Partitioning the effects of habitat fragmentation on rodent species richness in an urban landscape

Habitat fragmentation plays a major role in species extinction around the globe. Previous research has determined that species richness in fragments is affected by a number of characteristics. These include fragment age, size, and isolation, edge effects, vegetation coverage, habitat heterogeneity, and matrix content. Although most studies focused on one or a few of these characteristics, multiple characteristics work together to affect species richness, showing that the effects of habitat fragmentation are complex. The goal of our study was to partition the complex effects of habitat fragmentation by determining the direct, indirect, and cumulative effects of multiple habitat fragment characteristics on rodent species richness. In 2013, we determined rodent species richness in 25 habitat fragments within Thousand Oaks, California. In addition, we measured the following characteristics for each fragment: fragment age, area, isolation, shrub coverage, habitat heterogeneity, perimeter/area ratio, and percent non-urban buffer. Path Analysis was used to test the hypothesized model which described the direct, indirect, and cumulative effect of each habitat fragment characteristic on rodent species richness. Overall, the path model explained 67 % of the variation in rodent species richness among habitat fragments. Habitat heterogeneity had the greatest direct and total effect on rodent species richness. Fragment size had the next greatest total effect on rodent species richness but this was nearly entirely indirect through its influence on habitat heterogeneity, suggesting that large fragments containing the greatest diversity of habitats will support the most species. Our study shows that large habitat fragments support the greatest habitat diversity, which provides the highest likelihood of conserving rodent species richness in an urban landscape.     

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.     

Patterns of human disturbance and response by small mammals and birds in chaparral near urban development

We report on the extent of disturbance (including habitat alteration and road and trail proliferation) in chaparral near urban development and analyze the effects of disturbance on small mammal and resident bird species. Disturbance patterns were evaluated in a 6700 ha study area in southern California: effects on mammals and birds were investigated by analyzing relationships between vegetation structure and animal species richness and abundance. Disturbance was prevalent throughout the study area and included extensive human-altered habitat (from past human activities such as vegetation clearing, human-caused fires, refuse dumping, and vegetation trampling) and 157 km of roads and trails. A nonsignificant trend was found between human-altered habitat and proximity to development, but human-altered habitat was significantly associated with roadway proximity. Trails were also more frequent near urban development and roads. Small mammals responded strongly to disturbance-related vegetation changes, while birds showed little or no response. Mammals endemic to chaparral vegetation were less diverse and abundant in disturbed sites, whereas disturbance-associated species increased in abundance. Close proximity of urban development to natural areas resulted in alteration of natural habitat and proliferation of roads and trails. Variation in life history traits between birds and mammals may affect response to disturbance and influence persistence if disturbance continues. Conservation efforts must recognize the potential for habitat damage and associated declines in native animal species caused by disturbance near urbanization and implement strategies to reduce these threats.     

Suitability of biodiversity-area and biodiversity-perimeter relationships in ecology: a case study of urban ecosystems

Urban areas are probably the most fragmented environments with respect to the presence of semi-natural habitats and shape of these habitats may be significantly affected by urbanization. Patch perimeter in landscape or habitat studies is much less popular to study than patch area. The studied sites were situated in the industrial city of Pardubice, which is one of the ten largest cities in the Czech Republic with nearly 100,000 inhabitants. In total, 40 grasslands were studied within a circular area of 314 km². Butterflies and beetles with diurnal activity were studied during timed survey walks. A Simultaneous autoregressive model was used for test of the effect of biodiversity-area and biodiversity-perimeter relationships and for exclusion of potential bias caused by spatial autocorrelation. The models including patch perimeter performed better than those using patch area in explaining species richness, abundance and diversity of investigated organisms and were less influenced by spatial autocorrelation. The main conclusion and recommendation of the present study is that researchers should pay more attention to the possible influence of the patch perimeter as a potential predictor or co-predictor for landscape and habitat studies – especially in urban areas, where the negative effects of fragmentation might be much higher than in rural or more natural landscapes. Performing preliminary tests on comparisons between area and perimeter is highly recommended.     

