Short- and long-term effects of site factors on net N-mineralization and nitrification rates along an urban-rural gradient

Long- and short-term effects of urban site factors on net N-mineralization and nitrification rates were investigated in oak stands along an urban-rural land-use transect in the New York City metropolitan area. We used reciprocal transplants of undisturbed soil cores between urban and rural forests to determine the relative importance of long-term effects (mor vs. mull soils, quality of soil organic matter, and deposition of N) vs. short-term effects (soil temperature) of urban factors in controlling field N-transformation rates along the gradient. In addition, undisturbed soil cores from surface (A, Oe horizons) and subsurface (B horizon) soil were collected from urban, suburban, and rural stands and allowed to incubate in these respective sites to compare the net effect of all urban factors with transplanted-core results. The transplant experiment revealed that soil type (long-term) affected net N-mineralization and nitrification rates. Urban soils nitrified nearly 6.3 and 5.4 times more than rural soils incubating in urban and rural stands, respectively (p = 0.003 and p = 0.002, respectively). Similarly, in rural stands total accumulation of inorganic N was 87% higher in urban than in rural soils, whereas in urban stands, urban soils mineralized 83% more N than rural soils (p = 0.043 and 0.08, respectively). Comparing soils incubating in their native locations, urban soils incubating in urban stands mineralized more than 2.5 times the amount of N than rural soils incubating in the rural stands (p = 0.019). By contrast, urban soils incubating in urban stands exhibited a 8-fold increase in nitrification over rural soils incubating in rural stands (p = 0.008). As with the transplanted cores, the urban and suburban environments had a positive effect on net rates of N-mineralization and nitrification in both surface and subsurface layers of soil. The surface layer of suburban and urban stands had a 3- and 2.3-fold higher accumulation of net inorganic N than rural stands (ANOVA, p = 0.05). Similarly, in the subsurface layer both urban and suburban stands had 2.6-fold higher net N-mineralization rate than rural stands (ANOVA, p = 0.01). Along this urban-rural gradient, soils in oak stands exhibit higher net nitrification and, to a lesser extent, net N-mineralization rates in urban and suburban stands than in rural stands. Results from the transplant experiment and in situ measurements of surface and subsurface soil indicate that long-term effects (mor vs. mull soils, N deposition) contribute to the higher N-transformation rates in urban and suburban stands. As a result of these effects, urban and suburban stands have the potential for higher losses of N than rural stands.     

Potential nitrogen mineralization responses of urban and rural forest soils to elevated temperature in Louisville,KY

Soil nitrogen (N) mineralization is an important process determining terrestrial N availability, and evidence suggests elevated temperatures will enhance N mineralization rates. Along a 40 km urban-rural gradient of chestnut oak forest stands in Louisville, KY, we expected N mineralization rates would be higher in urban than in rural forests in part due to increased temperatures caused by the urban heat island. However, a 12-month field study along this Louisville gradient showed that annual N mineralization rates were lower in urban than in rural stands. Since variation in precipitation inputs and other factors across this land-use gradient may be influencing soil N mineralization rates, we conducted a three-month soil incubation experiment in the lab to determine the extent to which a + 2 °C temperature difference could affect soil N mineralization in urban and rural soils. Across the range of temperatures tested, rural soils mineralized N at twice the rate of urban soils under base (7.86 vs. 3.65 mg N kg^?1 AFDW soil d^?1) and elevated (9.08 vs. 4.76 mg N kg^?1 AFDW soil d^?1) temperatures (p < 0.01). A 2 °C temperature difference, did not significantly alter total inorganic N production in urban (p = 0.272) or rural soils (p = 0.293). The proportion of nitrate produced was lower in the urban (15.1 %) than in the rural soils (72.3 %; p < 0.01). These results suggest that differences in soil organic matter quality and potentially decomposer community composition are the primary explanatory factors for forests along this Louisville gradient.     

