Urbanization, Biodiversity, and Conservation

2002/01/01 by Michael L. McKinney, MICHAEL L. McKINNEY · 3,636 citations
Environmental Science · #Biodiversity #Biodiversity conservation #Biology #Conservation, Biodiversity, and Resource Management #Ecology #Geography #Land Use and Ecosystem Services #Urbanization #Wildlife-Road Interactions and Conservation

paper · pdf · doi:10.1641/0006-3568(2002)052[0883:ubac]2.0.co;2

published in BioScience 52(10), 883 (Oxford University Press)

openalex publication_date 2002/01/01 · openalex created_date 2025/10/10 · openalex updated_date 2026/08/05

Abstract

Among the many human activities that cause habitat loss (Czech et al. 2000), urban development produces some of the greatest local extinction rates and frequently eliminates the large majority of native species (Vale and Vale 1976, Luniak 1994, Kowarik 1995, Marzluff 2001). Also, urbanization is often more lasting than other types of habitat loss. Throughout much of New England, for example, ecological succession is restoring forest habitat lost from farming and logging, whereas most urbanized areas in that region not only persist but continue to expand and threaten other local ecosystems (Stein et al. 2000). Another great conservation challenge of urban growth is that it replaces the native species that are lost with widespread “weedy” nonnative species. This replacement constitutes the process of biotic homogenization that threatens to reduce the biological uniqueness of local ecosystems (Blair 2001). Urban-gradient studies show that, for many taxa, for example, plants (Kowarik 1995) and birds and butterflies (Blair and Launer 1997), the number of nonnative species increases toward centers of urbanization, while the number of native species decreases. The final conservation challenge of sprawl is its current and growing geographical extent (Benfield et al. 1999). A review by Czech and colleagues (2000) finds that urbanization endangers more species and is more geographically ubiquitous in the mainland United States than any other human activity. Species threatened by urbanization also tend to be threatened by agriculture, recreation, roads, and many other human impacts, emphasizing the uniquely far-reaching transformations that accompany urban sprawl. About 50% of the US population lives in the suburbs, with another 30% living in cities (USCB 2001). Over 5% of the total surface area of the United States is covered by urban and other built-up areas (USCB 2001). This is more land than is covered by the combined total of national and state parks and areas preserved by the Nature Conservancy. More ominously, the growth rate of urban land use is accelerating faster than land preserved as parks or conservation areas by the Conservancy (figure 1). Much of this growth is from the spread of suburban housing. It is estimated, for example, that residential yards occupy 135,000 acres in the state of Missouri (MDC 2002). This residential landscape represents nearly 1% of the total area of Missouri and is nearly three times the area occupied by Missouri state parks. Here I review the growing literature that documents how urban (and suburban) expansion harms native ecosystems. This knowledge can aid conservation efforts in two major ways. One is through the use of ecological principles—such as preserving remnant natural habitat and restoring modified habitats to promote native species conservation—to reduce the impacts of urbanization on native ecosystems. Rare and endangered species sometimes occur in urbanized habitats (Kendle and Forbes 1997, Godefroid 2001) and thus could be conserved there. Managing the large amount of residential vegetation (1% of the state area, as noted above) in ways that promote native plants and animals could also make a significant contribution to conservation. A second way in which the study of urban ecology can serve conservation is by helping to develop a more ecologically informed public. Providing a well-informed public could be the most important application of urban ecology, as a means of promoting effective conservation of native species (Kendle and Forbes 1997). Because 80% of the American public lives in or near urban areas, there are many opportunities for creating an informed public that can wield enormous economic and political pressure to promote conservation policies. People who live in urban environments often have a great appreciation of many urban species, such as birds (Clergeau et al. 2001). Indeed, residents of suburban and urban areas tend to place a much higher value on species conservation than those living in rural areas (Kellert 1996). This is reflected in voting behavior: Legislators from highly urbanized states and districts tend to be more supportive of strengthening the Endangered Species Act (Mehmood and Zhang 2001). Unfortunately, these conservation opportunities are hindered by the very poor ecological knowledge of typical American urbanites. A survey of Texas high school students, for example, showed that 60% of the students misidentified the opossum as a rodent and that ecological understanding of human effects on biota was even poorer; only 2% of the students knew that raccoons tend to benefit from many human activities (Adams et al. 1987). Urban-to-rural gradient studies examine changes in plants and animals along a transect from the inner city to surrounding, less-altered ecosystems; they also show what happens to surrounding native ecosystems as urban sprawl expands. General patterns that emerge from these studies are described below. Physical changes along the gradient strongly influence available habitat for native species. A number of reviews (Sukopp and Werner 1982, Medley et al. 1995, Pickett et al. 2001) show increases in these physical changes, as one moves toward the urban core, in such metrics as human population density, road density, air and soil pollution, average ambient temperature (“heat island” effect), average annual rainfall, soil compaction, soil alkalinity, and other indicators of anthropogenic disturbance. The percentage