of fear. Thus, the behavioural response to a perceived threat reflected the spatial mosaic of activity of humans in the city streets, their cultural practices of ritual-feeding, and their waste-management. For synurbic species, at the higher-end spectrum of adaptation to an urban life, human cultural practices and attitudes may well be the most defining dimensions of their urban niche. Our results suggest that, after initial urban colonization, animals may continue to adapt to the typically complex, heterogeneous environments of cities through fine-grained behavioural adjustments to human practices and activities.

Development of urban behaviour is associated with time since urbanization in a reed-nesting waterbird

Behavioural adaptations are recognized as crucial determinants for successful establishment and persistence of animal populations in a novel urban environment. Here, we examined mechanisms responsible for the development of urban behavioural type in a common waterbird, the Eurasian coot Fulica atra. We compared the behaviour of coots from a rural population and two urban populations that differed in the timing of colonization event (1960s vs. 2000s). We found that some behavioural characters associated with urban life (aggression during nest defence and boldness towards humans during foraging) were more strongly expressed in the older urban population when compared with the recently established urban population. By contrast, coots from the two urban populations showed a similar likelihood of exploiting human-derived food resources, as well as they showed similar levels of physiological stress. Urban coots were generally more aggressive, bolder, and less stressed than their rural conspecifics. Large behavioural and physiological divergence of coots from the recently established urban population and their rural conspecifics suggested that phenotypic plasticity and phenotype sorting may play a key role in the initial stages of urban colonization. On the other hand, increasing expression of boldness and aggression with the time since urbanization may suggest the role of microevolutionary adaptation in response to novel selective forces associated with the urban environment. Our results indicate that a combination of different processes (phenotypic plasticity, phenotype sorting, and microevolution) can determine successful colonization of urban areas by the Eurasian coots, and possibly other bird species.

Shyer and larger bird species show more reduced fear of humans when living in urban environments.

As the natural habitats of many species are degraded or disappear, there is scope for these species to be established in urban habitats. To ease the establishment and maintenance of urban populations of more species we need to better understand what degree of phenotypical change to expect as different species transition into urban environments. During the first stages of urban colonization, behavioural changes such as an increase in boldness are particularly important. A consistent response in urban populations is to decrease the distance at which individuals flee from an approaching human (flight initiation distance, or FID). Performing a phylogenetic generalized least-squares (PGLS) analysis on 130 avian species, I found that the largest changes in FID between rural and urban populations occur in species that are larger-bodied and naturally shy (higher rural FID), two phenotypic traits that are not normally associated with urban colonizers. More unlikely species may thus be able to colonize urban environments, especially if we design cities in ways that promote such urban colonizations.

Landscape Factors Associated with Subterranean Termite (Isoptera: Rhinotermitidae) Treatments and Colony Structure in Residential Subdivisions

Subterranean termites (Isoptera: Reticulitermes) are common structural pests, but it is not well known how landscape factors are associated with urban colonization. This study examined patterns of subterranean termite colonization in 13 mid-Missouri residential subdivisions. Ten- and 20-year-old homes built on historically agricultural and forested landscapes were inspected for treatment by termiticide application or bait stations. Contemporary and historical aerial imagery were analyzed using GIS software, and patterns of colonization were compared among subdivisions. The genetic structure of termite colonies collected in undeveloped landscapes and residential subdivisions was compared using microsatellite DNA. Twenty-year-old subdivisions had significantly higher treatment proportions than 10-year-old subdivisions. At year 10, historically forested subdivisions had a higher treatment proportion than historically agricultural subdivisions. By year 20, there was no significant difference in treatment proportion between historical landscape types, indicating that subdivisions built on agricultural landscapes eventually catch up to subdivisions built on forest landscapes. Although there was not strong statistical support, treated homes in historically agricultural subdivisions tended to be close to forest patches, but there was less of an association in historically forested subdivisions. Colonies in undeveloped landscapes were more inbred compared to colonies in residential subdivisions, indicating that colonies sampled in subdivisions had fewer secondary reproductive and were potentially younger than those sampled in undeveloped landscapes. This study provides some correlative support for the role of dispersing alates as urban colonizers, because treatments were often located at relatively long distances from undisturbed forest patches in historically agricultural subdivisions.