Litter decomposition and nitrogen mineralization in oak stands along an urban-rural land use gradient

We investigated litter mass loss and soil nitrogen (N)-transformation rates in oak stands along a 130-km, urban-rural transect originating in New York City to examine the relationship between changes in these parameters and previously documented differences in soil temperature, heavy metal and total salt concentrations, and soil biota. Reference litter from a rural site was placed in litterbags, and rates of mass loss and changes in N concentration in litter residues were measured over a 6-month period. Net N-mineralization and nitrification rates were measured in A horizon soils using laboratory incubations under constant moisture and temperature. Both mass loss (76%) and N release (65%) from litterbags reached their maximum in urban stands, whereas net N-mineralization rates were 2.3-fold higher than in rural A horizon soils. Litter fragmentation by earthworms and higher soil temperatures are potential causes of the higher mass loss rates observed in urban stands. The higher releases of N measured in the urban litterbags could be a result of their faster mass loss rates, exogenous inputs of N from atmospheric deposition, a relatively low heterotrophic demand for N, or a combination of these factors. The results of this study suggest that in comparison with rural stands, urban forests are characterized by comparatively high rates of litter decomposition, and may also be characterized by comparatively high rates of N mineralization. Additional studies are needed to test whether these effects are common to urban environments in general.     

Conifer growth and reproduction in urban forest fragments: Predictors of future responses to global change?

Global change has a large and growing influence on forests, particularly in urban and urbanizing areas. Compared to rural forests, urban forests may experience warmer temperatures, higher CO₂ levels, and greater nitrogen deposition, with exacerbated differences at urban forest edges. Thus, comparing urban to rural forests may help predict future effects of global change on forests. We focused on the conifer western red-cedar (Thuja plicata) to test three hypotheses: at urban forest edges, relative to rural forests and urban forest centers, trees experience 1) higher temperatures and nitrogen levels, 2) lower seedling recruitment, and 3) greater growth. We additionally tested anecdotal reports that 4) tree seedling recruitment in urban and rural forests is much lower than in ??pristine?? old-growth forests. To test these hypotheses, we quantified air temperature, soil nitrate, adult T. plicata growth and seedling recruitment in five urban and three rural parks at both forest edges and centers. We also quantified T. plicata recruitment at five old-growth ??pristine?? sites. Temperatures were highest at urban forest edges, and soil nitrate was highest in urban forests. In urban relative to rural forests, we observed greater T. plicata growth, but no difference in seedling densities. However, seedling densities were lower in urban and rural forests than in old-growth forests. In all, our results suggest urban influences enhance adult T. plicata growth, but not seedling recruitment. Recruitment in urban and rural forests was reduced compared to old-growth forests, implying that fragmentation and logging reduce T. plicata seedling recruitment.     

Rural land use bifurcation in the urban-rural gradient

The classic urban-rural gradient concept assumes a decline in land use intensity from an intensively developed urban core outward to residential suburbs, culminating in lightly developed rural areas. The concept provides a common framework directing urban socio-ecological research. Given that rural land use includes woodlands and croplands, and croplands appear as ecologically depauperate as urban lands, we investigated land-use patterns along urban-rural gradients for 30 large metropolitan areas in the eastern United States. We predicted a bifurcation at the rural end of the gradient between woodland and cropland land use that does not correspond with human population density (expected to be relatively low in rural areas regardless of land use type). Our data indicated that ‘rural’ was a poor substitute for ‘natural’ as the rural end of the gradients bifurcated at the rural end between woodland and cropland – croplands being demonstratively poor ecological habitats. Indeed, we found that when defined by habitat quality, the habitat known to be biotically homogenized (urban and cropland) remained steady along the urban-rural gradient. Our results do not undermine the utility of the urban-rural gradient framework, but do suggest that the gradient and/or human population density do not necessarily indicate shifts in habitat quality.     

Species change in an urban setting—ground and rove beetles (Coleoptera: Carabidae and Staphylinidae) in Berlin

The influence of environmental parameters on epigeic beetle communities of forest fragments in an urbanization gradient was studied in Berlin. Eight deciduous forests along a rural to urban gradient were sampled with pitfall traps. Species richness did not decline across the rural to urban gradient. As expected, impervious surface cover as an indicator of urbanization correlated not only with habitat fragmentation and heat island effect but also with altered soil properties. The proportion of forest specialist staphylinid species decreased with increasing urbanization. The differences between staphylinid communities of neighboring forest fragments were enhanced in the most urban parts, probably due to increased habitat fragmentation. Furthermore, the loss of flightless species with increasing habitat isolation emphasized the influence of habitat fragmentation. The carabid communities revealed the urbanization effects not as clearly as the staphylinid communities, but both taxa revealed that direct anthropogenic habitat alteration, indicated by removal of decaying wood, favors open-habitat specialists. The extent of the urbanization influence seems to vary seasonally. Environmental parameters associated with urbanization explain the ordination of species communities in the winter better than in the summer. Heat island effect is suggested as an explanation for this difference.     