of area that is impervious surface (pavement, asphalt, buildings) ranges from well over 50% at the urban core to less than 20% at the fringe of urban expansion (figure 2). In addition, the amount of subsidized energy and matter imported for use by humans and available to other species increases toward the urban center (Collins et al. 2000, Pickett et al. 2001). These physical changes produce a gradient of natural habitat loss that steepens from rural areas toward the urban center. As habitat is lost, it becomes increasingly fragmented into more numerous but smaller remnant patches (Medley et al. 1995, Collins et al. 2000). The lost natural habitat is then replaced by four types of altered habitat that become progressively more common toward the urban core. The four types of replacement habitat are listed below, in order of increasing habitability to most native species and decreasing proportion of coverage toward the urban core. The latter three types are based on Whitney (1985). Built habitat: buildings and sealed surfaces, such as roads Managed vegetation: residential, commercial, and other regularly maintained green spaces Ruderal vegetation: empty lots, abandoned farmland, and other green space that is cleared but not managed Natural remnant vegetation: remaining islands of original vegetation (usually subject to substantial nonnative plant invasion) It is probably intuitive to even the most casual observer that the increasing fragmentation of natural habitat by human disturbances in the direction toward urban centers will tend to reduce species richness (number of species) in that direction. There are, however, many variables that can affect the rate and consistency of species loss along the gradient, so empirical studies are crucial in measuring urban impacts. Many studies document that the lowest species diversities along the urban–rural gradient occur in the intensively “built” environments of the urban core. This has been shown for many taxa, including plants (Kowarik 1995), birds and butterflies (Blair 2001), many insects (Denys and Schmidt 1998, McIntyre 2000), and mammals (Mackin-Rogalska et al. 1988). In all these taxa, the number of species at the urban core is reduced to less than half of that found in the rural, more natural areas at the opposite end of the gradient (figure 2). Blair (2001), for example, found just 7 summer resident bird species in the central business district of Palo Alto, California, compared with 21 species that inhabited a natural area (preserve) outside the city limits. Similar reductions were found for birds and butterflies in other cities, as shown by Blair's (2001), and especially by Marzluff's (2001), comprehensive compilation of studies on urbanization impacts on birds. Much of the reduction in richness is obviously caused by the loss of vegetation. The number of species of animal taxa, such as birds (Shugart et al. 1975) and insects (Majer 1997), tends to correlate with the number of plants in an area. Also, area covered by vegetation is a good predictor of species numbers for birds (Goldstein et al. 1986); mammals, amphibians, and reptiles (Dickman 1987); and insects (McIntyre 2000). As over 80% of most central urban areas is covered by pavement and buildings (Sukopp and Werner 1982, Blair and Launer 1997), less than 20%, therefore, remains as vegetated area. Furthermore, the remaining vegetated habitat often contains low plant diversity as a result of erosion, trampling, pollution, invasion or cultivation of a few nonnative species, and many other human disturbances. Also, mowing, pruning, and other common landscaping practices further reduce the volume of the remaining vegetation (Gilbert 1989, Adams 1994). Some studies indicate that species richness tends to be higher in areas with low to moderate levels of human development (such as outlying suburban developments) than in more natural rural areas such as preserves. This suburban peak in species numbers is evident in many taxa, such as mammals (Racey and Euler 1982), birds and butterflies (Blair 2001), bumblebees (Pawlikowski and Pokorniecka 1990), ants (Nuhn and Wright 1979), lizards (Germaine and Wakeling 2000), and plants (Kowarik 1995). An explanation often suggested for this suburban peak (e.g., Blair and Launer 1997, Germaine and Wakeling 2000, Blair 2001) is the intermediate disturbance hypothesis. The initial human impacts of suburban sprawl are sometimes relatively mild, with only a few housing subdivisions in a matrix of largely natural or agricultural habitat. This promotes environmental heterogeneity, because different habitats occur alongside one another. Such habitat diversity is enhanced by the fact that individual homeowners often make individualistic choices in the plants that they cultivate (Henderson et al. 1998). In addition to providing spatial heterogeneity, these anthropogenic habitats are typically very productive (Falk 1976), being highly subsidized in scarce resources, ranging from water to nutrients (e.g., fertilizers). Cultivated plants include many ornamentals that often bear fruits and seeds that are utilized by animals, especially birds and bats (Munyenyembe et al. 1989, Adams 1994). Some animals have adapted to the direct consumption of human resources (Adams 1994) that are provided accidentally (garbage) or intentionally (bird food). In contrast to the above, other studies show that suburban areas have reduced species diversity compared to less-altered rural habitats (figure 2). For example, Marzluff's (2001) compilation of 51 bird studies found that 31 of the studies (61%) showed lower species richness in suburban and other areas of human settlement, compared with more natural rural areas. The remaining 20 studies reported either an increase or no change in diversity with increasing human settlement. The 51 studies covered a wide range of geographic and natural settings, so it is difficult to identify which variables determine whether a rise or fall of species richness occurs with increasing settlement and suburban development. Teasing apart these