Urban Ecosystems, 1997, 1, 229–246

Urban tree cover: an ecological perspective

WAYNE C. ZIPPERER,* SUSAN M. SISINNI AND RICHARD V. POUYAT
USDA Forest Service, 1100 Irving Avenue, Syracuse, NY 13210, USA

TIMOTHY W. FORESMAN
University of Maryland Baltimore County, Department of Geography, 5401 Wilkens Avenue, Baltimore, MD 21228, USA

Analysis of urban tree cover is generally limited to inventories of tree structure and composition on public lands. This
approach provided valuable information for resource management. However, it does not account for all tree cover
within an urban landscape, thus providing insufficient information on ecological patterns and processes. We propose
evaluating tree cover for an entire urban area that is based on patch dynamics. Treed patches are classified by their
origin, structure, and management intensity. A patch approach enables ecologists to evaluate ecological patterns and
processes for the entire urban landscape and to examine how social patterns influence these ecological patterns and
processes.
Keywords: tree cover; urban-to-rural gradient; patch dynamics; urban forest; ecological patterns and processes

Introduction
Urbanization modifies the extent and type of vegetation on a landscape through losses from deforestation
and fragmentation and gains from reforestation and afforestation (Godron and Forman, 1983; Sharpe et
al., 1986; Zipperer et al., 1990). With deforestation and fragmentation there are reductions of native flora
and fauna including keystone species (Murphy, 1988; Peter and Lovejoy, 1990), increases in non-native
species, increases in stream sedimentation and water pollution (Douglas, 1990; Roger, 1994), and
changes in air temperature, evapotranspiration, and albedo (Jager and Barry, 1990; Grimmond et al.,
1994; Heisler et al., 1994). Despite these impacts, ecologists know relatively little about ecological
processes of urban systems (Stearns and Montag, 1974; McDonnell and Pickett, 1990). This is borne out
further by a recent search for ecological papers highlighting ecological patterns and processes studies
conducted in the New York City metropolitan region (Flores et al., in review). The search included 2470
citations and identified only 145 citations on ecological processes.
Our paper argues for a new approach to studying urban ecological patterns and processes: identifica-
tion and evaluation of vegetation patterns through patch analysis. Little new data on vegetation are
furnished; rather, existing research is reviewed to provide a direction for future studies of urban veg-
etation. Assessments of community structure, vegetation dynamics, and ecosystem processes within
urban landscapes are discussed to support our approach.

Patch analysis: an alternative approach

Urban vegetation can be defined in two ways. First, it is defined as an assemblage of plant material above,
on, and below the ground surface within an urban landscape (Sanders, 1984). This definition includes
measurements of species structure, composition, and age; forest health; density; biomass; and leaf area.

* To whom correspondence should be addressed.