How birds colonize cities: genetic evidence from a common waterbird, the Eurasian coot

Much effort has been devoted to identify ecological and life‐history traits which facilitate urban colonization by wild avian species, but surprisingly little is known about the population‐level mechanisms of urbanization processes. In general, two different patterns of urban colonization have been proposed: 1) the model of independent colonization predicts that birds colonize cities independently in different geographical regions; 2) the model of leapfrog colonization assumes a single colonization event, while additional urban populations are established from the initial urban populations. The aim of this paper was to determine the pattern of urban colonization in a common waterbird, the Eurasian coot Fulica atra. For this purpose, we analysed microsatellite variation in three pairs of urban and rural coot populations from central Poland. We found that a newly‐established urban population was genetically more similar to neighbouring rural populations than to long‐established urban populations, as indicated by the analysis of fixation index, genetic distance and Bayesian assignment of individuals to genetic clusters. These results are consistent with the model of independent colonization, where neighbouring rural populations are a source of individuals that colonize new urban areas. However, our analysis also showed significant differentiation between long‐established urban populations and adjacent rural populations, suggesting that genetic connectivity between two types of habitat decreases with increasing time since urbanization. Our study shows high complexity of urbanization processes in wild animal populations, as well as it underpins utility of molecular tools in studying population‐level mechanisms of urbanization.

Niche segregation, competition, and urbanization.

Urbanization of species is an ongoing process where successful urban colonizers usually obtain large fitness benefits. Mechanisms proposed to explain associations between urbanization and life-history traits are based on behavioral flexibility in food and habitat use and reduced fear responses. We test the novel hypothesis that interspecific competition for proximity to humans is driving urbanization. We recorded the distance during the breeding season to human habitation for 50 pairs of closely related bird species, where one was closely associated with humans while the other species was not. The degree of urbanization was larger as was range size and abundance in the species more closely associated to humans. Flight initiation distance was shorter, and species closely associated with humans were more abundant in ancestral rural habitats. Likewise, species more closely associated with humans reproduced earlier and during longer periods. These results are consistent with the hypothesis that urbanization is promoted by interspecific competition. Resulting isolation by urban habitat may further facilitate contemporary adaptation to urban environments.

Functional trait changes in the floras of 11 cities across the globe in response to urbanization

Urbanization causes major environmental changes globally, which can potentially homogenize biota across cities through the loss and gain of particular types of species. We examine whether urban environments consistently select for plants with particular traits and the implications of such changes on the functional composition of urban floras. We classified plant recorded in 11 cities around the globe as species that have either colonized (arrived and naturalized), persisted or been lost (local extirpation) following urbanization. We analyzed how 10 traits previously linked with plant responses to environmental conditions explained membership of these three groups, by comparing colonisers with persistent and extirpated plants through individual city‐level Bayesian models. Then, we used meta‐analysis to assess consistency of traits across urban areas. Finally, we explored several possible scenarios of functional change using these results.On average, urban colonizers had heavier seeds, unspecialised nutrient requirements, were taller and were annual species more often, especially when compared to locally extirpated plants. Common trends of functional change in urban plant communities include shifts towards taller and heavier‐seeded plants, and an increased prevalence of the short‐lived species, and plants without mutualistic nutritional strategies. Our results suggest that plant traits influence the species that succeed in urban environments worldwide. Different species use different ecological strategies to live in urban environments, as suggested by the importance of several traits that may appear as trait constellations. Plant height and seed mass were the only traits associated with both colonizer and extirpated plant status in urban environments. Based on our data, predicting colonization in urban environments may be easier than identifying extirpation‐prone plants; albeit some regional variation, colonization seems strongly driven by environmental conditions common to most cities (e.g. altered disturbance regimes), whereas extirpation may depend more on processes that vary across cities.

Innovativeness and the effects of urbanization on risk-taking behaviors in wild Barbados birds.

The effects of urbanization on avian cognition remain poorly understood. Risk-taking behaviors like boldness, neophobia and flight distance are thought to affect opportunism and innovativeness, and should also vary with urbanization. Here, we investigate variation in risk-taking behaviors in the field in an avian assemblage of nine species that forage together in Barbados and for which innovation rate is known from previous work. We predicted that birds from highly urbanized areas would show more risk-taking behavior than conspecifics from less urbanized parts of the island and that the differences would be strongest in the most innovative of the species. Overall, we found that urban birds are bolder, less neophobic and have shorter flight distances than their less urbanized conspecifics. Additionally, we detected between-species differences in the effect of urbanization on flight distance, more innovative species showing smaller differences in flight distance between areas. Our results suggest that, within successful urban colonizers, species differences in innovativeness may affect the way species change their risk-taking behaviors in response to the urban environment.

