The re-emergence of indigenous forest in an urban environment, Christchurch, New Zealand

Christchurch, the second largest city in New Zealand is a planned city on a coastal plain on the east coast of the South Island. The birth of the city and the subsequent century of development was characterised by colonial values and tree and garden planting with familiar European species along with those from Australia, North America, and eventually all other continents. The image of an “English garden city” with classical parks of oaks and willow-lined rivers became the accepted norm and the way in which the city has been promoted to potential tourists. Gardening is one of the top two recreational activities and exotic species greatly outnumber native species in the flora and in gardens. This has had serious consequences for the highly fragmented and degraded indigenous vegetation and its co-adapted wildlife. A few hardy indigenous species continued to regenerate through this period, but since the 1970s, there has been a progressive change of attitude and interest in reclaiming the natural heritage of the city, manifest in widespread private and public planting of indigenous species and active habitat restoration. In this article we examine the indigenous and exotic shrub and tree components of the Christchurch flora as planted street trees, in domestic gardens, and in parks. We also present data on shrub and tree regeneration in parks and domestic gardens in the city. Indications are that the more sensitive, less intrusive management of urban environments, combined with the greater density of indigenous seed sources, has allowed regeneration of a wide range of indigenous species across a broad spectrum of habitats – from neglected gardens to pavement cracks to exotic plantations. This is despite the competition from the prodigious seed banks and density of exotic trees, shrubs, and ground covers and albeit minimal impacts of introduced browsing and seed eating mammals. If the present trends continue through appropriate management and facilitation, these tentative signs of native forest regeneration should eventually proliferate into a sustainable mixed origin urban forest that resurrects and preserves the natural character of the region.     

Distribution of invasive plants in urban environment is strongly spatially structured

Context

Urban environments create a wide range of habitats that harbour a great diversity of plant species, many of which are of alien origin. For future urban planning and management of the green areas within the city, understanding of the spatial distribution of invasive alien species is of great importance.

Objectives

Our main aim was to assess how availability of different ecosystem types within a city area, as well as several parameters describing urban structure interact in determining the cover and identity of invasive alien species.

Methods

We studied the distribution of chosen invasive plant species in a mid-sized city in the Czech Republic, central Europe, on a gradient of equal sized cells from the city centre to its outskirts.

Results

A great amount of variation was explained by spatial predictors but not shared with any measured variables. The species cover of invasive species decreased with increasing proportion of urban greenery and distance from the city centre, but increased with habitat richness; road margins, ruderal sites, and railway sites were richest in invasive species. In contrast, the total number of invasive species in cells significantly decreased with increasing distance from the city centre, but increased with habitat richness.

Conclusions

Our results suggest that different invasive species prefer habitats in the vicinity of the city centre and at its periphery and the spatial structure and habitat quality of the urban landscape needs to be taken into account, in efforts to manage alien plant species invasions in urban environments.

     

Response of forest soil properties to urbanization gradients in three metropolitan areas

We investigated the effects of urban environments on the chemical properties of forest soils in the metropolitan areas of Baltimore, New York, and Budapest. We hypothesized that soils in forest patches in each city will exhibit changes in chemistry corresponding to urbanization gradients, but more strongly with various urban metrics than distance to the urban core. Moreover, differences in parent material and development patterns would differentially affect the soil chemical response in each metropolitan area. Results showed that soil chemical properties varied with measures of urban land use in all three cities, including distance to the urban core, which was an unexpected result. Moreover, the results showed that the spatial extent and amount of change was greater in New York than in Baltimore and Budapest for those elements that showed a relationship to the urbanization gradient (Pb, Cu, and to a lesser extent Ca). The spatial relationship of the soil chemical properties to distance varied from city to city. In New York, concentrations of Pb, Cu, and Ca decreased to approximately background concentrations at 75 km from the urban core. By contrast, concentrations of these elements decreased closer to the urban core in Baltimore and Budapest. Moreover, a threshold was reached at about 75% urban land use above which concentrations of Pb and Cu increased by more than twofold relative to concentrations below this threshold. Results of this study suggest that forest soils are responding to urbanization gradients in all three cities, though characteristics of each city (spatial pattern of development, parent material, and pollution sources) influenced the soil chemical response.     