Forest leaf litter quantity and seedling occurrence along an urban-rural gradient

Leaf litter, by modifying microenvironmental conditions, can alter plant population distributions and is considered to be a major force in structuring many plant communities. Comparative studies of urban, suburban, and rural forests in the New York City (NYC) metropolitan area have identified numerous biotic and abiotic differences among these forests, including several involved in leaf litter decomposition. These differences in decomposition among the forests could result in differences in litter quantity on the forest floor and hence microenvironmental conditions and safe sites for germination of different plant species. We conducted a survey of forest floor leaf litter quantity in forests located along an urban-to-rural land-use gradient originating in urban NYC and extending to rural Connecticut. Mean litter depth, mass, and density increased significantly with increased distance of the forest from NYC.We also surveyed woody seedlings and compared the litter depth in which they naturally occurred to the mean litter depth of the surrounding forest. Seedlings of small-seeded species were much more likely to be located in litter shallower than mean forest litter depth than were seedlings of large-seeded species. Taken together, these results suggest that environmental changes associated with urbanization may have profound effects on long-term patterns of forest regeneration.     

The genesis,classification, and mapping of soils in urban areas

This paper discusses the concept of soil in both urban and rural environments, and along the urban-rural land use gradient, to illustrate the obvious need to increase our understanding of urban soils. Spatial variability of the urban landscape is illustrated with "Soil series - Urban land complexes" from Baltimore County, Maryland. The World Reference Base for Soil Resources (ISSS-ISRIC-FAO, 1994) proposed Anthrosol and Regosol major soil groups are discussed to show modern approaches to soil classification and to illustrate how the classification of urban soils is essentially undeveloped. Models of soil genesis help identify the processes and functions of the soil system. A conceptual model using Jenny's (1941) state factor approach for human impact on soil formation details the "anthroposequence." The benchmark anthroposequence model may be applied to studying soil systems along urban-rural land use gradients. The process of "anthropedogenesis" is supported to quantify the role of human activity in changing the "natural" direction of soil formation. Future directions of soil research in the urban landscape should involve large scale soil mapping (e.g. 1:6000), benchmark anthroposequences, improved soil classification, and refined characterization of the role of human activities in soil formation.     

Variation in urban forest productivity and response to extreme drought across a large metropolitan region

The growth and survival of urban trees and maintenance of urban forest canopy are important considerations in adaptation of urban regions to climate change, especially in relation to increasing frequency of extreme climatic events such as drought. However, urban forest growth and drought response may vary considerably within large urban landscapes across gradients in land use, urbanization, forest composition and structure, and environmental factors. We quantified urban forest growth and resilience and resistance to extreme drought in the greater Chicago metropolitan region based on patterns of annual basal area production from increment core analysis. We evaluated variation in growth and drought response in relation to a broad urban to rural gradient, land-use categories, local-scale environmental predictors, and forest community characteristics. Urban forest growth varied greatly among land-use classes and major genera. Plot-level variation in productivity was predicted most strongly (R² = 0.53) by total plot-level basal area, canopy height, species composition, soil and ground-cover characteristics, and position within the urban-rural gradient. Urban forest growth was strongly related to regional meteorological drought. In periods of extreme drought conditions growth declined in the year of the drought (i.e., was not resistant to drought effects), but was highly resilient to drought in the subsequent 5 year period. Drought response did not vary consistently across land-use classes or among major genera, and site or community characteristics had little explanatory power in predicting drought response. Improved understanding of factors driving variation in urban forest growth and drought response could help inform adaptation-focused urban forest management strategies.