Reproduction and survival in the city: which fitness components drive urban colonization in a reed-nesting waterbird?

Processes of adaptation to urban environments are well described for relatively few avian taxa, mainly passerines, but selective forces responsible for urban colonization in ecologically different groups of birds remain mostly unrecognized. The aim of this article is to identify drivers of recent urban colonization (Łódź, central Poland) by a reed-nesting waterbird, the Eurasian coot Fulica atra. Urban colonizers were found to adopt a distinct reproductive strategy by maximizing the number of offspring (carryover effects of higher clutch size), whereas suburban individuals invested more in the quality of the progeny (higher egg volume), which could reflect differences in predatory pressure between 2 habitats. In fact, reduced predation rate was strongly suggested by elevated hatching success in highly urbanized areas, where probability of hatching at least 1 chick was higher by 30% than in suburban natural-like habitats. Coots nesting in highly urbanized landscape had considerably higher annual reproductive success in comparison to suburban pairs, and the difference was 4-fold between the most and least urbanized areas. There was also a constant increase in size-adjusted body mass and hemoglobin concentration of breeding coots from the suburbs to the city centre. Urban colonization yielded no survival benefits for adult birds and urban individuals showed higher site fidelity than suburban conspecifics. The results suggest that the recent urban colonization by Eurasian coots was primary driven by considerable reproductive benefits which may be primarily attributed to: (1) reduced predation resulting from an exclusion of most native predators from highly urbanized zones; (2) increased condition of urban-dwelling birds resulting from enhanced food availability.

Behavioural and Ecological Keys to Urban Colonization by Little Ravens ()

Avian urban colonization is thought to be facilitated by a capacity for innovative feeding, ecological generalism and social foraging. However, the relative importance in exploiting urban resources and avoiding urban predators of being inherently ‘pre-adapted’ to the urban environment or adjusting to it through phenotypic plasticity requires more examination. These issues were explored in a native ‘urban adapter’, the Little raven Corvus mellori, by comparing its foraging ecology, group size and nest site use in Melbourne, Australia, and the surrounding exurban environment. Urban individuals manipulated human food waste and gleaned from sealed surfaces more than exurban conspecifics (suggesting behavioural flexibility), but foraging behaviour and substrate use were broadly similar in both environments (suggesting ‘preadaptation’). Little ravens foraged close to conspecifics and heterospecifics more frequently in the urban than the exurban environment, but some potential dietary competitors rarely foraged near urban Little ravens, possibly indicating some niche partitioning. Mean urban rate of agonistic interaction with other bird species was low (0.023 interactions per foraging raven observed). Although displacement of a raven &gt;10 m occurred in 61-70% of such interactions, the displaced individual usually rapidly resumed foraging nearby. Thus aggressive, interspecific interference competition for food appeared limited. Large groups of Little ravens were twice as common in the exurban as the urban environment, which was inconsistent with the hypothesis that social foraging facilitated urban colonization. Nest tree type (predominantly eucalypts), size and isolation were similar in urban and exurban environments, but urban nests were significantly more concealed. We suggest that ‘preadaptation’, behavioural innovation and a relative lack of significant, interspecific food competition have contributed to urban colonization by Little ravens.

Development and growth in synanthropic species: plasticity and constraints.

Urbanization poses serious extinction risks, yet some species thrive in urban environments. This may be due to a pronounced developmental plasticity in these taxa, since phenotypically, plastic organisms may better adjust to unpredictable urban food resources. We studied phenotypic plasticity in Nuctenea umbratica, a common European forest and urban vegetation spider. We subjected spiderlings to low (LF), medium (MF) and high (HF) food treatments and documented their growth and developmental trajectories into adulthood. Spiders from the three treatments had comparable numbers of instars and growth ratios, but differed in developmental periods. Longest developing LF spiders (♀ = 390, ♂ = 320days) had the smallest adults, but MF (♀ = 300, ♂ = 240days) and HF (♀ = 240, ♂ = 210days) spiders reached comparable adult sizes through shorter development. While males and females had comparable instar numbers, females had longer development, higher growth ratios, adult sizes and mass; and while males adjusted their moulting to food availability, female moulting depended on specific mass, not food treatment. We discussed the patterns of Nuctenea sex-specific development and compared our results with published data on two other Holarctic urban colonizers (Larinioides sclopetarius, Zygiella x-notata) exhibiting high plasticity and fast generation turn-over. We conclude that despite relatively unconstrained developmental time in the laboratory enabling Nuctenea to achieve maximal mass and size-main female fitness proxies-their relatively fixed growth ratio and long generation turn-over may explain their lower success in urban environments.