The importance of functional connectivity in the conservation of a ground-dwelling mammal in an urban Australian landscape

The distribution of the northern brown bandicoot (Isoodon macrourus), a medium-sized ground-dwelling marsupial, was examined in habitat fragments within the urban landscape of the city of Brisbane, Australia. From surveys conducted in 68 fragments, bandicoots were found to be present in 33 (49%) despite widespread habitat loss and fragmentation. Logistic regression analysis revealed that of 13 measured independent variables, functional connectivity was the only factor that significantly predicted the presence of bandicoots within fragments, with connectivity positively correlated with the likelihood of occupation. Functional connectivity was equated to the likelihood of bandicoot immigration into the focal fragment from the nearest occupied fragment, based on the estimated resistance to movement offered by the intervening matrix. Within Brisbane, riparian habitat fragments typically have a relatively high level of functional connectivity, as thin strips of vegetation fringing waterways serve as corridors between larger riparian areas and facilitate the movement of bandicoots between patches. Analyses based on the Akaike Information Criterion revealed that the optimal model based on landscape context variables was convincingly better supported by the data than the optimal model produced from fragment characteristics. However, it is important to examine both internal attributes of habitat fragments and external features of the surrounding landscape when modelling the distribution of ground-dwelling fauna in urban environments, or other landscapes with a highly variable matrix. As urban centres throughout the world expand, it is crucial that the ecology of local wildlife be considered to ensure functional connection is maintained between habitat patches, especially for the conservation of species that are highly susceptible to fragmentation.     

Small urban woodlands as biodiversity conservation hot-spot: a multi-taxon approach

To evaluate the importance of urban woodlands to serve as potential sites for biodiversity conservation, we analysed bird, carabid beetle and small mammal community responses to urbanisation at different spatial scales. We analysed the relationships between the variations of the structure (species richness S, diversity H′ and dominance D) of animal communities of woodlands distributed along a rural–urban gradient, and the variations along this same gradient of (1) the vegetation within woodlands, (2) the landscape at 100 m and (3) 600 m around the woodlands. We identified the spatial scales whose variations along the gradient most affected each animal community structure, and characterised community responses to these variations. Our results showed that urbanisation affected taxa differently according to their dispersal ability. Carabid beetles, less mobile, seem to be sensitive to increasing fragmentation and built surfaces from periurban to town centre which could make their movement within the urban landscape difficult. Birds, mobile species, seem to be more sensitive to variations of the vegetation structure within woodlands from periurban to town centre that could affect their capacity to maintain in habitat patches. Although our study did not allow relating the small mammal community structure to urbanisation, it suggests that this taxa is sensitive to urban local disturbances. A relevant management scale of woodlands can be specified for each taxa conservation. Urban woodlands accommodate over 50% of the species present in periurban woodlands, and effective management could enhance this number. Woodlands seem to be a good choice for promoting biodiversity conservation in towns.     

西安城市区野生草本花卉现状调研

西安地区的野生植物十分丰富,其中分布在城市区的草本植物种类繁多,对城市的环境及生物多样性保护具有重要的价值,同时也是城市景观的组成部分。一些科属植物具有较高的观赏价值和生态改善的作用。以西安城市区的野生草本花卉为研究对象,调查其植物种类及分布特点、形态特征,并分析了它们在城市园林景观中的应用价值。     

Interrelationship of birds and habitat features in urban greenspaces in Delhi, India

Biodiversity conservation in urban areas has become significant not only because of increasing human population in urban centers but also because it is one of the innovative ways to conserve biodiversity as suggested by various global environmental conventions. The present study was conducted with the purpose of assessing diversity and density of bird and woody species in some greenspaces of Delhi, the rapidly urbanizing capital of India. The landscape of Delhi consists of a broad spectrum of environments ranging from the city forests to highly modified artificial landscapes in certain parks. We assessed bird and woody vegetation in 20 m×50 m belt transects in each of the 19 randomly selected greenspaces of varying size. Results exhibit a negative relationship between the density of exotic woody species and bird diversity. Specifically, Prosopis juliflora – an exotic, which was the most abundant tree in our samples – exhibited a negative relationship with bird diversity. Principal component analysis (PCA) reveals that forest-preferring bird species increase with increasing greenspaces size, shrub diversity and shrub density. We conclude that maintaining larger greenspaces with high structural diversity may be effective in maintaining plant and bird diversity in the study area.     