1083-8155 © 1997 Chapman & Hall
230 Zipperer et al.
The second definition focuses on process and identifies those plant assemblages that are regularly
subjected to urban influences as urban vegetation (Sanders, 1984; McDonnell and Pickett, 1990). The
process definition includes structural components and processes within urban areas as well as areas
adjacent to or neighboring urban landscapes. Thus, a gradient of urban influences on vegetation exists
and can be evaluated from the urban core to rural areas (McDonnell and Pickett, 1990).
Analysis of vegetation in urban and urbanizing landscapes has been conducted at broad (Schmid, 1975;
Dorney, 1977; Brady et al., 1979; Rowntree, 1984; Sukopp and Weiler, 1988; Nowak, 1994b) and fine
(Lefkowitz and Greller, 1973; Stalter, 1981; Profous and Loeb, 1984; Rudnicky and McDonnell, 1989)
scales. At each scale, sampling is done by political or management units rather than ecological units.
Broad-scale analyses focus on either the land use or city-wide level and include descriptions and
measurements of vegetation structure that relate to species composition, percent canopy cover, tree
density, diameter distribution, and leaf surface area. Although this approach often assumes a homoge-
neity across a typology class (e.g. land use), it provides useful insights into tree management (e.g. Welch,
1994) or assessments of urban forest structure to improve air quality (Nowak, 1994a). Fine scale studies
tend to focus on natural areas or management units with historic significance. These studies often include
detailed analyses of site histories (Loeb, 1992, 1993) and community composition and structure (Rud-
nicky and McDonnell, 1989), and also providing a basis for site management (Sisinni and Anderson,
1993; Sisinni and Emmerich, 1995).
These approaches are well suited to meet their individual objectives; however, because of their
resolution (grain) and extent (Allen and Hoekstra, 1991), they may be insufficient to assess ecological
patterns or processes across an urban landscape. For example, understanding and managing hydrologic
cycles in urban and urbanizing landscapes require not only knowing the percent of canopy cover, often
the only vegetation parameter in hydrological models, but also specific information on location of tree
cover, site orientation, and vertical structure of vegetation for the entire watershed (Band et al., 1993;
Band and Moor, 1995; Neville, 1996). Similarly, location, distance between tree cover habitats, habitat
type, shape and size, and adjacent land cover are critical components for conservation strategies (Pickett
et al., 1992; Pickett and Parker, 1994; Forman, 1995). Since the collection of detailed data requires
additional time, energy, and funds, study objectives should determine the type of data needs.
An alternative approach is needed to investigate ecological patterns and processes within urban and
urbanizing landscapes. This approach should link ecological patterns and processes at broad spatial and
temporal scales and provide sufficient detailed information on community dynamics for fine scale site
management and ecological analyses (Turner and Gardner, 1990; Allen and Hoekstra, 1992; Yaro and
Hiss, 1996). In addition, because humans ultimately determine vegetation patterns, the approach should
link ecological and social patterns and processes.
A unit of measurement universal to all landscapes must be defined that effectively links, spatially and
temporally, both ecological and social patterns and processes. We recommend that that unit be a patch.
A patch is defined ‘‘as a relatively homogeneous area that differs from its surroundings’’ (Forman, 1995).
Aerial photographs of urban landscapes reveal natural and anthropogenic patches. Because anthropogenic
patches dominate the landscape, they are considered to be the background matrix. A background matrix
is ‘‘characterized by extensive cover, high connectivity and has major control of landscape dynamics’’
(Forman, 1995). Vegetation patches exist within this matrix and include grass, shrub, and tree cover
(treed patches). The term ‘‘treed’’ is used instead of forest because within urban landscapes tree cover
occurs in patches with a woody understory and developed forest floor (e.g. natural areas) and with a grass
or herbaceous understory (e.g. parks and yards; Fig. 1; Hobbs, 1988; Zipperer and Zipperer, 1992). A
patch also can be defined by other attributes than just by physical structure. For example, census data are
used to display social patterns spatially, and disparate areas can be viewed and classified (Grove and
Burch, 1997).
Urban tree cover 231

Figure 1. Treed patches represented at different scales. At the broad scale (aerial and remote sensing data), treed
patches are presented as distinct areas of tree cover. At the fine scale, treed patches are represented by individual trees
and understory structure. Information gathered at the fine scale can be aggregated to give broad scale representation;
however, broad scale information can not be deaggregated to give fine scale information. Patches A and B represent
different types of treed patch.

The patch, a fundamental unit of measurement for landscape analyses (Forman and Godron, 1981;
Turner, 1989; Forman, 1995), is selected because it: (1) implies a relatively discrete spatial pattern; (2)
implies a relationship of interactions and exchanges of one patch to another and to the surrounding
matrix; and (3) allows for a unit of management (Risser et al., 1984; White and Pickett, 1985; Forman
and Godron, 1986; Turner, 1989; Forman, 1995). Patches, however, are not spatially or temporally static.
Disturbances and biotic processes cause changes, thus creating a dynamic-heterogeneous mosaic of
patches (Pickett and Thompson, 1978; White and Pickett, 1985). Patch dynamics (i.e. how patches
change spatially, structurally, compositionally, and functionally; White and Pickett, 1985) influence
internal processes of the patch as well as interactions between itself and the adjacent matrix (Forman and
Godron, 1986). Understanding patch dynamics within urban and urbanizing landscapes will yield insights
into factors influencing how these landscapes function (McDonnell and Pickett, 1990).
For illustrative purposes, we focus only on treed patches. Similar characterizations can be made for
other patches of urban vegetation. When considering ecological processes, important patch attributes
include origin, configuration, vegetation structure, and management intensity. We classify patches into
three origin types: remnant, emergent, and planted. A remnant patch is defined as an area that was not
cleared during site development and existed before development. Because of their age, these patches
often are dominated by large diameter trees. Management varies from intensive to none. An emergent
patch resembles a remnant patch, but instead has regrown on a site previously cleared for development
(e.g. vacant lots and sites, fence rows, property boundaries). Structurally, these patches are characterized
by pioneer species and small diameter trees (<15 cm dbh; Dorney et al., 1984; Nowak, 1994b). Man-
agement varies from minimal to none. A planted patch is created by people planting trees (e.g. street and
232 Zipperer et al.
yard trees). Structurally, these patches are characterized by a range of diameters with large diameter trees
often located on older urban sites and smaller diameter trees on newly developed sites (cf. Nowak,
1994b). Three patch types are chosen to represent the array of possible structure, origin and management
combinations to examine ecological patterns and processes: remnant patches with large trees and un-
managed (i.e. having an understory), planted patches with large trees and intensively managed (i.e.
having grass cover), and emergent patches with smaller diameter trees and no management (Fig. 2). In
addition, unless stated, our discussion focuses on urban tree cover in the eastern United States.