     

Non-native plants in the understory of riparian forests across a land use gradient in the Southeast

As urbanization expands into rural areas, an increase in the number of non-native plant species at the urban-rural interface is expected due in large part to the increased availability of propagules from ornamental plantings. A study investigating the distribution of non-native plants in the understories of riparian forests across an urban-to-rural gradient north of Columbus, GA was initiated in 2003. A significantly greater number of non-native plant species occurred at the urban sites and at one site at the urban-rural interface, where 20 to 33% of the species encountered were non-native. In contrast, at the more rural sites non-native species comprised 4–14% of the total number of species. However, the importance values of non-native species as a whole did not change significantly across the land use gradient due to the high frequency and abundance of three non-native species (Ligustrum sinense, Lonicera japonica, and Microstegium vimineum) in the majority of the watersheds. Reductions in species richness and overstory reproduction associated with these non-natives could impact long-term forest structure and ecosystem function.     

Contrasting effects of urbanization on arboreal and ground-dwelling land snails: role of trophic interactions and habitat fragmentation

Urbanization generally reduces wildlife populations. Individual species responses, however, are often highly variable, and such variability can be explained by differences in species ecological traits. To examine this hypothesis, we focused on two co-occurring land snails, Ezohelix gainesi and Euhadra brandtii sapporo; the former is ground-dwelling and the latter is arboreal. We first estimated their population densities at nine sites distributed along an urbanization gradient: three were located in continuous natural forests, three at the edge of natural forests, and the rest in small isolated forests in urban areas. As a result, the ground-dwelling E. gainesi occurred at highest density in urban forests, followed by forest edges and continuous forests. By contrast, the arboreal E. b. sapporo occurred at highest density in continuous forests, but declined in forest edges and urban forests. We then conducted manipulative field experiments to quantify changes in predation pressure on these species. Ground-tethered E. gainesi and E. b. sapporo were repeatedly predated upon by forest-living mammals in continuous forests, but their survival rates increased in forest edges and urban forests. By contrast, canopy-tethered E. b. sapporo maintained high survival rates in all three forest types. The results indicate that a lack of mammalian predators enables ground-dwelling species to occur at high densities in urban forests, whereas the arboreal species was not affected by this predator relaxation effect. The combination of species-specific behavioural traits and changes in predator communities across an urbanization gradient has important effects on the biodiversity of urban ecosystems.

     

Understory plant species composition in remnant stands along an urban-to-rural land-use gradient

We examined the understory species composition of 24 remnant forest stands along an urban-to-rural gradient in the metropolitan Milwaukee, Wisconsin region to determine the relationships between plant community composition, human disturbance, and contrasting types of land use along a gradient of urbanization. A significant difference was found in shrub species community composition among three contrasting land-use categories but no significant difference was found in herbaceous community composition. Significant differences in human activity existed among rural, urban, and urbanizing land-use categories, but this index of disturbance was not significantly correlated to gradients in species composition. All stands in this study had been subjected to various types of human activity and environmental disturbances in the past. Our data suggest that differences in the relative importance of understory species exist among stands but these differences may not be caused by the impacts of urbanization alone. Changes in the natural disturbance regime of this landscape, along with the impacts associated with urbanization, have led to an individualistic response in the compositional dynamics of forest stands. This revised version was published online in July 2006 with corrections to the Cover Date.     

Silphids in urban forests: Diversity and function

Many ecologists have examined the process of how urbanization reduces biological diversity but rarely have its ecological consequences been assessed. We studied forest-dwelling burying beetles (Coleoptera: Silphidae)—a guild of insects that requires carrion to complete their life cycles—along an urban-rural gradient of land use in Maryland. Our objective was to determine how forest fragmentation associated with urbanization affects (1) beetle community diversity and structure and (2) the ecological function provided by these insects, that is, decomposition of vertebrate carcasses. Forest fragmentation strongly reduced burying beetle diversity and abundance, and did so far more pervasively than urbanization of the surrounding landscape. The likelihood that beetles interred experimental baits was a direct, positive function of burying beetle diversity. We conclude that loss of burying beetle diversity resulting from forest fragmentation could have important ecological consequences in urban forests.     