Short perceptual range and yet successful invasion of a fragmented landscape: the case of the red-bellied tree squirrel (<Emphasis Type="Italic">Callosciurus erythraeus</Emphasis>) in Argentina

Dispersal is a key element of the invasion process for introduced species, and is influenced by landscape connectivity. The red-bellied squirrel (Callosciurus erythraeus) was introduced to Argentina in 1970. Suitable forest habitat for this arboreal species is highly fragmented in a rural–urban matrix, but despite this, the squirrel population has spread. Squirrels disperse into new habitat patches using connective features such as forest corridors. They may also cross gaps but up to what extent is not known. Gap crossing success is influenced by perceptual range, which is the distance from which animals can perceive suitable habitat. Perceptual range has been previously estimated for vulnerable species, but not for introduced species. We used a model relating perceptual range to body mass to predict the perceptual range of the red-bellied tree squirrel in Argentina. We then tested our prediction of 202–221 m by releasing squirrels in an unfamiliar arable field at different distances (300, 200, 100 and 20 m) from woodland habitat. We assumed that if woodland could be perceived, squirrels would orientate toward it. We estimated perceptual range to be between 20 and 100 m, considerably lower than predicted. Our results indicate that squirrels can potentially cross small habitat gaps, but dispersal over greater distances lacking connectivity is less likely. Incorporating this information when modelling the spread of exotic squirrels in the Pampas Region can yield more accurate prediction of the invasion process and guide management practices to minimise their expansion.     

Urban-rural and temporal differences of woody plants and bird species in Harbin city,northeastern China

Urbanization has been greatly accelerated by the economic growth in China, while its possible effects on woody plants, bird species and their associations are not well defined yet. Here, we analyzed urban-rural gradients (landscape level: urban-farmland-forest-natural reserves; city level: ring road and urban build-up history) and temporal data (1955–1980–2014 for woody plants; 1980s–2010 s for birds) in Harbin city, China, to investigate the changes in the composition and diversity of woody plants and birds during urbanization. Both landscape gradient and temporal data confirmed that urbanization had the function of species conservations with sharp increases of alien species and tropical type plants. In the case woody species, 60-yr urbanization in Harbin had induced increases of 9 families and 17 genera, and there were 7–20 more families, 12–35 more genera, 1.6–2.6 higher Margalef richness in urban areas than those in nature reserves and local forest farms; Increases in alien species (4-fold in 60-yr urbanization; 21% in urban area vs <2% for non-urban region) and tropical type plants (1.6-fold in 60-yr urbanization; temperate/tropical ratio at 1.2 in urban area vs >1.6 in non-urban area) were mainly responsible for these compositional changes, which can be proved by their significant correlations. Moreover, moderate disturbance had peak values in alien species, tropical type plants, Shannon-wiener diversity, Margalef richness index and Pielou evenness index, and both ring road- and buildup history gradients showed the similar tendency. Compared with those in 1980s, forest- and eurytopic-habitats birds increased 9–11 species (23–39%), and omnivorous, insect-eating, and phytophagous bird increased 5–9 species (14.1–29.4%) in those in 2010s, indicating that bird temporal changes were closely related with the changes in urban forests owing to food supply and habitat provision. Our findings could provide data for biodiversity evaluation of urbanization effects, and is also useful for ecological re-construction of local cities in China.     

The utility of ancient forest indicator species in urban environments: A case study from Poznań, Poland

We aimed to assess the impact of land-use structure on AFIS occurrence and evaluation of these bioindicators in urban conditions. We compiled data about forest continuity based on archival maps and forest management plans; floristic records of 79 AFIS occurrences were collected within a 1 × 1 km grid and land-use form structure in Poznań (W Poland). We tested fidelity of AFIS using χ² tests and effects of land-use structure using random forest models. We also checked spatial autocorrelation and its impact on AFIS distribution patterns within old and recent forests, using spatially explicit generalized linear models. We found a strong relationship between AFIS number per grid square and fraction of forests and waters in land-use structure. Relationships between AFIS distribution and land-use shows that AFIS are a good proxy describing human impact or an urbanity gradient. AFIS number per grid square shows small, but significant positive spatial autocorrelation, which suggests possibilities of migration into new forests. AFIS, treated as a bioindicative group of species, may be a useful indicator of landscape and land-use structure transformation in urban environments. Occurrence of many AFIS that are endangered at the city scale indicates the need for conservation of old forests in urban areas, to preserve biodiversity in urban environments. Thus, AFIS may act as indicators of old forests with high conservation value in cities, which need special protection.     

