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Author(s): M. J. McDonnell and S. T. A. Pickett
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Source: Ecology, Vol. 71, No. 4 (Aug., 1990), pp. 1232-1237
Published by: Ecological Society of America
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1232
1232 SPECIALFEATURE
SPECIAL FEATURE Ecology, Vol.
Ecology, Vol. 71, No. 4
71, No.
TABLE1. Featuresof urbanization. 1986). The relative influences of urban and natural
environmental factors on ecosystem patterning, and
Structuralfeaturesof urbanization the extent to which ecosystem processes are also influ-
Dwellings enced, could be examined most easily along urban-
Factories
Officebuildings rural gradients, where human influences can be directly
Warehouses quantified.
Roads Likewise, a number of questions that fall within the
Pipelines framework of disturbance theory could be examined.
Power lines
Railroads In disturbance theory, manipulations of disturbance
Channelizedwaterways regimes are used to determine the significance of dif-
Reservoirs ferent disturbance types, intensities, and frequencies in
Sewagedisposal facilities
Dumps communities and ecosystems (Pickett and White 1985).
Gardens The study of the interactions between urbanization and
Parks disturbance regimes and their effects on ecological
Cemeteries
Airports properties provide an excellent opportunity to advance
Biota of urbanareas understanding in this general area.
One specific question that could be addressed using
Crops
Ornamentals changes in disturbance regimes along urban-rural gra-
Domestic pets dients is the balance between autogenesis and allogen-
Pests esis (Kolasa and Pickett 1990). If various disturbance
Disease organisms
and stress factors can be attributed to forces either
Socio-economicfactors within or outside the community, then the balance of
internal and external control of system organization
can be contrasted along the gradient.
Because urban areas appear so often as a dense, high- One additional area of ecological research that could
ly developed core, surrounded by irregular rings of benefit from studies along urban-rural gradients is that
diminishing development (Dickinson 1966), the gra- of species control on ecosystem fluxes. The simplifi-
dient paradigm is a powerful organizing tool for eco- cation of community composition and the introduction
logical research on urban influences on ecosystems. of new species in urban areas provides an opportunity
Like natural environmental gradients, urbanization to address questions concerning the mechanistic role
should present ecologists with a rich spatial array to of species in ecological processes on higher levels of
use in explaining or predicting environmental and eco- organization.
logical effects. Urban-rural gradients, moreover, pro- Finally, the intimate involvement of humans with
vide an opportunity to explicitly examine the role of the urban-rural gradient suggests that it would be an
humans. unparalleled situation in which to integrate humans as
subjects for ecological study. Human ecology is the
discipline that inquires into the patterns and process
WHY STUDY ECOLOGICALSYSTEMSALONG of interaction of humans with their environments
URBAN-RURAL GRADIENTS?
(Boyden 1977, Boyden and Millar 1978, Vayda 1983).
From an ecologist's perspective, urbanization pro- Human values, wealth, life-styles, resource use, and
duces a variety of unprecedented and intense "exper- waste, etc. must affect and be affected by the physical
imental manipulations." Examples include changes in: and biotic environments along urban-rural gradients.
(1) disturbance regimes, (2) biota, (3) landscape struc- The nature of these interactions is a legitimate ecolog-
ture, (4) physiological stresses (e.g., air pollution), and ical research topic and one of increasing importance.
(5) cultural, economic, and political factors. In most Clearly, the interactions among various anthropo-
cases, both the spatial extent and magnitude of the genic factors and between anthropogenic and natural
manipulations are greater than those that ecologists are variables make urban-rural gradients potentially com-
typically able to produce. plex. These interactions must be assessed before anal-
The coarse-scale, anthropogenic manipulations of yses such as those suggested above can be carried out.
ecological systems along urban-rural gradients provide Furthermore, it is certain that urban-rural gradients
an opportunity to address basic questions at various are not appropriate for all ecological questions. Never-
spatial scales. For example, questions related to hier- theless, we believe that such gradients do provide new
archy theory could be addressed. The central problem and sometimes unique opportunites for ecologists to
in this theory is to determine at what scale ecological test assumptions and predictions of many ecological
processes and patterns uniquely appear (O'Neill et al. theories.
1234
1234 SPECIALFEATURE
SPECIAL FEATURE Ecology,
Ecology, Vol.
Vol. 71, No.
No. 4
Plot
Vegetation Plot
NY
NYC /Lir U^
SPECIFIC ALONGTHEURBAN-RURAL
QUESTIONS McGhie and Posner 1981) or new anthropogenically
AN ILLUSTRATION
GRADIENT: derived sources? How does the pattern of hydropho-
We have indicated how the study of ecological sys- bicity vary in time and space? Does it limit resource
tems along urban-rural gradients could be used to ad- availability to plants (e.g., by reducing N mineraliza-
dress topics of general ecological interest. In this section tion)? How does it affect litter decomposition rates and
we will discuss several specific uses of an urban-rural belowground processes? Does it affect gas fluxes from
soil to the atmosphere? Is it amplified by other stresses
gradient, drawing in part on our study of ecosystems
in the New York City Metropolitan area. The region and disturbances? Although these questions are gen-
includes a readily measurable gradient of land use ra- erated by the pattern of contrasting hydrophobicity
diating from the New York City urban core to suburban along the urban-rural gradient, they are relevant to
and rural areas at increasing distances from the city larger concerns of biogeochemical fluxes in a broad
(Fig. 1). Remnant forest patches still exist in the city range of ecosystems.