variables, such as the role of the natural setting, is clearly a priority for further work on urban–rural gradients. Bell (1986), for example, has suggested that urbanization in a tropical rain forest may have different effects on local species richness than urbanization in other natural settings, because rain forest birds have exceptional difficulty adapting to human settlements. Areas of active development tend to have low biodiversity because of the devastating impact on native species of most residential and commercial development methods. Before construction of most residential and commercial buildings, it is common for developers to remove most vegetation and even topsoil (Sharpe et al. 1986). This reduces construction costs by allowing equipment ready access to the construction site. A study of the of natural vegetation urban development in found that only of the original vegetation was not (Sharpe et al. 1986). The loss of native vegetation (and total vegetated has a impact on native animal species richness in the of housing construction to in (Vale and Vale and 1994). construction is some of the area is which it as habitat for nearly all species. In Palo Alto, California, for example, of the area of residential is covered by pavement (Blair and Launer another 20% of the area is covered with housing. the remaining much is with (usually and and 2000). conservation can and (figure 2). The most effective (and in the is to as much remnant natural habitat as Many studies how native species richness in a remnant habitat increases with the area of that habitat. This is for many taxa, including birds mammals (Dickman and plants 1995). One way to in housing is to vegetation. A number of such as The and 2000), have the of vegetation for conservation this of construction is by most residential to costs of more vegetation is less in the et al. and is by many homeowners and 2000). A major influence on natural is the or the of that are often in a highly matrix that also as a of nonnative species. A major challenge is that remnant habitats are to by nonnative species of plants and animals such as and 2001). These nonnative and can reduce the of the remnant habitat to native species, especially birds. In the of population these become population that are to of the native species. can also on restoring native species in managed and In natural biotic succession increases the number of plant and animal species a disturbance et al. 2000). This is also of and managed habitats that for succession to studies have how succession increases species diversity in and managed for example, plant diversity in urban 1979), diversity in (Majer 1997), and bird species richness in residential (Vale and Vale 1976, et al. 1989, Luniak 1994). As a residential areas (usually the urban tend to have higher species richness than (e.g., et al. The studies show that the rate of species succession is very and is the few and especially the from increasing total ecological succession also often reduces the diversity of native species in an area et al. 2000), many of which on disturbance to 1997). Another to increase native biodiversity in managed habitats is to cultivate a of plant species. with native plant species may benefit not only native plant but also native animal For example, native bird species richness in (Munyenyembe et al. and and tends to correlate with the volume and species diversity of native vegetation. the percentage of native species in a has been found to correlate with the percentage of native plant species et al. 1998). with native plants can benefit many local native bird species 1997). Species in to to the often physical changes along the urban–rural gradient (Gilbert 1989, Adams 1994). there are probably many ways to these changes in species many bird (e.g., et al. and Blair 2001) and (e.g., and studies have that species along the gradient can be for into three to human Blair's (2001) these are and (figure 2). birds are the for work on urban–rural these three have also been for work on butterflies (Blair and Launer and lizards (Germaine and Wakeling 2000). These show that, even in highly modified species are in ways that in of these has a of ecological that the impacts of urban sprawl on native species. One of the most important that the three is the extent to which species on resources to in an area 2001). As subsidized resources increase toward the urban core, there is a increase in species that are that are on human are to but are in that they also use natural tend to only on natural resources 2001). Because mammals, to a plants are the along urban–rural they will be the major are species that are very to human and habitat disturbances. The species to in the of humans are large mammals, especially because they are relatively and have low and were the to settlement and urban include species adapted to the of such as and many birds that are very to the of humans and et al. and 1982, and Adams 1994). species that are very to human activities include and plants (Stein et al. 2000), the loss of which is to to and for agricultural and settlement are often found in the matrix of human land that occur in suburban For species are common in managed suburban such as residential yards and commercial as well as habitats (e.g., These plants include species by humans (e.g., and as well as species that are common in managed and suburban The most common species are (e.g., and (e.g., that on suburban are often as or et al. This is an landscape for most (Henderson et al. 1998). Among animals, urban typically include many species often to as which are adapted to forest and surrounding areas et al. Adams 1994). These animals many including such as plants and The great of such subsidized is one these animal urban often an and that is much than in natural areas (Adams 1994, Marzluff 2001). Another is that natural of these animals are by human activities and Blair 1999). For urban include a high proportion of These include and such as the American and many such as and such as et al. and 1982, and Adams 1994, 2001). of