Diversity and distribution of the urban tree population in ten major Nordic cities

In order to have a healthy and sustainable urban tree population, a high diversity of species and genera is needed. This study examined (1) the diversity and distribution of genera and species of urban trees in the Nordic region; (2) the diversity in different sites of the city, distinguishing between street and park environments; and (3) the presence of native versus non-native tree species in urban environments in the Nordic region. The analysis of tree diversity was based on urban tree databases comprising a total of 190 682 trees in 10 Nordic cities – Aarhus and Copenhagen in Denmark; Espoo, Helsinki, Tampere and Turku in Finland; Gothenburg, Malmo and Stockholm in Sweden; and Oslo in Norway. The tree databases for Copenhagen, Espoo, Helsinki, Stockholm and Tampere only record street trees, while the remaining databases also include park trees. Tilia was the most dominant genus in Arhus, Copenhagen, Espoo, Gothenburg, Helsinki, Oslo and Stockholm, while Sorbus was the most dominant in Malmo and Betula in Tampere and Turku. Tilia × europaea was the most common species, comprising 16.0% of the total number of tree species. There was a higher proportion of species in parks than in street environments. The number of non-native species was higher than the number of native species in both street and park environments. However, the number of individuals belonging to native species was higher than the number of non-native individuals in all cities and environments except park environments in Arhus. The concluding recommendation from this study regarding greater diversity of genera and species is to exploit local experiences of rare species from local urban tree databases. After appropriate evaluation, urban tree planners can evaluate these rare species in larger numbers for e.g. street environments, where the need is greatest.     

A multiscale assessment of the diversity of New Zealand’s nursery trees

Nurseries play an important role providing trees for a variety of managed environments including urban forests. The diversity of urban forests and forest restoration projects are influenced by nursery species availability, and as such, there is a need to better understand tree species diversity at nurseries. We collected tree species lists from 75 nurseries throughout New Zealand, which were used to describe species richness (alpha diversity) and to examine similarity in the composition of native and non-native species assemblages among nurseries (beta diversity) at three spatial scales: island, region, city. Together, the nurseries grew 863 species, 174 of which were native to New Zealand, from 312 genera and 130 families. Nurseries grew significantly more non-native species (μ = 63.5, σ = 60.6) than native species on average (μ = 31.7, σ = 22.7) (t = 2.99, df = 48.45, p = 0.004). Beta diversity for native and non-native tree species were only significantly different at the scale of cities or regions, not at the larger scale of islands. Few species were grown in all cities or all regions and the majority of those that were common were native species. In contrast, non-native species dominated the unique species at all spatial scales, (i.e., species uniquely grown in one city, region, or island). By quantifying tree species diversity in New Zealand’s nurseries, this research provides a basis to better understand the influence that nurseries have on urban and peri-urban tree diversity, and ultimately how that diversity impacts resilience and the provision of ecosystem services.     

Potential hazard characteristics of Tilia, Betula, and Acer trees removed in the Helsinki City Area during 2001–2003

In order to study decay, and to improve the management and protection of old urban trees, a total of 256 felled urban trees were examined during 2001–2003: 95 Tilia spp., 74 Betula spp., and 87 Acer spp. Most of the trees (73%) were located in the main parks and along the main streets in the downtown area of Helsinki City, Finland. The mean age of the trees was over 60 years, and the majority (64%) were old park trees. Poor condition and increasing risk of failure were the main reasons for felling in 82% of the cases. Thirty three percent of these trees were degenerated or dead, but the amenity value of 14% of the risk trees was still high. The latter were old, big trees which posed a potential hazard, but had a vital and balanced crown.Some characteristic profiles for potential failure were identified for each of the tree species studied: Ganoderma lipsiense in the butts and hollows in the stems of Tilia spp., weak fork formations together with Rigidoporus populinus on Acer spp., and degeneration together with decay in the stem on Betula spp.Decay fungi most commonly identified were R. populinus, G. lipsiense, Inonotus obliquus and Piptoporus betulinus. In addition, Kretzschmaria deusta was very common in three of the parks, and on every one of the tree species investigated.     

Promoting and preserving biodiversity in the urban forest

Efforts at mitigating global biodiversity loss have often focused on preserving large, intact natural habitats. However, preserving biodiversity should also be an important goal in the urban environment, especially in highly urbanized areas where little natural habitat remains. Increasingly, research at the city/county scale as well as at the landscape scale reveals that urban areas can contain relatively high levels of biodiversity. Important percentages of species found in the surrounding natural habitat, including endangered species, have been found in the urban forest.