How small cities affect the biodiversity of ground-dwelling mammals and the relevance of this knowledge in planning urban land expansion in terms of urban wildlife

Fundamental principles regarding urban biodiversity are based on studies conducted in large cities. However we cannot know whether the same phenomena occur in smaller cities or how small cities affect biodiversity. Small cities are an inherent element of urbanization and in the future, most global urban growth is expected to take place in small and medium-sized cities. Understanding the effects of small cities on biodiversity will be an important aspect in planning urban land expansion. Our study examined the effects of a small city on communities of small ground-dwelling mammals on 41 sites arranged in a four step gradient of urbanization. In 6700 trap-days, we caught 2333 individuals comprising 15 species. In the downtown area the same phenomena as those described for large cities were observed: a reduction in species richness and diversity, a decline in the abundance of urban sensitive species and an increase in synurbic species. However, in contrast to large city studies, green areas outside the downtown area did not differ from rural sites in small mammal population parameters. This phenomenon of relatively unchanged fauna outside the downtown area shows that small cities have the potential to maintain a high level of diversity of small ground-dwelling mammals if appropriate planning of further building expansion is implemented. More studies of small cities are needed to better assess their impact on biodiversity. This knowledge can then be applied in better planning for urban wildlife. Generalizations based solely on large city studies are inadequate and may lead to incomplete or inappropriate conservation strategies for small cities.     

Lentic and lotic odonate communities and the factors that influence them in urban versus rural landscapes

Habitat alteration via urbanization has very different effects on even closely related taxa. Most research investigating the ecological effects of urbanization has focused on birds or mammals, resulting in a relatively poor understanding of how the species richness and community composition of invertebrates may change. We quantified differences in species richness of adult odonates (dragonflies and damselflies) at lentic and lotic sites in urban and rural landscapes, and we examined environmental factors that might drive the differences in community composition that we observed. For lotic sites, species richness did not differ between urban versus rural sites for either dragonflies or damselflies. For lentic sites, urban and rural sites contained similar dragonfly species richness, but damselfly species richness was significantly lower at urban sites than at rural sites. Differences in lentic odonate community composition were associated with the amount of urban development within 150 m of each site, mean algal coverage, and distance to the urban center. At lotic sites, water temperature and distance to the urban center were correlated with differences in odonate community composition. The differing responses to urbanization observed in this study were probably a consequence of differences between lentic versus lotic ecosystems and between dragonflies versus damselflies in dispersal capability and habitat specificity. Given that different environmental factors affected these taxa differently in lentic and lotic sites, maintaining the highest level of odonate diversity possible across a landscape will require the use of different management practices for each ecosystem type.     

Urbanization and the abundance and diversity of Prairie bats

The effects of urbanization on biodiversity are generally considered to be negative, but the potential for landscape context to modulate these effects has not been adequately examined because most urban ecology research has been conducted in one biome: the temperate forest. This bias also applies to studies of the urban ecology of bats, whose diversity is correlated with habitat heterogeneity. We investigated the hypothesis that in the fairly flat, homogeneous Prairies, urbanization, by creating structurally complex islands, benefits bats by increasing access to the vertical landscape elements (buildings and trees) in which they roost. From 2006 to 2008, we surveyed bat assemblages in and around Calgary, Alberta, using mist nets to capture them and bat detectors to record their echolocation activity. Our data supported the prediction that urbanization increases the abundance of Prairie bats, but not the prediction that it increases their diversity. Instead, the urban bat assemblage was less diverse, and exhibited decreased species evenness compared to non-urban assemblages. Although Myotis lucifugus dominated bat assemblages throughout our study area, this was most evident in the city, and this species drove the increased urban abundance of bats. Ultimately, we reject our hypothesis and conclude that urbanization in the Prairies may create attractive habitat for one synanthropic bat, but is detrimental to others.     