Ecological pattern
Morphology, environment, management
Three primary factors influence the occurrence of treed patches: urban morphology, the environment, and
vegetation management decisions (Stearns and Montag, 1974; Dorney et al., 1984; Sanders, 1984;
Richards et al., 1984; Nowak et al., 1996). Urban morphology, the built structure and the pattern of
development, determines the availability and configuration of growing space (Sanders, 1984). From a
landscape perspective, urban morphology is the background matrix, and the urban vegetation composes
the patches and corridors within the matrix (cf. Forman, 1995). In general, moving along an urban-to-
rural gradient, the availability of growing spaces tends to increase as development density decreases
(Whitney and Adams, 1980; Sanders, 1984; Nowak, 1994c).
Both management decisions and the natural environment affect the establishment, growth, maturity,
reproduction, and senescence of a tree. This effect occurs across both broad-regional and fine-site scales.
Regions can be characterized by climate (moisture and thermal) and landform patterns to predict species
occurrence (Küchler, 1969; Bailey, 1983; Sanders and Rowntree, 1983; Nowak et al., 1996). An analysis
of canopy cover by ecoregions in the United States shows that forested areas have the highest average
canopy cover (31%) with the highest percent covers occurring in the Southeast and the Pacific Northwest
(Nowak et al., 1996). In contrast, arid and desert areas have the lowest average canopy cover (10%;
Nowak et al., 1996).

Figure 2. Patch origin by management intensity to identify their probable locations within an urban landscape.
Definitions of patch origins are given in the text.
Urban tree cover 233
At the finer site scale, physical parameters (e.g. soil moisture and type, and temperature) play a critical
role in species occurrence. In addition, species characteristics such as seed production, dispersal, estab-
lishment, growth, maturity, reproduction, and historic presence in remnant patches also play a significant
role in species occurrence (Pickett et al., 1987a; Pickett and McDonnell, 1989).
Despite urban morphology and environmental factors, humans ultimately decide tree cover patterns in
urban landscapes through active and passive management decisions (Fig. 2). Humans decide which
vegetation remains (remnant), when and what sites are cleared for development, which species are
planted and where (planted), and which areas are allowed to develop naturally (emergent). Whereas
management of urban vegetation and its consequent benefits have been the traditional focus of urban
forestry (Grey and Deneke, 1986; Miller, 1988), a call for a systems approach to inventory and manage
the resource has occurred for several decades (Stearns and Montag, 1974; Dorney et al., 1984; Richards
et al., 1984). Only recently, however, has there been a shift from single tree management to an ecosystem
or landscape approach to account for interactions within and among patches (Zipperer et al., 1995).