as well as elsewhere on the land-use gradient, providing
an excellent opportunity to investigate long-term hu- Community organization
man impacts on forest ecosystems. We know already that forests in highly urban envi-
ronments differ in both subtle and patent ways from
Soil resources those in the surrounding countryside (Bagnall 1979,
A preliminary study of the physical and chemical Airola and Buchholz 1984, Hobbs 1988, Rudnicky and
properties of forest soils along a land-use gradient from McDonnell 1989). For instance, urban and suburban
New York City to rural Dutchess County revealed that forests have a conspicuous proportion of exotic and
the soils at the urban end of the gradient were more naturalized species (Bagnall 1979, Airola and Buchholz
hydrophobic than rural sites (White and McDonnell 1984, Hobbs 1988, Rudnicky and McDonnell 1989),
1988, unpublished manuscript). This novel pattern pro- and frequently a lower representation of certain native
vides a stimulus to address several ecological ques- species. However, little is known about the functional
tions, including the following: Is the formation of the importance of the differences in composition. Like-
hydrophobic soil the result of natural processes (Adams wise, the structure of urban and rural forests differ
et al. 1970, DeBano 1971, Reeder and Jurgensen 1979, (Rudnicky and McDonnell 1989). The canopy height
August 1990
August 1990 URBAN GRADIENTS
URBAN GRADIENTS 1235
CAUSESAND ECOLOGICAL
ANTHROPOGENIC EFFECTS
ALONGURBAN- RURALGRADIENTS
A B C
FIG.2. A composite model of the effectsof urbanizationon ecologicalphenomena.The threecolumns are abstractedfrom
the details of Tables 1 and 2, and the arrows indicate causal linkages between the featuresof urban areas (column A), as
inputs,and the ecologicalphenomena(columnsB, C) as results.The focus of the researchprogramwould be on the phenomena
representedby rows 1 and 2, althoughthe results would be helpful in decisions concerningsocietal phenomenarepresented
by row 3. Feedbacksfrom columns B and C to A would receive attentionin researchbuildingon that underthe scope of this
program.
is more uneven and the number and connectedness of Indeed, such questions may have a stimulatory effect
treefall gaps differs from rural forests in the region. on landscape ecology as a whole. In the spirit of an
Furthermore, the fauna and its interactions (e.g., pre- exploratory analysis, assume that the elements of a
dation, Churcher and Lawton 1987) differ in urbanized landscape are more distinct toward the urban end of
areas. These conditions indicate that organization and the gradient (Godron and Forman 1983). Note, how-
dynamics of the forests should differ along the gradient. ever, that the pattern of distinctness can be determined
Once the composition, size structure, and stand ar- by direct measurement of aerial photographs and maps,
chitecture are adequately documented, attention can so that whether distinctness of patches increases or
turn to questions such as these: Is the regeneration of decreases monotonically, or is humped in some middle
current or prior dominants limited in forests at the ur- distance on the gradient, is a simple empirical matter.
ban end of the gradient? What are the compositional Illustrative questions that may follow upon the pattern
or architectural correlates of regeneration success or analysis include the following: Is dispersal of plants,
failure? What are the abiotic, biotic, and anthropogenic over equivalent distances, more limited in urban than
causes of regeneration patterns? Do changes in the an- rural ranges of the gradient? Is any such change a direct
imal community have direct or indirect effects on the result of the activities of people (including vehicles,
plant community dynamics by changing herbivory, pets, and pests) or some indirect effect of the size, shape,
predation, or dispersal relations of the plants? How do and arrangement of landscape elements? How does
human perceptions, values, and behaviors affect the size, shape, distance, and arrangement of patches relate
dynamics and persistence of forest communities along to their species composition and to ecosystem pro-
the gradient? cesses within them?
TABLE 2. An elaborationof the environmentallyand biot- tures of urbanization (Fig. 2a) as well as their effects
ically relevanteffectsof the featuresof urbanization(Table (Fig. 2b, c) is an underutilized area for ecological re-
1).
search, but one of increasing importance given the ex-
tension and magnitude of anthropogenic effects today.
Physicaland chemical environment
Local climate
Air pollution CONCLUSION
Hydrologicchanges The growth of metropolitan areas in North America
Waterpollution
Soil changesand earth movement and indeed worldwide indicates that knowledge of eco-
Populationand community characteristics systems under the influence of urbanization can only
Altereddisturbanceregimes become increasingly important. The magnitude and
Introducedspecies nature of the change in the physical, chemical, and
Increasein morbidity biotic environments that are associated with urban-
Alteredassimilation ization provide an unprecedented suite of "experi-
Alteredreproductivestatus
Changesin growthrates mental manipulations" that ecologists can utilize. We
Phenologicalchanges propose a framework to guide the design and integra-
Reduced longevity tion of ecological studies along urban-rural gradients
Social and behavioralchanges
Genetic drift and selection and indicate its utility for addressing basic ecological
Populationsize and structure questions. Finally, we suggest that the study of urban-
Altered successions rural gradients provides a new context in which to
Reduced richness
Landscapefragmentation integrate humans as critical components of ecological
systems. The results of these studies will not only con-
Ecosystemstructureand function
tribute to our understanding of basic ecological prin-
Debris dams
Forest floor ciples, but are critical to the ecologically sound man-
Sediment loading agement of human-dominated ecosystems.
Patchiness
Layeringof vegetation ACKNOWLEDGMENTS
Productivity
Nutrient flux We would like to thank J. Brown, G. Guntenspergen,J.
Decomposition Koch, P. Matson, P. Vitousek, and W. Zipperer for their
Nutrient retention comments and suggestionson the manuscript.G. Likens, G.
Loss of redundantpathways Lovett, and R. Pouyat made helpful suggestionson earlier
Loss of compartments versions of the manuscript.Contributionto the programof
Alterationof equilibria the Instituteof EcosystemStudies,The New York Botanical
Managementand capital apportionment Garden.
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