Influence of urban landscape structure on bird fauna: a case study across seasons in the city of Valencia (Spain)

Studies on bird fauna of urban environments have had a long history, but the potential of studies mapping the distribution of birds in cities probably has not fully developed. The bird fauna of the municipality of Valencia (Spain) was studied to determine the influence of urbanization on bird species richness and abundance. Birds were censused during winter and the breeding season of years 1997–1998 in 197 squares measuring 49 ha each from a rural and an urbanized area. Across seasons the number of species decreased around 40% in the city compared with the rural landscape surrounding it. Such pattern could be attributed to the low number of farmland species capable to use the habitats inside the city, and the limited ability of urban parks in attracting woodland species. In the urban landscape, the influence of the dimensions and spatial arrangement of habitat patches was outweighed by the amount of each habitat per square. Bird richness and the abundance of most species were negatively related with the amount of built-up habitat per square and positively with the amount of urban parks, and of habitat diversity. Conversely, bird fauna was largely independent of mean park size per square especially during winter, indicating that at the landscape scale even small patches of habitat could play an ecological role. Conservation of urban bird diversity could benefit of two complementary strategies: (i) the protection of the surrounding rural landscape from urban development; (ii) habitat enhancement within the city. Particularly, a proper design and habitat management of urban parks could improve their suitability for urban bird fauna.     

Trends in bird species richness,abundance and biomass along a tropical urbanization gradient

Impacts of urbanization on biodiversity are commonly studied using urbanization gradients which provide a space-for-time substitution in estimating consequences of urban expansion. Rates of urbanization and human population growth are high in tropical countries of the developing world, which also hold most of the world’s biodiversity hot-spots, yet few studies have considered biodiversity trends along urban gradients in these regions. Bird communities across a gradient of nine sites in Uganda, from the city centre of Kampala to outlying rural locations, were studied over a six year period. These sites were ordered along an urbanization gradient using Principle Components Analysis based on habitat variables estimated at each site. Bird species richness showed a decrease from rural to urban sites, a trend especially evident in forest birds. There was no clear pattern in total abundance, total biomass or biomass per individual along the gradient. However, this latter result was heavily influenced by a colony of Marabou Storks at one site. When this species was omitted, there was evidence of a positive trend with urbanization, showing that as species richness decreased, the bird community was increasingly dominated by larger species with increasing urbanization, which were mainly scavengers able to exploit human refuse. These results provide further support for the negative impacts of urbanization on species richness, but also demonstrate trends in abundance and biomass are variable across different regions. In particular, the increasing dominance of larger species in urban areas may be relevant to certain geographic and/or socioeconomic contexts.     

Urbanization affects the trophic structure of arboreal arthropod communities

Urbanization is one of the most significant causes of habitat fragmentation on the planet, resulting in substantial losses of biodiversity and disruptions to ecological processes. We examined the effects of urbanization on the diversity and abundance of arboreal invertebrates in a dominant tree species (Angophora costata) in a highly urbanized landscape in Sydney, Australia, identifying the potential ecological consequences of shifts in diversity. We hypothesized that trophic structure would be influenced by landscape context with a greater richness and abundance of invertebrates in small remnants and edges. Canopy arthropods were sampled via beating from trees in 15 sites in three landscape contexts; five large patches of continuous vegetation, five edges of large patches and five small urban remnants. Trees in large patches supported fewer individuals compared to trees in small urban remnants and edge sites. The composition of assemblages and overall trophic structure also differed between edges and large patches, with a greater abundance of grazing insects in edges. No differences were detected between small urban remnants and edges, suggesting that observed differences might be attributed to an edge effect as opposed to an area effect per se. These changes in trophic structure, revealing a greater abundance of grazing herbivores and a reduced abundance of predators and parasitoids in edge sites, are consistent with work describing elevated levels of herbivory in edges of remnant vegetation. Future management of remnant urban vegetation and associated biodiversity requires not only an understanding of how trophic status influences the extent of responses by arboreal invertebrate communities, but also how these will affect ecosystem functioning.     