Anthropogenic environments exert variable selection on cranial capacity in mammals

It is thought that behaviourally flexible species will be able to cope with novel and rapidly changing environments associated with human activity. However, it is unclear whether such environments are selecting for increases in behavioural plasticity, and whether some species show more pronounced evolutionary changes in plasticity. To test whether anthropogenic environments are selecting for increased behavioural plasticity within species, we measured variation in relative cranial capacity over time and space in 10 species of mammals. We predicted that urban populations would show greater cranial capacity than rural populations and that cranial capacity would increase over time in urban populations. Based on relevant theory, we also predicted that species capable of rapid population growth would show more pronounced evolutionary responses. We found that urban populations of two small mammal species had significantly greater cranial capacity than rural populations. In addition, species with higher fecundity showed more pronounced differentiation between urban and rural populations. Contrary to expectations, we found no increases in cranial capacity over time in urban populations-indeed, two species tended to have a decrease in cranial capacity over time in urban populations. Furthermore, rural populations of all insectivorous species measured showed significant increases in relative cranial capacity over time. Our results provide partial support for the hypothesis that urban environments select for increased behavioural plasticity, although this selection may be most pronounced early during the urban colonization process. Furthermore, these data also suggest that behavioural plasticity may be simultaneously favoured in rural environments, which are also changing because of human activity.

Candidate gene polymorphisms for behavioural adaptations during urbanization in blackbirds

Successful urban colonization by formerly rural species represents an ideal situation in which to study adaptation to novel environments. We address this issue using candidate genes for behavioural traits that are expected to play a role in such colonization events. We identified and genotyped 16 polymorphisms in candidate genes for circadian rhythms, harm avoidance and migratory and exploratory behaviour in 12 paired urban and rural populations of the blackbird Turdus merula across the Western Palaearctic. An exonic microsatellite in the SERT gene, a candidate gene for harm avoidance behaviour, exhibited a highly significant association with habitat type in an analysis conducted across all populations. Genetic divergence at this locus was consistent in 10 of the 12 population pairs; this contrasts with previously reported stochastic genetic divergence between these populations at random markers. Our results indicate that behavioural traits related to harm avoidance and associated with the SERT polymorphism experience selection pressures during most blackbird urbanization events. These events thus appear to be influenced by homogeneous adaptive processes in addition to previously reported demographic founder events.

The Hong Kong Fever of 1843: Collective Trauma and the Reconfiguring of Colonial Space

AbstractThis paper adds to a relatively new line of research investigating inadvertent transformations of urban colonial space generated by collective trauma. To the now classic iterative, and racial dynamics of Neild's ‘accommodation’ in the development of Madras,1 and of Yeoh's ‘contesting’ of the built environment in Singapore2—specifically in the tugging and pulling between local and colonial influences within the spatial discourse of colonial port cities—needs to be added that of single or multiple-event collective trauma. Such trauma, perceived as brought upon by unexpected external causes, might consolidate, perhaps accelerate, or even sever a previous sequence of spatial negotiation, particularly if that sequence was politically vulnerable or immature. The paper is a focused account of such an occurrence: the small-scale yet intensely traumatic events of Hong Kong Island while still in its colonial infancy in 1843, the year of the ‘Hongkong Fever’. It argues that a new conception of malaria—considered then a miasma—now linked both to location and construction, led to the first reactive, yet decisive, reconfiguring of a previously improvised urban colonization process, consequently salvaging Hong Kong's position within a wider imperial context.