Forest development
Before we discuss community dynamics for treed patches, a general discussion of forest development
following a disturbance is provided for comparison. Forest development has four phases: establishment,
thinning, transition, and steady-state (Bormann and Likens, 1979; Oliver, 1980; Oliver and Larson,
1990). The establishment phase is characterized by low competition and numerous tree seedlings. The
thinning phase represents the reduction of stem density as competition between individuals increases. The
transition phase is characterized by the formation of canopy openings and the development of an
understory. The final phase, steady-state, represents a balance between small and large diameter trees,
and between closed-canopy and gap environments (Bormann and Likens, 1979; Oliver, 1980; Oliver and
Larson, 1990; Peet, 1992).
Forest development of urban-treed patches depends on patch origin and management. For planted
vegetation, the establishment phase consists of clearing or near clearing of all vegetation and subse-
quently planting ‘‘new’’ vegetation (Sharpe et al., 1986; Zipperer and Zipperer, 1992). Remnant veg-
etation ranges from isolated individuals to large natural areas. The establishment phase for emergent treed
patches is projected to be similar to that of a natural forest because these patches result from the
colonization, establishment, and persistence of species reaching unmanaged sites.
The thinning phase represents a reduction of individuals. However, in urban landscapes, the loss of
individuals may be from factors other than competition. Planted vegetation losses occur principally from
selecting the wrong species for site conditions and mortality because of urban stress (e.g. urban heat
island; Grey and Deneke, 1986; Miller, 1988). For remnant individuals, losses result from changes in
environmental conditions created by fragmentation, enclosing savanna trees by planted vegetation, root
damage during construction and from compaction, and altering water tables and other hydrologic pro-
cesses (McBride and Jacobs, 1976; McBride and Jacobs, 1986). For emergent patches, the thinning phase
may represent a reduction of individuals from competition; however, since these patches have a high
proportion of edge conditions, less loss may occur and a higher stem density is maintained (cf. Ranney
et al., 1981).
During the transition phase, planted trees are maturing and possibly senescing. Active management
may be required to establish and maintain continuous canopy cover because site mowing precludes
natural regeneration (Richards, 1983). Like planted trees, remnant patches without an understory also
may require management to maintain the cover type. For remnant patches with an understory, data
indicate that the transition phase may represent a loss in structural complexity and a lack of self-
perpetuation of existing species composition (Rudnicky and McDonnell, 1989). Emergent patches are
expected to mirror community dynamics of a natural forest; however, additional data are needed to better
understand forest regeneration and development in both remnant and emergent patches.
234 Zipperer et al.
Species richness
Comparisons of presettlement with postsettlement tree cover identifies increases in tree species richness
regardless of ecoregion (McBride and Jacobs, 1986; Zipperer et al., 1991; Nowak, 1993a). This increase
results from introduced species through planting and afforestation practices of public and private land
owners. Distinct differences are hypothesized when tree species richness curves for the entire urban
landscape are compared with a natural forest (Fig. 3; cf. Nowak, 1993a).
A number of different species richness curves have been described for successional development after
site establishment in natural forests (Peet, 1992). These different curves indicate that species richness
does not exhibit a simple and consistent response to any one environmental factor (Peet, 1992). In
general, species richness increases to a peak late in succession and then declines as early succession
species are lost (Auclair and Goff, 1971). In terms of forest development phases, species richness reaches
a peak during the establishment phase, and then declines during the thinning phase as less competitive
species are lost. Richness peaks again during the transition phase as canopy openings occur and then
fluctuates during the steady-state (Peet, 1992).
For the urban landscape, there may be a decline in tree species richness from initial site clearing for
development, but richness often can increase quickly as new species are planted. This increase is
hypothesized not to peak as in a natural forest but rather to continually increase as new species are
planted, thus offsetting species losses caused by environmental and biotic influences (Nowak, 1993a; Fig.
3A).

Figure 3. Comparison of tree species richness curves for a natural forest and the entire urban landscape (A) and for
patch origin (B) within an urban landscape.
Urban tree cover 235
Tree species richness is hypothesized to be different depending on the origin and management of urban
vegetation (Fig. 3B). Planted treed patches (e.g. street, yard, and park trees) are predicted to have the
greatest richness because of human influences. The rate of increase after development depends on the
number of species that are planted and survive. A continued increase will depend on replacement of
species after storms, from losses to pathogens (e.g. Dutch elm disease), and changes in landscape design.
Emergent patches are predicted to be the least species rich of the three origins because of vegetation
dynamics. Emergent vegetation occurs where management activities are insufficient to halt succession
(Fig. 2). Species colonizing and persisting at these sites are generally early successional species (Nowak,
1994b). In contrast, intermediate values are projected for remnant patches. This condition results from
non-native species colonizing the site and native species persisting on the site. For both emergent and
remnant patches, increases in tree species richness depend on site availability for germination, the
dispersal of neighboring species, and species performance on the site.

Canopy cover
Regardless of ecoregion, tree cover changes as a site is urbanized (McBride and Jacobs, 1986). In
forested ecoregions, canopy cover decreases from nearly 100% to an average of 31% (Nowak et al.,
1996), and postsettlement cover is fragmented into smaller patches (Fig. 4; Sharpe et al., 1986; Zipper-
er et al., 1990). Projected physical effects of fragmentation include increased solar radiation reaching
the ground, higher diurnal temperatures because of increased solar heating, increased desiccation and
wind throws along newly created forest edges, and decreased evaporation and transpiration (Saunders

Figure 4. Projected development of tree cover by patch origin for three ecoregions – forest, prairie/savanna, and
desert – during different phases of urban development.
236 Zipperer et al.
et al., 1991). Biotic effects include isolation and increased habitat loss and edge (Saunders et al.,
1991).
In contrast, for prairie/savanna and desert ecoregions, canopy cover actually increases with urbaniza-
tion (McBride and Jacobs, 1976; Nowak et al., 1996) because of planting, fire suppression, and watering.
Unlike the original cover in prairie and savanna ecoregions, where individual trees are scattered, the
postsettlement pattern has areas of single trees and patches of emergent and planted vegetation. The
increase in tree cover may negatively affect remnant individuals through competition for resources
(McBride and Jacobs, 1976). In contrast to forested and savanna ecoregions, emergent patches of trees
will not occur on abandoned sites in desert urban landscapes without watering.