Patterns of pollinator turnover and increasing diversity associated with urban habitats

The role of urban expansion on bee diversity is poorly understood, but it may play an important role in restructuring pollinator diversity observed in rural regions at the urban perimeter. We studied bee communities in two habitats essential for pollinators (residential gardens and semi-natural areas) at 42 sites situated at the edge of greater Montreal, Canada. Bee species richness, abundance and functional diversity all increased with urbanization in both habitat types, but gardens and semi-natural areas supported distinct bee communities with unique responses to urbanization in terms of species turnover. Compared to semi-natural sites, residential gardens supported bees that foraged from a greater number but a lower proportion of available plant species. Bees did not discriminate between exotic and indigenous plant species in either gardens or semi-natural sites and were attracted to flowers in either habitat irrespective of their origins. Protecting semi-natural ruderal areas and providing residential garden habitats for pollinators are both effective means of promoting regional bee diversity in urbanizing regions.     

Urban and wildland herpetofauna communities and riparian microhabitats along the Salt River, Arizona

Metropolitan areas are continually expanding, resulting in increasing impacts on ecosystems. Worldwide, riverine floodplains are among the most endangered landscapes and are often the focus of restoration activities. Amphibians and reptiles have valuable ecological roles in ecosystems, and promoting their abundance and diversity when rehabilitating riparian systems can contribute to reestablishing degraded ecosystem functions. We evaluated the herpetofauna community by measuring abundance, richness, diversity, and species-habitat relations along three reaches (wildland, urban rehabilitated, and urban disturbed reaches) varying in degree of urbanization and rehabilitation along the Salt River in central Arizona. We performed visual surveys for herpetofauna and quantified riparian microhabitat along eight transects per reach. The wildland reach had the greatest herpetofauna species richness and diversity, and had similar abundance compared to the urban rehabilitated reach. The urban disturbed reach had the lowest herpetofauna abundance and species richness, and had a similar diversity compared to the urban rehabilitated reach. Principal Component Analysis reduced 21 microhabitat variables to five factors which described habitat differences among reaches. Vegetation structural complexity, vegetation species richness, densities of Prosopis (mesquite), Salix (willow), Populus (cottonwood), and animal burrow density had a positive correlation with at least one herpetofauna community parameter, and had a positive correlation with abundance of at least one lizard species. Rehabilitation activities positively influenced herpetofauna abundance and species richness; whereas, urbanization negatively influenced herpetofauna diversity. Based on herpetofauna-microhabitat associations, we recommend urban natural resource managers increase vegetation structural complexity and woody debris to improve herpetofauna habitat when rehabilitating degraded riparian systems.     

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.     

Urbanization impacts on the structure and function of forested wetlands

The exponential increase in population has fueled a significant demographic shift: 60% of the Earth's population will live in urban areas by 2030. While this population growth is significant in its magnitude, the ecological footprint of natural resource consumption and use required to sustain urban populations is even greater. The land use and cover changes accompanying urbanization (increasing human habitation coupled with resource consumption and extensive landscape modification) impacts natural ecosystems at multiple spatial scales. Because they generally occupy lower landscape positions and are linked to other ecosystems through hydrologic connections, the cascading effects of habitat alteration on watershed hydrology and nutrient cycling are particularly detrimental to wetland ecosystems. I reviewed literature relevant to these effects of urbanization on the structure and function of forested wetlands. Hydrologic changes caused by habitat fragmentation generally reduce species richness and abundance of plants, macroinvertebrates, amphibians, and birds with greater numbers of invasives and exotics. Reduction in soil saturation and lowered water tables result in greater nitrogen mineralization and nitrification in urban wetlands with higher probability of NO^– ₃ export from the watershed. Depressional forested wetlands in urban areas can function as important sinks for sediments, nutrients, and metals. As urban ecosystems become the predominant human condition, there is a critical need for data specific to urban forested wetlands in order to better understand the role of these ecosystems on the landscape.     