This contribution concisely highlights some examples of urban biodiversity research from various areas of the world. Key issues involved in understanding the patterns and processes that affect urban biodiversity, such as the urban–rural gradient and biotic homogenization, are addressed. The potential for urban areas to harbor considerable amounts of biodiversity needs to be recognized by city planners and urban foresters so that management practices that preserve and promote that diversity can be pursued. Management options should focus on increasing biodiversity in all aspects of the urban forest, from street trees to urban parks and woodlots.     

Isolation of arbuscular mycorrhizal fungi and Azotobacter chroococcum from local litchi orchards and evaluation of their activity in the air-layers system

Soil samples were collected from rhizosphere of litchi-growing areas of North-Western Himalayan Region (NWHR) of India, for finding qualitative and quantitative differences in arbuscular mycorrhizal (AM) fungi and Azotobacter chroococcum. These samples were taken from plants being grown in different cultivation types namely, weed control with weedicides or tillage; orchard floor either clear or with cover crops; intercropping with cereals and legumes. Qualitative and quantitative differences were noticed with different cultivation types and a marked reduction in the AM fungi was observed in orchards where chemicals were used for weed control and intensive farming system was used on the orchard floor. AM fungi were generally abundant in the soils with range pH 5.5–6.6. Among different AM fungi retrieved from the soils, Glomus spp. was most dominant. Fifteen AM fungal species were isolated, identified and characterized and along with their ability to colonize the roots. In the soil samples, a marked variation in viable bacterial count of A. chroococcum was also noticed due to varied physico-chemical characteristics of the orchard soils. The changes in AM fungal species composition can be attributed to changes in soil chemical properties resulting from cultural practices such as ploughing, application of chemical fertilizers and weedicides. An experiment was also conducted to study the comparative efficacy of four dominant and frequently occurring indigenous AM species namely, Glomus fasciculatum (Thaxter sensu Gerdemann), G. magnicaulis (Hall), G. mosseae (Nicol. & Gerd.), Gigaspora heterogamma (Nicol. & Gerd.) and two A. chroococcum strains viz., AZ₁ and AZ₂ singly and in dual combination to evaluate their effect in air-layers system. Dual inoculation of G. fasciculatum and AZ₁ increased total root length of air-layered shoots by 81.39% over uninoculated control. These studies indicated that indigenously isolated AM fungal species and A. chroococcum strains can be used for air-layering for better adaptation under specific agro-climatic and ecological zone conditions.     

Biotic homogenization of China’s urban greening: A meta-analysis on woody species

Human preferences for the species in urban greening have greatly facilitated the spread of non-native species, resulting in the homogenization of urban plant communities across spatial scales. We selected 11 major cities along the Yangtze River in China and examined the species composition in their urban plant communities. We found that China’s urban plant communities are becoming homogenized, as urban communities of different cities are highly similar to each other despite the geographical separation. Meanwhile, these artificial communities we investigated have diverged greatly from the natural communities at both the city and the geographic scale. We recorded a total of 91 woody species that have been used in urban greening in all 11 cities. Of those species, 27% were cultivars and introduced species, and 25% were being used outside of their native distribution ranges in China. This may be explained by the market and urban planners who tend to favor greening plants that are highly profitable and have aesthetic ornamental traits, rather than spending time introducing and acclimatizing the native species in each city. Given the current trend of homogenization, measures that recognize the importance of native species should be emphasized with comprehensive urban planning strategies.     

Multi-scale responses of bird species to tree cover and development in an urbanizing landscape

Landscape change is an ongoing process for even the most established landscapes, especially in context to urban intensification and growth. As urbanization increases over the next century, supporting bird species’ populations within urbanizing areas remains an important conservation challenge. Fundamental elements of the biophysical structure of urban environments in which bird species likely respond include tree cover and human infrastructure. We broadly examine how tree cover and urban development structure bird species distributions along the urban-rural gradient across multiple spatial scales. We established a regional sampling design within the Oak Openings Region of northwestern, Ohio, USA, to survey bird species distributions across an extensive urbanization gradient. Through occupancy modeling, we obtained standardized effects of bird species response to local and landscape-scale predictors and found that landscape tree cover influenced the most species, followed by landscape impervious surface, local building density, and local tree cover. We found that responses varied according to habitat affiliation and migratory distance of individual bird species. Distributions of short-distance, edge habitat species located towards the rural end of the gradient were explained primarily by low levels of urbanization and potential vegetative and supplemental resources associated with these areas, while forest species distributions were primarily related to increasing landscape tree cover. Our findings accentuate the importance of scale relative to urbanization and help target where potential actions may arise to benefit bird diversity. Management will likely need to be implemented by municipal governments and agencies to promote tree cover at landscape scale, followed by residential land management education for private landowners. These approaches will be vital in sustaining biodiversity in urbanizing landscapes as urban growth expands over the next century.     