these three to be to different of human impacts. The highly productive and plant a of and plant (Falk for while bird and many plants that produce seeds (Adams 1994). of the many areas, including over suburban habitats and the high of many especially those that are to and are also common urban 2001). As most mammals the high of by in suburban environments different urban are to from human as well as of provided by humans and and and 1987). One of finds through and are of to human in these animals much from the of suburban including growing and (Falk Another of species that forest (e.g., in and for (Dickman 1987). These species typically for in surrounding areas. Some are raccoons and that in and other resources provided by are such as and that a wide of As with of large addition to subsidized to very high population of urban species and 1999). often (e.g., and Werner 1982, et al. 2001), are very on human The of urban is not the amount or types of vegetation and and et al. 2001). The of such as the of and with to enormous population and Adams 1994). probably the most of the (Blair 2001). urban which are largely of species from urban are of a very of the these are well adapted to modified urban environments humans the (Adams 1994, Marzluff 2001). environments typically have more in common with other cities than with natural ecosystems (Sukopp and Werner 1982), so urban are often not native to a region (Adams 1994, Kowarik 1995, Blair 2001), but tend to from city to and the are found in all cities in (Mackin-Rogalska et al. and (Adams 1994). This is also for urban plants Among urban tend to be species that can high levels of especially and reviews in and Werner 1982, Whitney Kowarik 1995). include that abandoned and commercial and plants that can in and that are typical of plants include to high levels of air and and and urban are often species adapted to areas and are to the of very urbanized areas and Adams 1994). include the and Another of of species that are to human include the and urban tend to be or and Adams 1994). urban in human and the in or near they are (Adams 1994) and include such species as the or and including a of species. nonnative species toward the Many studies have found that the number (and of nonnative species tends to increase along the urban–rural gradient, toward the urban center. In the proportion of species that is nonnative from less than a few in rural areas to over 50% at the urban core. These to plants in the United States and (Kowarik 1995) and birds in the United States (Blair 2001). The population of nonnative mammals (Mackin-Rogalska et al. and birds tends to increase the they are to the urban core. The increase in nonnative species toward the urban core a number of human One is that higher human population the urban core produce increasing of nonnative species, for example, the cultivation of nonnative plants (Mackin-Rogalska et al. Kowarik 1995). Another cause is the increasing amount of habitat toward the urban core, which opportunities for nonnative species of plants (Kowarik 1995, and animals (Adams 1994, Marzluff 2001) that can the In Nature and Forbes that, as highly urbanized areas are occupied by species that in the of there will be relatively few native species of conservation in areas of high human population review some however, of species of insects and plants found in highly urbanized habitat conservation and could be for that such species. most species in urbanized areas are found that have development 2001). species most occur include city vegetated areas and other public that are from development (Gilbert 1989, and Forbes 1997). from the conservation of native species, knowledge of the species of urban biodiversity can be very as an to the natural An enhanced appreciation of by the 80% of the American public that lives in this could promote more effective political and economic of such knowledge include of the public in the natural of local species and with nonnative species (Kendle and Forbes 1997). is a growing cause of many environmental (Benfield et al. 1999). The impact of urbanization is in the growing literature on the urban–rural These studies show changes in species richness and species along the Species richness of many often along the gradient, with the lowest richness to be found in the urban core. ways to biodiversity as cities expand and natural habitat. Such efforts most on preserving as much remnant natural habitat as as to most current land development which remove most natural vegetation land development has native animal biodiversity can be by with a diversity of native plant species. this habitat from disturbance to ecological succession will not only plant and animal diversity but also tend to reduce the diversity of nonnative species. Unfortunately, most current landscaping tends to with nonnative plant species in spatial (Henderson et al. 1998, and and succession through the of those ecosystems great and Forbes 1997). Species also changes along the urban–rural is that nonnative species become more common toward the urban core. include native species such as large and birds that in the initial of suburban is to large of native habitat and reduce human of species. mammals and birds that are adapted to forest and areas, in suburban especially subdivisions ecological succession has and are very important for biodiversity because half of the American public lives in a suburban (USCB 2001). biodiversity be most effective on these suburban and species to promote an understanding of such as ecological succession and the role of native plants in promoting native animal Because of its enormous and political a more ecologically informed suburban population could the for conservation of native species in all ecosystems. of land covered in the lower by are from of the United States for the for Nature Conservancy which is from and colleagues This is a very and of changes in surface area, species and as from a number of in the conservation with to urban sprawl are shown at the

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