Urban areas may serve as habitat and corridors for dry-adapted,heat tolerant species; an example from ants

We collected ants from six urban and one forest land-use types in Raleigh, NC to examine the effects of urbanization on species richness and assemblage composition. Since urban areas are warmer (i.e., heat island effect) we also tested if cities were inhabited by species from warmer/drier environments. Species richness was lower in industrial areas relative to other urban and natural environments. There are two distinct ant assemblages; 1) areas with thick canopy cover, and 2) more disturbed open urban areas. Native ant assemblages in open environments have more southwestern (i.e., warmer/drier) distributions than forest assemblages. High native species richness suggests that urban environments may allow species to persist that are disappearing from natural habitat fragments. The subset of species adapted to warmer/drier environments indicates that urban areas may facilitate the movement of some species. This suggests that urban adapted ants may be particularly successful at tracking future climate change.     

Richness, composition and trophic niche of stingless bee assemblages in urban forest remnants

Few studies directly address the consequences of habitat fragmentation for pollinating insect communities, particularly for the neotropical key pollinator group of stingless bees. Most studies on bees have defined habitat fragments as remnant patches of floral hosts or forests, overlooking the nesting needs of bees. Their conclusion is that habitat fragmentation is broadly deleterious; however, there are contrasting results in the literature. Insightful studies on habitat fragmentation and bees should consider fragmentation, alteration and loss of nesting habitats—not just patches of forage plants –, as well as the permeability of the surrounding matrix to interpatch movement. Here we investigated the effects of fragmentation caused by urbanization on stingless bee species’ composition and richness, as well as the permeability of the surrounding matrix. We collected bees from flowering plants and recorded phytosociological variables of five forest remnants (ranging from 64 ha to 900 ha) in southeastern Brazil. Large fragments did not contain more species per unit area than smaller ones; in fact, we found more species in small fragments, most of which were generalist bees. The presence of more habitat generalist stingless bee species was also correlated to the structure of vegetation in these fragments. In conclusion, the quality of the habitat within a fragment (structure of vegetation) as well as the quality of the matrix has a direct relation to the bee species composition. This can be seen in the direct relationship between structural diversity of the environment and age of the fragments. The matrix that holds the most recent fragments, probably due to the sprawl of the city, is more heterogeneous than the one with the oldest fragments. The most heterogeneous matrices have a certain balance between the trees, buildings and bare soil or herbaceous vegetation coverage, making the array less impermeable to bees.     

Diversity,abundance, and species composition of ants in urban green spaces

Urbanization threatens biodiversity, yet the number and scope of studies on urban arthropod biodiversity are relatively limited. We sampled ant communities in three urban habitats (forest remnants, community gardens, vacant lots) in Detroit and Toledo, USA, to compare species richness, abundance, and species composition. We measured 24 site characteristics to examine relationships between richness and composition and habitat patch size, vegetation, and urban features. Ant richness was higher in forests (26) than in gardens (14) and intermediate in vacant lots (20). Ant richness in gardens and vacant lots negatively correlated with abundance of an exotic ant species (Tetramorium caespitum); thus this ant may affect native ant richness in urban habitats. Ant composition differed with habitat type, and abundance was lowest in forests. Site characteristics varied with habitat type: forests were larger, had more woody plants, higher woody plant richness, more branches, and leaf litter whereas lots and gardens had more concrete and buildings. Vacant lots had taller herbaceous vegetation, and gardens had higher forb richness, density, and more bare ground. Differences in vegetation did not correlate with ant richness, but several vegetation factors (e.g. patch size, number and size of trees, leaf litter, and amount of concrete and buildings) correlated with differences in ant species composition. Additional factors relating to soil, nests, or microclimatic factors may also be important for urban ant communities. Implications for biodiversity conservation in urban ecosystems are discussed.     