Biomass
Several descriptions of biomass accumulation in natural forests have been described (Peet, 1981; Peet,
1992). The simplest is that biomass increases in a smooth asymptotic fashion. A second hypothesis
follows forest development after a disturbance. Biomass accumulation peaks late in the thinning phase,
then declines to an intermediate level during the late thinning stage as gaps are formed and the canopy
is no longer composed of a population of synchronously large trees (Peet, 1981; Peet, 1992).
Biomass curves have not been developed for urban vegetation. A pattern similar to a natural forest, but
with less biomass accumulation, is hypothesized for urban landscapes (Fig. 5). A comparison of stored
carbon in plant biomass in urban landscapes and nonhuman dominated forests supports this difference.
Approximately 11 and 17 metric tons/ha of carbon are stored in Oakland and Chicago forests, respec-
tively (Nowak, 1993b, Nowak, 1994b), compared with 55 metric tons/ha for natural forests (Birdsey,
1990). This difference occurs because there are less trees, and down material (i.e. branches, leaves, twigs,
and fruits) is often removed from the site.
Of the different treed patches, remnant vegetation is projected to have the greatest accumulated
biomass per hectare, and most similar to nonhuman dominated forests. In some remnant patches, biomass
accumulation may be less because fallen material such as trees and logs are removed (Barlow et al., 1987;
Cramer, 1993; Sauer, 1994). In addition, remnant patches may be going through a structural shift. Large
diameter trees, lost because of environmental and biotic factors, may not be replaced by similar physi-
ognomic species (e.g. Rudnicky and McDonnell, 1989), perhaps resulting in a loss in standing biomass.
For planted trees, the biomass curve is expected to be similar to a natural forest; however, because of
intensive management practices with pruning and the removal of deadwood from the site, the accumu-

Figure 5. Projected biomass accumulation curves for a natural forest and by patch origin.
Urban tree cover 237
lation curve is less. The drop in biomass accumulation is expected for a similar reason as the drop in a
natural forest; the shift from a synchronous population of large trees to a population of all sized
individuals (Richards, 1983). In addition, some species may not grow to their genetic potential in urban
areas because they often are planted on less than optimal sites. Emergent patches are projected to have
similar curves to remnant patches but with a faster rate of accumulation because they are dominated by
young trees and early successional species.

Ecological process
Vegetation dynamics
In the previous section on ecological patterns, we examined how remnant, emergent, and planted treed
patches vary structurally. In this section we examine how these structural differences influence the
movement of species and nutrients.
No treed patch within the urban landscape escapes human influences. Humans decide what species to
plant, when and where, what site to manage, and what cover a site will attain. In addition, there are
indirect effects (e.g. air pollution) that influence a treed patch. Consequently, analysis of vegetation
dynamics within an urban landscape must consider human influences. The proposed patch analysis
enables us to examine the interactions between vegetation dynamics and human influences.
Vegetation dynamics encompass site availability, species availability, and species performance (Pick-
ett et al., 1987a; Table 1). Site availability refers to sites available for establishment, colonization, and
persistence of vegetation. This component varies depending on urban morphology, management inten-
sity, and disturbance regime. In highly managed sites, succession is controlled. Seeds may reach available
sites, but because of management intensity, their establishment and persistence are prevented. In contrast,

Table 1. A Summary of the Causes, Process and Factors of Vegetation Dynamics (from Pickett et al.
1987a)