Forest soils adjacent to urban interstates: Soil physical and chemical properties,heavy metals,disturbance legacies,and relationships with woody vegetation

While urban ecology is an expanding field of study, some natural areas within the urban environment remain under-examined. These include naturally regenerating forest communities adjacent to urban interstates. In addition, the status of interstate soils and their relationships with the community composition of forested interstate verges has received little ecological study. The purpose of this study was to examine variation in soil conditions along forested interstate corridors in Louisville, KY and to explore the extent to which soil characteristics (e.g., bulk density, pH) and heavy metals (e.g., Pb, Zn) vary with respect to three factors: interstate (e.g., traffic density), surrounding urban environment (e.g., industrial land use), and interstate construction legacies. Additionally, we explored the relationships between several edaphic factors and woody vegetation structure in these forested verges. We found that the degree and direction of the slope of land towards the interstate and the distance to the interstate pavement were strong determinants of soil characteristics and heavy metal concentrations, suggesting that the movement of de-icing salts, heavy metals, and other pollutants from the interstate was important in determining forest soil conditions along urban interstates. Since within our study area these highways did not extend into rural lands, variation in urban land uses and cover within 26 km of the city center was not large enough to explain variation in soil characteristics or heavy metals, except for a positive correlation between chromium and surrounding industrial land use. We did find that past physical soil disturbance caused by interstate construction (e.g., imported fill) left an important legacy on soil characteristics, heavy metal retention, and woody plant growth patterns in forests adjacent to urban interstates. The legacy of interstate construction on the current forest community structure (e.g., lower species richness) and the future forest (e.g., reduced tree regeneration) may further alter ecosystem productivity and ecosystem services provided by these forests and their soils.     

A comparison of carbon and nitrogen stocks among land uses/covers in coastal Florida

Coastal areas are rapidly developing due to population growth and the appeal of coastlines. In order to gain insight into how land use/cover affects carbon (C) storage in a coastal context, we examined soil and vegetation C and soil nitrogen (N) across land uses near Apalachicola, FL. Forested wetlands had the greatest soil C and N storage, while natural pine forests and pine plantations had the least. In paired plots, urban lawns had significantly greater mineral soil N content compared to urban forest remnants. Total ecosystem C (soil + vegetation) was higher in forested wetlands than all other land uses/covers combined due to the high organic content of those wetland soils. Urban forest remnants and lawns had greater total ecosystem C than natural pine forests and pine plantations, which likely reflects the differential influence of prescribed fire and less frequent anthropogenic disturbances between the rural and urban areas, respectively. Projections of land use change in Franklin County, FL combined with these data suggest that increases in C storage are possible with continued urbanization along the Gulf Coast, if forest remnants are left and lawns are incorporated in built-up areas. However, this study does not account for C emissions during land conversion, or any emissions associated with maintaining urban built-up and residential areas. A better understanding of land use/cover influences on C pools has applications for planning and development, as well as ecological and environmental protection in the region.     

The effects of urbanization on ant communities and myrmecochory in Manitoba,Canada

With the increase in urbanization globally, there is an increased need to understand the ecology of forest fragments in urban and urbanizing landscapes. Although urban forests are known to be relatively lacking in plants whose seeds are dispersed by ants, little is known about the effects of urbanization on the community composition and behaviour of forest dwelling ants. Ant communities in forest fragments along an urban–rural gradient were described using a rapid quadrat search technique and multivariate analysis. Interactions between the ants and seeds of the myrmecochorous Viola pubescens within a subset of these forests were described using a series of cafeteria experiments. Urbanization was found to be associated with changes in microhabitat characteristics and a concomitant simplification of the ant community. Despite this, the removal rate of V. pubescens seeds actually increased in urban forests, which may be a result of the foraging behaviour of the remaining species.

Emerging dragonfly diversity at small Rhode Island (U.S.A.) wetlands along an urbanization gradient

Natal habitat use by dragonflies was assessed on an urban to rural land-use gradient at a set of 21 wetlands, during two emergence seasons (2004, 2005). The wetlands were characterized for urbanization level by using the first factor from a principal components analysis combining chloride concentration in the wetland and percent forest in the surrounding buffer zone. Measurements of species diversity and its components (species richness and evenness) were analyzed and compared along the urbanization gradient, as were distributions of individual species. Dragonfly diversity, species richness, and evenness did not change along the urbanization gradient, so urban wetlands served as natal habitat for numerous dragonfly species. However, several individual species displayed strong relationships to the degree of urbanization, and most were more commonly found at urban sites and at sites with fish. In contrast, relatively rare species were generally found at the rural end of the gradient. These results suggest that urban wetlands can play important roles as dragonfly habitat and in dragonfly conservation efforts, but that conservation of rural wetlands is also important for some dragonfly species.     

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

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