Dispatches

Dispatches

Tolerance of Auditory Disturbance by an Avian Urban Adapter, the Noisy Miner

AbstractUrbanization creates challenges for wildlife, most notably through changes in resource availability and the frequent occurrence of sensory disturbance. Some native species, however, have been able to exploit and thrive in urban environments. Research, in this regard, has mostly focused on the ecological conditions that have allowed such species to prosper. In contrast, less attention has been devoted to evaluating how they cope with human proximity and disturbance. In a field experiment on a successful Australian ‘urban adapter’, the Noisy miner, Manorina melanocephala, we compared tolerance of a loud, startling sound stimulus by urban and rural individuals. We found group size differences between birds occupying urban and rural sites: more urban birds came into the testing area in response to the initial alarm‐call playback compared with rural birds. Urban and rural birds also differed significantly in their behavioural response profile to the test sound stimulus. Nearly half (47.5%) of the rural, but only 22.5% of the urban birds took flight and of those that did, only 1 of 9 urban individuals retreated &gt;5 m, compared with 13 of 19 rural birds. About one‐third of urban, but only 5% of rural individuals responded to the sound stimulus with aggressive displays. The most frequent response to the stimulus, irrespective of habitat type, was to remain near the sound source and engage in visual surveillance. The high frequency of this response in both urban and rural individuals suggested that most noisy miners were quite ‘bold’, a temperament trait that is likely to be important in successful urban colonization by birds.

Regional Decline of an Iconic Amphibian Associated with Elevation, Land-Use Change, and Invasive Species

Ecological theory predicts that species with restricted geographic ranges will have the highest probability of extinction, but species with extensive distributions and high population densities can also exhibit widespread population losses. In the western United States populations of northern leopard frogs (Lithobates pipiens)-historically one of the most widespread frogs in North America-have declined dramatically in abundance and geographic distribution. To assess the status of leopard frogs in Colorado and evaluate causes of decline, we coupled statewide surveys of 196 historically occupied sites with intensive sampling of 274 wetlands stratified by land use. We used an information-theoretic approach to evaluate the contributions of factors at multiple spatial extents in explaining the contemporary distribution of leopard frogs. Our results indicate leopard frogs have declined in Colorado, but this decline was regionally variable. The lowest proportion of occupied wetlands occurred in eastern Colorado (2-28%), coincident with urban development and colonization by non-native bullfrogs (Lithobates catesbeianus). Variables at several spatial extents explained observed leopard frog distributional patterns. In low-elevation wetlands introduced fishes, bullfrogs, and urbanization or suburbanization associated negatively with leopard frog occurrence, whereas wetland area was positively associated with occurrence. Leopard frogs were more abundant and widespread west of the Continental Divide, where urban development and bullfrog abundance were low. Although the pathogenic chytrid Batrachochytrium dendrobatidis (Bd) was not selected in our best-supported models, the nearly complete extirpation of leopard frogs from montane wetlands could reflect the individual or interactive effects of Bd and climate patterns. Our results highlight the importance of considering multiple, competing hypotheses to explain species declines, particularly when implicated factors operate at different spatial extents.

Independent colonization of multiple urban centres by a formerly forest specialist bird species

Urban areas are expanding rapidly, but a few native species have successfully colonized them. The processes underlying such colonization events are poorly understood. Using the blackbird Turdus merula, a former forest specialist that is now one of the most common urban birds in its range, we provide the first assessment of two contrasting urban colonization models. First, that urbanization occurred independently. Second, that following initial urbanization, urban-adapted individuals colonized other urban areas in a leapfrog manner. Previous analyses of spatial patterns in the timing of blackbird urbanization, and experimental introductions of urban and rural blackbirds to uncolonized cities, suggest that the leapfrog model is likely to apply. We found that, across the western Palaearctic, urban blackbird populations contain less genetic diversity than rural ones, urban populations are more strongly differentiated from each other than from rural populations and assignment tests support a rural source population for most urban individuals. In combination, these results provide much stronger support for the independent urbanization model than the leapfrog one. If the former model predominates, colonization of multiple urban centres will be particularly difficult when urbanization requires genetic adaptations, having implications for urban species diversity.

Évolution des paysages et émergence de risques environnementaux : l'exemple des dortoirs urbains d'étoumeaux en Bretagne

Change in landscape structure involve environmental problems : case of urban roosts of starlings in Brittany. — The distribution of starling roosts have recently changed in several regions of West Europe. A research in Brittany shows a trend to increase of roost sites in the cities. An analysis conducted around Rennes, reveals potential roost sites (indeciduous hedges, waste lands) which play a role in urban colonization process by starling and may involve medical problems.