General causes Contribution processes

of succession or conditions Modifying factors
Site availability Coarse-scale Size, severity, time, dispersion
disturbance
Differential species Dispersal Landscape configuration, dispersal agents
performance
Propagule pool Time since last disturbance, land use treatment
Resource availability Soil condition, topography, microclimate, site
history
Differential species Ecophysiology Germination requirements, assimilation rates,
performance growth rates, population differentiation
Life history Allocation pattern, reproductive timing,
reproductive mode
Environmental stress Climate cycles, site history, prior occupants
Competition Hierarchy, presence of competitors, identity of
competitors, within-community disturbance,
predators, herbivores, resource base
Allelopathy Soil chemistry, soil structure, microbes,
neighboring species
Herbivory, predation, Climate cycles, predator cycles, plant vigor, plant
and disease defenses, community composition, patchiness
238 Zipperer et al.
emergent patches represent sites with minimal to no management which may provide conditions for the
establishment of vegetation.
An important aspect of site availability is the disturbance regime (Pickett et al., 1987b). A disturbance
regime is the collective set of disturbance types and their extent, time, magnitude, and dispersion (White
and Pickett, 1985; Glenn-Lewin and van der Marrel, 1992). It creates heterogeneity at the landscape level
and available sites for colonization at the community level. Although disturbance regimes have not been
described for urban landscapes, they clearly consist of both anthropogenic and natural events. In fact, site
conversions to an urban land use may predispose sites to disturbances such as flooding, fire, and
windthrows. Analysis of anthropogenic disturbances within remnant patches shows a higher frequency
of dumping, recreational activities (footpaths, bike trail, and horses), earth moving, and mowing, than
rural sites (Sharpe et al., 1986) (Table 2). Unfortunately, little information exists on the size, frequency,
and magnitude of these and other disturbances within an urban landscape and their influences on
vegetation dynamics.
The colonization of a site initially depends on species availability and dispersal (van der Valk, 1992).
Landscape configuration, seed banks, and seed predation influence species availability and dispersal
(Pickett et al., 1987b). Urban landscapes are highly fragmented because of land use conversions (Sharpe
et al., 1986; Zipperer et al., 1990), thus limiting dispersal (Saunders et al., 1991). Because remnant
patches often are the only source of many native species (Greller, 1975), limited dispersal may inhibit
colonization of available sites by native species. Likewise, we know very little about metapopulation
dynamics in urban landscapes (Opham, 1991). Metapopulation is defined as a ‘‘population consisting of
spatially-separate subpopulations that are connected by the dispersal of individuals’’ (Forman, 1995).
Unanswered questions – such as which areas act as sinks, which areas serve as sources, and how do
disturbances change the metapopulation status of a site – need to be answered to better understand
population interactions among patches.
An urban landscape configuration includes the vegetation patches and the background matrix com-
posed of buildings, roads, and other anthropogenic structures. Studies have shown that buildings can
enhance or buffer winds, thus changing wind patterns (Heisler et al., 1994). These effects subsequently
may influence the dispersal of wind-dispersed species. For example, the dispersal distances from parent

Table 2. Percentage of forest areas affected by disturbance for three urbanizing landscapes and five
rural landscapes in southeastern Wisconsin (more than one type of disturbance may occur in a forest
area; from Sharpe et al. 1986)

Difference
Disturbance Rural Urban (Urban-Rural)
Houses and yard 3.8 25.1 21.3
Dumping 8.6 18.8 10.2
Logging 30.1 25.7 −4.4
Footpaths 29.5 53.9 24.4
Roads 34.5 20.8 −13.7
Grazing 14.2 4.3 −9.9
Mowing (parks, institutions, etc.) 0.4 5.0 4.6
Earth moving (quarries, etc.) 2.7 5.4 2.7
Bike trails 11.3 1.8 −9.5
Horses 1.8 3.9 2.1
Pigs 0.4 0.0 −0.4
Farm equipment 1.9 0.0 −1.9
None 22.9 8.6 −14.3
Urban tree cover 239
plants may be quite different in an urban landscape as compared with a rural landscape. In fact, a
continuum of wind dispersal distances may exist as building and infrastructure configurations (i.e.
orientation, density, and height) change along an urban-to-rural gradient.
Upon reaching a site, a seed will remain dormant, germinate, or be eaten (van der Valk, 1992). Only
a limited number of studies have been conducted on seed banks in urban landscapes. Analysis of forest
interiors of urban remnant forest patches (>100 ha) in New York City revealed seed banks similar to rural
forests (Kostel-Hughes, 1995). Other studies of urban remnant forest patches have indicated a higher
density of seedlings of non-native species than native species (Rudnicky and McDonnell, 1989). Infor-
mation on seed banks in emergent patches is lacking. Additional studies are necessary to understand the
dynamics of site colonization by both native and non-native species.
Urban landscapes may have higher population densities of seed predators than rural landscapes (Nilon,
1996). Current studies in natural areas within New York City are being conducted to assess the impact
of seed predators on forest regeneration and to determine whether there is a preference for native or
non-native seeds (Nilon, 1996). Replicated studies need to be conducted in natural areas in other cities.
Seed predation, however, cannot be examined independent of seed dispersal (van der Valk, 1992). Seed
predators may cache their seeds in adjacent locations, thus assisting dispersal and germination away from
the parent plant.
With the successful dispersal to an available site, germination depends on environmental and biotic
conditions (Pickett et al., 1987a). Seed germination is just one aspect of species performance (Table 1).
Other considerations include: how does the urban environment (heat island effects, air pollution, human
activities, non-native species) affect native species performance? A 10-year study along an urban-to-rural
gradient from New York City to Litchfield County, Connecticut has identified changes in soil conditions,
nutrient cycling, and decomposition when comparing urban and rural stands (for a review, see McDon-
nell et al., 1997). These changes may have significant influences on native species performance and
rehabilitation efforts in natural areas in urban landscapes (McDonnell, 1988; Pouyat and Zipperer, 1992;
Pouyat et al., 1995; Zipperer and Pouyat, 1995). In addition, changes in the disturbance regime and
community structure, high densities of non-native species, and human activities may create conditions
better suited to non-native species in remnant and emergent patches (McDonnell and Roy, 1997).