Dispersion of kangaroo rat mounds at multiple scales in New Mexico,USA

Burrowing mammals create disturbances that increase the ecological heterogeneity of landscapes. In desert systems, banner-tailed kangaroo rats (Dipodomys spectabilis) construct large mounds that greatly influence the spatial patterning of soils, plants, and animals. The overall effects of the patches generated by D. spectabilis depend on the dispersion patterns of the mounds; these patterns may be sensitive to scale and landscape position. We examined the distribution of D. spectabilis mounds across multiple scales in four 40-ha grassland plots in New Mexico, USA. We used Ripley's K-function for our point-pattern analysis. The dispersion patterns of mounds were generally scale-sensitive but depended somewhat on plot-level densities, which were related to topographic position and grazing history. Mound spacing was either regular or random at small scales (0–50 m), random or aggregated at intermediate scales (50–300 m), and aggregated at large scales (300–3000 m). This scale-dependency of pattern reflected spatial domains in which different biotic (territoriality, dispersal, grazing) and abiotic (soil texture and drainage) factors exerted strong influences. How other organisms perceive the spatial patterning of mounds will depend on the extent of their movements. Patches may appear regular to one species but aggregated to another. The dispersion of D. spectabilis mounds also has implications for the spatial structuring of desert rodent communities. D. spectabilis excludes smaller species of kangaroo rats from areas around their mounds; they create spatial heterogeneity in behavioral dominance that may influence the distribution of subordinate species at multiple scales.     

Road verges as invasion corridors? A spatial hierarchical test in an arid ecosystem

Disturbed habitats are often swiftly colonized by alien plant species. Human inhabited areas may act as sources from which such aliens disperse, while road verges have been suggested as corridors facilitating their dispersal. We therefore hypothesized that (i) houses and urban areas are propagule sources from which aliens disperse, and that (ii) road verges act as corridors for their dispersal. We sampled presence and cover of aliens in 20 plots (6 × 25 m) per road at 5-km intervals for four roads, nested within three localities around cities (n = 240). Plots consisted of three adjacent nested transects. Houses (n = 3,349) were mapped within a 5-km radius from plots using topographical maps. Environmental processes as predictors of alien composition differed across spatial levels. At the broadest scale road-surface type, soil type, and competition from indigenous plants were the strongest predictors of alien composition. Within localities disturbance-related variables such as distance from dwellings and urban areas were associated with alien composition, but their effect differed between localities. Within roads, density and proximity of houses was related to higher alien species richness. Plot distance from urban areas, however, was not a significant predictor of alien richness or cover at any of the spatial levels, refuting the corridor hypothesis. Verges hosted but did not facilitate the spread of alien species. The scale dependence and multiplicity of mechanisms explaining alien plant communities found here highlight the importance of considering regional climatic gradients, landscape context and road-verge properties themselves when managing verges. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The use of gradient analysis studies in advancing our understanding of the ecology of urbanizing landscapes: current status and future directions

Over the past decade, the urban–rural gradient approach has been effectively used to study the ecology of cities and towns around the world. These studies have focused on understanding the distribution of plants and animals as well as ecosystem processes along gradients of urbanization that run from densely urbanized inner city to more rural exurban environments. We reviewed 300 papers investigating urbanization gradients that were published in peer-reviewed journals between 1990 and May 2007. Sixty-three percent of the papers investigated the distribution of organisms along urbanization gradients. Only five papers addressed the measures used to quantify the urbanization gradient itself. Within the papers addressing the distribution of organisms, 49% investigated the responses of birds to urbanization gradients, and <10% of the papers investigated more cryptic organisms. Most of these studies utilized a variety of broad measures of urbanization, but future advances in the field will require the development of some standardized broad measures to facilitate comparisons between cities. More specific measures of urbanization can be used to gain a mechanistic understanding of species and ecosystem responses to urbanization gradients. While the gradient approach has made a significant contribution to our understanding of the ecology of cities and towns, there is now a need to address our current knowledge gaps so that the field can reach its full potential. We present two examples of research questions that demonstrate how we can enhance our understanding of urbanization gradients, and the ecological knowledge that we can obtain from them.