Reduction by half: the impact on bees of 34 years of urbanization

Urban ecosystems are growing rapidly and urbanization is an important cause of the loss of biodiversity. Bees are declining in abundance worldwide, including urban areas, and this decline is alarming because of their global importance as plant pollinators. Here we examine that decline by comparing a bee assemblage sampled in the 1980s and again in 2015, in an urban area of the city of Curitiba. Both studies sampled assemblages with hand-nets every two weeks during one year of study. Bee species richness has declined by 45% (112 species then, 63 today). Two species that have disappeared, Gaesischia fulgurans (Holmberg, 1903) and Thectochlora basiatra (Strand, 1910), have also disappeared elsewhere in the city. Also, relative abundances of species have changed, notably with the increase of social bees. Large bees that nest in cavities have also increased relative to small bees that nest in the ground. These findings are similar with previous reports indicating the sensibility of bees to urbanization. The increase in paved areas, in urban population and in exotic plants are all probably responsible for the sharp decline in bee diversity and abundance.     

Determinants of species richness within and across taxonomic groups in urban green spaces

Urban green spaces provide habitat for numerous plant and animal species. However, currently we have little knowledge on which determinants drive the species richness within and across taxonomic groups. In this paper we investigate the determinants of total, native, and endangered species richness for vascular plants, birds, and mammals within and across taxonomic groups. We examined a stratified random sample of 32 urban green spaces in Hannover, Germany. Species inventories for plants and birds were generated on the basis of line transect surveys. Mammals were surveyed by means of point counts using camera traps. Using a principal component analysis and multiple regression models, we tested 10 explanatory variables for species-area effects, distance effects, and the effects of habitat structure of green spaces on species richness. When analyzing single explanatory variables, we determined that the species richness of all groups was significantly positively correlated to patch area, number of habitat types, and a short distance to the nearest green space. Testing combined effects of variables showed that patch area in combination with habitat heterogeneity was most important for plants (total, native, and endangered), birds (total and native), and overall species richness. This emphasizes the importance of the species-area effect and the effects of habitat structure on species richness in urban green spaces. We conclude that, in the context of urban planning, it is important to conserve large green spaces that include a high diversity of habitats to maintain high species richness.     

Local and landscape drivers of arthropod abundance,richness, and trophic composition in urban habitats

Urban green spaces, such as forest fragments, vacant lots, and community gardens, are increasingly highlighted as biodiversity refuges and are of growing interest to conservation. At the same time, the burgeoning urban garden movement partially seeks to ameliorate problems of food security. Arthropods link these two issues (conservation and food security) given their abundance, diversity, and role as providers of ecosystem services like pollination and pest control. Many previous studies of urban arthropods focused on a single taxon (e.g. order or family), and examined either local habitat drivers or effects of landscape characteristics. In contrast, we examined both local and landscape drivers of community patterns, and examined differences in abundance, richness, and trophic structure of arthropod communities in urban forest fragments, vacant lots, and community gardens. We sampled ground-foraging arthropods, collected data on 24 local habitat features (e.g., vegetation, ground cover, concrete), and examined land-cover types within 2 km of 12 study sites in Toledo, Ohio. We found that abundance and richness of urban arthropods differed by habitat type and that richness of ants and spiders, in particular, varied among lots, gardens, and forests. Several local and landscape factors correlated with changes in abundance, richness, and trophic composition of arthropods, and different factors were important for specific arthropod groups. Overwhelmingly, local factors were the predominant (80 % of interactions) driver of arthropods in this urban environment. These results indicate that park managers and gardeners alike may be able to manage forests and gardens to promote biodiversity of desired organisms and potentially improve ecosystem services within the urban landscape.