Nutrient and carbon cycling

Extensive literature exists on nutrient and carbon budgets for forest ecosystems in nonhuman dominated
watersheds. Nutrient inputs include the atmosphere, nitrogen fixation, rock weathering, and hydrologic.
Nutrient losses include hydrologic, gaseous, and particulate losses to the atmosphere, fire, and forest
harvesting. In the urban landscape, fertilizers and high levels of NOx and SOx supplement nutrient inputs,
whereas removal of trees, branches, leaves, shrubs, fruits, and grass clippings increase nutrient losses.
Unfortunately, only a limited number of studies have assessed nutrient cycling within the urban landscape
and these studies have focused on individual components of the landscape (i.e. lawns and remnant forest
tracts) rather than the entire urban watershed.
In a recent paper, McDonnell et al. (1997) discussed how litter decomposition and nitrogen cycling
within remnant oak patches varied along an urban-to-rural gradient extending from New York City to
Litchfield County, Connecticut. They found that urban forests exhibited faster litter decomposition and
nitrification rates than rural forests even though the urban forest had reduced fungal and microarthropod
populations and poorer leaf litter quality than rural forests. Decomposition and nitrification differences
were attributed to two anthropogenic causes: increased average temperatures caused by the urban heat
island effect and the successful colonization of earthworms (Pouyat et al., 1995; McDonnell et al., 1997).
Using the same oak stands, Groffman et al. (1995) evaluated carbon dynamics and separated soil carbon
into four pools: (1) readily mineralizable carbon with a turnover time of days to weeks; (2) labile carbon
with a turnover time of weeks to months; (3) potentially mineralizable carbon with a turnover time of
240 Zipperer et al.
months to years; and (4) passive carbon which is very recalcitrant with a turnover time of years to decades
and possibly centuries. Analyses revealed that urban forests had lower labile carbon and higher total
passive carbon than rural forests (Groffman et al., 1995). These studies need to be replicated in other
urban landscapes and extended to other types of treed patch.
An important element with respect to nutrient cycling is the effect of vegetation structure on the
movement of nutrients within and through a treed patch. Structural features such as canopy size and leaf
area have not been measured from an ecological context (e.g. patch type), but have been measured from
a management context (i.e. land use; Nowak, 1994b). For the City of Chicago, average leaf area index
(excluding grass) for tree-covered areas was 6.0; a value toward the lower end of the range (5–8) for
deciduous forests (Barbour et al., 1980). This low value was attributed to an understory of grass and
impervious surfaces. The highest percent of total leaf area for Chicago was observed on vacant lots
(8.4%), institutional lands (e.g. parks, cemeteries, and golf courses; 37.8%), and residential areas (49.7%;
Nowak, 1994b).
For the different treed patches, remnant, with its understory of saplings, shrubs and herbaceous plants,
is projected to have the highest leaf area index and is predicted to have an index similar to that of a
deciduous forest. Even though emergent patches are dominated by small diameter trees, they also have
an understory and are predicted to have a leaf area equal to or slightly less than planted patches on a per
area basis. Planted trees often are characterized by large diameter trees (Nowak, 1994b). Street trees, for
example, because of their size (50% of the street tree population was $31 cm dbh), accounted for 24.0%
of the total leaf area in Chicago and 43.7% of the leaf area in residential areas (Nowak, 1994b).
Although canopy size and leaf area measurements are used to assess ecological function such as
transpiration and carbon sequestration, another important consideration is tree physiology (Waring and
Schlesinger, 1985). Within an urban environment, ambient temperatures are 5–10°C higher than rural
temperatures (Ackerman, 1985). During summer months, ambient temperatures may exceed the optimum
temperature range for photosynthesis, thus reducing CO2 uptake and transpiration. Consequently, urban
trees may have lower CO2 uptake and transpiration rates than rural trees during those months; however,
this difference may be offset during the spring and fall when warmer temperatures may enhance pho-
tosynthesis and transpiration rates (Waring and Schlesinger, 1985).
Microclimates created by the size, shape, and type of treed patch may differ among treed patches and
subsequently influence CO2 uptake and transpiration rates. For example, the microclimate created by
remnant and emergent patches may be cooler and more humid than that created by a planted patch of
similar size and shape. This difference may influence the duration individual trees, within a patch, can
transpire and uptake CO2 for photosynthesis. Obviously, additional research is needed to determine a
carbon budget for an urban landscape, how different treed patches influence that budget, and how it
differs from carbon budgets in rural landscapes.
The interception of pollutants also may differ among patch types. Deposition of pollutants occurs
laterally, from the side, and vertically, from above the canopy (Pouyat et al., 1995). In patches with an
understory, lateral deposition generally occurs within 30 m from the patch edge, a zone spatially defined
as ‘‘road side environment’’ (Airola, 1984). Depending on patch width and edge type, remnant and
emergent patches, with their understory, should intercept more lateral deposition than a planted patch. In
contrast, a planted patch initially may intercept more vertical deposition because of its large canopy and
leaf area (Nowak, 1994b); however, more vertical deposition actually may be filtered out by remnant and
emergent patches because the pollutant passes through the canopy, sapling, and shrub strata. In fact,
optimum configurations involving spacing, patch shape, and vertical structure probably exist that maxi-
mize evapotranspiration rates, rain, and particulate matter interception, and carbon sequestration for
different urban landscapes. More research is needed to determine how landscape configuration of patch
types influences these processes.
Urban tree cover 241
Eventually, particulate matter will reach the ground surface through rain fall and litter. Within remnant
patches, particulate matter, such as heavy metals, accumulate in the forest floor (Pouyat and McDonnell
1991). This accumulation affects both nutrient cycling, carbon storage, and soil fauna (Pouyat et al.,
1994; Groffman et al., 1995; Pouyat et al., 1995). Because of a developed forest floor, accumulation of
pollutants is hypothesized to be the greatest in remnant patches and the least in planted patches. Overall,
because of their ability to possibly intercept and accumulate more pollutants, remnant and emergent
patches are hypothesized to have a significant effect on air and water quality within urban landscape. The
extent of the effect depends on the areal coverage of these patch types. This is not to say planted patches
are unimportant within an urban landscape, but rather, from an ecological perspective, remnant and
emergent patches seem to significantly influence ecological processes.

Conclusion
Urban areas are a mosaic of physical, ecological, and social patches. How these patches interact to control
fluxes of water, nutrients and carbon, and biodiversity is one of the principal goals for the long term
ecological research program in Baltimore, Maryland (Fig. 6). As a step toward achieving this goal, we
propose a scheme of identifying and describing treed patches that is different from current vegetation

Figure 6. A composite model of the effects of urbanization on ecological phenomena modified from McDonnell and
Picket (1990) and McDonnell et al. (1997). The arrows indicate causal linkages and feedbacks between the com-
ponents of urban areas and ecological phenomena. Social-economic policies and decisions are the principal drives
of the system; however, a feedback from ecosystem effects influences these decisions and subsequently urban biota
and urban structural features. The italic text represents further stratification of a component based on experimental
design and results (see McDonnell et al., 1997).
242 Zipperer et al.
inventorying techniques used in urban landscapes. This work, coupled with patch analysis of social and
physical patterns (Grove and Burch, 1997; Foresman et al., 1997), will allow us to produce high
resolution data that are necessary for simulation models to estimate ecological and socioeconomic fluxes
for an entire watershed and city. Likewise, the proposed scheme is applicable to both human and
nonhuman dominated landscapes, thus enabling the evaluation of urban effects on ecological processes
across a gradient of urbanization from urban to rural. Adding data from historical records and aerial
photographs will enable us to test hypotheses about how social and ecological patterns have changed and
how they may change in the future.

Acknowledgments
We would like to thank Dave Nowak, Mark Walbridge, and two anonymous reviewers for their help in
improving the manuscript. Funding support was provided by USDA Forest Service Research Unit
NE-4952.

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